S1DA CONTRIBUTIONS TO BoTANy WAS FOUNDED BY Lioyo H. SHINNERS (LEFT) IN 1962. INHERITED BY WILLIAM F. MAHLER (RIGHT), DIRECTOR EMERITUS oF BRIT in 1971, AnD SINCE 1993, IT HAS BEEN PUBLISHED BY Brit Press. Barney L. Lipscoms, EDIToRr Botanical Research Institute of Texas 509 Pecan Street Fort Worth, Texas 76102-4060, US, 817 332-444] / 81 ’ 332-4112 FAX Electronic mail: sida@brit.org Home page at the URL: http://www.brit.org/sida/ W. {JOHN W. THIERET, ASSOCIATE EDITOR Dept. of Biological Sciences Northern Kentucky University Highland Heights, KY 41099 USA FELIX LLAMAS, CONTRIBUTING SPANISH EDITOR Dpto. de Botanica, Facultad de Biologia Universidad de Lean E-2471 Leon, SPAIN The views ex reflect those of the editors or of the Botanical Research Institute of Texas Guidelines for contributors are available upon request and on our SIDA home page as well as the last pages of each volume. Subscriptions for year 2006: $39. Individual $75. USA Institutions $85. Outside USA numbers issued twice a yeal < N zx BOTANICAL RESEARCH INSTITUTE OF ISSN 0036-1488 yressed in this journal do not necessarily TABLE OF CONTENTS DEDICATION John W. Thieret (1926-2005) RALPH L. THOMPSON John W. Thieret, the curator Rosert F.C. NAczi John W. Thieret, a student's perspective (1967-2005) Rosert R. HAYNES John W. Thieret, valuable botanical friend (1965-2005) RONALD L. STUCKEY John W. Thieret, colleague and editor friend (1969-2005) WILLIAM F. MAHLER John W. Thieret, associate editor of Sida (1972-2005) BARNEY LipscOMB SYSTEMATICS A new narrowly endemic species of Clematis (Ranunculaceae: sub- genus Viorna) from northeastern Texas Dwayne ESTES Hill cane (Arundinaria appalachiana), a new species of bamboo (Poaceae: Bambusoideae) from the southern Appalachian Mountains J.K. TRripcett, A.S. WEAKLEY, AND L.G. CLARK Revision of Bauhinia subgenus Bauhinia section Amaria (Cercideae: Caesalpinioideae: Fabaceae) RICHARD P. WUNDERLIN Dendrophorbium restingae (Asteraceae: Senecioneae). anew species from Sao Paulo, Brazi AristOnio M. TELEs, Jimi N. NAKAJIMA, AND JOAO R. STEHMANN Novedades en Gentianaceae para América del Sur Eva M. FicippA AND GLORIA E. BARBOZA Amelichloa: a new genus in the Stipeae (Poaceae) Mirta O. ARRIAGA AND Mary E. BARKWORTH Taxonomy of the Cuscuta pentagona com plex (Convolvulaceae) in North America Minal Costed, Guy L. NESOM, AND SASA STEFANOVIC Taxonomy of the Cuscuta salina-californicacom plex (Convolvulaceae) Minal Costea, Guy L. NESoM, AND SASA STEFANOVIC Taxonomy of C tag d Cuscuta umbrosa(C olvulaceae) Minal Costa, Guy L. NESoM, AND SASA STEFANOVIG Taxonomy of the Cuseuta indecora (Convolvulaceae) complex in North America Minal Costea, Guy L. NEsoM, AND SASA STEFANOVIC 21 25 33 55 65 97 123 129 145 A new species of Pediomelum (Fabaceae) from the lower Piedmont Plateau of Georgia and South Carolina JAMES R. ALLISON, MICHAEL WAYNE Morris, AND ASHLEY N. EGAN Big-fruited buckthorn, Sideroxylon macrocarpum (Sapotaceae), a long-forgotten Georgia endemic JAMES R. ALLISON Sexual and apomictic prairie {leabane (Erigeron strigosus) in Texas: geographic analysis and a new combination (Erigeron strigosus var. traversii, Asteraceae) RicHARD D. Noyes, HEIDI GERLING, AND CARLA VANDERVOORT More American black sapotes: new Diospyros (Ebenaceae) for Mexico and Central America MITCHELL C. PROVANCE AND ANDREW C, SANDERS Taxonomy of North American species of Oldenlandia (Rubiaceae) Epwarb E. TERRELL AND HAROLD ROBINSON New taxa and combinations in cultivated bamboos (Poaceae: Bam busoideae C.M.A. STAPLETON Estudios en las Apocynaceae neotropicales XXIV: ees eee deae: Echiteae), un desapercibido nuevo género endémico de Bahia, Brasil J. FRANCISCO MORALES Estudios en las Apocynaceae neotropicales XX V: novedades y nuevos reportes en las Apocynaceae (Apocynoideae: Rauvolfioideae) de Venezuela J. FRANCISCO MORALES Taxonomic overview of the Heterotheca villosa complex (Asteraceae: Astereae) Guy L. Nesom Taxonomy of Lantana sect. Lantana (Verbenaceae): |. correct application of Lantana camara and associated names ROGER W. SANDERS Crataegus series Parvifoliae and its putative hybrids in the south- eastern United States J.B. PHipps AND K. Dvorsky A new species of Stenanthium (Melanthiaceae) from Tennessee, U.S.A. B. EUGENE WOFFORD A new species of Dodecatheon (Primulaceae) from the northern coast range of Oregon and Washington KENTON L. CHAMBERS The lectotypification and 19" century history of Croton alabamensis (Euphorl biaceae s.s. ) KENNETH J. WURDACK Two new species of Elymus (Poaceae) in the southern U.S.A. and other notes on North American Elymus species JULIAN J.N. CAMPBELL Anew hybrid genusand 12 new combinationsin North American grasses Mary E. BARKWORTH Quadruple, triple, double, and simple pappi in the goldenasters, subtribe Chrysopsidinae (Asteraceae: Astereae) JOHN C. SEMPLE Notes on types in Apocynaceae: Asclepiadoideae in Cuban herbaria and four lectotypifications in West Indian Gonolobinae ALEXANDER KRINGS AND PAUL R. FANTZ Comments on the Gerbera-complex (Asteraceae: Mutisieae) Hans V. HANSEN Validation of the name Dahlia sublignosa (Asteraceae) DayLe E. SAAR AND PAuL D. SORENSEN A new combination in Polytrichastrum (Bryophyta: Polytrichaceae) G.L. SMITH MERRILL A new combination in Tetraphis (Bryophyta: Tetraphidaceae) JupitH A. HARPEL A new combination in the Bromus catharticus complex (Poaceae: Bromeae sect. Ceratochloa) ANA MariA PLANCHUELO CHROMOSOME NUMBERS Chromosome numbers for western and arctic North American species of Antennaria (Asteraceae: Gnaphalieae) JERRY G. CHMIELEWSKI ANATOMY AND MORPHOLOGY Root-shoot anatomy and post-harvest vegetative clonal develop- ment in Lophophora williamsii (Cactaceae: Cacteae): implications for conservation MArtTIN TERRY AND JAMES D. MAUSETH ETHNOBOTANY Taxonomy and conservation of medicinal plants in canal-irrigated areas of Punjab, Pakistan KHALID FAROOQ AKBAR AND MOHAMMAD ATHAR BOTANICAL HISTORY Notes on the life and work of James Brigham McFarlin, Florida botanist TOM PALMER FLORISTICS, ECOLOGY, AND CONSERVATION Floristic diversity and composition of terra firme and seasonally inundated palm swamp forests in the Palma Real Watershed in lower Madre de Dios, Peru FERNANDO H. CorRNEJO VALVERDE, JOHN P. JANOVEC, AND MATHIAS W. TOBLER A vascular plant inventory of Starkey Wilderness Preserve, Pasco County, Florida EMILY FERGUSON AND RICHARD P WUNDERLIN Vascular floras of Sonoita Creek State Natural Area and San Rafael State Park: Arizona’s first natural-area parks STEVEN P. MCLAUGHLIN Annotated checklist of the vascular flora of the Big Sandy Creek Unit, Big Thicket National Preserve, Texas Larry E. Brown, BARBARA R. MAcRoserts, MICHAEL H. MACROBERTS, Pau. A. HARCOMBE, WARREN W. Pruess, I. SANDRA ELSIk, AND STANLEY D. JONES The north-south transition of flora across Arkansas: a preliminary phytogeographical analysis MicHAEL H. MACROBERTS AND BARBARA R. MACROBERTS High levels of seed inviability among seven populations of the en- dangered Short’s goldenrod (Solidago shortii: Asteraceae) PATRICK J. CALIE, ROBERT EC. NAcCZ1, CHRISTINA SHACKLEFORD, AND JESSICA CAICEDO Naturalization and extirpation of water hyacinth (Eichhornia crassipes, Pontederiaceae) in southwestern Arkansas, U.S.A. RENN TUMLISON AND BRETT SERVISS Fungi associated with Borrichia frutescens (Asteraceae): insect galls and endophytes DIANE TE StRAKE, AMY HADDOCK KEAGY, AND PETER D. STILING Type localities of vascular plants first described from Ohio: supplement JAMES S. PRINGLE Notes on southwestern Moraceae ALAN T. WHITTEMORE Exotic plant introduction in Kansas, two new species IRALEE BARNARD New and disjunct records of Eleocharis liesneri (Cyperaceae) from South America Davip J. ROSEN New and significant records of vascular plants for Florida and for Collier County and Lee County, Florida GEORGE J. WILDER AND MArtHA R. McComprs Carex bicknellii (Cyperaceae) new to Arkansas PauL M. McKenzie, C. THEO WITSELL, AND JOE WoOoLBRIGHT Green fringed orchid (Platanthera lacera, Orchidaceae) in south- ern Louisiana CHARLES ALLEN, SARA THAMES, HowARD ANDERSON III, Bit NEWTON IIL, RHONDA HAMPTON, AND GEORGE FISHER 805 Seymeria falcata (Scrophulariaceae), a new record for Texas and the United States JOSELYN FENSTERMACHER 811 Cayratia japonica (Vitaceae) new to North Carolina and an updated key to the genera of Vitaceae in the Carolinas ALEXANDER KRINGS AND Rosert J. RICHARDSON 813 New and noteworthy collections for Arkansas JAMES H. PECK AND BreTT E. SERVIss 817 Clematis morefieldii (Ranunculaceae) new to Tennessee Dwayne ESTES AND CHRIS FLEMING 821 Gratiola brevifolia (Plantaginaceae) new to the flora of Delaware, the Delmarva Peninsula, and the Mid-Atlantic WESLEY M. KNAPP AND DWAYNE ESTES 825 Hedyotis australis (Rubiaceae) new to Missouri and Florida and related species in the south-central United States WALTER H. LEwis 831 Book reviews and notices 144, 150,176, 196, 208, 226, 242, 330, 354, 366, 422, 446, 460, 468, 484, 502, 532,538, 544, 546,548, 564, 614, 634, 660, 764, 768, 776, 780, 800, 810, 816, 830, 837 Announcements 78, 96, 724 Index to new names and new combinations in Sida 22(1), 2006 Achnatherum arnowiae (S.L. Welsh & N.D. Atwood) Barkworth, comb. nov-—496 Amelichloa Arriaga & Barkworth, gen. nov—146 Amelichloa ambigua (Speg.) Arriaga & Barkworth, comb. nov—147 Amelichloa brachychaeta (Godr.) Arriaga & Barkworth, comb. nov—147 Amelichloa brevipes (E. Desv.) Arriaga & Barkworth, comb. nov—147 Amelichloa caudata (Trin.) Arriaga @ Barkworth, comb. nov—148 Amelichloa clandestina (Hack.) Arriaga & Barkworth, comb. nov.—148 Ammophila breviligulata subsp. champlainensis (F Seym.) Walker, Paris & Barrington ex Barkworth, comb. nov—496 Arundinaria appalachiana Triplett, Weakley & L.G. Clark, sp. nov—88 Bahiella J.F Morales, gen. nov.—342 Bahiella blanchetii (A. DC.) J.F Morales, comb. nov—342 Bahiella infundibuliflora J.F. Morales, sp. nov—345 Bauhinia amatlana Wunderlin, sp. nov—99 Bauhinia arborea Wunderlin, sp. nov—102 Bauhinia ayabacensis Wunderlin, sp. nov—105 Bauhinia geniculata Wunderlin, sp. nov—111 Bauhinia petiolata var. caudigera (S.F Blake) Wunderlin, comb. nov—115 Borinda angustissima (T.P. Yi) Stapleton, comb. nov—332 Borinda contracta (T-P. Yi) Stapleton, comb. nov—332 Borinda nujiangensis (TP. Yi) Stapleton, comb. nov—332 Borinda utilis (TP. Yi) Stapleton, comb. nov-—332 Bromus catharticus var. elata (E. Desv.) Planchuelo, comb. nov-—556 Clematis carrizoensis D. Estes, sp. nov—67 Cuscuta indecora var. attenuata (Waterfall) Costea, comb. & stat. nov—216 Dahlia sublignosa (PD. Sorensen) D.E. Saar & PD. Sorensen, comb. & stat. nov—545 Dendrophorbium restingae A. Teles, ].N.Nakaj. & Stehmann, sp. nov—123 Diospyros costaricensis M.C. Provance & A.C. Sanders, sp. nov—291 Diospyros gomeziorum M.C. Provance @ A.C. Sanders, sp. nov.—282 Diospyros tuxtlensis M.C. Provance G A.C. Sanders, sp. nov—295 Dodecatheon austrofrigidum K.L. Chambers, sp. nov—462 — Drepanostachyum falcatum var. sengteeanum Stapleton, var. nov—332 5 Ink ° ob NIWIT | , J 4 10 BRIT.ORG/SIDA 22(1) great passion was plants. Although John was focused on plant taxonomy and learning new plants his entire life, he had other diverse interests including clas- sical music, particularly opera. He also enjoyed sharing his knowledge far be- yond botany and the natural sciences through verbal discourse about diverse topics in history, literature, art, religion, and the social sciences. Scientists often are judged by the number of articles by them or about them. If this is any measure of a person, John stands taller than a coastal redwood. Dur- ing his professional career, John authored at least 157 refereed journal articlesand book articles. Forty-two articles appeared in Sida, Contributions to Botany, which was his favorite journal. He also regularly published in Bartonia, Canadian Field- Naturalist, Castanea, Economic Botany, Journalofthe Arnold Arboretum, Rhodora, Taxon,and Transactions/Journalofthe Kentucky Academy of Science. In addition to books, journal articles, and 19 nomenclatural com binations, John wrote 136 book reviews, 65 articles for Encyclopaedia Britannica, 46 articles for Encyclope- dia Americana, and many popular science articles. John served several significant editorships and advisory roles during his professional career. He wasa Member of the Editorial Board of Economic Botany from 1959-1965, Book Editor of Economic Botany from 1959-1984, Editor from 1986-1990, and Associate Editor from 1992-2002. He was a founding Member of the Editorial Board for the Vascular Flora of the Southeastern United States project from 1981-2005, and wrote seven southeastern generic flora treatments. John served as the Associate Editor of Sida, Contributions to Botany, from 1972-2005, and contributed to its excellence and prestige through his dedicated work. Barney L. Lipscomb, current Editor of Sida, Botanical Research Institute of Texas, notes, ‘John wasa towering lighthouse to the editors of Sida. His stead- fast editorial, ‘light, was a never ending source of guidance and navigation in keeping Sida on a positive course.” From 1981-1995, John served as Abstract Editor of Transactions of the Ken- tucky Academy of Science (TKAS). He then assumed the position of Editor of TKAS in 1996 and changed the title from Transactions to Journal of the Ken- tucky Academy of Science JKAS) in 1998 to better reflect its content. Through his efforts, John greatly enhanced the stature of the JKAS as a multi-disciplin- ary journal of state and regional scientific literature. During 1983-2005, one of his most significant roles was serving asa Mem- ber of the Editorial Committee of the monumental multi-volume project, Flora of North America North of Mexico. He edited the first 10 published volumes and prepared 25 family and generic treatments. John also was an Advisor in Botany tor Encyclopaedia Britannica from 1959-2005 and a Member of the Advisory Committee at the Lloyd Library in Cincinnati from 1992-2005. John loved editing and his ability as an editor was extraordinary. He was a perfectionist and a superlative editor second to none, but his efforts always brought out the best in authors. Those who submitted manuscripts for John’s em THOMPSON, JOHN W. THIERET: HIS LIFE AND CAREER 11 editorial scrutiny often found them returned with a profusion of red ink on the printed text. When his recommended changes were made, the greatly improved manuscripts always told the story better. John received many awards and honors during his career. One of his most esteemed honors was having a mint in the Lamiaceae that he discovered in Loui- siana named for him. In 1964, botanist Lloyd H. Shinners named this new spe- cies Scutellaria thieretii Shinners. He received the 1984 Distinguished Kentucky College/University Scien- tist Award from the Kentucky Academy of Science for his significant academic research and teaching contributions to the Commonwealth. John was recently presented the 2005 Outstanding Academy Service Award from the Kentucky Academy of Science for his outstanding editorial contributions to the JKAS. To commemorate his contributions to the Northern Kentucky University Herbarium, it was officially renamed The John W. Thieret Herbarium by the Northern Kentucky University Board of Regents on 22 March 2000. In 1994, he established The John W. Thieret Research Award as an annual award to the NKU biological sciences student who accomplished the most sig- nificant research. The Thieret family would like to continue this student honor, and have asked that donations in his memory be sent to the Northern Kentucky University Foundation, designated to that research award. John is survived by his devoted wife of 55 years, Mildred Thieret, his five children, Robert, Nancy, and Jeffrey in Minnesota, Richard in China, and Jenni- fer in Kentucky, seven grandchildren, and five great-grandchildren. John Thieret was one of the patriarchs of North American plant taxonomy, and one of the last great field naturalists of the 20th century. His death marks the ending of a botanical era, but his legacy continues through the work of many former students and colleagues. John was an inspiring and stabilizing mentor who enriched the lives of those who knew him. He will be greatly missed by all. ACKNOWLEDGMENTS lam especially grateful to Mildred Thieret for graciously sharing her memories about John and providing photographs of her husband. lalso acknowledge con- tributions from Barney L. Lipscomb, Botanical Research Institute of Texas, Ronald L. Jones, Eastern Kentucky University, David M. Brandenburg, Dawes Arboretum, James O. Luken, Coastal Carolina University, and Debra K. Pearce, Northern Kentucky University. REFEREED PUBLICATIONS Bairp, J.R.and J.W. THierer. 1985. Notes on Themeda quadivalvis (Poaceae) in Louisiana. |selya 2:129-137. Bair, J.R.and J.W. THierer. 1988.The bur gherkin (Cucumis anguria var.anguria, Cucurbitaeae). Econ. Bot. 42:447-451. 12 BRIT.ORG/SIDA 22(1) Bairo, J.R.and J.W. THierer. 1989. The medlar (Mespilus germanica, Rosaceae) from antiquity to obscurity. Econ. Bot. 43:328-372 Baird, JR. and JW. Tuierer. 1993. Spartina. In: J.C. Hickman, ed. The Jepson manual: higher plants of California. University of California Press, Berkeley and Los Angeles. Pp. 1296- 1297. Beat, E.O. and J.W. THieret. 1986. Aquatic and wetland plants of Kentucky, Kentucky State Nature Preserves Commission, Scientific and Technical Series, Number 5, Frankfort. BranbensurG, D.M., W.H. BLackwelt, and J.W. THiereT. 1991. Revision of the genus Cinna (Poaceae). Sida 14:581-596. Branoensure, D.M., J.E. Estes, $.B. Russet, and J.W. THieret. 1991. One-nerved paleas in Cinna arundinacea L. (Poaceae). Trans. Kentucky Acad. Sci. 52:94-96. BraNnbenBurG, D.M., J.E. Estes, and J.W. THierer. 1991.Hard grass (Sclerochloa dura, Poaceae) in the United States. Sida 14:369-376. BranbensurG, D.M. and J.W. THieret. 1996, Sclerochloa dura (Poaceae) in Kentucky. J. Ken- tucky Acad. Sci. 57:47-48. stp D.M. and J.W. THierer. 2000. Cinna and Limnodea (Poaceae): not congeneric. Sida 19:195-200. ree D.M. and J.W. THieret. 2003. Epipactis helleborine (Orchidaceae) in Kentucky, with overview of literature on biology of the species. J. Kentucky Acad. Sci. 64:55-74. Buopett Il, G.F.and J.W. THieret. 1985. Notes on Frigenia bulbosa (Apiaceae). Bartonia 51:69-76. Bupbett Il, G.F. and J.W. THieret. 1997. Saururaceae. In: Flora of North America Editorial Commitee, eds. Flora of North America, Vol. 3. Oxford Univ. Press, New York and Oxford, Pp. 37-38 Buooett II,G.F.and J.W. THierer. 2004.Lobed leaves in Salix exigua, sandbar willow (Salicaceae), in Kentucky. J. Kentucky Acad. Sci.65:51. Caraway, M.L. and J.W. THieret. 1985. Amphibromus scabrivalvis (Gramineae) in Louisiana. Sida 11:207-214. Crark, H.L. and J.W. THierer. 1968. The duckweeds of Minnesota. Michigan Bot. 7:67-76. Crark, R.C., R.L. Jones, TJ. Weckman, R.L. THomeson, J.W. THieReT, Kentucky State Nature Preserves Commission, and K. Feeman. 2005. State records and other noteworthy collections for Ken- tucky. Sida 21:1909-1916. CrANFILL, R. and J.W. Trieret. 1981. Thirty additions to the vascular flora of Kentucky. Sida 9:55-58. DetaHoussaye, AJ, and J.W. THieret. 1967. Cyperus subgenus Kyllinga (Cyperaceae) in the continental United States. Sida 3:128-136. Eckenwatoer, J.E. and J.W. THieret. 1993. Keys to gymnosperm families. In: Flora of North America Editorial Committee, eds. Flora of North America, Vol. 2. Oxford Univ. Press, New York and Oxford. Pp. 345-346. Evers, R.A.and J.W. THieret. 1957.New plant records: Illinois and Indiana. Rhodora 59:181. Hawt, D.W.and J.W. THierer. 2003. Chrysopogon. in: Flora of North America Editorial Commit- tee, eds. Flora of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Pp. 633-636. HARTMAN, R.L., J.W. THieret, and R.K. Raseter. 2005. Paronychia. |In:Flora of North America Edito- THOMPSON, JOHN W. THIERET: HIS LIFE AND CAREER 13 rial Committee, eds. Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Pp. 30-43. Hits, M.H., J.W. THieret, and J.D. MorerieLo. 2003. Sarcobatus.|n: Flora of North America Edito- rial Committee, eds. Flora of North America, Vol. 4. Oxford Univ. Press, New York and Oxford. Pp. 387-389. Kartesz, J.T., P Atten, and J.W. THieret. 1997. Epilobium brachycarpum (Onagraceae) in Ken- tucky. Trans. Kentucky Acad. Sci. 58:99. Kartesz, J.T. and J.W. THieret. 1991. Common names for vascular plants: guidelines for use and application. Sida 14:421-434. Lanory, G.and J.W. THiereT. 1973./soetes louisianensis Siana. Sida 5:129-130. Luken, J.O. and J.W. THierer. 1987a. Linum grandiflorum (Linaceae), Papaver dubium (Papaveraceae), and Salvia pratensis (Labiatae): additions to the Kentucky flora. Trans. Kentucky Acad. Sci. 48:26. Luken, J.O.and J.W. Tuierer. 1987b.Sumac-directed patch succession on northern Kentucky roadside embankments. Trans. Kentucky Acad. Sci. 48:51—54. LukeN, J.O. and J.W. Tierer. 1988. A life-form spectrum for Ohio. Trans. Kentucky Acad. Sci. 49:38-39. Luken, J.O. and J.W. THieret. 1995. Armur honeysuckle (Lonicera maackii; Caprifoliaceae): its ascent, decline, and fall. Sida 16:479-503. Luxen, J.O.and J.W. THierer. 1996. Amur honeysuckle, its fall from grace. BioScience 46:18—24. LuKen, J.O. and J.W. THieret. 1997. Amur honeysuckle, its fall from grace. Arnoldia 57:2-12. Luken, J.O. and J.W. THigret. 2001. Floristic comparisons of mud flats and shorelines at Cave Run Lake, Kentucky. Castanea 66:336-351. Luken, J.O. and J.W. THieret (eds.). 1997. Assessment and management of plant invasions. Springer-Verlag, Inc., New York, NY. Luken, J.O., J.W. THieret, and J.R. Karresz. 1993. Erucastrum gallicum (Brassicaceae): invasion and spread in North America. Sida 15:569-582. Meotey, M.E., H. Bayan, J. MacGrecor, and J.W. THieret. 1985.Achyranthes japonica (Miq.) Nakai (Amaranthaceae) in Kentucky and West Virginia:new to North America. Sida 11:92-95. Meotey, M.E., R. Craneitt, and J.W. THieret. 1983. Vascular flora of Kentucky: additions and other noteworthy collections. Sida 10:114-122. Meotey, M.E. and J.W. THierer. 1991. Ulmus parvifolia (Ulmaceae) naturalized in Kentucky. Sida 14:610-613. MoHtensrock, R.H. and J.W. THieret. 1987. Trees: a quick reference guide to trees of North America. Collier Books, Macmillan Publishing Company, New York, NY. Naczi, R.F.C. and J.W. THieret. 1996a. Addition to the flora of Bradford County, Pennsylvania. Bartonia 59:81—-85. Naczi,R.F.C.and J.W. THieret. 1996b.Invasion and spread of Coincya monensis (Brassicaceae) in North America. Sida 17:43-53. Naczi, R.F.C. and J.W. THieret. 1996c. The gold-cone tamarack (Larix larcina forma lutea, Pinaceae) in Pennsylvania. Bartonia 59:123-124. Isoetaceae), a new species from Loui- — 14 BRIT.ORG/SIDA 22(1) Naczi, R.F.C. and J.W. Tuierer. 2000. Additions to the flora of Potter County, Pennsylvania. Bartonia 60:117-120. Nienaser, M.A. and J.W. Tuieret. 2003. Phytolaccaceae. In: Flora of North America Editorial oo eds. Flora of North America, Vol.4.Oxford Univ. Press, New York and Oxford. Po Pearce, D. . eae JW. Tuieret. 1991. Japanese-quince (Chaenomeles speciosa, Rosaceae), a dual-use shrub. Econ. Bot. 45:285-288. Pearce, D.K.and J.W. Tuieret. 1993. Persimmon (Diospyros virginiana, Ebenaceae) and may- apple (Podophyllum peltatum), Berberidaceae): proximate analysis of their fruits. Trans. Kentucky Acad. Sci. 54:30-31. Raseter, R.K.and J.W. THieret. 1988.Comments on the Caryophyllaceae of the southeastern United States. Sida 13:149-156. Raseter, R.K.and J.W.THierer. 1997. Sagina (Caryophyllaceae) range extensions in Canada:S. japonica new to Newfoundland, S.procumbens, new to the Northwest Territories. Canad. Field-Naturalist 111:309-310. Reece, W.D. and J.W. Tuieret. 1966. Botanical study of the Five Islands of Louisiana. Castanea 31:251-277. Smith, C.E, Jr.and J.W. THieret. 1959a.An English obituary account of Thomas Nuttall. Bartonia —s Smith, C.E., Jr.and J.W. THieret. 1959b. Thomas Nuttall (1786-1859): an evaluation and bibli- ography. Leafl. West. Bot. 9:33-42. SpactH, J.P. and J.W. THieret. 2004. Notes on “coffee” from the Kentucky coffeetree (Gymnocladus dioicus, Fabaceae). Sida 21:345-356. Spuut, R.W. and J.W. Tuierer. 1989. Confusion between multiple and aggregate fruits. Bot. Rev. 55:53-72. Tuieret, J.W. 1953. Gross morphology of the seeds of the Scrophulariaceae and classifica- tion of the family. Ph.D. dissertation. The University of Chicago. THieret, J.W. 1953. Dipsacus laciniatus in Illinois. Rhodora 55:268. THiereT, J.W. 1954. The tribes and genera of Central American Scrophulariaceae. Ceiba 4: 164-184. THieret, J.W. 1955a. The seeds of Veronica and allied genera. Lloydia 18:37-45. THiereT, J.W. 1955b. The status of Berendtiella A. Gray. Ceiba 4:304—-305. THieret, JW. 1956a. Bryophytes as economic plants. Econ. Bot. 10:/5-91. THieReT, JW. 1956b. Nardoo. Amer. Fern J. 46:108-109. ] ] ] — THieret, J.W. 1956c. Stenardrium Nees versus Gerardia L. Taxon 5:58-59. THiereT, J.W. 1957. Plants new to Illinois and to the Chicago region. Rhodora 59:289. Tuieret, J.W. 1958a.Agalinis Rafinesque versus Chytra Gaertn. Taxon 7:142-143. THieret, J.W. 1958b. Castilleia Mutis ex L. versus Bartsia L. Taxon 7:83-84. THiereT, J.W. 1958c. Economic botany of the cycads. Econ. Bot. 12:3-41. Tuieret, J.W.1959a. Grassland vegetation near Fort Providence, Northwest Territories. Canad. Field-Naturalist 73:161-167 TuiereT, J.W. 1959b, Scrophulariaceae. In: J. Angely (ed). Catalogo e estatastica dos géneros botanicos fanerogamicos, Vol. 49. THOMPSON, JOHN W. THIERET: HIS LIFE AND CAREER 15 THiereT, J.W. 1960a. Calamovilfa longifolia and its variety magna. Amer. Midl. Nat. 63: 169-176. THiereT, J.W. 1960b.The formaldehyde method of collecting plant specimens. Turtox News 38:114-115 THieret, J.W. 1961a. A collection of plants from the Horn Plateau, District of Mackenzie, Northwest Territories. Canad. Field-Naturalist 75:77-83. THierET, J.W.1961b.New plant records for southwestern District of Mackenzie.Canad. Field- Naturalist 75:111-121. THieret, JW. 1961¢. The Scrophulariaceae-Buchnerae of Central America. Ceiba 8:92-101. THiereT, J.W. 1961d.The specific epithet of the pecan. Rhodora 63:296. TuiereT, J.W. 1962a. Exceptional height for Rhododendron lapponicum. Canad. Field-Natu- ralist 76:123. Tiere, J.W. 1962b. New plant records from District of Mackenzie, Northwest Territories. Canad. Field-Naturalist 76:206-208. TuiereT, J.W. 1963a. Additions to the flora of Louisiana. Castanea 28:169-1 70. TuiereT, J.W. 1963b. Additions to the flora of the Northwest Territories. Canad. Field-Natu- ralist 77:126. Tuieret, J.W. 1963c. Botanical survey along the Yellowknife Highway, Northwest Territories, Canada. |: Catalogue of the flora. Sida 1:117-170 THierET, J.W. 1963d. Life-forms in the plains flora of southern Mackenzie, Northwest Territo- ries. Rhodora 65:149-157. TuiereT, J.W. 1963e. The correct name for the watermelon. Taxon 12:37. Tieret, J.W. 1964a. Botanical survey along the Yellowknife Highway, Northwest Territories, Canada. II: Vegetation. Sida 1:187-239. TuiereT, J.W. 1964b. Eriogonum annuum (Polygonaceae): biennial in Nebraska. Sida 1:382. Tuieret, J.W. 1964c. Fatoua villosa (Moraceae) in Louisiana: new to North America. Sida 1:248. Tuieret, J.W. 1964d. Lysimachia japonica (Primulaceae) and Clinopodium gracile (Labiatae) in Louisiana: new to the United States. Sida 1:249-250. TuiereT, J.W. 1964e. More additions to the Louisiana flora. Sida 1:294-295. TuHiereT, J.W. 1966a. Additions to the Louisiana flora. Sida 2:264—-265. THiereT, J.W. 1966b. Habit variation in Myrica pensylvanica and M. cerifera. Castanea 31: 183-184. THiereT, J.W. 1966c. Seeds of some United States Phytolaccaceae and Aizoaceae. Sida 2: THiereT, J.W. 1966d. Synopsis of the genus Calamovilfa (Gramineae). Castanea 31:145-152. THieRET, J.W. 1967a.Life-forms in the flora of Minnesota. J. Minnesota Acad. Sci.34:251-277. Tuierer, J.W. 1967b. Neogaerrhinum kelloggii (Greene) Tuieret, comb. nov. (Scrophulariaceae) Sida 3:187, THieret, J.W. 1967c. Supraspecific classification in the Scrophulariaceae: a review. Sida 3: 87-106. Tuieret, J.W. 1967d. Thirty additions to the Louisiana flora. Sida 3:123-127. BRIT.ORG/SIDA 22(1) THierET, J.W. 1968. Additions to the vascular flora of Louisiana. Proc. Louisiana Acad. Sci. 31:91-97 THieret, J.W.1969a. Baptisia lactea (Rafinesque) THieret, comb.nov. (Leguminosae). Sida 3:446. THiERET, J.W. 1969b. Notes on Epifagus. Castanea 34:397-402, THiereT, J.W. 1969c. Rumex obovatus and Rumex paraguayensis (Polygonaceae) in Louisi- ana: new to North America. Sida 3:445-446. THieret, J.W. 1969d. Sagittaria guayanensis (Alismataceae) in Louisiana: new to the United States. Sida 3:445. Tiere, J.W. 1969e. Trifolium vesiculosum (Leguminosae) in Mississippi and Louisiana: new to North America. Sida 3:446-447, THiereT, J.W. 1969f. Twenty-five species of vascular plants new to Louisiana. Proc. Louisiana Acad. Sci. 32:78-82. THiereT, J.W. 1970a. Bacopa repens (Scrophulariaceae) in the conterminous United States. Castanea 35:132-136. THIERET, J.W. 1970b. Nemophila microcalyx, an incorrect name. Rhodora 79:399-400. TuiereT, J.W. 1970c. Orobanchaceae. In: C.L. Lundell, ed. Flora of Texas, Vol. 2. Pp. 331-337. THiereT, JW. 1970d. Sere Df yhulariac eae—fiqwort family.In:E.A Menninger. ed Flowering vines of the world. Pp. 316-321. THIERET, J.W. 1971a. Additions to the Louisiana flora. Castanea 36:219-222. THiereT, J.W.1971b. Eriocaulon cinereum R.Br. in Louisiana. Southw. Nat. 15:391. THiereT, J.W.1971¢.Observations on some aquatic plants in northwestern Minnesota. Michi- gan Bot. 10:117-124. THierer, J.W. 1971d. Physalis lagascae (Solanaceae) in Louisiana: new to the conterminous United States. Sida 4:277. THiereT, J.W. 1971e. Quadrat study of a bottomland forest in St. Martin Parish, Louisiana. Castanea 36:174-181. THieret, JW. 1971f. The genera of Orobanchaceae in the southeastern United States. J. Arnold Arbor. 52:404-434. THieret, JW. 1972a. Aquatic and marsh plants of Louisiana: a checklist. J. Louisiana Soc. Hort. Res. 13:1-45. : THiereT, J.W.1972b. Checklist of the vascular flora of Louisiana: Part 1. Fern and fern allies, gymnosperms, and monocotyledons. Lafayette Nat. Hist. Mus, Techn. Bull. 2. THierET, J.W. 1972c. Rotala indica (Lythraceae) in Louisiana. Sida 5:45. THiereT, JW. 1972d. Synopsis of Hemichaena, including Berendtiella (Scrophulariaceae) Fieldiana Bot. 34:89-99, THieret, JW. 1972e. The Phrymaceae in the southeastern United States. J. Arnold Arbor. 53:226-233. THieRET, J.W. 1972f. Zeuxine strateumatica in Louisiana. Amer. Orch. Soc, Bull. 41:413. THiereT, J.W. 1973. Sex and the angiosperms. Sida 5:59-60. THieret, J.W.1974.Allium ampeloprasum (Liliaceae) and Trifolium vesiculosum (Leguminosae) in Oklahoma. Sida 5:286-287. THOMPSON, JOHN W. THIERET: HIS LIFE AND CAREER 17 THieret, JW. 1975a. Hemigraphis reptans (Acanthaceae), a greenhouse weed in Louisiana. Sida 6:115. THiereT, J.W. 1975b. The Mayacaceae in the southeastern United States. J. Arnold Arbor. 56:248-255. THiereT, J.W. 1976a. Floral biology of Proboscidea louisianica (Martyniaceae). Rhodora 78: 169-179. TuiereT, J.W. 1976b. Vascular plants new to Ohio. Castanea 41:181-183. THieret, J.W.1977a. Cyperus louisianensis (Cyperaceae), a new species from southern Loui- Siana. Proc. Louisiana Acad. Sci. 40:23-26. THiereT, J.W.1977b. Juvenile leaves in Oklanoma Marsilea (Marsileaceae). Sida 7:218-219. THieret, J.W. 1977c. Life-forms in the Michigan flora. Michigan Bot. 16:27-33. THiereT, J.W. 1977d. The Martyniaceae in the southeastern United States. J. Arnold Arbor. THieRET, J.W. 1979. Hyptis mutabilis (Labiatae) in southeastern United States. Sida 8: 202-209. TiereT, J.W. 1980. Louisiana ferns and fern allies. Lafayette Natural History Museum. Pub- lished in conjunction with The University of Southwestern Louisiana, Lafayette. TuiereT, J.W. 1982. The Sparganiaceae in the southeastern United States. J. Arnold Arbor. 63:341-355. THierET, J.W. 1988. The Juncaginaceae in the southeastern United States. J. Arnold Arbor. 69:1-23. TuierET, J.W. 1989. Picea abies (Pinaceae) naturalized in southeastern Minnesota. Sida 13:505. THieRET, J.W. 1993a. Calocedrus. In: Flora of North America Editorial Committee, eds. Flora of North America, Vol. 2. Oxford Univ. Press, New York and Oxford. P. 412. TuHiereT, J.W. 1993b. Pinaceae. In: Flora of North America Editorial Committee, eds. Flora of North America, Vol. 2. Oxford Univ. Press, New York and Oxford. Pp. 352-354. THIERET, J.W.1993c. Psilotaceae. In: Flora of North America Editorial Committee, eds. Flora of North America, Vol. 2. Oxford Univ. Press, New York and Oxford. Pp. 16-17. THieRET, J.W. 2003a. Arthraxon. In: Flora of North America Editorial Committee, eds. Flora of North America, Vol. 25.Oxford Univ. Press, New York and Oxford. P.677. TuHieret, J.W. 2003b. Calamovilfa. In: Flora of North America Editorial Committee, eds. Flora of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Pp. 140-144. Tuieret, J.W. 2003c. Coix.In: Flora of North America Editorial Committee, eds. Flora of North America, Vol. 25.Oxford Univ. Press, New York and Oxford. Pp. 703-704. THiereT, JW. 2003d. Eremochioa. In: Flora of North America Editorial Committee, eds. Flora of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Pp. 688-690. THierET, J.W. 2003e. Hackelochloa. In: Flora of North America Editorial Committee, eds. Flora of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Pp.691-693. THieReT, J.W. 2003f. Microstegium. In: Flora of North America Editorial Committee, eds. Flora of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Pp.623-624. THierer, JW. 2003g. Monoanthochloé. |n: Flora of North America Editorial Committee, eds. Flora of North America, Vol. 25. Oxford Univ. Press, New York and Oxford. Pp. 28-30. 18 BRIT.ORG/SIDA 22(1) TuiereT, J.W. 2005. Agrostemma. Pp. 214-215 In: Flora of North America Editorial Commit- tee, eds. Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Tuieret, J.W. and C.M. Atten. 1974. Setaria pallide-fusca (Gramineae) in Louisiana. Castanea 39:290-291. THieret, J.W. and J.R. Bairo. 1985. Thalaspi alliaceum (Cruciferae) in Kentucky and Indiana. Trans. Kentucky Acad. Sci.46:143-145. Tieret, J.W. and D.M. Branbensure. 1986. Scaevola (Goodeniaceae) in southeastern United States. Sida 11:445-453. Tuieret, J.W. and R.A. Evers. 1957. Notes on Illinois grasses. Rhodora 59:123-124. Tiere, J.W.and S.F.Gtassman. 1958. Grasses new to Illinois and the Chicago Region.Rhodora 60:264. THiereT, J.W., R.L. HARTMAN, and R.K. Rasecer. 2005. Herniaria. In: Flora of North America Edito- rial Committee, eds. Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Pp. 43-45. THieret, J.W.and J.T. Kartesz. 1997. Lardizabalaceae.In:Flora of North America Editorial Com- mittee, eds. Flora of North America, Vol. 3. Oxford Univ. Press, New York and Oxford. Pp. 293-294. THiereT, J.W. and B.L. Lipscome. 1985. Scaevola sericea Vahl var. taccada (Gaertn.) THiereT and Lipscomb. Sida 11:103. Tuieret, J.W.and J.O. Luxen. 1996. The Typhaceae in the southeastern United States. Harvard Pap. Bot.8:27-56. THiereT, J.W. and R.S. Maptes. 1979. Thelypteris interrupta (Polypodiaceae) new to Louisiana. Iselya 1:55, Tuieret, J.W.,W.A. Nierinc, and N.C. Oumsteao. 2001. National Audubon Society Field Guide to North American wildflowers: eastern region, revised edition. National Audubon Soci- ety, Alfred A. Knopf, Inc., New York, NY. THieRET, J.W. and R.K. Raseter. 2005a. Corrigiola. In: Flora of North America Editorial Commit- tee, eds. Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Pp. 48-49, THiereT, J.W. and R.K. Rasever. 2005b. Polycarpaea. In: Flora of North America Editorial Com- mittee, eds. Flora of North America, Vol. 5.Oxford Univ. Press, New York and Oxford. Pp. 23-25. THieret, J.W.and R.K. Rasever. 2005c. Polycarpon.|n: Flora of North America Editorial Commit- tee, eds. Flora of North America, Vol.5.Oxford Univ. Press, New York and Oxford. Pp. 25-26. THiereT, J.W.and R.K. Raseter. 2005d. Saponaria.In: Flora of North America Editorial Commit- tee, eds. Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Pp. 157-158. Tuieret, JW. and R.K Raseter. 2005e. Scleranthus. In: Flora of North America Editorial Com- mittee, eds. Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. Pp. 149-151. THiereT, J.W.and R.K. Ragecer. 2005f. Vaccaria.|n:Flora of North America Editorial Committee, eds. Flora of North America, Vol. 5. Oxford Univ. Press, New York and Oxford. P. 156. THOMPSON, JOHN W. THIERET: HIS LIFE AND CAREER 19 Tiere, J.W.and R.L. THompson. 1984. Cleome ornithopodioides (Capparaceae):adventive and spreading in North America. Bartonia 50:25-26. THieret, J.W. and S.B. Younc. 1988. The Kerguelen-cabbage, Pringlea antiscorbutica (Brassicaceae). Econ. Bot. 42:288-291. THompson, R.L.and J.W. THieret. 1986.Alopecuru dinaceus (Poaceae) established in Ken- tucky. Trans. Kentucky Acad. Sci.47:138. Vincent, M.A. and J.W. THieret. 1987. Thymelaea passerina (Thymelaeaceae) in Ohio. Sida 12:51-54. Wesset, M.V. and J.W. THierer. 2000. Agrimonia (Rosaceae) in Kentucky with notes on the genus. J. Kentucky Acad. Sci.61:146-162. Woopwaro, R.W. and J.W. THieret.1953.A genetic study of complementary genes for purple lemma, palea, and pericarp in barley (Hordeum vulgare L.). J. Agronomy 45:182-185. POPULAR SCIENCE ARTICLES Branbensure, D.W. and J.W. THierer. 1999. Phytological peregrinations in West Virginia. The Dawes Arboretum Newsletter 33(1):2-3. BranbensurG, D.W.and J.W. THieret. 2000. Hobnobbing with the Egyptians. The Dawes Arbo- retum Newsletter 34(10):3. BranbdensurG, D.W. and J.W. THieret. 2001. Rafinesque and us. Lloydiana 6(1):4—-9. Reprinted with photographs in: Daweswood 1(2):20-28, 2004. Buppett, Il, G.F. and J.W. THieret. 1981. Squawroot: it toils not. Explorer 23(2):28-29. Buooett, Il, G.F. and J.W. THieret. 1981. Harbinger-of-spring. Explorer. 24(1):24—26. Tiere, J.W. 1955. Dragon's blood. Nature Mag. 48(7):372-374, 388. TuiereT, J.W. 1974. The papaw, Ohio's “tropical” fruit. Explorer 16(3):17-19. TuiereT, J.W. 1975. Long live the seeds. Explorer 17(1):4-7. Tiere, J.W. 1975. Beechdrops—benign parasite. Explorer 16(3):12-14. THierET, J.W. 1977. Consider the foxglove. Explorer 19(1):4—7. THieret, J.W. 1978. Red cedar. Explorer 20(4):26-29. Tiere, J.W. and D.R. Bezanson. 1983. Dutchman’s-breeches and squirrel-corn. Explorer 25(1):8-13. Tiere, J.W. 1996. Incredible edibles: the ginkgo. Lloydiana 1(1):14-16. Tiere, J.W. 1996. Incredible edibles: lupine seeds. Lloydiana 1(3):9-12., THiereT, J.W. 1997. Frankincense and myrrh. Lloydiana 1(4):7-9. Tiere, J.W. 1997. Incredible edibles: durian. Lloydiana 2(2):12-17. Tuieret, JW. 1997. Incredible edibles: insects. Lloydiana 2(4):9-12. THiereT, J.W. 1998. Incredible edibles: teff. Lloydiana 3(2):4-7. THiereT, J.W. 1999. Incredible edibles: corn smut. Lloydiana 4(2):12-16. Tuieret, J.W. 1999. On the trail of Rafinesque. Flora of North America 13(3-4):13. THiereT, J.W. 2005. Kentucky's tropical fruit, the papaw. The Lady-Slipper 20(3):2-3. Ware, T.R.and J.W. THierer. 1987. Boomer of the prairie. Explorer 29(1):7-10. pay JOHN W. THIERET, THE CURATOR Robert F.C. Naczi Claude E. Phillips Herbariur Department aan & nie ee Delaware State University pee Se 9901, U.S.A. raczi@desu.edu John Thieret’s complex and remarkable personality hada way of leaving its mark on all who encountered him; those who met John and came to know him even briefly have rich and lasting memories of him. 1am the one who, in many ways, had to step into John’s shoes soon after his retirement from Northern Kentucky University, where | began my academic career in 1992. Of course, I could never fill John’s shoes, and I soon learned to tell my new-found NKU colleagues on first meeting them that | was occupying the position John had vacated, but | was not John’s “replacement.” Most of the time, I received knowing chuckles in response to such a self-introduction. Occupying the academic appointment John had held also meant I was the first toassume a curatorship for the herbarium that John built. After his arrival at NKU in 1973 as Chairman of the Biology Department, John lost no time in establishing a herbarium there. During my tenure as curator at NKU, John re- mained very active—in effect, John and I served as co-curators of the herbarium for nine years, to my great benefit. This relationship makes me uniquely quali- fied to remember John The Curator, a side of John to which | will limit the rest of my remarks. John strove for excellence in everything concerning the NKU Herbarium. He put his heart and soul into collecting, identifying, labeling, mounting, and filing plant specimens in the herbarium. Though he collected far more speci- mens than most botanists (in the several tens of thousands), he was never in competition with others to beat their collecting records. Rather, he was inter- ested, above all, in quality over quantity. Instead of complaining about the te- dium of pressing plants, typing labels, or gluing specimens on mounting paper, John took sincere pleasure in executing these tasks that many field botanists foist on assistants. Once a specimen was in the herbarium, he insisted on the highest standards for its curation. Many times when I hosted visitors to the NKU Herbarium, or | traveled to use other herbaria in Kentucky, | heard the remark that the NKU Herbarium was the best curated herbarium in the state of Ken- tucky. Within a short time, | came to see for myself that this statement was true. John had a photographic memory, which served him very well as curator. | can best illustrate John’s memory by relating an experience that continues to SIDA 22(1): 21 — 23. 2006 22 BRIT.ORG/SIDA 22(1) amaze me. In the late spring of 1996, John entered the herbarium the day after I had returned from a long field trip through the southeastern United States for my research. John found me puzzling over a specimen I had collected. Though I was familiar with most flowering plants present in the springtime in decidu- ous forests of the Southeast, | had found something in one Alabama forest that completely stumped me. | had never seen this species, didn’t know its family, and couldn’t even be sure it was a monocot, though | somehow thought so. Turn- ing to John, | admitted that | didn’t know where to begin with this one, but suspected it was something “really good.” After all, what else could I say to cloak my ignorance? John gazed at my find, hesitated only a moment, didn’t say a word, and marched to one of several bookcases present in the herbarium. There, he removed one issue of one of the many scientific journals for which we had long series. Soon, he produced a photograph, and asked me, “Is this your plant?” With that photograph (Whetstone 1984: 133), I realized 1 had found the rare Croomia pauciflora(Nutt.) Torr. of the obscure family Stemonaceae. In response to my query if he had seen the plant, John responded, “No, but I remembered seeing that photograph appear in this paper many years ago.” When it came to desiderata for the herbarium, John was utterly ecumeni- cal. Whether the lowliest, most inconspicuous, most downtrodden weed or the loftiest tree, John was interested in learning its name, how to identify it, whether it was poisonous, and if he had any specimens of it in the herbarium. His en- thusiasm was infectious and quickly endeared him to other naturalists, par- ticularly in the field. When he encountered a plant species he hadn't seen in several years, he would treat the plant as if he’d run into a long-lost friend. My most recent field trip with John was last April, when he came to Delaware fora brief visit. linsisted on taking him toa nearby racetrack that was full of vernal weeds in all their unmown glory. Within minutes, John showed me Sibara virginica (L.) Rollins, a mustard species he knew well from Kentucky. Further investigation showed it to be previously undocumented for Delaware, though botanists have been hunting green treasure in this state since the earliest 1800s. Such was all in a day’s work if your combination of vast knowledge, keen eye, and unclouded memory were what made John Thieret the unparalleled botani- cal talent he was. — — John was very generous with imparting knowledge it had taken him years to amass. From him, | learned a tremendous amount, and have benefited im- mensely. Though John could be demanding, uncompromising, and prickly at times, he was also uncommonly human. So often when | expressed frustration at ill-prepared students, particularly ineffectual bureaucracy, or the insuffi- ciency of a 24-hour day, John would sigh and remark, “In a hundred years, no one will care a thing about it.” Yet, he cared about curation, cared about her- baria, and cared especially deeply for the NKU Herbarium, one of John Thieret’s great legacies. Hopelully, in a hundred years, people will care about John. NACZI, JOHN W. THIERET, THE CURATOR 23 John Thi llecting herbari i Elko County, Nevada, 11 August 1993. REFERENCE WuetsTone, R.D. 1984. Notes on Croomia pauciflora (Sstemonaceae). Rhodora 86:131-137. JOHN W. THIERET, A STUDENT’S PERSPECTIVE (1967-2005) Robert R. Haynes' Department of Biological Sciences Box 870345 Tuscaloosa, Alabama 35487-0345, U.S.A. It was a sad day 7 December 2005 when I was going through my e-mail and had two from Ron Jones. The first one indicated that John Thieret had suffered a massive brain hemorrhage, and the second one indicated that he had passed away. limmediately called Mrs. Thieret to offer her my condolences. I pointed out that I am now retired and live in rural North Louisiana. Consequently, | check my e-mail less often than while I was still a professor. Thus | had just learned of his illness. That was, by the way, the day he had passed. My first time to meet John Thieret was March 1967. I was in my final se- mester at Louisiana Tech, and he and | had agreed for me to attend The Univer- sity of Southwestern Louisiana (now University of Louisiana at Lafayette), where he was then employed, and to study under his direction for a master’s degree in plant systematics. Dr. Thieret at the time had a grant from NSF to prepare a Flora of Louisiana. Howard Clark, a graduate student at the time, and Dr. Thieret came to Louisiana Tech and took me on a weekend field trip to Northeastern Louisiana. We met about 6:00 a.m. at Carson Taylor Hall, the building that housed the Botany and Microbiology Department. Dr. Thieret had a truck with a camper that would sleep three people, which he had purchased with the NSF monies. In the rear of the camper were a few plants of flowering Sanguinaria canadensis (Papaveraceae), which they had collected the previous afternoon. That was my first time to see the species other than by herbarium specimen. During our weekend field trip, we visited quite a few beautiful deciduous forests, most with spring wildflowers. In particular we found Trillium (Liliaceae) (I cannot remember the species, and | am sure that John Freeman has since named that one diff ly than what Thieret would have called it). Again, that was my first experience with Trillium. The manner we used for locating good collecting habitat was to examine topographic maps and find areas with con- siderable relief. | thought at the time that we were quite fortunate in locating habitat. Since then I have learned much about field work, and | know that an excellent field botanist, such as John Thieret, has a sense for determining from topographic maps where good habitat might be. I began to learn the personality of John Thieret that weekend. He was con- Present Address: PO. Box 146, Athens, Louisiana 71003, U.S.A. SIDA 22(1): 25 — 31. 2006 BRIT.ORG/SIDA 22(1) stantly smiling, making jokes, telling stories, and teaching. From that weekend to the day he died, he was always teaching whenever | was in his presence. George Fisher, who was a graduate student at Tech in 1967, and | went to Lafayette later that spring for a collecting trip to the Florida parishes with Thieret. | realized more about his practical jokes early on that trip. We followed his directions to the USL campus and Biology Building where he was awaiting our arrival. He asked us to follow him to his house where his cam per/truck was parked. He started home and took us through more back roads (all dirt) than one can imagine. When we arrived at his house (on a paved road of course), he got out of his car laughing saying he just wanted to have some fun. By following the paved road, one could get from the USL campus to his house in half the a time it took us! The Florida parish trip had a profound effect on my botanical career. In western Washington Parish, we collected a pondweed that Thieret called Potamogeton nodosus (Potamogetonaceae). | did not question him at all. During my first semester at USL, Dr. Thieret brought out the specimen and asked me to determine it. | said you called it P nodosus. He said that now he was not certain about that determination. I keyed out the specimen and consistently arrived at Potamogeton epihydrus, which was not supposed to be in Louisiana. I looked at the distribution, and it was known no closer than North Carolina at the time. | went to him, and he said that is what he determined the specimen to be also. That began a lifelong study of the genus Potamogeton. The fall of 1967 entered the graduate program at USL. Dr. Thieret had three other graduate students at the time, Howard Clark, Alex Lasseigne,and Billy Dan Hinton. Howard was working on the Lemnaceae of Louisiana, Alex was study- ing the Fabaceae (we called it Leguminosae at the time) of Louisiana, and Billy Dan was studying Parietaria (Urticaceae). | have always been interested in aquatic vascular plants, and my plans were to undertake a study of Utricularia (Lentibulariaceae) of Louisiana. Dr. Thieret began going through his literature and correspondence and learned that Peter Taylor of Kew had undertaken a study of Utricularia of North America and he, Dr. Thieret, did not want me to duplicate any study underway. Therefore, my future in the Lentibulariaceae came to an abrupt end. We talked about several genera, none of which really interested me, and we finally settled on Conopholis (Orobanchaceae), a genus as far from the aquatic environment as possible. As everyone who knows me is aware, I never really left the aquatic environment, however. Dr. Thieret had all of his graduate students work in the herbarium, and, current graduate students would find this hard to believe, without pay. We mounted and filed specimens, pulled specimens for loans, etc. As it turned out, | became the unofficial collections manager. | had closer interactions with Dr. Thieret than others for that reason. Of course, lalways received the blunt of any rage if things did not go just right, even if it was totally out of my control. a HAYNES, JOHN W. THIERET, A STUDENT’S PERSPECTIVE 27 John Thieret loved classical music! He had a phonograph (tapes and CD's did not exist then!) that played in the herbarium much of the time, and he often placed it on one of the herbarium carts and played it in class. Thieret could whistle quite loudly, and he often was going around the biology building whis- tling classical music. Thieret suffered from occasional migraines, and, as lam sure most people with migraines, he was quite moody when he had a migraine. Whenever he was having a mood swing asa result of a migraine, he whistled exceptionally loudly. We graduate students learned to get out of the way whenever we heard that extremely loud whistling. In fact, if | was unusually late arriving in the morning and heard that whistle, I just went right on out of the building to a grill that was just off campus. All other graduate students had learned the rou- tine, and we all met at the grill for coffee until we thought the whistling had ceased. (I told him about our coffee excursions during a trip he and I made out West in 1998. He was totally unaware of these excursions, or so he said.) Without question, John Thieret was an editor, an excellent editor. I never will forget the first draft of my thesis. 1 proudly gave it to him, and it soon re- turned looking as if a chicken had stepped into red ink and then walked all over my manuscript! I learned from him to be much more careful when writ- ing and speaking. He had many favorite sayings, but one that | remember very well was related to editing. It is “consistency is next to godliness.” He said, be consistent; if you are incorrect at least be consistently incorrect! [learned much botany from John Thieret, but one thing in particular was that he did not like the term “pistil.” He said that it was an ambiguous term and that carpel was a much better term. | adopted his argument and never used or taught pistil after that. In courses that I taught, | always had to explain to the students why I did not use the term pistil regardless that it was in the text. John Thieret eventually became an editor for Flora of North America, and the edito- rial board adopted the term pistil over carpel. 1am sure it was difficult for him to accept such a change, but he did. As it turned out, he was the taxon editor for the families that I contributed to FNA. He had to fight with me to get me to use the term pistil. I told him why, he agreed, but he said his hands were tied. I told him I was not going to type in the word pistil, but he was welcome to replace carpel with pistil in my manuscripts, which he did reluctantly. One of the courses that Thieret taught at USL was Plant Ecology. I had pre- viously taken such a course, so I did not take it from Thieret. He decided to take the students on a field trip to Western Texas and Eastern New Mexico. He had never been to that area, but I had been there several times, as my in-laws lived in Eastern New Mexico at the time. He asked me where to see habitats, and | told him several areas that would be good. Rather than attempt a trip on his own, he asked me to serve as a guide on the trip. I agreed, and we headed west. We were in the desert scrub west of Sanderson, Texas one afternoon where he 28 BRIT.ORG/SIDA 22(1) was talking to the class about Ephedra and other desert scrub plants. | had moved away from the class and was examining closely some Ephedra plants, when he started the class back toward the car. He was moving at his normal pace “full steam ahead and damn the torpedoes” when I heard an awful scream from him. I was 25 feet or more behind the class when | caught him running! He had walked upon a Western Diamond-back Rattlesnake! It was coiled un- der some scrub, and he had almost stepped on it. We stood still (after I settled his nerves) and watched the snake for a few minutes. It finally began to move, and crawled across our path. That was the longest snake that I have ever seen, or so it seemed at the time. Dr. Thieret wanted to show the ecology class, in addition to desert scrub, mountain vegetation, terminating with tree line and above. He asked if such existed in southern New Mexico, and I said I could take him above tree line. He thought it was too far south, and, as he often did, would occasionally throw some joking jabs into the conversation indicating his disbelief. We started to- ward Ruidoso, New Mexico, where | knew there wasa ski resort and the moun- tain went above timberline. As we turned a curve and the mountain came into view, with the alpine tundra and snow cap, he immediately said, “I told you it was here, you just would not believe.” Of course it was a joke, and he used this phrase very often. Unfortunately, the ski lift was closed for the season, so we had to climb up one of the ski runs to timberline. We did so, and enjoyed collecting in the alpine tundra, the first time that | actually ever went above timberline. Howard Clark and Billy Dan Hinton graduated my first year at USL. David Dike came to USL asa graduate student during my second year, and he chose to study the life history of Ottelia alismoides (Hydrocharitaceae), a species intro- duced from Southeast Asia. The species was known to occur in The Pool. Lacassine National Wildlife Refuge, south of Lake Charles, LA. Dr. Thieret, David, and I went to Lacassine to plan his study. We were in a small boat in The Pool and were being blown rapidly by the wind. 1 saw something in the water that | did not recognize and made a grasp at it as we were blown by. Remarkably, | grabbed the plant and enough of it was uprooted to make a couple of speci- mens. | looked at the specimen and had no idea what it was. | gave it to Dr. Thieret, and he, too, had no idea what it was. We placed it ina bag and took it back to the lab, where, using Aquatic Plants (of India), I quickly determined it as Blyxa aubertii. Hydrocharitaceae). We had gone to study one introduced aquatic from Southeast Asia and had found a second. We prepared a manuscript to report the species new to North America, which appeared in SIDA later that year. Dr. Thieret sent a duplicate to MICH asa gift, and Ed Voss sent back a letter stating that was the first record from the Western Hemisphere of Blyxa in their herbarium. Thieret immediately sent back a letter indicating that it was our first record also (always a practical joker!). John Thieret mastered a technique for rapidly pressing specimens and re- a HAYNES, JOHN W. THIERET, A STUDENT'S PERSPECTIVE 29 cording the data. While he was working on a Flora of Louisiana, he was collect- ing abundantly all over the State. He had us graduate students (usually me) prepare his pressing papers. All papers were separated and carefully folded. Any that had color printing must be discarded as the color could get on the plants and affect the corolla color. He had the carpentry shop on campus prepare him a box with two compartments, each compartment slightly larger than the size of a folded sheet of newspaper. This box was a permanent fixture in the rear of the camper mentioned above. We graduate students were to divide the folded newspaper into two stacks, one on which we stamped his collection numbers and the other without num- bers. We used the herbarium numbering machine to stamp his collection num- bers on the paper. The numbered stack went into one side of the box, and the unnumbered stack went into the other side. In the field we would collect a batch of specimens and stack them beside the box. Thieret always pressed his collections within a few minutes following col- lecting. After a short time following collection, he would say, “we have a press- ing engagement.” That meant it was time to press plants. He hada tape recorder that he used to record the data, and this tape recorder always rode in the box, with the newspaper serving as padding for it. He would stand in front of the box of papers, record the collecting locality and habitat data, get a numbered paper from top of the stack, read that number into the recorder, record the spe- cies name, and finally record any specific information about the specimen. The first individual was pressed in the numbered paper, and all duplicate speci- mens were pressed in unnumbered papers. It was always this sequence (consis- tency!), so keeping duplicates with the correct number even though they were in unnumbered papers was easy. The field pressing was in a field press, com- posed of six or seven cardboards cinched with a press strap. The full field press stayed in the truck the remainder of the trip, and we transferred specimens toa permanent press upon returning to the lab, often four or five days later. For the evening, we would park wherever he could find acceptable and spend the night. This place may be a roadside park (illegal in Louisiana, but with State plates he always got by with it), state park, etc. One of his favorite places to spend the night was a rural cemetery. He said that no one visited these cemeteries at night, and he would not be bothered. I do not think he ever was. Each night in the field, all of the recorded data were transcribed. He had a por- table typewriter that stayed in the camper, and he would type all data of that day into his loose-leaf field notebook. The recorder then was ready for the next day. Once back at the lab, he had a secretary type labels using his typed field notes. All duplicate labels were photocopied, a practice that probably was not advisable with technology of the day. Also once back at the lab the process of moving specimens from field press to permanent press began. Again, consis- tency was important here! One would always begin at the TOP of the field press, 30 BRIT.ORG/SIDA 22(1) never the bottom. (Remember here the numbered paper went into the field press first, followed by unnumbered papers.) Starting from the top, then the unnum- bered papers would go into the press first for a particular collection, with the final sheet of a collection being numbered. Consequently it went on top for that collection number. After all specimens were transferred to permanent presses, then the presses were placed on the driers, which were in the work area of the greenhouse. Removing the specimens from the permanent press also required consistency. Always start from the TOP never the bottom. As the press was dis- assembled, the first paper out was numbered, usually followed by one or more unnumbered papers. These unnumbered papers went inside of the numbered one until another numbered paper arrived. This process was repeated until all specimens were out of the press. Using this procedure, regardless if the same species immediately followed itself, the duplicates always stayed together. Af- ter the presses were broken down, the specimens were all placed in a cabinet for unmounted specimens. Collections he made for the Flora of Louisiana project were used as ex- change specimens for the USL herbarium (LAF). Once the unmounted cabinet was full or nearly so, which was once or twice a year, he and I would begin the process of deciding where to send duplicates. He exchanged with 15 or 20 her- baria. We would spread out the numbered paper followed by all of the unnum- bered ones of a collection. The best specimen would go into the LAF stack, and then all others were sorted into one, two, three, four, etc. stacks, one sheet per number per stack, one stack per exchange herbarium. The original label went in the sheet for LAF, and photocopies went with the duplicates. It was my re- sponsibility to get the duplicates all boxed, shipping labels and invoices pre- pared, and get the specimens to the post office. With us sending out thousands of exchange specimens each year, we also received thousands of exchange specimens in return. Thieret and I often made a game of opening the exchange boxes. We would turn the stack so that the labels were face down and both stand there in anticipation. One of us would quickly open the top newspaper, and our game was to see which one could say the name first. He usually won, but I did occasionally - he probably just let me win, however. After I graduated from USL, l attended The Ohio State University, where I earned my Ph. D. under the direction of Dr. Ronald L. Stuckey. While I was at Columbus, Dr. Thieret was hired by Northern Kentucky University, in Alexan- dria, KY to be Chairman of Biological Sciences. John Thieret believed in precision. He always said he was easily pleased with perfection. Once I had completed my Ph.D., Dr. Thieret invited me down to give a seminar at NKU. Since he had known my wife and son while we were in Lafayette, he invited us to stay at his home in Alexandria. He sent directions that stated after crossing the Ohio River in Cincinnati continue south until in- HAYNES, JOHN W. THIERET, A STUDENT’S PERSPECTIVE 31 tersect State Route 27. Then follow State Route 27 to Alexandria. After we crossed the Ohio River, we noticed U. S. 27 a mile or so from the bridge. | told my wife that Dr. Thieret was so precise that he certainly did not mean U.S. 27 since he clearly stated State Route 27 (we had his directions in handl!). We continued south and finally gave up, deciding that he must have meant U.S. 27, which he did. We turned around, found U. S. 27 and followed his directions onto campus. He was outside the Biology Building frantically awaiting our arrival, since it was less than 30 minutes before the seminar was scheduled to begin. His his- tory of precision almost upset the applecart. I saw Dr. Thieret only occasionally after I left Lafayette, although we al- ways kept in touch. Following the start of FNA, where I ran across him mostly was at the Missouri Botanical Gardens, as he was always there just prior to the annual Systematics Symposium for a FNA editorial meeting. In July 1998, he and I made a two week collecting trip to Utah and Nevada. We agreed that I would drive my Jeep SUV and that I would pick him up at the St. Louis airport. I was to bring all of the collecting equipment and floras since | had much more room than he would have. When I met him at the airport, I was surprised to see that he had one small shoulder bag for a two week trip. | asked about his lug- gage. He said this is all I have. Needless to say, I could not image how he was going to get along for two weeks with just that in the small bag (certainly no larger than 3 inches by 9 inches by 15 inches). He said it was less expensive for him to fly from Lexington to St. Louis than from Cincinnati to St. Louis, regard- less that the plane leaving Lexington stopped in Cincinnati prior to arriving at St. Louis and similarly on return. So Mrs. Thieret drove him to Lexington where he caught the plane and flew to St. Louis. His plans were to exit the plane in Cincinnati on the way home rather than go to Lexington. He obviously could not get his luggage if it was checked to Lexington, so he had to take aboard everything he needed. I stillam amused about his reasoning, although it worked. We had a wonderful time in the mountains and deserts. | collected many aquat- ics, and he collected many grasses (all under my numbers). I bet my friends who know | almost never collect grasses could not imagine why so many ap- peared under my numbers! Regretfully, the last time I ever saw him was when | left him at the airport in St. Louis after that wonderful two weeks. | did talk with him occasionally, and we did often exchange e-mails, but | was never in his presence again. [regret that we were unable to get together for eight years prior to his death. We had planned another collecting trip, this one to Northern Minnesota. My mother, however, became seriously ill, and we had to cancel that excursion. Few people have affected my life the way John W. Thieret did. We certainly had some rocky times when I was handling the herbarium at USL, but we had many, many wonderful times together, which completely obliterate any rocky times we might have had. JOHN W. THIERET, VALUABLE BOTANICAL FRIEND (1965-2005) Ronald L. Stuckey Emeritus Professor of Botany Herbarium, Museum of Biological Diversity Ohio State University 1315 Kinnear Road Columbus, Ohio 43212-1192, U.S.A. John W. Thieret was one of my valuable botanical friends, and his loss through death is now clearly becoming a reality. Il received notice of his death on 7 De- cember 2005 from a short morning telephone conversation from his zoological colleague, Tom Rambo of Northern Kentucky University. Tom has been known to me since his graduate school days when he was studying ornithology. We were together during summers at The Ohio State University’s Franz Theodore Stone Laboratory Biological Field Station in western Lake Erie on Gibraltar and South Bass Islands, at the town of Put-in-Bay, Ohio. My recollections of John Thieret are based on facts that I have obtained from my correspondence with him, students of mine that have taken his Field Botany class that he taught at the Stone Laboratory, my interactions with him as an editor for Sida and book review editor for Economic Botany, and other recollections I have of him during our 40 years of working together. In 1965, | had completed my doctoral dissertation on The Taxonomy and Distribution of the Genus Rorippa (Cruciferae) North America, a group of yel- low-petaled plants in the mustard family, commonly referred toas marsh cress. I received the Ph. D. degree at graduation on 1 May 1965 at the University of Michigan, Ann Arbor, and arrived in Columbus in September of that year to begin my career as an assistant professor of botany at The Ohio State Univer- sity. | believe I had heard of the name John W Thieret, as a botanist who had been in the Chicago area, but now was teaching and conducting research at the University of Southwestern Louisiana at Lafayette. John W. Thieret, Rorippa, and Ronald L. Stuckey (1965-1966) A letter dated 3 December 1965 arrived addressed to me, the short text which stated: We note with interest that you are carrying on work with Rorippa. May we send to you our speci- mens of this taxon for annotation? Most of the material in our herbarium consists of Louisiana plants collected during extensive field work over the past several years in connection with our state flora project SIDA 22(1): 33 — 49. 2006 34 BRIT.ORG/SIDA 22(1) I responded on 7 December (exactly 40 years earlier on the same day of his death) telling him that I would annotate his specimens of Rorippa, and on 28 January 1966 | signed the loan form for 31 specimens and one to retain. Upon returning the loaned specimens, I wrote in my letter to him of 1 March 1966 offering to write a key to the species of Rorippa that grow in Louisiana, as John was pre- paring a flora of that state. | also noted that my dissertation was not yet pub- lished, and therefore some new names and/or combinations were not yet valid. John’s immediate reply of thanks for my annotations on the specimens was re- ceived a few days later. The letter concluded with “Let us know if we can ever be of help to you. We are always willing to loan or to try to collect specimens for those who ask...” John W. Thieret and Robert R. Haynes (1968-1969) Robert R. Haynes was a graduate student of who worked in the Herbarium un- der John Thieret’s direction. Bob also earned a masters degree at the University of Southwestern Louisiana, Lafayette with a thesis, titled “A Monograph of the Genus Conopholis (Orobanchaceae),” completed in May 1969. Robert was re- ally more interested in aquatic vascular plants, and he wanted to study pond- weeds in the genus Potamogeton but, of course, no Ph. D. program was offered in biology or botany at the University of Southwestern Louisiana. He sought Thieret’s advice, who told him that a young professor by the name of Ronald L. Stuckey in the Department of Botany at The Ohio State University might be a place where he could study these narrow-leaved pondweeds. Robert R. Haynes, Potamogeton, and Ronald L. Stuckey (1968-1973) Bob’s first letter dated 12 October 1968, came to me with the following request: A f This spring I hope to receive a Master of Sci the University of Southwestern Louisi- ana.I plan to begin work this summer toward a Detar of Philosophy degree in plant systematics and am considering Ohio State University as a possible school. My main interests lie in aquatic plants. = Pret — | would like to do some biosystematic work in the genus Potamogeton subsection Pusilli m writing to inquire if there is any possibility of working under your direction and of oe oe the financial aid that will be necessary if lam to continue my studies By this time Bob had in press his paper on the “Potamogeton in Louisiana” (Proc. Louisiana Acad. Sci. 31: 82-90. 1968). The first paragraph of my reply to Bob on 22 October 1968 stated: Itisa pleasure to learn of your interest in beginning a program toward a Doctor of Philosophy degree in plant systematics at The Ohio State University. | am also pleased to learn of your interests in aquatic plants, particularly in the ge- nus Potamogeton. My letter continued explaining that | was “primarily interested in floristic and phytogeographical problems in aquatic and shore plants.” | acknowledged that his interest in aquatic plants would fit into our program. I mentioned the avail- STUCKEY, JOHN W. THIERET, VALUABLE BOTANICAL FRIEND 35 ability of teaching assistantships, and asked Bob that he send a summary of his course work, his graduate Record Examination Scores, and asked the Graduate School Admissions Office to send him application forms and instructions. John W. Thieret’s letter of recommendation for Bob’s graduate work has not been lo- cated and probably no longer is extant. Bob's application for Graduate School, after review by the Graduate Committee in the Department of Botany, initially was not favorable for accepting him. John W. Schmit, chairman of the committee and I, with the permission of the Committee, agreed to give Bob a trial period, and asked him to enroll in the 10-week summer field program, taking a maxi- mum of four courses at the Franz Theodore Stone Laboratory. Bob completed this work during the summer of 1969 and earned Ass in all four courses. He then re- turned to Louisiana where he was a teacher at Cecila Jr High School in Cecila. In the fall of 1969, Department Chairman Schmitt and College Dean Rich- ard H. Behning agreed that I needed more permanent help with my responsi- bilities as Curator of The Ohio State University Herbarium. The position of Assistant Curator was established. The first person appointed to the position had to leave at the conclusion of the fall 1969 quarter, and with the position vacant, I called Robert R. Haynes to see if he would accept that responsibility. From February 1970 to June 1971 he was the Assistant Curator in the OSU Her- barium, and from September 1971 to June 1973 he held teaching assistantships. Bob completed his dissertation in June 1973 on “A Revision of North American Potamogeton Subsection Pusilli (Potamogetonaceae),” which was published in Rhodora 76: 564-649. 1975. Bob remained in Columbusat The Ohio State Univer- sity for the academic year 1973-1974 and was an instructor in botany teaching General Botany for three quarters and Local Flora during the summer quarter. Bob developed a career as a professor teaching plant taxonomy, advising graduate students, curating a herbarium—all of which concluded with his re- tirement in May 2005 from the University of Alabama, Tuscaloosa. His major work on Potamogeton and related genera is published in the Flora of North America North of Mexico, Volume 22: 3-94. 2000. John W. Thieret and the Stone Laboratory (1970-1971) In 1956 a new summer teaching program was developed at the FT. Stone Labo- ratory under newly appointed Director Dr. Loren S. Putnam, ornithologist of The Ohio State University. In the new system, Dr. T. Richard Fisher of the De- partment of Botany taught two 5-week courses, Field Botany during the first term and Higher Aquatic Plants during the second term. I had the privilege of attending the Stone Laboratory and took the Field Botany course from him in the summer of 1959. Since that time I kept in contact with Dr. Fisher, who later was responsible for my joining the faculty of The Ohio State University in the fall of 1965. Among my responsibilities eventually was to teach at the Stone Laboratory, and that opportunity surprisingly came the following summer 36 BRIT.ORG/SIDA 22(1) when I taught Field Botany. During the 1967 season I taught the Aquatic Plants course. Then in 1968 Fisher left the University to become chairman of the Bio- logical Sciences Department at Bowling Green State University in Ohio. Dr. Put- nam, who hada budget for the courses at the Laboratory, and in agreement with chairman Schmit of the Department of Botany, chose to hire a non-OSU teacher for the Field Botany course in 1969 and 1970. That arrangement was to allow me to have more time to develop a research program studying the aquatic vas- cular plants of western Lake Erie, instead of teaching both courses as Dr. Fisher had done since 1956. Putnam also preferred to have non-OSU professors teach- ing two consecutive summers and then change to a different teacher. Putnam relied on my advice in these situations and asked whom we might hire to teach the Field Botany course during the summer of 1971. At the time Robert Haynes was the Assistant Curator of the OSU Herbarium, and | asked him for sugges- tions. Naturally he suggested John W. Thieret, and I reported to Putnam with this recommendation From Putnam | soon knew that Thieret was hired to teach Field Botany, and a letter of 9 November 1970 came to me from Thieret: Ive just spoken with Dr. eee epenaine Put-in-Bay next summer. Lasked him to suggest someone to whom I could writ rsyll abus or what-have e-you u for Botany 610, Field Botany; the text or texts ‘ised: class ne pies ocalities; etc.; etc.; etc. In response to my question, he suggested your name. Thus you are the chosen victim. John followed with four questions that concerned the course outline, the text- book used, class procedure, and good field localities to see and collect plants. In regard to the latter item he wrote: | must admit that lam a bit panicked by the thought of trying to find enough such localities on a one island. (A bit more limited area than lam used to at Lake Itacsa, you know. | mean “Itasca.”).. n certainly looking forward to trying my hand at Put-in-Bay. Maybe I'll even be given the shandees meet you. Best wishes. And please give my regards to Bob Haynes. My three and one-half page letter of 17 November 1970 followed. It began: lam very pleased to learn that you are going to be with us teaching Botany 610 (Field Botany), this summer at Stone Laboratory. Your letter of 9 November, full of questions is at hand, and I'll try to answer them as best I can. I took the field botany course at Stone Lab in 1959 and taught it once, in 1966. Since then I have been developing and teaching the aquatic plant course which meets every other summer and to develop a research program around the theme of the changing flora of the islands and of the aquatic situations that we find in the area. My letter continued by giving John some statements about the kinds of students that take the course and their knowledge of plants. I noted that the course prima- rily concerned the flowering plants, that Clara G. Weishaupt'’s Vascular Plants of Ohio was the book used, and that I had an outline of the course with various procedures that I would mail him. Then I wrote a long statement about class field — STUCKEY, JOHN W. THIERET, VALUABLE BOTANICAL FRIEND 37 trips, assuring him that he would have plenty of plants and ample places to take students. With reference to field work, ladded a statement about taking field trips by boat to the various islands in Lake Erie and to the Ohio mainland. My letter concluded with comments on the available library resources, labo- ratory supplies for the course, materials for pressing and drying collected plants, and the information sheets, more commonly known then as “handouts,” that I used in the 1966 Field Botany course. The latter I offered to mail to him. I con- cluded the letter about my own projected plans for the up-coming summer. I will be at the Laboratory all summer myself. ie Hise term neal be for research, at which time I will be spending most oot myt time writing. Hopefully, he “Flora of the Erie Islands,” and | the “Origin, Changes, and Geo- graphical Relationships of the Acuane and Matsh Flora of Nestea a Erie and Northwestern Ohio.” During the second term I will k | lam looking forward very h to your coming to Stone Lab next summer. Please do not hesitate to write if you have further questions. I hope the above comments will be helpful to you. John did not hesitate to send a return letter, dated 20 November 1970. It began, “I certainly thank you for your long letter—and all the useful data therein. Be- lieve me, your help is deeply appreciated.” After commenting on the various items | had mailed him, he concluded his letter saying, “After I've gone over all the material thoroughly, I'll let you know if any other questions arise.... Thanks a lot for all your fine help.” John continued his pursuit concerning the 1971 summer program at the Stone Laboratory, and on 19 January 1971, he wrote to Director Putnam, greet- ing him with: Dear Puttie: I note in your bulletin for summer 1970 that you had, in 1969 a series of talks—a “Special Lecture ae 25 Ca ee ay am not “outstanding” (except in my unrivalled lam noscientist, I still intend, via this letter, to offer to give some sort of a talk this coming summer. I have done quite a bit of botanical field work in Canada’s District of Mackenzie—in the Great Slave Lake region. Asa result of the work, I have a set of slides that some people find interesting, especially when the slides are accompanied by the delight- ful eee presented by humble John. If _ wish such a presentation—let’s call it a travelog— I'd be glad to import my slides to Put-in-Bay this coming June. I'm not at all certain that a travelog by as ee be worth a mention in your 1972 eae of it might fill in an otherwise empty evening on Gibraltar. And besides, I like to talk..Let me know. This letter reveals much of the kind of humor John displayed about himself. Regarding these special lectures, beginning in the summer of 1970, Director Putnam asked me to make arrangements for the special lecture series at the Laboratory. | contacted the potential speakers, prepared the program, hosted their visit, and introduced them when they made their presentations on Thurs- day evenings. Scheduling a special lecture by John W. Theiret was easily ac- complished, and I distributed the lecture schedule with Thieret listed for 13 38 BRIT.ORG/SIDA 22(1) July 1971 with the title “The Yellowknife Highway Region, Canada: Flora and Vegetation.” John’s wish to present a lecture was fulfilled, and his lecture title later was listed in the 1972 admissions bulletin. Again, Thieret, Rorippa, and Stuckey (1966-1972) When I completed my dissertation on the genus Rorippa in 1965, it was diffi- cult to locate a journal that would publish a large taxonomic monograph as a single paper or issue of a journal. My advisor, Edward G. Voss, had anticipated that I could retain the entire dissertation as one unit if it were published in the Contributions from the University of Michigan Herbarium. This publication had been inactive for some years, and editor Rogers Mc Vaugh reaffirmed that funds were not available at that time to reactivate it. The conclusion was that 1 would have to look elsewhere to publish my dissertation. Edward G. Voss held in high regard the taxonomic knowledge of Lloyd H. Shinners of Southern Methodist University. When Lloyd taught the field tax- onomy course in the summer of 1952, at the University of Michigan Biological Station, Pellston, Ed was his graduate teaching assistant and from then on grew a close botanical relationship and great respect for each other. In 1962, at SMU Shinners began publishing a private taxonomic journal named Sida, and Ed suggested that | might contact Dr. Shinners regarding publication of my Rorippa monograph. I then proceeded to correspond with Shinners about my study of Rorippa, and during 1966 I extracted and published two short papers taken from my dissertation. They were “The distribution of Rorippa sylvestris (Cruciferae) in North America” (Sida 2:361-376. 1966), and Rorippa walteri and R. obtusa synonyms of R. teres (Cruciferae)” (Sida 2:409-418. 1966). During 1967 through 1969, I revised my monograph on Rorippa, and on 5 December 1969, I wrote to Dr. Shinners asking if he would publish my Rorippa study in Sida. He enthusi- astically responded on 9 December 1969, and told me to send the manuscript to him after Christmas. A letter of 26 March 1970 confirmed that he had accepted the manuscript, whose “bulk is a bit appalling.” | heard nothing further from Shinners until I learned of his death of 16 February 1971. The future status of Sida was in doubt, and therefore the publication of my Rorippa monograph also was in doubt. Later on 15 March 19711 wrote, I wrote Dr. William FE Mahler, who was Shinners replacement, thanking him for his letter of 3 March 197] which indicated that publication of Sida would be continued and that addi- tional work was to proceed on my Rorippa monograph. “I am pleased that you have selected Dr. Thieret to go over my manuscript in preparation for its publication...As you may know Dr. Thieret will be at our University for 5 weeks this summer teaching at Stone Laboratory. | will be there at the same time and can work with him on the manuscript.” In April 1971, John and I began a correspondence on the preparation of the Rorippa monograph for publication in Sida. Together, John and I discussed the STUCKEY, JOHN W. THIERET, VALUABLE BOTANICAL FRIEND 39 Rorippa manuscript while both of us were at the Stone Laboratory. Later in the year John wrote me letters on the progress of the editing which continued into 1972. John’s letter of 11 January 1972 stated: When I finish with the ms., I shall return it to Mahler. I have just been asked to serve as Associate Editor of Sida—which is just what I’ve been doing all along! By April John had sent the edited Rorippa manuscript to Mahler, and the latter replied to me, 13 April 1972, that my “manuscript is already at the printers.” Thieret received the galley proof sheets and then sent them on to me for read- ing. Together that summer at Stone Laboratory, John and I read the galley sheets. Page proofs came on 25 July 1972, according to Mahler's letter to me, on 8 Au- gust 1972. lreturned the final page proofs, and by 27 September, | was complet- ing the transaction to purchase 200 copies of my study of Rorippa. My mono- graph, “The Taxonomy and Distribution of the genus Rorippa (Cruciferae) in North America” appeared in Sida 4(4):279-430. 1972. John W. Thieret was listed as the associate editor and William F. Mahler as the copyright owner. Had it not been for Thieret entering the situation, my monograph might never have been published. I owed hima great debt of gratitude for his unselfish help in editing my publication. Again, Thieret and Stone Laboratory (1972) In the summer of 1972, Thieret returned to the Stone Laboratory and again taught Field Botany. This summer he had prepared a book of mimeographed course handouts that were to be given to each student. Nearly everything that John wrote had some humor associated with it. At the bottom of the Table of Contents page was the notation, HAVE YOU THANKED A GREEN PLANT TODAY? y masters student David L. Moore took the course from Thieret that sum- mer. At that time also, David was studying the changes, since 1895, in the aquatic vascular plant-flora of East Harbor State Park, Ottawa County, Ohio (Completed 1973; portion published in Ohio J. Sci. 76:78-86. 1976). David later in 1976 earned the Ph.D. degree from The Ohio State University by studying the distribution of freshwater algae in northeastern Ohio with Dr. Clarence E. Taft. Since then David has taught botany courses in the Department of Biology at Utica College of Syracuse University, Utica, New York. Beginning in 1993, he returned to the Stone Laboratory and has taught the aquatic plants course every summer ex- cept 2005. The course was not taught that summer. In response to my request for information about John Thieret’s 1972 Field Botany class, David provided the following three paragraphs: As | retrieved an OSU soft cover binder from my office shelf, | was reminded of the passage of 34 years by the yellowed, brown- edged, | hand-written pa ages! ies summer 1972 Field B otany Course W ith Dr. Joka ae came pac tome as | iad SS the pages of notes. We learned a total of 19] taxa in those five 25 plant families. Indeed it was the first time I had ever seen floral formulae and floral ae My favorite plant was Matricaria matricarioides, the Pine- 40 BRIT.ORG/SIDA 22(1) apple Weed, which Dr. Thieret picked by the docks and gleefully crushed in his palm so we could sniff the fragrance of fresh pineapple. At the first class session Dr. Thieret went over the syllabus. Everyone laughed when he referred to examinations as “mental enemas.” I will always remember the cover of his mimeographed course handouts which he provided to us that first day. The front page pictured Shiva Nataraja, the Hidu Lord of the Dance ona Nuphar advena leaf which protruded from the fruit of Nelumbo lutea. In the right rear hand was a leaf of Rhus toxicodendron (Poison Ivy) while in her left rear hand was an Aescylus glabra leaf (Buckeye). The afternoon of July 8 was my first trip to Pelee Island. It was probably one of the highlights of the session, and where I first saw Populus grandidentata and Diplotaxis. | still don’t know if it was D. tenuifolia or D. muralis. At the end of the session, just before the final exam, Dr. Thieret read a poem he had composed on the porch of Bayview House where he and his family lived. In it he dedicates a verse to each of the students in the class. After he read it he gave each of us a two page mimeographed copy on yellow paper. I kept my copy in my notebook after we finished the course. | retyped it with the same spelling, punctuation and format that he used except that I placed it in two columns. There is one misspelling left intact - “cannister”- and the two lower case letters which should have been capitalized. All of his “poetic license was retained). | have selected the following six verses of John’s poem, titled “Botany 610 In RHYM, Summer 1972,” for publication here: BoTANy 610 IN RuHyYM SUMMER 1972 This is the forest primeval That lovely young Madchen Miss Schiller The murmuring Acer and Celtis is is the class that beneath it Romped cheerfully after their Leader. This is the class below pictured In verses rhythmic, poetic. ‘Twas a pleasure to watch Mr. Moore Whose knowledge of plants was not poor. Like an eye-talian barber He combs over east Harbor And there gave Fearless Leader a tour. And now it is time for Miss Kneller [tried hard this plant world to sell ‘er. By selling my wares lincreased her cares. So often “Work hard, dear” Id tell ‘er. With plant names | really did fill ‘er. But a turtle so cool She found ina pool Was really, | think, what did thrill ‘er. There goes that bold man Rick McGill Out He works day and night Wit And does get his plant press quite fill. ora botanical kill. vallof his might And last, but not least, Fearless Leader. Whose disciples learned oak, elm, and cedar. How oft, on The Rock He did gather his flock And exhort them there, much like old Peter. STUCKEY, JOHN W. THIERET, VALUABLE BOTANICAL FRIEND 41 At the conclusion of the poem, John wrote, “... presented with thanks and appreciation to my Stone Lab Students. Long may they flower!” The verse pertaining to Rick McGill is noteworthy as he was working with me for his master’s degree making acom- parison of the vascular plant flora of the two lakes in northern Champaign County, Ohio (completed 1973; not pub- lished). Field Botany, Course Handouts Summer 1993, John W. Thieret, Tom Duncan, and the “Flora of the Erie Islands” (1972-2005) Tom Duncan, an undergraduate major- ing in education and working asa teach- ing assistant in chemistry at the The Ohio State University, introduced him- self to me during his sophomore year in the early spring of 1968. Later that spring he accompanied me on field trips learning the local flora, and in the spring of 1969 he began working as an assistant under my direction in the OSU Her- barium. That summer I invited Tom to serve as my research assistant at the FT. Stone Laboratory. He continued to serve part-time as my research assistant on the Flora of the Erie Islands until he graduated at the end of the fall quarter 1970. Tom continued working on the Flora manuscript from 1971 through 1973 while he was a graduate student at the University of Mighican. He earned his Ph. D.in Botany there in 1976 and was employed as an assistant professor botany beginning 1 July 1976 at the University of California, Berkeley. He was granted tenure there in 1982, was the Director of the Herbarium 1982-1991, then be- came Director of the Museum Informatics Project until his retirement in 2002 as an associate professor emeritus at age 54. I first met John W. Thieret in July 1972 at the Franz Theodore Stone Labo- ratory on Gibraltar Island in Lake Erie. John was at Stone Laboratory for the summer to teach Field Botany. I was visiting Stone Laboratory for a few days to discuss with Ronald L. Stuckey progress on the preparation of our manuscript titled The Flora of the Erie Islands: Its Origin, History, and Change. Stuckey initiated a project on the flora of the Erie Islands in the mid-60s. In 1969, I undertook a project as his research assistant on the changes in the flora of the seven small islands in western Lake Erie (The Michigan Botanist 9:175- 200. 1970), and was invited to collaborate on the more inclusive project in 1970. Cover page of John W. Thieret’s Field Botany, Course Handouts, Summer 1972. 42 BRIT.ORG/SIDA 22(1) By 1972 we were in the initial stages of preparing together a manuscript de- scribing the results of our research. John Thieret was spending his second summer as an instructor at Stone Laboratory. He had become intensely focused on the flora of the islands and was interested in discussing with me the progress to date on my collaboration with Stuckey in this regard. One area we discussed was the lack of information in previous floras about cultivated plants on the islands. Cultivated species are of interest because they are an important source for the adventive and naturalized flora of the islands, are a conspicuous aspect of the flora, and are an important feature of the flora for the residents of the is- lands. John and | concluded that we would undertake a survey of gardens on South Bass Island and compile a list of the cultivated flora for inclusion in the manuscript Stuckey and | were preparing on the total flora. We spent a day wandering the roads of South Bass Island stopping at houses and asking local residents if we could examine and record the names of the plants in their gardens. With a copy of Liberty Hyde Bailey’s Manual of Culti- vated Plants (1949), in hand, we were able to prepare a preliminary list of plants. Ultimately this list included 258 taxa. John often stated that our field trip on South Bass Island that day was one of his favorite memories of his time at Lake Erie. After our initial meeting in 1972 we continued to correspond and talk on the telephone about the progress of the flora manuscript. In 1982 John brought the nomenclature of our list of cultivated plants up to date using Hortus Third (1976), the nomenclatural ref- erence for cultivated plants at that time. In his letter of 3 February 1982 he con- cluded by saying, “best wishes from John. I wish we could wander over South Bass Island again. Fun, fun, fun.” Whenever | think of these events, lam always reminded of John’s charm- ing personality, his appreciation of natural history, and his joy in studying plants. His interest in the flora of the Erie Islands continued for over 30 years until the end of his life. Stuckey and I completed our manuscript in 1976 and submitted it to the Ohio Biological Survey for publication. The Survey accepted the manuscript for publication and planned for publication in 1979. John volunteered to serve as editor for the Survey and worked extensively on the manuscript. Unfortu- nately, the Ohio Biological Survey could not publish the manuscript at that time, partially because of the lack of funds, and Stuckey’s involvement with other projects for the Survey. For the next 20 years the manuscript and three edited copies resided in Stuckey’s office with an additional copy in the Franz Theodore Stone Laboratory library. During this period, John continued his interest and frequently asked both Stuckey and me if we would complete the manuscript and find a way to render itin a form for wide distribution. During a visit to The Ohio State University in — STUCKEY, JOHN W. THIERET, VALUABLE BOTANICAL FRIEND 43 1999, Stuckey and I discussed what might be done to resurrect the manuscript, complete it with all editorial suggestions incorporated, and either publish it or distribute it electronically. John was enthusiastic about this turn of events and offered his services to help us in any way he could. Progress in this direction was made but no definite arrangements about a means of distribution had been made by the time of John’s death. We are investigating the possibility of distributing it through the digital Knowledge Bank at The Ohio State University. Ronald L. Stuckey and | are de- termined tocomplete this task to honor John’s long-standing interest in the Erie Islands, and we plan to dedicate the completed flora to him. Thieret and Stuckey’s Promotion (1972) Later in the year 1972, John made an important contribution to my career. He wrote a letter, dated 19 October, supporting my promotion to full professor in The Ohio State University. Addressed to Botany Department Chairman John A. Schmitt, it was short and to the point, quoted here in its entirety: It seems to me that Dr. Stuckey has shown himself to be a competent and indefatigable researcher. His frequent publications attest to his capabilities not only as an “original researcher” but asa skilled synthesizer of the work of others. [PS] John: This is a trite, trivial, and, for me, traumatic letter. | simply cannot rise to greater heights. Even though the subject is worthy of those greater heights. Jlohn Again, Thieret and Stone Laboratory (1974, 1977, 1981, 1983, 1986) After 18 years as Director of the Stone Laboratory, Putnam retired following the summer session of 1973, and College Dean Richard H. Bohning appointed Dr. Charles E. Herdendorf to succeed him. Now Professor and Director Emeritus, Herdendorf, on 23 March 2006, provided his remembrances of John Thieret, briefly reviewing the succeeding summers John taught Field Botany at the Labo- ratory—a total of seven summers over a 15 year period. I first met Dr. John W. Thieret, in 1971 at the Stone Laboratory where he was teaching a summer course in Field Botany, and I was beginning to organize the newly founded Center for Lake Erie Area Research (CLEAR) at the Laboratory. The next year we both served on the summer faculty of Stone Laboratory, Dr. Thieret again teaching Field Botany, while I taught Physical Limnology. Finan- cial constraints in 1973 caused the cancellation of classes at the Laboratory, but by 1974 I had assumed the duties of Laboratory Director, and Dr. Thieret was invited again to teach field botany. In 1974, Dr. Thieret and his class also par- ticipated in a CLEAR research project titled Ecological Analysis Related to the Proposed Seawall Improvement and Ground Rehabilitation Project at Perry’s Vic- tory Monument, Put-in-Bay, Ohio. Dr. Thieret returned to Stone Laboratory to teach Field Botany in 1977, 1981, 1983, and 1986. On June 30, 1983, he also par- ticipated in the Stone Laboratory Guest Lecture Series by presenting a seminar titled The Muskegand |: Exploring for Plants in Canada’s Northwest Territories. 44 BRIT.ORG/SIDA 22(1) Other than his account of the boreal plants along the new Yellowknife High- way, one of his most memorable descriptions was that of the voracious infesta- tion of black flies encountered around Great Slave Lake. John Thieret is fondly remembered for his dry sense of humor and fascinating stories of field experi- ences. Some of his students may have found him to be excessively demanding, but all received a quality introduction to the study of Field Botany. I was not present at the Stone Laboratory in the summer of 1977, but my masters graduate student John R. Wehrmeister attended and took the Field Botany course from Theiret. Student Wehrmeister was researching the ecologi- cal life cycle of the pondweed Potamogton crispus in North America (completed 1978; published in part The Michigan Botanist 31:3-16. 1992). In 1981, Mr. Wehrmeister earned an M.D. degree from the Medical College of Ohio in To- ledo, and he has had a successful career as a physician in the practice of inter- nal medicine in the Maumee-Toledo, Ohio, area. In reply to my request, he wrote the following commentary: lam pleased to offer thoughts about Dr. Theiret. His personality and teaching style has had a lasting 1m act on me. | had heard Dr. Thieret give a lecture on the botany of the Northwest Territories,Canada = > pane was a student at the University of Michigan Biological Station at Pellston. Dr. Theiret was a visiting faculty member there in 1975, but I did not get to know him that summer. Later in 1977, while studying at The Ohio State University, I took his Field Botany course at the Stone Laboratory. Following the course, in late August and early September, my friend Ed Toth, and | accompanied Dr. Thieret on a plant collecting trip to Louisiana, and upon returning, visited with him at his home in Highland Heights, Kentucky. Thoughts of him are full of fond memories. He was a delightful man, a favorite but serious teacher, and a scholar with a great sense of humor. Memory of his facial expression includes what seemed to be a permanent grin. “Dr. Thieret anecdotes” were the order of the day among his students. Good-natured though he was, in contrast he was strongly serious in the classroom. With some embar- rassment | remember being admonished by him for the poor effort | showed in mounting herbarium specimens for an early class project. At my request Dr. Theiret wrote a letter of recommendation for some job or school position to which I'd applied. He thereupon sent mea purported carbon copy of the letter which, perhaps needless to say, was far from complementary. Luckily this was one of his pranks. The real letter contained the usual laudatory comments. A few additional observations of Theiret’s summers at the Stone Labora- tory are noteworthy. By 1981, he had modified the “Course Handouts” book. On the cover that summer was a branch showing the leaves of the hackberry tree, one of the most common trees that grew on the shallow limestone and dolo- mite bedrock. On page 4 was the quotation, “The beginning of wisdom is to call things by their right names.—Oriental proverb.” In his course the students only were required to learn the scientific names of the plants. I did not agree with him; rather I taught students to learn both the STUCKEY, JOHN W. THIERET, VALUABLE BOTANICAL FRIEND 45 John W. Thieret’s Field Botany Class at the F.T. Stone ii etal eke rarer in oe Dare eee quarry on Kelley ‘s Island, Erie County, Ohio, by Ronald L. Stuckey, Ju uly hn W. Thieret, George F, Buddell, IL, assistant; five men students, in alphabetical ne not matched to individual i in the ie Ron Baker, Doug Brewer, John George, Mark Harrell, and Dan Wojta scientific and common names, which would allow them to communicate with persons of many different educational backgrounds and professions. His edito- rial style gave me concern also, but I never discussed it with him. He continu- ally overused the dash, which therefore changed my writing style. I also be- came annoyed when he was editing the “Flora of the Islands.” | often listed several plants in sequence. He would change the sequence by aligning them in alphabetical order. My choice of sequence usually was to list them from the most common or typical to the less common in a particular habitat. My align- ment was ecological related as to where the plants grew. Plants do not grow in places alphabetically, but he apparently was not aware that my order of the plants also had a meanin John liked to talk as he himself has written. During two other summers he gave a presentation in the Guest Lecture Series, on 19 July 1977 and 30 June 1983. Both times, his title was “The Muskeg and I,” which was just as delightful as it was the first time with the more formal title, when he gave it in the sum- mer of 1971. Andrea Wilson, now Mrs. Matthew Schlageter in the Southeastern Devel- opment Office of The Ohio State University, Bonita Springs, Florida, came in 46 BRIT.ORG/SIDA 22(1) the summer of 1982 as Director Herdendorl’s Laboratory Secretary and Admin- istrative Assistant. She knew John Thieret from the 1983 and 1986 summers at the Laboratory. Her statement sent to me 24 March 2006 concisely summa- rizes John’s presence there. | vividly recall Dr. John Thieret as a very kind and thoughtful man. He al- ways had an encouraging word and a smile for everyone. He was devoted to his ne touched. — teaching and research and was well respected by all whose lives His memory will continue to live not only in his publications, but also in the knowledge he shared with his students. Thieret, Book Reviews, and Stuckey (1973-1979) During the five year period of 1973-1979, Thieret sent me books to review for the journal Economic Botany. For 25 years, 1959-1984, John voluntarily served as book editor for Economic Botany. John’s strive for perfection gave the journal “a reputation as having one of the most comprehensive and best edited book review sections of any professional journal. When at times the acid pen of a reviewer might be too harsh on an author, John would always suggest a critical but more even-handed approach.” (W. Hardy Eshbaugh, Economic Botany 38(4): 498. 1984). John usually sent me local flora or plant taxonomy books to review. Ini- tially he sent me two books with a letter dated 24 September 1973. The book titles were A Flora of the White Mountains, California and Nevada by Robert M. Lloyd and Richard S. Mitchell 1973) and The Genus Lesquerella (Cruciferae) in North America by Reed C. Rollins and Elizabeth A. Shaw (1973). Thieret’s suggestions were: QQ Probably the best thing to do so far as the flora is concerned is simply to produce a “notice” of the work. A full-blown “review” is not necessary. If you can get the notice and the review to me in about 3 months or less, I'd be ecstatic. Allot ner 150- 200 words for the flora, about 300 for the mustard opus. Of course, if you fee you really need more—or do not need so many—the final decision is yours. | wrote 10 book reviews for Economic Botany. They and all of my other book reviews are available in my archives at the Hunt Institute for Botanical Docu- mentation, Pittsburgh, Pennsylvania. John wrote his own review about his 25 years as Book Review editor. He considered it as “one singularly devoid of ex- citement.” One colleague, with perhaps a touch of jocularity, called him the “dean among U.S. book review editors” (Economic Botany 40:24-26. 1986). Thieret, Aquatic Plants, and Stuckey (1982-1983) [had forgotten that during the summer of 1983 when John was teaching at the Stone Laboratory, he asked me to review the book manuscript he was complet- ing for the late Ernest O. Beal on the “Aquatic and Wetland Plants of Kentucky.” On 26 August 1983 John wrote: | hope you remember that you have gone over both major sections of the book, ie., the dicots (sum- STUCKEY, JOHN W. THIERET, VALUABLE BOTANICAL FRIEND 47 mer 1982) and the monocots (summer 1983). The comments you made on the monocots were (as were those on the dicots) gratefully peceveds Most of them have been incorporated into the book, | improving same! | appreciate your r prose. I’m working hard on the book, hop- ing to meet a deadline of | November 1983 ibe getting he book to the publisher. Best wishes. I further helped John with the book by writing on 2 September 1983 a support- ing letter for its publication to Richard Hannan of the Kentucky Nature Pre- serves Commission: “Dr. John Thieret has asked me to write in behalf of his manuscript co- authored with the late Dr. Beal on the aquatic and wetland plants of Kentucky. This manuscript is very carefully prepared in all aspects. Dr. Thieret has gone to great lengths to check the accuracy of identifications of specimens seen, the reliability of literature records and citations, and the conciseness and correct usage of technical terminology. His work in this regard is very well done, and adds great credibility to a work of this type. Very few states have books on their aquatic plants, and this book will be a welcome addition to those books on aquatic plants. Perhaps, more important, however, is that this book will be extremely valuable in a state which has no “modern” general flora. Consequently, this book will be a valuable contribu- tion to that “modern” general flora that needs to be generated. Those needing floristic information will at least be able to get current information on aquatic and wetland species. I trust that you find this contribution worthy of publica- tion, as lanxiously look forward to seeing it in print In the book’s acknowledgements by Thieret, my name, along with many others, appears among those who “gave aid during the preparation of this book.” Thieret and Stuckey Since Summers at Stone Laboratory (1996-2005) Since the summer days of the 1970s and 1980s at the Stone Laboratory, most of my contacts with John Thieret have been by telephone, except the following interactions which are noteworthy. When John became editor of the Transac- tions of the Kentucky Academy of Science, [thought he could help me with pub- lishing some of my writings that were nearly completed on the controversial North American botanist Constantine Samuel Rafinesque (1783-1840). | had published several papers on Rafinesque’s botanical studies earlier in my career. Since Rafinesque had held the position of Professor of Botany and Natural His- tory at Transylvania University in Lexington, Kentucky, from 1819-1826, my contributions were appropriate for the Kentucky Academy’s publication. With the assistance of my botanical friend, James S. Pringle of the Royal Botanical Gardens in Hamilton, Ontario, Canada, we published, with John’s editorial as- sistance, “Common names of vascular plants reported by C.S. Rafinesque in an 1819 descriptive outline of four vegetation regions of Kentucky” Clrans. Ken- tucky Acad. Sci. 58:9-19. 1997). At the Bicentennial Celebration of Rafinesque’s 200 birthday held on the campus of Transylvania University, 21-22 October 48 BRIT.ORG/SIDA 22(1) 1983, | spoke on “Opinions of Rafinesque expressed by his American botanical contemporaries” (Bartonia 52:26-41. 1986; reprinted with some changes in Charles Boewe (Profiles of Rafinesque. Chapter 10. 2003). Following that event, and as a contribution toward my studies of the pioneer botanists of the Ohio River Valley, | wrote an extensive manuscript on “Rafinesque’s Botanical Pur- suits in the Ohio Valley 1818-1826).” With John Theiret’s editorial skills, 14 years later this manuscript was completed and published by the Kentucky Academy of Science in its newly titled Journal of the Kentucky Academy of Science (59(2):111-157. 1998). John appreciated and was very supportive of my scholarship on the his- tory of nineteenth century North American plant taxonomists and their work. He asked me for a short project he could contribute about Rafinesque in Ken- tucky. | suggested he return to the localities in the state where Rafinesque ob- tained plants that he named as new to science. He was delighted with the idea, and by automobile during a week in July 1999, John and his friend, David M. Brandenburg of the Dawes Arboretum, Newark, Ohio, retraced a botanical ex- cursion that Rafinesque took during 1823 in central and south-central Kentucky, Rafinesque’s itinerary was outlined in his autobiography, Life of Travels 1836), which took him two months to complete on foot. A short description of the Thieret-Brandenburg trip appeared in the Flora of North America Newsletter (13:;G-4). 1999. July-December). David and John wrote the full story of the trip under the title, “Rafinesque and Us” (Lloydiana 6():4-9. 2001). During retirement, my friend George W. Paulson, emeritus professor of medicine in the Medical Center of The Ohio State University, has been writing about early medical practices of the pioneer physicians in the central Ohio area. Among these physicians was Samuel Thomson (1769-1843) who discovered that ear — chemical extracts from the leaves and seeds of the plant, Lobelia inflata, had medical properties that potentially would cure headaches, tinnitus, syphilis, and especially asan expectorant to loosen bronchial congestion. Paulson wanted to write a paper on Thomson, and asked me to help with it since | would know about the Lobelia plant. Most of the paper was written by George, and I edited it, but George could not locate a journal for its publication, so he asked for my help. [thought Lloydiana, a quarterly periodical of articles for the popular au- dience published by the Lloyd Library in Cincinnati, would be an ideal place for its publication. I talked to John Thieret on the telephone and asked him for his review of the article and editorial assistance. John,a member of the Board of Trustees of the Lloyd Library, helped with revision and editing of the article, which appeared in Lloydiana (8(3):2-6. 2004. Summer). Lloyd Library Direc- tor and editor of the periodical noted in her column, “From the Director,” that the Thomson-Lobelia paper was indeed an “interesting article.” John Thieret so ably helped make it possible to be printed in the Lloyd Library newsletter. Nearly every time John and [hada telephone conversation since our teach- —_—_ STUCKEY, JOHN W. THIERET, VALUABLE BOTANICAL FRIEND 49 ing days at the Stone Laboratory, he always asked about progress on the Flora of the Erie Islands, and he wanted me to finish it. | always had to think of some excuse, mostly that I was involved in other more worthwhile projects. Some relief came for me when Tom Duncan returned to my office in 1999, and he agreed to take all of the various components related to the project with him to Berkeley, California. There he has worked on the Flora to reflect our knowledge of it, as it was in the 1970s. Tom wrote of his role in this effort in this paper. For 30 years, from about 1971 to 2001, | was at work preparing another kind of book about South Bass and Gibraltar Islands. Titled “Lost Stories,” the book includes Theresa Thorndale’s “Island Jottings” about selected places on these islands of the 1890s that she published in the Sandusky Register. My contribu- tion describes these same places on the islandsa century later during the 1990s. This book won the 2003 Florence Roberts Head Book Award given by the Ohioana Library Association of Columbus, Ohio. In his last known note to me, John Thieret wrote, “Many thanks for the info[rmation] on your award for the Put-in-Bay book. ’'m so glad that you did such a book. You certainly deserve the award.” I sometimes think that for John, it may have been a consolation prize for the Flora of the Erie Islands. John Thieret was a most unusual botanist. I know of no one else like him. His dedication, his kindness, his sense of humor, his field knowledge of plants, and the numerous and varied ways he helped me through my career has been a most worthwhile and enjoyable adventure. ACKNOWLEDGMENTS AND SOURCES OF INFORMATION lam especially grateful and thankful to Kathy Royer for preparing the final typewritten manuscript of this paper for publication in Sida. lalso extend grate- ful thanks to David lL Moore, Tom Duncan, Charles E. Herdendorf, John R.Wehrmeister, and Andrea (Wilson) Schlageter for their written contributions to this paper. Notes made during telephone conversations with the above indi- viduals and the letters referenced in this paper are filed under the title of this publication in the Archives of Ronald L. Stuckey and in the Archives of the Hunt Institute for Botanical Documentation, Pittsburgh, Pennsylvania. As an added note, former Stone Laboratory Director Loren S. Putnam died during the writing of this paper, 19 April 2006, at age 92.5 years. JOHN W. THIERET, COLLEAGUE AND EDITOR FRIEND (1969-2005) William F. Mahler 204 W Clara lowa Park, Texas 76367-1304, U.S.A. John Thieret’s reputation as a dedicated botanist and unique individual pre- ceded our actual meeting, which occurred sometime in the late sixties. Over the years we drifted apart, but the impact John made on my professional career and my success with Sida, Contributions to Botany remains strong. Although he spent the night at our house only a couple of times when he visited the SMU herbarium, the field trips we took in my 1969 red Ford pickup were the high- lights of our botanical relationship. Our biggest collecting trip was to Guadalajara, Mexico in April of 1970. We traveled by way of the Chihuahuan Desert and then returned through the Sonoran Desert. The pickup was the perfect mode of transportation for a long- range collecting trip because of the saddle tanks and the camper shell. We could travel six hundred miles between fill-ups and could sleep in the back if we couldn't find lodging. Also, there was plenty of room for all our collecting sup- plies. The decision to take this particular trip was made on a lark. We'd heard of others doing this and so we thought we'd do the same. As it turned out, we actu- ally did find quite a few noteworthy specimens. When Rupert Barneby found out where we were going, he suggested we visit some locations for the legumes that he had collected some years back. He needed some flowering specimens. We were successful in finding some specimens north of Zacatecas, which he cited in his monograph on Dalea. The trip was far from boring. North of Guadalajara, John injured his ankle while trying to get a specimen from the branch of a tree. He knew he had sprained his ankle, but he was afraid he might have broken it, too. An x-ray ina hospital in Guadalajara showed no apparent broken bones, but judging by the amount of pain and the routine swelling by the end of the day, John felt that it was more than just a sprain. However, with my expertise in speaking Spanish, | managed to find ice at motels to treat the swelling. We were able to continue with our collecting trip. John was always the teacher. To pass the time when driving, we would grill each other on botanical glossary terms and other aspects of botany. I had never taken an economic botany course, but I survived the equivalent of one after experiencing the tropics with him. He was the opposite of me, since | eat to live SIDA 22(1): 51 — 54. 2006 52 BRIT.ORG/SIDA 22(1) John Thieret, Chihuahuan Desert, Mexico, April 1970. — and he lived to eat. In the restaurants, meals were scrutinized very diligently ean and I was often quizzed on the content. He ate and sampled many kinds of fruits from the markets, many of which | had never heard of. In the field, he was al- ways tasting and smelling the fruits, leaves, etc. and discussing the results a spitting out the remains. His knowledge of tropical fruits was uncanny. We followed the coastline on our way home. As we neared the border, we drove past a bakery in a small town. John’s acute sense of smell overcame him and | had to turn around so he could stock up on some bread before we left Mexico. My common sense told me to refrain since we had come that far with no digestive problems. As it turned out, he did get a light case of diarrhea, but | did not pursue the issue. Ss ter At Mazatlan, we bought more newspapers (by the kilos) for our collections at the local newspaper office, and we spent our last night in Mexico south of Nogales, Arizona. U.S. Customs wanted time to go through our specimens look- ing for rusts for one of their projects. We went to dinner Cunch) and when we returned, customs had just finished. We headed back to Texas with our many collections and John’s foot still in pain. Once we got on the road, John got anxious to get home and check out his injury. We drove all night and stopped in Abilene, Texas for gas the next morn- ing. At Love Field in Dallas, I let him out at the airport where he caught a plane MAHLER, JOHN W. THIERET, COLLEAGUE AND ED 53 John Thieret, Chihuahuan Desert, Mexico, April 1970. i and I went on home. I arrived home about five o'clock in the afternoon after being up and at the wheel for thirty-six hours. John told me later that they had found a hairline crack in one of his ankle bones in addition to the sprain. Although John was not one for attending meetings, he agreed to accom- pany me in the red Ford pickup to the Southwestern Association of Naturalists meeting in Tucson, Arizona. During registration, he headed over toa young man and shook hands. The fellow seemed confused and asked John if they had met before. John informed him that they had been officemates for ten years back in Chicago. John’s appearance was much different because he had shaved his beard that he had always had. Everyone had a good laugh about it, especially John. The field trips were productive and entertaining, but John’s most valuable contribution to my career was his involvement in Sida. Lloyd Shinners had told me that I needed to meet John when I first arrived at SMU, and he was right. When I inherited Sida, he, along with others, encouraged me to continue its publication, as opposed to others who felt | should drop the journal. With the help of Mrs. Helen Koresh, a sister of Lloyd's, | was able to clear the financial obligations Lloyd had encumbered with the authors of unpublished manu- scripts. [sent John some articles for him to review that I had written about Lloyd. John really “cleaned them up’ and returned them with more red ink than black. 54 BRIT.ORG/SIDA 22(1) This was the beginning of his “internship” and status as Associate Editor of Sida that he held until his death. | have always considered myself lucky to have associated with many indi- viduals whose knowledg furthered my career goals. John was one of those individuals. The wide range e and insight enriched my professional knowledge and of things I learned from John—both professionally and asa friend—will always figure significantly in my lile. JOHN W. THIERET, ASSOCIATE EDITOR OF SIDA (1972-2005) Barney Lipscomb Botanical Researcl h Institute of Texas 509 Pecan Street Fort Worth, Texas 76102-4060, U.S.A. John W. Thieret was an extraordinary and innovative teacher who shared his knowledge freely. Without doubt he was an influential botanist and one of the most distinguished editors of the 20th century at least in North America. “Big John,” as sometimes called him, was my steady and faithful cheerleader, Sida supporter, and crusader extraordinaire. He was always there whenever I needed advice or help with Sida or for that matter anything. Notwithstanding John’s many obligations, he called me after receiving each and every issue of Sida. And the voice I heard each time was inspirational, always filled with praise for the good but soft and tender on the good that could be better. John’s source of inspiration was perhaps from the many years of editorial experience with Eco- nomic Botany, Flora of North America, and the Journal of the Kentucky Acad- emy of Science. John knew first hand the amount of work involved and maybe sensed there was always room fora little more praise than was given [to editors everywhere]. “Keep up the fine work good ole fellow! How are you doing? How are you holding up? Let’s go on a field trip! Now is there anything you would like for me to do? Edit a manuscript? Review a book and write a review? What- ever it is just send it my way and [ll promptly return it to you.” John Thieret was forever supportive and never failed to do what he said he would do. He was always thinking of how he could help the other person and I always felt like | was that other person. John so loved Sida that he gave nearly 40 years of his life. To John be the glory—great things he has done for Sida, Contributions to Botany! This is my tribute to him. — Highlights of John Thieret’s contributions to Sida Ralph Thompson's (2006) excellent tribute provides a detailed account of the great service John Thieret rendered to botany and academia, his chosen fields. My tribute will recount his great and lasting contributions toa journal he cared so much about, Sida, Contributions to Botany. John’s love and early relationship with Sida was tied closely to that of his colleague and friend Lloyd H. Shinners at Southern Methodist University. Therefore, I pick up John’s trail of communi- cation with Shinners even before Shinners started Sida in 1962. 1961—According to Ginsburg (2002), “Thieret was on staff of the Chicago Natural History Museum when he wrote to Shinners in 1961 to inquire about academic SIDA 22(1): 55 — 63. 2006 56 BRIT.ORG/SIDA 22(1) positions for botanists. Shinners was very helpful, and when Thieret moved to the University of Southwestern Louisiana [Lafayette] the two became friends and close collaborators.” Thieret wrote, “I really think that a private line ought to be established between your herbarium and ours...” Music was another shared enthusiasm. 1962—L loyd H. Shinners alone wrote, edited, and published the first two issues of Sida which were distributed on November 23rd. He said that he would have to publish his own journal because his own articles would never be published otherwise. So it was quite a tribute when he published others’ papers. 1963—With the publication of the third issue of Sida on December 13, John Thieret became the fourth author aside from Shinners to have a paper published in the journal, behind W. Phillip Osborne, Walter H. Lewis, and Arthur Cronquist. John’s 54 page Yellowknife Highway paper followed Arthur Cronquist’s paper, The taxonomic significance of evolutionary parallelism. — Botanical survey along the Yellowknife Highway, Northwest Territories, Canada. |: cata- logue of the flora. Sida 1(3):117-170. 1964—Five of John’s manuscripts were submitted and published in 1964. The first issue contained the second part of John’s botanical survey along the Yellowknife Highway. In addition, Sida 1, no. 4 contained another significant and memorable event for John Thieret. A new flowering plant species was named in his honor for the first time ever. Lloyd Shinners (1964) named Scutellaria thieretii (Labiatae) for John. Lloyd wrote, “The Louisiana plant is considered to be one more in a group of very closely related species, and in honor of an ener- getic and productive collector it is named... am indebted to Dr. Thieret for sup- plying the two SMU collections, and for the loan of mounted specimens from the University of Southwestern Louisiana.” And so the relationship between John and Lloyd strengthens. Botanical survey along the Yellowknife Highway, Northwest Territories, Canada. Il: veg- etation. Sida 1(4):187-239 Shinners began work on Sida 1(5), but it was a struggle to get the manuscripts ready for the printer (Ginsburg 2002). Thieret apparently offered help with is- sue no. 5, but | could find no correspondence indicating whether or not Shinners ever took John up on his offer. Maybe John helped by submitting well-written and superbly edited manuscripts for Lloyd to publish. Fatoua villosa (Moraceae) in Louisiana: new to North America. Sida 1(4):248. Lysimachia japonica (Primulaceae) and Clinopodium gracile (Labiatae) in Louisiana: new to the United States. Sida 1(4):249-250. More additions to the Louisiana flora. Sida 1(5):294-295. Eriogonum annuum (Polygonaceae): biennial in Nebraska. Sida 1(6):382. a LIPSCOMB, JOHN W. THIERET, ASSOCIATE EDITOR 57 1965—There was a break in the action for John as far as submitting papers to Sida. John did not have a single paper in either of the two Sida issues published in 1965. Regardless, John apparently stayed in close contact with Lloyd. pal 1966—The next documented account of cor Lloyd took place on January 26. I ce between John and “Lam sending, for possible publication in SIDA, a manuscript entitled “Seeds of Some United States Phytolaccaceae and Aizoaceae.” The article was published on September 27. Seeds of some United States Phytolaccaceae and Aizoaceae. Sida 2:352—360. Sometime in 1966, Shinners had his first “all out” case of insulin shock and after hearing about that, Thieret wrote, “I was appalled to hear your account of the insulin shock.” I suspect John once again offered Lloyd help with Sida but ap- parently no letter with such words are in Shinners’ archives. Shinners recov- ered from the insulin attack and published a fourth issue of Sida in 1966: vol- ume 2, number 6 was published on December 30. 1967—According to Ginsburg (2002) Shinners was probably exhausted after publishing four issues of Sida in 1966 and even more exhausted after his second NSF grant ended in May 1967. Thieret apparently kept in close enough contact to know the details. According to Ginsburg (2002), “He [Thieret] was willing to come to Dallas for a weekend to help in getting an issue together, or he would read and edit anything that Shinners m ight send.” Ina letter to Shinners, Thieret once again offered assistance with Sida. “Sida is a most worthy cause. To help out is a privilege...” John not only continued to correspond with Lloyd, but he also submitted manu- scripts to Sida and Lloyd published four of John’s papers in 1967. Supraspecific classification in the Scrophulariaceae: a review. Sida 3(2):87-106. Thirty additions to the Louisiana flora. Sida 3(3):123-127. Cyperus subgenus kyllinga (Cyperaceae) in the continental United States. Sida 3(3):128- 136. [with AJ. Delahouhoussaye] Neogaerrhinum kelloggii (Greene) Thieret, comb. nov. (Scrophulariaceae). Sida 3(3):187, 1968—John Thieret wrote Lloyd on April 18, 1968. Glad to hear, also, that SIDA is such a well-going concern. Once again I make an offer to come to Dallas for a weekend if I can be of help—real help—in getting together an issue. Also, I'll be glad to read and edit anything you send over. Even that Czech paper—which you specifically said not to edit, you will recall.” Two issues of Sida were published in 1968 but John did not have any papers in either issue. 1969—Shinners published two issues of Sida in 1969: 3(5) and 3(6), August and 58 BRIT.ORG/SIDA 22(1) November respectively. In between those two issues John sent Lloyd a letter dated September 9, 1969: “| was surprised to get the Bigelowia [Anderson] opus. Have you decided to establish an editorial board to “protect” your contributions against non-promotion? (Ea!) After hearing about another round of Shinners’ health problems, Thieret again offered to help proof read Sida. “You are willing | recall to have me take care of the index lor vol. 4 ol Sida. Right?” Meanwhile John submitted four manuscripts for publication. Undoubtedly they were submitted in excellent shape and Lloyd probably had to do very little work on these papers to get them ready for publication. ; Sagittaria guayanensis (Alismataceae) in Louisiana:new to the United States. Sida 3(6):445. Rumex obovatus and Rumex paraguayensis (Polygonaceae) in Louisiana: new to North America. Sida 3(6):445-446. actea (Rafinesque) Thieret, comb. nov. (Leguminosae). Sida 3(6):446. — — Baptisia Trifolium vesiculosum (Leguminosae) in Mississippi and Louisiana: new to North America. Sida 3(6):446-447. 1969-1970—William FE Mahler was appointed three-quarters time assistant professor in the Biology Department at SMU and one-quarter time with the herbarium. Shinners’ Herbarium Report involved mostly Mahler's activities; Sida activities were still under the control of Shinners. Once again John offered Lloyd help in another letter probably dated sometime in April or May 1970 (see also Mahler's tribute). “Went ona collecting foray this past weekend—the first extensive one since I got back from Mexico. The ankle still gives trouble... Indeed, by the time the collecting trip was over, I was limping some- thing awful (). You were willing, I recall, to have me take care of the index for volume 4 of SIDA. Right???... If you are still, | shall get onto the task, a bit ata time. Shaw's paper alone is a challenge.” 1971—Lloyd Shinners published Sida 4(2) on January 15, 1971 and one month later Shinners passed away on February 16, 1971. William F. Mahler inherited Sida from Lloyd’s sister Helen Koresh and he immediately became editor and publisher. February 17,1971, the very day after Lloyd’s death, John Thieret com- posed and mailed a letter to Mahler at SMU offering personal assistance and moral support. pele | “Just a note to reaffirm my offer to do what | can do to help keep SIDA a going thing. The journal a memorial to its founder. should certainly not be allowed to die but should be kept viable as, perhaps hel J; LF s for specific things could do, Icertainly would be glad ting. | would be willing to come over there every once in a while to work closely with you or with anyone else interested. Editing, reading proof, indexing—all would suit me fine. Speaking of indexing, Lloyd already gave me the go-ahead on doing the index for volume 4. Certainly you are thinking of an article on Lloyd, in SIDA, with possibly a list of his publica- tions. I'd be glad to help here, especially on the bibliographic work involved. LIPSCOMB, JOHN W. THIERET, ASSOCIATE EDITOR 59 eats yi offers b have ee made, It is now up to you to see that they are taken u ne to urge in to do all _ can to ee SIDA going?’ John Thieret was concerned about the future of Sida. Here is his letter to SMU urging Southern Methodist University’s President Dr. Willis M. Tate and the Dean of Graduate School, Dr. CC. Albritton, Jr, to continue the publication of Sida. “The name of the president I got from the 1969-70 edition of the WORLD OF LEARNING. Iassume that ie is ae the man. The Albritton part is for certain OK 1 ] iat botan I sht (or something like that) and that I strongly urge SMU to aor all te can to continue the Seat of SIDA, a journal that is an excellent one that has filled a need. Etc. Etc. Ete. Signed John T. Even with all the encouragement from a number of Lloyd's colleagues, South- ern Methodist University never committed to preserving the journal or the col- lections that really made a name for the university in the botanical sciences. Other scientific journals were also born there but either died there or moved on to more supportive organizations. The dedication and hard work that Lloyd Shinners devoted in championing the cause of botany at SMU was kept alive through colleagues and the Botanical Research Institute of Texas (BRIT). Will- iam Mahler committed 21 years of personal time and money in keeping Sida, Contributions to Botany successful. In 1987, Mahler along with colleagues Lip- scomb and Andrea McFadden were instrumental in the establishment of the free-standing research institution BRIT. The core of the BRIT collections is the Lloyd H. Shinners Collection in Systematic Botany, originally at Southern Meth- odist University. Since Sida was privately published by Mahler it too joined BRIT. Mahler officially transferred ownership of Sida to BRIT in 1993. Mahler (1971) published Sida 4(3) with a tribute to Lloyd and John had one paper in the same issue: Physalis lagascae (Solanaceae) in Louisiana: new to the conterminous United States. Sida ADT F John’s offer of help was at last accepted. Mahler officially asked him to serve as associate editor of Sida in late 1971 or early 1972. One of the first papers John edited as associate editor was a large manuscript on the genus Rorippa sent to Dr. Shinners (early 1970) by Dr. Ronald L. Stuckey, associate professor of botany at Ohio State University. Dr. Stuckey expressed a desire to help with Sida ina letter dated February 24, 1971 to Dr. Mahler and Dr. Thieret. It has come to my attention that Professor Lloyd Shinners has passed away. I talked with John Thieret er]. My interest is that over a year ago I 2 by phone soy and he suggested that I write to you[Ma sent my d rtation, a sae aS peers on ties eeu ion apa to Dr. Shiners for publi- cation in Sida, which he ag nadditio enses for publi- cation, and this d to Sida i f $250.00 each, one in eee 1969 and one in December 1970. aie I would hope that you and Dr. Thieret, or anyone else, could see to it that the present manuscript on hand and/or in press for Sida could be published, and certainly as a memo- 60 BRIT.ORG/SIDA 22(1) rial to Dr Shinners. | would be willing to do most anything, at this distance, to help tl se al | suspect financing is the big problem, and | would be willing to contribute some more funds, if | knew for sure my eee Male be published in Sida. Furthermore, I suspect some editing or deletions of my | dhe ge, elp. It was my understanding that the specimen citations would =~ not be pu plished. t also say that Mr. Robert Haynes, our assistant curator of the Herbarium, and former student of Dr. Thieret, has his manuscript on the genus Conopholis being published in Sida. [ would also like to see this fine journal for taxonomic botany and related interests be contin- ued either by Southern Methodist University or by a federation of the south-central botanists, for example. Mahler's letter (3 Mar 1971) back to Dr. Stuckey and again Dr. Thieret is involved: “In regard to your letter of 24 February 1971, we are planning to continue ea and follow Dr. Shinner’s plants for Vol. 4 (3) and (4). Issues 3 and 4 will be the paper lescribed in his letter of 26 March 1970. You are being credited with a $500 contribution towards the publication of your MS. At this time, | cannot say what the financia — situation will be in relation to SIDA and its $12 page costs (re Shinners’ letter, 9 December 1969), but hopefully we can continue as leas by Dr. Shinners. My continuing to work like Dr. Shinners did would be impossible with the other activities (du- ties). He did not get a chance to go over the MS. in detail and I would like to send it toa reviewer such as Dr. Thieret. Perhaps you would like to make the changes Dr. Shinners suggested first, and then send it to Dr. Thieret. We have plenty of time in this respect as | do not expect No. 3 to come our till summer (71) at this time. [appreciate the briefing (previous correspondence) as it has been very helpful. | wish that ev- eryone would do it this way. Perhaps they will as I have one other individual who has brought me up to date in the same manner. The Conopholis paper of Haynes is still in the mill and status at this time unknown.” Dr. Stuckey’s response: Lenclose Xerox ¢ sof two letters which show that Dr. Reed C. Rollins has read and reviewed my Rorippa manuscript. | am eae that you have selected Dr. Thieret to go over my manuscript in ' g preparation for its publication. It would be best if you transmitted the manuscript directly to him, rather than sending it back to me. The revision that Dr. Shinners mentioned involves a small item of about one page. | can do this and send the revision to Dr. Thieret. As you may know Dr. Thieret will be at our University for 5 weeks this summer teaching at Stone Laboratory. | will be there at the same time and can wor < with him on the manuscript at that time.” Mahler's letter (23 Mar 1971) to Dr. Stuckey and Dr. Thieret is mentioned: found your MS. yesterday (sti [have not located the plates, but will in time) and received your letter today. Dr. Shinners had discussed this MS. with me and stated that it would have to be trimmed to ca. the length of Kral’s paper 9ca. 175 pp.). As suggested by Shinners to you, ca. 150 pp. can be eliminatec readily (the citation of specimens). That still leaves ca. 100 pp. (70 pp. of MS. + 30 maps, illus.) to be eliminated. Thisi is one of the items of “reviewing” that | was referring to in regard to John Thieret. If u wish to do this yourself and possibly you should as it’s your MS., John Thieret might be helpful in short cuts so that you will not leave out any essential ena) The discussions, etc., are rather ver- bose and the same ideas could be expressed in fewer words.’ a 1972—Thieret’s letter January 11, 1972) to Dr. Stuckey announcing his associ- ate editorship: LIPSCOMB, JOHN W. THIERET, ASSOCIATE EDITOR 61 SIDA, founded by Lloyd H. Shinners, is privately published by Wm. F. TAXONOMY AND DISTRIBUTION OF THE Mahler, SMU Herbarium, Dallas, Texas, 75222, U.S.A. Subscription price $8 GE es RORIPPA (CRUCIFERAE) (U.S.) per volume NORTH AMERICA RONAL DL. Such EY Columbus, Ohio 43210 Rorippa consists of those yellow-petaled, numerous-seeded, readily- dehiscent-fruited Ley and shore “plans in the Cruciferae Distributed Berea in temperate the genus ccurs every continent except Antarctica, but the Species are more iber may increase to 60 or 70 an expanded knowledge Tonal “(a968) jeanatderedtt here to be about 70 species, some imperfectly known. One ea eeue Sees, R. palustris, is known fom eac me continent where t . Busch (1915) a Eurasia; s Warncinaencs (1960) ha eight species for Middle Europe; and, in Flora Europaea, yea “uss included 10 specie: ASSOCIATE EDITOR and So America. I 7 cal ones) as native to North America; seven species are con: nsidered as John W. Thieret itoce The evolutionary relationship of these foreign members to the orth America, Rorippa is most common and abundant, both as to inva and t F aiene Arte irs of lakes te ce banks rea and rivers, and in marshes bee swales; or in wet inca Pinte sites ong roadside, drainage, and irrigation ditches in ni fallow or cultivated fields and gardens, along railroad grades and ditche: HISTORY OF THE GENUS rippa begins with Scopoli (Fi Carn, ed. 1, 520. 1760), who wrote the first, but brief, eneric diagnosi three species were named. The first contains the exact phraseology as that provided by Lin- naeus (Sp. Pl. 675. 1753) for attire sylvestre. Scopoli cited ee as one of five sources; on this basis, Rorippa sylvestris (Linnaeus) Besser s taken as the type of the genus. Abrams (1944) listed eae am- pat il fulfillment of the of Michipina Aaa SIDA Contributions to Botany, Volume 4 Number 4, pages 279—430 Cop. >) aa The Herbarium ight 1972 by wm, F. Mahler Left: Copyright page of Sida 4(4), the first issue of Sida in which John Thieret was appointed associated editor. Right: ope £D lac. I J. Davt co Look e.g rr “Thanks for l our Rorippa ms. Mahler is really the one to talk to about any financial matters. When I finish with the ms., | shall return it to Mahler. From then on, any dealings will be conducted by him. I have just been asked to serve as Associate Editor of Sida— which is just about what I've been doing all along.” John Thieret came to the editorial rescue as associate editor of Sida in 1972 and for the next 34 years, John proudly and faithfully served the journal in so many different ways. Mahler along with John’s superb editorial knowledge and skills and the help of many other colleagues, kept Sida going strong. Mahler passed the editorial baton to Lipscomb in 1983 but remained publisher until 1993; John was associate editor from 1972-2005. 1972-2005—Not only was John an active associate editor of Sida, he was also contributing manuscripts. Twenty-seven of John’s papers were published in Sida during his 34 years of associate editorship. This represents almost 25% of all the papers John published in that same time frame. In fact, of all the scientific papers listed in Ralph Thompson’s tribute (2006), 26% were published in Sida, more than three times that of the next closest journal. The amount of work he contributed is enormous and the success, the foundation that John Thieret (and Wm. F. Mahler) made to Sida cannot be underestimated. We all have been for- 62 BRIT.ORG/SIDA 22(1) tunate to have had John’s involvement with the journal but my (Lipscomb) ca- reer was truly impacted by this good friend. John’s influence on my (Lipscomb) personal career In the fall of 1975 my botanical career began as herbarium botanist at Southern Methodist University. There I worked in the great herbarium and botanical li- brary built by the prolific researcher, writer and founder of Sida, Lloyd Shinners. My first two years of work at SMU were strictly herbarium related and | casu- ally observed the editorial work on Sida by Wm. F Mahler. In the beginning I did not know who John Thieret was but it didn’t take me long to find out. Tra- dition is wonderful. Shinners told Mahler he needed to meet John and Mahler told me I needed to meet John. In 1977, two years into my job at SMU, I was appointed assistant editor of Sida by my mentor, friend, and colleague Bill Mahler. It was around this time—pre-email days—that | first began to commu- nicate with “Big John” Thieret over the phone and through letters. Over time | began to interact with him more and more on Sida’s issues and editorial mat- ters. It would be years later before | actually met the man behind the red pen. | had no idea John was the master of editing. | was too young to envision the immediate and future impact this one man would have on my life. He genu- inely cared for me as an individual and shared his expertise and knowledge with me freely. Through John, a whole new world of wisdom and knowledge was available at my doorstep. John Thieret took me in almost as if | was one of his own children. He (and Mahler) helped me to learn the role of an editor one red pen at a time. A few years passed before | finally got to meet John in person. He was ev- erything that I envisioned. | enjoyed one full week with John and his lovely wife Mildred back in the late 1990s. | stayed in their home in Alexandria not far from Northern Kentucky University (Highland Heights) where John worked. It was in their home that I had my first meal of tef (Eragrostis tef (Zucc.) Trot- ter). It wasalso at the Thieret home that John taught me the word “kex” or “kexes” for a dead stalk or stalks of herbaceous plants, “especially of large umbellifer- ous plants” (Oxford English Dictionary). | enjoyed four wonderful days with John traveling and botanizing through northern and eastern Kentucky. One of the many botanical things I learned from John on that trip was the edible fruits of Rhus typhina (Staghorn Sumac). He told me the drink from the sweet fruits was called “rhusade.” After returning to the Cincinnati area from out botanical field trip, John and I drove to one of his fun places to botanize called Jungle Jim’s International Farmers Market in Fairfield (Cincinnati), Ohio. One quote | found on the web said, “It is the ultimate food shopping adventure.” We walked into this humongous store and headed straight to the produce section and did we ever have a great time identifying plants and plant parts. It was an awesome experience with John, the economic botanist. And we visited another of John’s mo LIPSCOMB, JOHN W. THIERET, ASSOCIATE EDITOR 63 favorite plant places, Spring Grove Cemetery in Cincinnati. What an incredible place to study and learn about the many different woody plants and what a wonderful teacher I had. For 33 years John Thieret served Sida, Contributions to Botany as associate editor. His crusading editorial and scientific contributions to Sida are beyond measure! He was a good friend and advocate of Lloyd Shinners and Sida; he was a solid anchor of support to Wm. F Mahler in the storm of uncertainty that followed Lloyd's death; and he was a radiant lighthouse of editorial help to me as a young editor. He was the guiding light of editorial excellence and author- ity. Sida is today in large part because of John Thieret. His 33 years of editorship with Sida stands longer than anyone else associated with the journal. John Thieret was my rock and my anchor. His editorial energy and enthu- siasm and loyalty to Sida will forever be remembered and deeply appreciated! Did I ever say thank you John? I’m sure I did many times but let me say it in print in Sida one last time for everyone to see, Thank you John for all that you've done and continue to do through your teachings. Postscript: Changes in the journal, Sida no more I'm glad these tributes to John Thieret could be published in the journal that he loved so very much. Soon the journal Sida, Contributions to Botany will no longer be published by that name but instead will be called the Journal of Botanical Research Institute of Texas. The new journal will have a new look but generally the same format, botanical focus, and editorial direction will remain. The lead- ership of the Botanical Research Institute of Texas has decided the name “Sida” is offensive and has too much of a negative connotation because of its similar- ity to the word meaning AIDS in Spanish, French, Italian, and Portuguese. Per- haps the name change is a good thing but I am personally saddened to see Shinners’ original and long-standing name displaced. ACKNOWLEDGMENTS Ithank Gary Jennings (library /archive help), Ruth Ginsburg (archive help and reading the ms), and Ralph Thompson and Guy Nesom for helpful comments. REFERENCES BoTANICAL RESEARCH INSTITUTE OF TEXAS ArcHives. 2006. Lloyd Herbert Shinners archive. Botanical Research Institute of Texas library, Fort Worth. Ginspurc, R. 2002. Lloyd Herbert Shinners: by himself. Sida, Bot. Misc. 22 Matter, W.F. 1971.Lloyd Herbert Shinners 1918-1971. Sida 4:228-231. SHINNERS, L.H. 1964. Scutellaria thieretii (Labiatae), a new species from coastal Louisiana. Sida 1(4):251-252. THompson, R.L. 2006. John W. Thieret (1926-2005). Sida 21:3-19. A NEW NARROWLY ENDEMIC SPECIES OF CLEMATIS (RANUNCULACEAE: SUBGENUS VIORNA) FROM NORTHEASTERN TEXAS Dwayne Estes abe of Tennessee Heal ium CERN Department of Ecology and Evolutionary Biology vines of Tennessee Knoxville Tennessee 37996, U.S.A. tnplants@yahoo.com ABSTRACT Clematis carrizoensis, a new ese! endemic species of subgenus Viorna, is described from the Carrizo Sands of northeastern Texas. This species belongs to a complex of similar species character- ized by glaucous and glabrous oe leaf surfaces and stems. Four additional species comprise this com ee C. addisonii, C glaucophylla C. texensis. and o versicolor denen carriZoensis 1s iS LOED NO: logi ically LLLUSL similar | to bluist it ] C. versicolor. It differs from those two coe in having ei cream ee lightly lilac- need se- pals. From C. addisoniiand C. glaucophylla, C. carrizoensis differs in sepal color and in havi = thicker and more reticulate peaves: All ns species of the complex are allopatric and occupy somewhat dif- ferent habitat Further discussion of their See eee differences, Poreene a@iabaren ane habitat preferences is provided. RESUMEN Se describe Cl ti Oe endo a de Carrizo Sands L oO pl ] en el noreste de Texas. Esta especie pert tener los tallos y el envés glaucos y glbros FI | complejo tae tras Caddicantt C glaucophylla, C. texensis, y C. versicolor. rizoensis es la mas fe ee a . Lees ep los escarlata ie C. ve mei sépalos que van del rojo purpura al lavanda azulado. ener los sépalos amarillo crema ligeramente tenidos de lila. De C. addisonii y C. aaeoonsi ite. se eee C. carrizoensis por el color de los sépalos y por tener las jas mas ee y mas Saaeconee . as cinco cece coBpSe son ope acaS y FOSpan morfoldgicas, Giseibucion peop y preferencias de habitat. In North America, 18 species of Clematis (Ranunculaceae) belong to the subge- nus Viorna (Pringle 1997). Of these, eight currently recognized species were included in the formerly recognized Clematis section Viorna subsection Viornae (Erickson 1943; Dennis 1976; Kral 1987): C. addisonii Britton, C. glaucophylla Small, C. morefieldii Kral, C. pitcheri Torr. & A. Gray, C. reticulata Walter, C. texensis Buckley, C. versicolor Small, and C. viorna L. These species are peren- nial vines characterized by relatively large (1-3 cm) urceolate flowers with thick leathery sepals that are mostly arranged in solitary or few-flowered cymes borne SIDA 22(1): 65 — 77. 2006 66 BRIT.ORG/SIDA 22(1) on axillary peduncles subtended by leafy bracts (Dennis 1976). These taxa can be delineated into two groups based on leaf vestiture (Dennis 1976). The first group including C. morefieldii, C. pitcheri, C. reticulata, and C. viorna all have variously pubescent abaxial leaf surfaces and stems. The species of the second group, characterized by glabrous and glaucous abaxial leaf surfaces and stems, include C. addisonii, C. glaucophylla, C. texensis, and C. versicolor. These species form a complex here referred to as the C. glaucophylla complex. The species of the C. glaucophylla complex can be distinguished from each other by relatively few qualitative morphological characteristics. Often, a com- bination of features, including morphology, habitat, and distribution, must be used to identify taxa. Several structures including the caudex, roots, inflores- cences, and achenes offer little or no taxonomic value in separating most spe- cies of the complex (Erickson 1943; Dennis 1976). The most taxonomically use- ful traits are leaf texture, degree of leaf vein reticulation, and sepal color, although even these characters exhibit considerable variation and intergrada- tion (Erickson 1943: Dennis 1976). For instance, leaf thickness is useful when extremes are exhibited, but Dennis (1976) noted that factors such as leaf age and environmental conditions often make it a difficult character to assess. Sim1- larly, degree of vein reticulation is a useful character when found in its extremes but it can be confounding when trying to evaluate the continuum between ex- tremes (Dennis 1976). Erickson (1943) concurred, noting that while reticulate venation is a valuable taxonomic character, “it is not often possible to distin- guish between degrees of reticulation consistently.” Dennis (1976) considerec sepal color to be important for distinguishing species of the complex; for ex- ample thin-leaved specimens of C. versicolor can only be distinguished from C. glaucophylla by sepal color. Other characteristics, including degree of leaf dis- section, leaflet shape, flower shape, and stamen vestiture, are only generally useful for distinguishing species and are apparently never diagnostic. While the species of the C.glaucophylla complex may be difficult to differentiate mor- phologically, all members of the complex exhibit specific physiographic pref- erences (Dennis 1976) and have non-overlapping ranges (Fig. 1). Clematis texensisand C. versicolor are completely allopatric with the former restricted to the Edward's Plateau of Texas and the latter confined to the Inte- rior Highlands (Arkansas, Missouri, Oklahoma) and Interior Low Plateaus (Ala- yama, Kentucky, Tennessee). They differ consistently in flower color and gen- erally in leaf dissection and leaflet shape. Both species usually have relatively thick and conspicuously reticulate leaves. Erickson (1943) called attention to possible intermediate populations occurring in the region between the Interior Highlands and the Edward's Plateau noting that some specimens from this re- gion have been difficult to assign to a particular species. Erickson arbitrarily referred questionable specimens from southwestern Arkansas and southeast- ern Oklahoma to C. versicolor and specimens from northeastern Texas (Smith — en ees ESTES, A NEW SPECIES OF CLEMATIS FROM TEXAS 67 Legend Clematis spp. @ = §=C. addisonii @ = $C. carrizoensis = ~=6C. glaucophylla C. texensis 4 ~~ C. versicolor Co TkKilometers Fic. 1.G hi f the fi i ftha Cf tie al hull | fr aera ) i 1976) County) to C. texensis. Dennis (1976) contended that the Texas specimens were not C. texensis because the sepals were white to pale lavender rather than the bright red characteristic of that species. Interestingly, Dennis (1976) referred the Texas plants to C. versicolor in spite of the fact that they also differed from that species in sepal color and habitat and occurred ina different physiographic region ca. 200 km from the nearest known C. versicolor population. Additiona study of herbarium specimens from this region coupled with field work has revealed that the Texas populations previously referred to C. versicoloractually represent a new species described below. Clematis carrizoensis D. Estes, sp. nov. (Figs. 2, 3, 4B). Type: U.S.A. TEXAS: Smith Co. 1.5 mi NW of Lindale, sandy soil, vine growing along fence near roadside, 15 May 1949, ].F Hennen 408, consists of 2 sheets (HOLOTYPE: SMU). jen A Clematide versicolore sepalis e lilacinis flavidis-cremeis, foliis plerumque dissectioribus, foliolis saepiuscule emarginatis ad apices profundius cordatis basibus differt. Perennial trailing or scandent vines to 3 m long, from a woody caudex bearing numerous smooth fleshy apparently unbranched roots. Stems stout proximally, slightly flexuous distally, 6 angled, 12 ribbed, glabrous and glaucous, reddish- brown to pale brown. Leaves oppositely decussate, imparipinnate to pinnate- 68 BRIT.ORG/SIDA 22(1) THE UNIVERSITY OF TENNESSEE 1 Small W. Michael Dennis December , 476 PLANTS OF TEXAS Clematis reo ulata Valte | HOLOTYPE OF: | Smith Co.1 1.5 miles nortivest of Lindale. Sanly soil. Vine growing along; fence near roadside. | Ctesniit ; sis D. Estes Sheet 1 of 2 J. Ps Hennen 1,08 May 159 1oL9 Fic. 2. Holotype (UJ. Hennen 408, SMU) of Clematis carrizoensis. ESTES, A NEW SPECIES OF CLEMATIS FROM TEXAS 69 Fic.3. Clemati ji is. A. Portion of plant showing g| baxial leaf surfaces. B. Close-up of abaxial surface of | ins). D. Fruit. Photos above leaflet showing raised-reticulate venation. C. Flower, fi iew ( isy I f | H] 1 i D. Estes 07032 P ternate, largest at about mid-stem, spreading, to 2.0 dm long, longer than wide to as wide as long; petiole stout, equaling or more frequently shorter than the lowest leaflets, to 6.0 cm long, glabrous and glaucous, base of opposing petioles usually connate; rachis geniculate; leaflets symmetrical or asymmetrical, paired, 8-10 on imparipinnate leaves, to 25 on proximal-most pinnate-ternate leaves, spreading to erect, reduced distally on rachis, the distal-most often form- 70 BRIT.ORG/SIDA 22(1) ing a short simple tendril, proximal-most entire or 2-3-lobed to trifoliate, ovate, broadly ovate, or subrotund, leaflets to 7.0 cm long and to 6.0 cm wide, coria- ceous, base shallowly to deeply cordate, rounded, or truncate, margins slightly revolute, leaflet apices and lobe apices rounded, retuse, or occasionally acute, mucronate, adaxial and abaxial surlaces strongly reticulate with all vein or- ders conspicuously raised, forming an intricate reticulum, light green adaxially, paler abaxially, drying pale olive-brown to olive-green, glabrous and glaucous on both surfaces but more conspicuous on abaxial surface; petiolules 1.5-3.0(-7 cm) long, one-fourth to one-third length of leaflet or rarely one-half to as long as leaflet, stout, glabrous and glaucous. Flowers solitary on long axillary pe- duncles or rarely axillary and terminal, nodding, ovoid or urceolate; peduncles erect, curved or frequently abruptly recurved at apex, 7.5-17.0 cm long, stout, glabrous and glaucous, with 2 foliose bracts; bracts sessile or subsessile with petioles to 4.0 mm, positioned mostly near the middle of the peduncle, base subcordate to deeply cordate, margins entire or 2-3-lobed, otherwise similar in shape, size, texture, and vestiture to leaflets. Sepals erect, connivent, ovate-lan- ceolate to oblong-ovate, apices reflexed, 1.8-3.0 cm long, 0.7-1.3 cm wide, coria- ceous, ca. 12 nerved, abaxially yellowish-cream, or greenish-yellow and often faintly tinged with lilac proximally, adaxial surface similar in color to the abaxial surface at least distally, mostly drying tan or light brown, edges nar- rowly crispate, white-tomentulose, abaxial and adaxial surfaces glabrous. Sta- mens linear, 1.8-2.4 cm long, the filaments flattened, in dried material dark brownish-red in the proximal half, densely pilose from below middle to apex, anthers 6.0 mm long, connective extended 1.75-2.0 mm beyond anthers, pilose. Fruit tightly clustered in spherical heads 4.5-8 cm in diameter, light brown, body rhomboidal-ovate, 6.0-9.0 mm long and 6.0-7.0 mm wide, acuminate at both ends, compressed, marginally thickened, sericeous; style curvate, 3.0-5.5 cm long, with a yellowish-brown plumose coma, hairs of coma spreading to scending, to 3.0 mm long. Chromosome number unknown. aoa specimens ene TEXAS: Cherokee Co.: ca. 9.5 mi E of Troup, ca.6 mi W ol Price, ca. ni N of Hwy 13, sandylands, 12 Jul 1977, G. Bee 50. 38 (BRIT). Smith Co.: sandy woods east ol wee 16 May 1902, J. Reverchon ee [two sheets] (MO); on R.R. [railroad] near Lindale, Apr 1923, J. Reverchon s.n. (MO); western Tyler along Cotton Belt Railway, east end of Pine Burr Rd., infrequent at edge of woods along railway, 21 May 1949, V.L. Cory 56212 (SMU); western Tyler, S of Pine Burr Rd, infrequent in sandy woods, 10 Aug 1950, V.L. Cory 57860 (SMU); just N of Interstate 20 in woods E ol Old Tyler-Mineola Rd., 20 Oct 1965, D.S. Correll & H.B. Correll 32041 (LL); sandy field with scattered post oak in SW angle where US Rte. 69 and Loop 323 intersect, north edge of Tyler, 28 Aug 1966, D.S. Correll 33513 (LL); pure sand in field on Loop 323 near intersection with Rte. 69, Tyler, 6 Jun 19609, D.S. Correll 37401 (LL); in SW angle of intersection of US 69 and Loop 323, N edge of Tyler, under oaks next to 12 Oaks Motor Hotel, growing in sandy field under Quercus incana, Q. margaretta, with Opun- tia, Yucca, 22 Jun 1975, WM. Dennis 5I718 (TENN); NW portion of city of Tyler, 0.75 km WSW of junction ol TX State Hwy 323 Loop and US Hwy 69, ca. 20 m SW of junction of Hwy 323 Loop and Silver Creek Dr, 32°22'42.1"N, 95°2020.6"W, along N edge of scrubby oak-hickory woods and in a small opening nearby, rooted in deep sand and scram acorer small shrubs and herbs, with Quercus ESTES, A NEW SPECIES OF CLEMATIS FROM TEXAS 71 \“}. spp., Carya, Rhus aromatica, Prunus gracilis, and Toxicodendron pubescens, 9 May 2005, D. Estes 07032 (GH, NY, TENN, VDB). Van Zandt Co.: 2.3 mi E of Ben Wheeler, sandy loam, 6 May 1951, R. Van Vleet 1257 (SMU). Distribution and Habitat.—Clematis carrizoensis is endemic or nearly endemic to the Carrizo Formation, a narrow zone (19 km wide) of relatively deep depos- its of coarse to find sand that trends southwest to northeast for ca.720 km from south-central to northeastern Texas (Macbryde 1933; Sorrie and Weakley 2001). Sorrie and Weakly (2001) identified an additional 12 taxa endemic or nearly endemic to this formation. Within the Carrizo Sands, C. carrizoensis has only been collected from three contiguous counties, Cherokee, Smith, and Van Zandt, all in the northeastern section of the formation. Clematis carrizoensis grows in or along the edges of open, well-drained, prairie-like areas in full sun or partial shade. Within these communities, C. carrizoensis trails along the ground or Over low herbs, shrubs, or fences, apparently rarely climbing higher into low branches of trees. Oak-hickory woodlands or thickets border these sites. Domi- nant tree species associated with one population of C. carrizoensis in Tyler, Texas (site of D. Estes07032 cited above) include Quercus incana Bartr., Q. marilandica Muench., 9. nigra L., Q. stellata Wang., Carya texana BuckL., Sassafras albidum (Nutt.) Nees, and Ulmus crassifolia Nutt. Frequent small trees, shrubs and woody vines were Opuntia sp., Prunus caroliniana (P. Mill.) Ait. P gracilis Engelm. & A. Gray, Rhus aromatica Ait. R.copallinum L., R.glabraL., Rubus trivialis Michx., Stillingia sylvatica Garden ex L., Toxicodendron pubescens P. Mill., Vaccinium arboreum Marsh., Viburnum rufidulum Raf., Vitis mustangensis Buckl., V. rotundifolia Michx.and Yucca sp. Baptisia sp., Cnidoscolus texanus(MuellL.-Arg.) Small, Croton argyranthemus Michx., Mimosa sp., Paronychia drummondii Torr. & A. Gray, Pteridium aquilinum (L.) Kuhn, Rumex hastatulus Baldw.,, — 72 BRIT.ORG/SIDA 22(1) Schizachyrium scoparium (Michx.) Nash and Talinum rugospermum Holzinger were closely associated herbaceous taxa. Phenology.—Clematis carrizoensis flowers from early May to early August: fruits are produced beginning in late June and persist until November. Etymology.—The specific epithet “carrizoensis” was selected in honor of the Carrizo Sands, the geologic formation to which C. carrizoensis is apparently restricted. Common Names.—Carrizo sands leather-flower; sand clematis Conservation Significance.—Clematis carrizoensis is a rare species. | have seen only 12 specimens from three counties in northeastern Texas; 10 of these are from just four sites in a small area of Smith County. Label data from her- barium specimens also indicate that this species is uncommon where it occurs. Most of the specimens examined for this study were collected over 30 years ago possibly indicating that this species is declining or simply overlooked. Most of the populations represented by collections are located near or in the city of Tyler, Texas. These populations may be subject to extirpation by future development. This species should be searched for in other sandy sites, particularly those as- sociated with the Carrizo Formation, in northeastern Texas as well as in sandy habitats in nearby portions of Miller County, Arkansas and Caddo Parish, Loui- endangered and should be afforded KL siana. This species should be considerec protection at both the state and federal level. Species of the Clematis glaucophylla Complex Five species comprise the C. glaucophylla complex. While all species differ in rather subtle morphological characteristics, they are all allopatric and mostly inhabit different plant communities in different physiographic provinces. The complex is in need of a detailed phylogenetic study. Clematis addisonii, a narrow endemic of a four county area in the Ridge and Valley of western Virginia (Fig. LD), inhabits dry calcareous woods and out- crops (Kral 1983). Itis characterized by a mostly erect, non-twining habit, mostly simple, thin, and inconspicuously reticulate cauline leaves, and solitary termi- nal {lowers that have purple to reddish-purple sepals with whitish apices. Clematis glaucophylla ranges {rom northwestern Florida west to southeast- ern Mississippi and north to southeastern Tennessee (Fig. L); it is reportedly dis- junct to McCurtain County, Oklahoma (Dennis 1976). Reports of C.glaucophylla from Kentucky Jones 2005) are based on misidentified specimens (D. Estes, pers. obs.). Throughout most of its range in the Piedmont and Gulf Coastal Plain, C. glaucophylla grows in sandy neutral to slightly acidic soils along streams (Dennis 1976), but at some sites in Alabama and southeastern Tennessee, C. glaucophylla grows at the edges of dry rocky calcareous forests (D. Estes, pers. obs.). Clematis glaucophylla hasa oe habit and compound cauline leaves that have mostly thin and inconspicuously reticulate leaflets. The proximal leat- ESTES, A NEW SPECIES OF CLEMATIS FROM TEXAS 73 lets are often deeply lobed, divided, or ternate. Flowers of C. glaucophylla are borne on axillary peduncles and have sepals that are cherry-red or reddish- purple abaxially and light colored near the abaxial tip and on the whole sur- face adaxially. Clematis texensis is endemic to the Edward’s Plateau of Texas (Fig. 1) where itis often associated with calcareous woodlands, edges, cliffs, and stream banks. The species hasa viny habit and compound cauline leaves with relatively thick and conspicuously reticulate leaflets. The proximal leaflets vary from simple to lobed or ternate. The flowers are borne on axillary peduncles and the sepals are scarlet or rose-red abaxially and near the tip adaxially (Fig. 4). Clematis versicolor has a bicentric distribution with part of its range cen- tered in the Interior Highlands of southwestern Missouri, western Arkansas, and eastern Oklahoma and the other portion centered in the Interior Low Pla- teau of central Tennessee, extreme northern Alabama, and south-central Ken- tucky (Fig. 1). It is usually associated with calcareous bluffs, dry woods, and roadsides with exposed limestone. Clematis versicolor has a viny habit, relatively thick and conspicuously reticulate compound cauline leaves with the proxi- mal leaflets entire or rarely lobed or divided, and flowers borne on axillary peduncles. The abaxial surfaces of the sepals of C. versicolor are mostly pur- plish-red proximally and often are cream-colored or greenish distally (Fig. 4). Clematis carrizoensis is restricted to sandy prairie-like areas among oak- hickory woodlands in three counties in northeastern Texas (Fig. 1). The species hasa viny habit, thick and conspicuously reticulate compound leaves that have proximal leaflets frequently lobed, divided, or ternate, flowers borne on axil- lary peduncles, and sepals that are al lly yellowish-cream and faintly tinged with lilac proximally (Fig. 4). Characters Useful for Distinguishing Members of the Clematis glaucophylla Complex The members of the C. glaucophylla complex can be distinguished by differ- ences in habit, leaf dissection, leaf texture, leaflet shape, degree of vein reticu- lation, flower position, sepal color, habitat, and geography (Table 1; Fig. 1). In terms of habit, C. addisonii is the only erect non-viny member of the complex. It has stems that are mostly less than | m tall whereas the other spe- cies in the complex have viny stems mostly greater than 1 m in length. Dennis (1976) noted that while C. addisonii is the only species in the complex that ex- hibits this growth form, occasional individuals of C. addisonii bear apical branches that often become entangled and approach a viny habit. Leaf dissection is a character that is only generally useful for distinguish- ing species of the complex, the exception being C. addisonii. Clematis addisonii has leaves that are mostly simple but occasionally vary to pinnate with up to six leaflets. The rest of the species of the complex have leaves that are pinnate 74 BRIT.ORG/SIDA 22(1) Taste 1.Characteristics of the Clematis glaucophylla complex, compiled from data pr ted in Dennis (1976) and Pringle (1997). texensis carizoensis versicolor addisonii glaucophylla iny>1m viny,> 1m viny,> 1m erect,< 1m viny,> 1m Leaf Dissection |-pinnate to ]-pinnate to 1-pinnate, simple, or distal 1-pinnate to pinnate pinnate- proximal leaves 1-pin- —_ pinnate- ternate, ternate, leaflets nate ternate, proximal leaf- — proximal leaf- ee proximal lets commonly lets commonly unlobec leaflets lobed lobed dk commonly or 3-foliate or 3-foliate 2-3 lobed lobed or 3-foliate Leaf Texture coriaceous coriaceous coriaceous thin thin Venation reticulate, reticulate, reticulate, not reticulate, — not reticulate, raised raised raised not raised not raised Flower Position mostly axillary mostly axillary mostly axillary mostly terminal mostly axillary Flower Color scarlet to yellowish pure redto purple to cherry red to e-red cream, ee os nder, purplish-red, — reddish- lilac-tinged creamy OF whitish distally purple proximally greenish distally to pinnate-ternate with more than six leaflets. The degree of compounding of the proximal leaflets is a useful yet non-diagnostic character that according to Erickson (1943) “must be used with judgment.” For example, Erickson (1943) noted that leaf dissection may generally be used to separate C. texensis and C. versicolor. Clematis glaucophylla, C. texensis,and C. carrizoensis frequently have lobed, divided, or ternate proximal leaflets whereas the proximal leaflets of C. versicolor are usually unlobed or rarely lobed or ternate. Erickson (1943) considered leaflet shape to be of general use in delimiting taxa, especially the leaflet apices. Generally, C. addisonii, C. glaucophylla, and C. versicolor have broadly to narrowly obtuse leaflet apices. Clematis texensis and C. carrizoensis have leaflet apices that are more frequently emarginatec apically but that vary to obtuse. Leaflets of C. carrizoensis, C. texensis, and C. versicolor are almost always mucronate at the apex. Among the species of the C. glaucophylla complex, two groups can be de- lineated based on leaf texture and degree of leaf vein reticulation. Clematis glaucophyllaand C. addisonii belong toa group characterized by relatively thin and non-reticulate leaves. The other group, consisting of C. carrizoensis, C. Wu ~ texensis and C. versicolor, has relatively thick and conspicuously reticulate leaves. Once again, these characters are relative and are hard to differentiate when only one species is encountered and can sometimes be difficult to assess ESTES, A NEW SPECIES OF CLEMATIS FROM TEXAS 75 from herbarium material. Furthermore, shade plants of all species of the com- plex may be thin and non-reticulate. Flower position is a character that is of limited taxonomic value in the C. glaucophylla complex. Most species have flowers borne on axillary peduncles, however, C. addisonii has flowers that are mostly borne on terminal peduncles. Perhaps one of the most useful characters for distinguishing the members of the C. glaucophylla complex is sepal color (Table 1, Fig. +). Unfortunately, it is also the character that is most difficult to preserve and interpret on herbarium specimens. In C. glaucophylla, the sepals are more frequently cherry red to red- dish-purple abaxially except distally they are often light colored. Clematis texensis has sepals that are generally scarlet or rose-red, and the red coloration is present on both the abaxial surface and on the distal portion of the adaxial surface. Compared to C. glaucophylla and C. texensis, the sepals of C. addisonii are more purplish, varying from light purple to purplish-red with whitish api- ces. Typical specimens of C. versicolor have sepals that are purplish-red or blu- ish-lavender proximally and cream-colored or greenish distally. The sepals of C.carrizoensis are yellowish-cream and lightly tinged with lilac proximally. Habitat can be used to distinguish the species of the C. glaucophylla com- plex to some extent. Clematis addisonii, C. texensis, and C. versicolor are appar- ently obligate calciphiles. Clematis glaucophylla is a facultative calciphile. Clematis carrizoensis is not associated with calcareous rock outcrops but rather is restricted to deep sandy soils. All of the species seem to grow best in edge habitats: however C. addisonii, C. texensis, and C. versicolor are sometimes found in closed-canopy forests. When incomplete or non-flowering specimens of species of the C. glaucophylla complex are encountered, the most reliable character that can be used to distinguish the species is geography. All of the taxa are allopatric. The ranges of C. versicolorand C.glaucophylla abut one another in portions of south- eastern middle Tennessee and possibly extreme northern Alabama. Clematis texensisand C. carrizoensisare restricted to Texas and C. addisoniti is restricted to Virginia. Clematis versicolor is found only in portions of Oklahoma, Arkan- sas, Missouri, Kentucky, Tennessee, and Alabama; it should be also searched for in extreme northeastern Mississippi (Tishomingo Co.) and in extreme south- ern Illinois. Clematis glaucophylla occurs in Mississippi, Alabama, Georgia, Florida, and Tennessee. Alabama and Tennessee are the only two states that support more than one species of the complex. KEY TO THE SPECIES OF THE CLEMATIS GLAUCOPHYLLA COMPLEX (ADAPTED FROM ERICKSON 1943, DENNIS 1976, AND PRINGLE 1997) 1. Plants ascending to erect, sometimes branched above, rarely viny; middle and lower cauline leaves mostly simple; flowers mostly terminal; restricted to Ridge and Valle Province of western Virginia C. addisonii Britton 76 BRIT.ORG/SIDA 22(1) 1. Plants scrambling or climbing, viny; middle and lower cauline leaves compound; lowers axillary; species not erviginles. 2. Leaves usually thin,ad without raised secondary and ae veins C. glaucophylla Small 2. Leaves usually thick, adaxial surface with moderately to prominently raised secondal y al id tertiary veins 3. Abaxial surface of sepals scarlet to rose-red or PSOE C AID and one lilac tinged; leaflets mostly ovate to subrotund, base mostly subcordate to deeply cordate, apex usually Ue lower leaflets commonly 2-3- lobed or 3-foliate; plants restricted to Tex 4. Sepals scarlet to rose-red abaxially; stamens not densely or uniformly pubescent, connective mostly glabrous or with a few stiff trichomes; plants of limestone uplands of the Edwards's Plateau in south-central Texas C. texensis Buckley 4. Sepals yellowish-cream abaxially and lightly tinged with lilac; stamens densely and uniformly pubescent, connective moderately to densely pubescent; plants of deep sandy habitats of northeastern Texas __ C. carrizoensis Estes 3. Abaxial surface of sepals purplish-red or bluish-lavender proximally and cream-colored or greenish distally; leaflets mostly ovate to elliptic, base obtuse to subcordate, apex mostly obtuse; lower leaflets mostly unlobed and undivided, infrequently lobed or ternate; plants of the Interior High- lands of Missouri, Oklahoma, and Arkansas and the Interior Low Plateau of Tennessee, Kentucky, and Alabama C. versicolor Smal! ACKNOWLEDGMENTS I would like to thank the faculty and staff at the following herbaria for loaning specimens or facilitating visits during this study: BRIT, MO, SMU, TENN, TEX- LL. Gratitude is also extended to R.L. Small, B.E. Wofford, and J. Beck for review- ing earlier versions of this article, to R. Naczi for preparing the Latin diagnosis, to J. Singhurst for assistance with field work, to C. Fleming for preparing the map, to S. Baskauf for use of his photo of C. versicolor,and to WC. Cart for use of his photo of C. texensis. REFERENCES Dennis, W.M.1976.A biosystematic study of Clematis, sect. Viorna, subsection Viornae PhD. Dissertation. The University of Tennessee. Knoxville Erickson, R.O. 1943. Taxonomy of Clematis section Viorna. Ann. Missouri Bot. Gard. 30:1-62. Jones, R.L. 2005. Plant life of Kentucky: an illustrated guide to the vascular flora. The Uni- versity Press of Kentucky, Lexington. Kral, R. 1983. A report on some rare, threatened, or endangered forest-related vascular plants of the South. USDA Forest Service, Southern Region, Tech. Publ, R8-TP2. 2 vol- umes. KRAL, R. 1987. A new “Viorna” Clematis from northern Alabama. Ann. Missouri Bot. Gard. 74:665-669. ESTES, A NEW SPECIES OF CLEMATIS FROM TEXAS 77 McBryoe, J.B. 1933. The vegetation and habitat factors of the Carrizo sands. Ecol. Monogr 3:249-297 Prinate, J.S. 1997. Clematis. In: Flora of North America north of Mexico. Volume 3. Magnoliophyta: Magnoliidae and Hamamelidae, ed. Flora of North America Editorial Committee. Oxford University Press, NY. Pp. 158-176. Sorrie, B.A.and A.S.Weaktey. 2001. Coastal Plain vascular plant endemics: phytogeographic patterns. Castanea 66:50-82. 78 BRIT.ORG/SIDA 22(1) ANNOUNCEMENT The Rupert Barneby Award The New York Botanical Garden is pleased to announce that Rodrigo Duno de Stefano, of the Centro de Investigacion Cientifica de Yucatan A.C. (CICY), is the recipient of the Rupert Barneby Award for the year 2006. He will be studying the family Leguminosae in the Yucatan Peninsula Biotic Province (YPBP), Mexico. With about 60 genera and more than 260 species there, the Leguminosae are one of the most important plant elements of the Yucatan region. This study will also contribute toa revision of four legume genera for the “Illustrated Flora of the Yucatan Peninsula” (G. Carnevali, general editor). The New York Botanical Garden now invites applications for the Rupert Barneby Award for the year 2007. The award of US$ 1,000.00 is to assist research- ers to visit The New York Botanical Garden to study the rich collection of Leguminosae. Anyone interested in applying for the award should submit their curriculum vitae, a detailed letter describing the project for which the award is sought, and the names of 2-3 referees. Travel to the NYBG should be planned for sometime in the year 2007. The application should be addressed to Dr. James L. Luteyn, Institute of Systematic Botany, The New York Botanical Garden, 200% Street and Kazimiroff Blvd., Bronx, NY 10458-5126 USA, and received no later than December |, 2006. Announcement of the recipient will be made by De- cember 15¢, Anyone interested in making a contribution to THE RUPERT BARNEBY FUND IN LEGUME SYSTEMATICS, which supports this award, may send their check, payable to e New York Botanical Garden Dr. James L. Luteyn Institute of Systematic Botany The New York Botanical Garden 200th St. and Kazimiroff Blvd. Bronx, NY 10458-5126 USA SIDA 22(1): 78. 2006 HILL CANE (ARUNDINARIA APPALACHIANA), A NEW SPECIES OF BAMBOO (POACEAE: BAMBUSOIDEAE) FROM THE SOUTHERN APPALACHIAN MOUNTAINS JK. Triplett! A.S.Weakley Department of Ecology, Evolution Univ. of North Carolina Herbarium (NCU) and Organismal Biology North Carolina Botanical Garden lowa State University University of North Carolina Ames, lowa, 50011-1020, U.S.A. Campus Box 3280 Chapel Hill, North Carolina, 27599-3280, U.S.A. L.G. Clark Department of Ecology, Evolution and Organismal Biology lowa State University Ames, lowa, 50011-1020, U.S.A. ABSTRACT A newly recognized Sees of A Arundinaria from the southern Appalachian Moun ae! is nla illustrated, and compared with the related A.giganteaand A. tecta.A is distinguished by a combination of eecetiye morp hological characters hn eatures of branching ane as TORO By) a anatomy, and ecology. Recognition of this species is consistent monophy ly of the species and its sister relationship with A. te a A key for the eoaaienen of Arundinaria species in North America is included along with a comparative table based on morphology, leaf anatomy, and ecology. Key worps: bamboo, Arundinaria, eastern U.S., Appalachian Mountains RESUMEN — a ee pecie de Arundinaria del sur de la Cogulers ae ies saan y se ; | : laci las A gigantea y A.tecta.S r z= S por una 1 aad g vegetati 70. I] P le | if morfologia y anatomia de las hojas, y ecologia F] a : A dar £ genéticos que apoyan la monofilia de la especie nueva y su posicion como especie hermanaa A. tecta. Se incluye una clave para la identificacion de especies de Arundinaria en América del Norte y una tabla comparativa basada en la morfologia, anatomia de las hojas, y ecologia. Pacapras claves: bambu, Arundinaria, este de los Estados Unidos, la Cordillera Appalachiana INTRODUCTION Arundinaria Michx. is a genus of north temperate woody bamboos (Poaceae: Bambusoideae) with a complex taxonomic history involving numerous enti- ties that have been placed within it at one time or another over the past two ‘Author for correspondence (triplett@iastate.edu) SIDA 22(1): 79 - 95, 2006 80 BRIT.ORG/SIDA 22(1) hundred years (McClure 1973; Li 1997; Judziewicz et al. 1999), with older treat- ments including upwards of 400 heterogeneous species from Asia, Africa, and the Americas. Currently Arundinaria is treated in a restricted sense to include only those species endemic to the eastern United States (Ohrnberger 1999), but debate continues regarding the inclusion of certain Asiatic taxa (e.g., Pleioblastus Nakai, Pseudosasa Makino ex Nakai, Bashania P.C. Keng & Yi, and Oligostachyum Z.P. Wang & G.H. Ye) that share key morphological features with the North American species (Li 1997; Judziewicz et al. 1999). However, because Arundinaria gigantea (Walter) Muhl. is the type species for the genus, its ge- neric placement is secure. As such, Arundinaria represents the only bamboos native to North America and the only temperate bamboos (the North Temper- ate clade of Clark et al., in press; Zhang & Clark 2000; Kelchner & Clark 1997) native to the New World, and provides another example of the classic disjunc- tion pattern in the flora of eastern Asia and eastern North America (Wen 1999). With a species richness ratio of approximately 20:1 up to 90:1, depending on which Asian group is used for comparison within the North Temperate bam- boo clade, Arundinaria and allies also provide an example of the intriguing asymmetry within this disjunction discussed by Guo and Ricklefs (2000). Arundinaria s.s. encompasses arborescent or subarborescent woody bam- boos with leptomorphic (running) rhizomes, persistent to deciduous and mostly glabrous culm leaves, and leaves at the tip of new shoots crowded into a dis- tinctive fan-shaped cluster or top knot with blades expanded as on foliage leaves. Branch complements typically have | primary branch and 0-2 subequal sec- ondary branches arising from shortened internodes at the base of the primary branch, which rebranch to produce up to 40 or more secondary branches on older culms. The culm and foliage leaves bear fimbriae and usually also au- ricles. Synflorescences in this group are determinate, open, and racemose or paniculate, with 6-12 laterally compressed florets per spikelet. Spikelets have 1-2 glumes and 3 stamens per floret. Like most temperate bamboos, Arundinaria has a basic chromosome number of x = 12 and presents several enigmatic characteristics including delayed flowering and monocarpy; repro- duction is primarily vegetative and seed production is infrequent and unpre- dictable. Arundinaria ss. is confined to southeastern portions of the continen- tal United States (specifically Alabama, Arkansas, Delaware, Florida, Illinois, Indiana, Kentucky, Louisiana, Maryland, Mississippi, Missouri, New Jersey, North Carolina, Ohio, Oklahoma, South Carolina, Tennessee, Texas, Virginia, and West Virginia), occurring from the Coastal Plain from New Jersey south to Florida and west to eastern Texas, and inland through the Piedmont to moder- ate elevations in the Appalachian Mountains. Arundinaria once formed exten- sive and dense canebrakes (with or without tree canopy) covering vast areas of fertile river bottomland often described by early explorers in the pre-colonial U.S. (West 1935), but has been greatly reduced in extent and abundance from TRIPLETT ET Al 81 its historical range by grazing and fire suppression (Hughes 1951, 1957, 1966; Platt & Brantley 1997; Judziewicz et al. 1999). Two species (Arundo gigantea Walter, Arundo tecta Walter) were originally described by Walter (1788) and represent extremes of morphological types found among inland and coastal forms, sometimes referred to as “Mississippi- type” and “Atlantic-type’ aan 1943), pees eeae ee (river cane or giant cane) forms extensive nies in low woods, moist ground, and along riverbanks from the lowlands east tof the Appalachians, west to Mis- souri, up the Mississippi Valley to southern Illinois and up the Ohio River to southern Ohio. Arundinaria tecta (Walt.) Muhl. (switch cane) forms colonies in non-alluvial swamps, moist pine barrens, live oak woods and along sandy margins of streams, preferring moister sites than A. gigantea. It is restricted to the Coastal Plain of the southeastern U'S., from southern Maryland to Alabama and Mississippi (McClure 1973; Hitchcock 1951). Within Arundinaria s.s., complex population-level variation has contin- ued to be problematic for taxonomists and field botanists (McClure 1973; Judziewicz et al. 1999; Platt & Brantley 1997). Phenotypic diversity among North American cane populations has inspired diverse taxonomic interpretations, generally with 1-3 taxa recognized at either specific or subspecific levels (Gilly 1943; Young & Haun 1961; Voight & Mohlenbrock 1964; Radford et al. 1968; Hitchcock 1971; McClure 1973; Campbell 1985; Tucker 1988; Platt & Brantley 1997). McClure (1973) published the most recent exhaustive treatment of Arundinaria and took a conservative approach, recognizing a single polymor- phic species (A. gigantea) and three subspecies, one of which [A. gigantea ssp. macrosperma (Michx.) McClure] is a catch-all for putative hybrids derived from the introgression of the other two. However, McClure acknowledged that fur- ther studies, particularly genetic-based studies, were necessary to clarify the phylogeny and taxonomy of this group. In the Southern Appalachians, astute botanical observers have long ques- tioned the identity of a curious short-statured cane that typically occupied sites away from streams and rivers. Among the diverse North American populations of Arundinaria, a variety with deciduous foliage was recorded by botanist C.D. Beadle in Western North Carolina (Beadle 1914; see also Young 1945). Beadle recognized this form as A. tecta var. decidua, not ruling out the possibility that it might in fact be a distinct species. It is unclear why Beadle associated this deciduous variety with A. tecta, although perhaps it was because of the small stature typically associated with A. tecta. In the first half of the nineteenth cen- tury, two botanists affiliated with the University of North Carolina Herbarium, William Willard Ashe and William Chambers Coker, made notes on specimens collected near Highlands, North Carolina indicating that the short, delicate, deciduous cane of the mountains might be a distinct taxon. Roland Harper (1928) was also intrigued by cane specimens occurring on bluffs in northern 82 BRIT.ORG/SIDA 22(1) Alabama and thought that an unrecognized taxon might be present. In the lat- ter half of the twentieth century, botanists and ecologists have informally con- sidered the small, upland cane as “hill cane” and have been unconvinced that it could be assigned to either A. gigantea or A. tecta. Hill cane is often common in mesic and submesic Sones and upland woodlands. Moreover, because of over- lapping morphological characteristics, floristic descriptions of A. tecta and A. gigantea may be confounded by this distinct form of cane. For example, Arundinaria tecta has been described as occurring along river branches 450 to 580 feet above sea-level on the southeast slopes and along the courses of moun- tain streams and shady mesic hillsides in the foothill region, well back from water (Harper 1928; Peattie 1929), but these inland and upland habitats almost certainly are populated by hill cane, not switch cane. In our field work on Arundinaria, we located several widespread populations of hill cane along the southern Appalachian Mountain chain, extending the range of the form that Beadle described. Field work has been complemented by herbarium studies to produce our current understanding of the range for this entity. Species limits within Arundinaria s.s. have not been examined previously ina phylogenetic framework or with molecular tools. Our investigation of hill cane is part of a larger study of the phylogenetic history of the North Temper- ate bamboo clade (in collaboration with the Bamboo Phylogeny Group), and as part of that study we are reconstructing the phylogeny of river cane, switc cane, and hill cane utilizing Amplified Fragment Length Polymorphism (AFLP) data to test the monophyly of putative species and to correlate the results with morphological and geographical characters, with the goal of producing a re- vised treatment of the genus. Ordination analyses (PCA) of mot phological char- acters and AFLP studies will be presented in a later publication. Preliminary cladistic analysis of AFLP data (Triplett & Clark, in prep.) dem- onstrated that the three types of cane form separate monophyletic lineages en- compassing two previously recognized entities (river cane and switch cane) and one entity encompassing those plants recognized as hill cane. Moreover, hill cane specimens from a wide geographic range cluster as the sister clade of A. tecta, rejecting the hypothesis that hill cane is an ecologically induced form of A. tecta and instead suggesting that it is a distinct lineage with a unique evolu- tionary history. These results prompted a reevaluation of diagnostic characters within Arundinaria s.s. Preliminary ordination (PCA) analyses of morphologi- cal characters similarly identified three non-overlapping entities and allowed us to recognize the most important diagnostic vegetative features. These groups correspond precisely with the three lineages derived from the AFLP data. We therefore propose the recognition of each entity at the species level: A. gigantea (river cane), A. tecta (switch cane) and a previously undescribed species (hill cane). In advance of the publication of the Flora of North America, we here describe and illustrate the new species, A. appalachiana Triplett, Weakley, & —_ =) TRIPLETT ET AL., A NEW SPECIES OF 83 L.G. Clark, from the southern Appalachian Mountains, and compare and con- trast it with its congeners A. gigantea and A. tecta. MATERIALS AND METHODS Field studies of natural populations conducted in October 2003 and July- October 2005. Standard bamboo collection procedures were followed (Soderstrom & Young 1983); bulky specimens of rhizomes, branch comple- ments, and culm nodes and internodes were made for all collections. Herbarium specimens from A, F, GA, GH, ISC, NCU, and US (herbarium acronyms following Holmgren & Holmgren 2006) were examined. While our taxonomic circumscription of Arundinaria s.s. is based on an approach com- bining morphological and molecular data, we have relied upon morphological characters to provide identifications of the specimens examined. Complete specimens, including culm leaves, buds, branch complements with foliage leaves, and synflorescences were rarely available, and some herbarium speci- mens of Arundinaria could not be conclusively assigned to species. Only one flowering specimen was located among the specimens identified as A. appalachiana. Specimens were measured for a variety of morphological characters, in- cluding foliage leaf length and width, inner ligule length, inflorescence length, spikelet length, and lengths of spikelet bracts (glumes I-IV, lemma and palea). Top knot (the cluster of leaves at the tip of new shoots) and foliage leaf lengths were measured from the base of the pseudopetiole to the tip of the blade. Leaf width was measured at the widest point. Primary branch length was measured from the point of origin at the node to the end of the branch axis. Synflorescence length was measured from the base of the basalmost branch to the apex of the main axis. Spikelets were removed from specimens and softened using a modi- fied Pohl’s solution (Pohl 1965: 750 ml distilled water, 250 ml 1-propanol, 2 ml liquid dish soap), dissected, examined, and measured for floral characters us- ing a dissecting microscope equipped with a micrometer. Anatomical charac- ters of leaf blades (both epidermal micromorphology and cross sections) were obtained using light microscopy of sections made following standard proto- cols for free hand sectioning and epidermal peels (Clark 1986; Ellis 1976, 1979). RESULTS AND DISCUSSION Our decision to recognize this taxon at the species level is based upon the com- bination of phylogenetic and morphologic analyses with careful consideration of the decisions made in the past regarding the North American Arundinaria species complex and the ability to diagnose monophyletic units. This interpre- tation follows from morphological (ie., diagnostic characters) and phylogenetic (ie. unique ancestry) species concepts (Olmstead 1995; Sites & Marshall 2003): The features discussed below are those identified as the most diagnostic based 84 BRIT.ORG/SIDA 22(1 on observations made during field work and morphological ordination analy- ses (PCA). The most consistent differences among the North American species are seen in vegetative characters including features of the rhizomes, culm in- ternodes and culm leaves, branching, and top knot and foliage blades, described below and summarized in Table 1. Distribution & ecology Arundinaria appalachiana is indigenous to the southern Appalachian Moun- tains where it occurs in the southern Blue Ridge, Blue Ridge /Piedmont Escarp- ment, upper Piedmont, and Ridge and Valley physiographic provinces (Fig. 1). The full extent of its distribution is still poorly known, because of the infre- quent collection of bamboos in eastern North America and the of ten poor qual- ity of the existing specimens; for this reason we have chosen to supplement vouchered specimens with additional county records based on what we con- sider reliable sight records of this new species (these counties should be veri- fied with vouchers). Hill cane iscommon in oak-hickory forests and woodlands on mesic, submesic, and xeric slopes and uplands, sometimes occurring as well in hillside seepages, but nearly always on slopes, bluffs, and ridges away from perennial streams, in contrast to the geographically sympatric A. gigantea. As- sociated overstory species include Quercus montana Willd., 9. coccinea Mtnchh., Q. alba L., Q. rubra L., Q. falcata Michx., Pinus echinata Mill. P virginiana Mill, Carya alba (L.) K. Koch, Carya glabra (Mill.) Sweet, and Nyssa sylvatica Mar- shall. Although these slope and ridge forests are well-drained, the annual rain- fall amounts in this region are high and equably distributed. In the escarpment gorges of the Chattooga River, Whitewater River, Thompson River, Horsepasture River, Toxaway River, Horsepasture River, and Eastatoe Creek, the highest an- nual rainfalls in eastern North America (exceeding 80 inches a year) maintain higher then expected moisture levels even in topographic situations that tend tocreate xeric communities (ridgetops and convex upper slopes and side ridges) (Robinson 2000). Still, there is no question that hill cane occupies unusually dry and upland habitats compared to its congeners. Arundinaria gigantea typi- cally occurs on the floodplains of large to small rivers, sometimes edging onto lower portions of mesic slopes, whereas A. tecta typically occurs along small to medium blackwater rivers, in swamps, on deep peat in pocosins, and in small seepages with organic soils. Arundinaria dppalachiana is sympatric (at least in the broadest sense) with both of its congeners, in that it occurs in the same counties and within a few kilometers of populations of both A. gigantea and A. tecta. Within the re- gion occupied by A. appalachiana however, A. gigantea occurs only along the upper reaches of major rivers, notably the Little Tennessee and French Broad Rivers, while A. tecta also occurs at lower elevations and in different topographic and ecological situations. TRIPLETT ET AL., A NCW SF EXITED UF Taste 1. Morphological comparisons of A dinaringn palachiana,A. tecta, and A. gigantea. Character A. appalachiana A. tecta A. gigantea Rhizome air canals present or absent — present absen ulcus usually absent usually absent et present Culm leaf duration persistent persistent eciduous Culm leaf auricles absent present, deciduous present, deciduous Top knot number of leaves 6-12 9-12 - Top knot leaf blade length (cm) 9-22.5 20-30 16-24 Compressed basal internodes 2-5 2-4 0-1 n primary branch 1° branch basal nodes: 2° branches absent present,subequal — present, subequal Primary branch length (cm) 7-33 usually >50 15-25 Foliage leaf blade length (cm) 5-20 7-23 8-15 Foliage leaf blade width (cm) 8-2 1-2 0.8-1.3 Foliage leaf vestiture pilose or glabrous densely pubescent densely pubescent or glabrous or glabrous Foliage leaf duration deciduous evergreen evergreen Foliage leaf texture chartaceous coriaceous subcoriaceous Foliage leaf abaxial tessellation weakly tessellate strongly tessellate | 1] DUUT ly LESSEN GLO The species biology of A. appalachiana is poorly understood. Its congeners are long-lived monocarpic perennials, and this appears to be the case with A. appalachiana as well. It has been seen flowering and fruiting even more infre- quently than its congeners; judging from its habitat, its flowering and fruiting may be stimulated by fire (like A. tecta), and the paucity of fertile specimens may reflect the general suppression of fire in its habitat through the twentieth century. Field observations suggest that clones of A. appalachiana are slow- growing and very long-lived, certainly persisting for decades and likely for cen- turies. Morphology Rhizomes.—The rhizomes of A. appalachiana are leptomorphic, a characteristic they share with other north temperate woody bamboos; however, in many cases the growing tips of new rhizomes travel only a short distance before turning up to form a new culm, thus presenting a sympodial branching pattern. This pat- tern also occurs in A. tecta but has not been confirmed for A. gigantea. An inter- esting characteristic of this species is the variability in air canal development (McClure 1963). Air canals are present in some specimens but not others, and may in fact be longitudinally and peripherally discontinuous in rhizomes of A. appalachiana. Air canals are ntly continuous. Culm internodes.—Although the culm internodes of A. appalachiana can lyf present in A. tecta and desta Be 86 BRIT.ORG/SIDA 22(1) Ree Py ae aie. ee Ceae ar co aewen aaa Syste ea NI Oy anes serene Ee to ee sta oo KD RES be somewhat flattened behind the branch complement, the internodes lack a prominent groove or sulcus. This is consistent with A. tecta but contrasts with A. gigantea, which typically has internodes that are prominently sulcate. Branching.—In bamboos, the morphology and architecture of the set of branches arising from culm nodes (the branch complement) is a source of nu- merous taxonomically useful characters. In Arundinaria, the pattern of short- ened or compressed internodes at the base of primary branches and the extent and pattern of secondary branching are especially valuable. The branch comple- ment of A. appalachiana is characterized by 2-5 shortened or compressed in- ternodes at the base of the primary branch, without rebranching in this basal area. The first elongated internode above the shortened ones is typically con- strained to ~30% the length of distal internodes. In contrast, A. tecta, while having a similar pattern of compressed internodes, typically will produce buds and branches from the nodes in the area of compression, creating subequal branches from the base of the primary branch. Arundinaria gigantea typically has only one (or no) compressed basal internode, but if present, this node may produce a secondary branch. Primary branches in A. appalachiana are usually TRIPLETT ET Al 87 less than 35 cm long. In contrast, A. tecta produces long primary branches usu- ally >50 cm. Culm leaves.—The culm leaves of A.appalachiana are typically shorter than their associated internodes at the base of the plant, becoming progressively longer towards the top knot. At midculm they are approximately the same length as the associated internode. In contrast, midculm culm leaves of A. tecta are longer than their associated internodes, and those of A. gigantea shorter. Arundinaria appalachiana and A. tecta have persistent culm leaf sheaths, whereas A. gigantea has deciduous sheaths. The culm leaf sheaths of A. appalachiana are tessellate; however, their tessellation is not as pronounced as itisin A. tecta. The culm leaves lack well-developed, prominent auricles, unlike A. gigantea and A. tecta. Top knot and foliage leaves.—In Arundinaria, leaves at the tip of new culms are crowded into a distinctive fan-shaped cluster or top knot, with their blades expanded as on foliage leaves. The top knot leaf blades of A. appalachiana are typically 9-22.5 cm in length, while A. tecta typically has larger blades (20-30 cm long); those of A. gigantea are typically 16-24 cm in length. The foliage leaf blades of A.appalachiana are deciduous; bladeless branches persist on the plants after leaf drop, often with the older sheaths still intact. The blades are chartaceous, presumably, since the blades are deciduous, the plant invests less energy in producing them by producing less sclerenchyma. In contrast, the leaves of A. gigantea are persistent and subcoriaceous, while leaves of A. tecta are persistent and coriaceous. The abaxial surfaces of the leaf blades of A. appalachiana are weakly tessellate, whereas in A. gigantea and A. tecta they are strongly tessellate. The abaxial and adaxial leaf surfaces are typically sparsely to more or less densely pilose (glabrous in some populations) in A. appalachiana. Leaf blades of A. gigantea are typically densely pubescent with short, soft hairs on abaxial surfaces, while the blades of A. tecta are densely pubescent on both surfaces; neither of these pubescence patterns has been seen in A.appalachiana It is important to note that Arundinaria gigantea and Arundinaria tecta can both survive in deeper shade in the forest, albeit in stunted conditions, and thus it is possible to find diminutive or depauperate plants of both that super- ficially resemble hill cane, mostly in stature. However, a combination of char- acters such as the branch complement, leaf texture, and leaf pubescence are usually sufficient to distinguish these stunted forms of river and switch cane from hill cane populations and from each other TAXONOMIC TREATMENT KEY TO THE SPECIES OF ARUNDINARIA SENSU STRICTO 1. Primary branches with 0-1 compressed basal internodes; culm internodes usually sulcate; culm leaves deciduous A. gigantea 88 BRIT.ORG/SIDA 22(1 1. Primary branches with 2-5 compressed basal internodes; culm internodes usually terete; culm ies Salsa to oy cele UOUS. 2. Foliage bl Is, persiste f ly pubescent or gla- brous, elk tessellate; tae ae eae usually more than 50 cm long, basal nodes developing secondary branches; top sus blades a. 30. cm long A.tecta 2: ieee we hartaceous, deciduous, al | glabrot Is. Weakly branches usually less than 35cm ong. basal ode: not devel- oping secondary branches; top knot blades 9-22.5 cm long A. appalachiana Arundinaria appalachiana Triplett, Weakley @ L.G. Clark, sp. nov. (Figs. 1-5). TYPE: Ue STATES. ALABAMA: Dekalb Co.: Desoto State Park by Rt. 89 (34° 30' N Lat; 85° 38' W Long.), ~ ca. 515 m, 25 Jul 2005, Triplett & Ozaki 99 (HOLOTYPE: ISC; ISOTYPES: MO, NCU, UNA, U Rhizomata leptomorpha, strat l eri i lum praesenti.Culmi 2-6 mm diametro, 0.5-1.8 m alti, omnino glabri, erecti. nternodia 4.5-12 cm ronga te teretia ailgine sulco). Vaginae culmorum 5.5-11(-15) cm longae, persistentes, sine auriculis. Culmi j inali(5-)6-12 foliorum, laminis (5-)9-22.5 cm longibus, 1.4-2.8 cm latibus, linear bee vel lineari-lanceolatis vel ovati- lanceolatis, pilosis vel interdum glabris. Rami pew ad nodos medianos c I 3 en longi, internodiis abbreviatis sine gemmis basi 2-5. Folia eae complementi 3-7. Laminae foliorum (3-)5-20 cm longae, 0.5-2 cm latae, shanties ae, deciduae, pilosae vel interdum glabrae, abaxialiter infirme tessellatae. Synflorescentiae 7-11.5 cm ee 2-5 cm latae, determinatae racemosae apertae; pedicelli 4-25 mm longi, spiculis 6-8. Spiculae 3-5.5 cm longae, glumis 1(-2), flosculis basilibus interdum sterilibus, 5-8 flosculis fertilibus continentibus et anthoeciis rudimentalibus terminalibus 1-3; glumae inaequales, 5-nervatae, attenuatae, glabrae; gluma I 3-6.5 mm longa; gluma II 5.5-9 mm longa; lemmata fertilia 11-16 mm longae, 7-1l-nervata, apicibus acutis vel acuminatis, abaxialiter glabris; palea LO-13 mm longa, 8-10-nervata. Stamina 3; anthera 5-7 mm Woody bamboo. Plants of diffuse to (pluri-) caespitose habit. Rhizomes leptomorphic, usually horizontal for a only short distance before turning up at the apex to form a culm (therefore often presenting a sympodial branching pattern), hollow (with a small central lumen), peripheral air canals sometimes present (if so, apparently discontinuous longitudinally and/or peripherally). Culms 2-6 mm in diameter, 0.4-L8 m tall, erect, tillering; internodes 4.5-12 cm long (progressively shorter towards culm apex), terete, hollow, glabrous, flat- tened behind the branch complement on larger culms but the sulcus not promi- nent; nodes solitary, the nodal line horizontal, supranodal ridge not aoe bud one per node (single) on a slight promontory, triangular, the shoulders of the prophyll ciliate. Culm leaves persistent, approximately equaling associated internodes at midculm, typically shorter than associated internodes at the culm base, becoming proportionally longer towards the culm apex; sheaths 5.5-11 (-15) cm long, shortest on lower nodes, becoming progressively longer towards the culm apex, glabrous, margins ciliate; blades 0.5-1.4 cm long, triangular to lanceolate, reflexed to erect, glabrous, deciduous, intergrading into top knot leaves; auricles absent; fimbriae 1-9 mm long, ascending to erect; inner ligules ca. 0.1 mm long, a fringe of short cilia; outer ligule absent. Top knot leaves in an apical cluster of (5-)6-12; sheaths glabrous, margins ciliate; auricles absent; fim- 90 BRIT.ORG/SIDA 22(1) Fic. 3. Arundi hiana. A. Partially di I florets. B. Synfl with five spike- lets. Scale bar = 1 cm. m. Drawings based on Ahles & Leisner 15147. (Illustrations by J. Triplett) briae 1-9 mm, ascending to erect; blades (5-)9-22.5 cm long, 1.4-2.8 cm wide, L:W = 8.3-9.3, linear, linear-lanceolate or ovate-lanceolate, chartaceous, pilose or glabrous, abaxially weakly tessellate, apices acuminate, bases attenuate to cuneate, midrib + centric. Branching infravaginal (rarely extravaginal); primary branches | per node, 7-33cm long, with 2-5 compressed basal internodes, basal nodes not developing secondary branches; first elongated internode shorter than subsequent ones (~30%); higher order branches present on older plants, reiter- ating the 1° branch (ie., with the same pattern of compressed basal internodes and branching). Foliage leaves 3-7 per complement; sheaths glabrous, margins ciliate, weakly tessellate; auricles absent; fimbriae 1-9 mm, ascending to erect; inner ligule glabrous or ciliate, fimbriate or lacerate; outer ligule present as a minute rim: blades linear, linear-lanceolate, or ovate-lanceolate, chartaceous, deciduous, surfaces pilose (sometimes glabrous), abaxially weakly tessellate, apices acuminate, bases attenuate to cuneate, midrib + centric; primary branch foliage leaf blades (4-)9-20 cm long, (0.5-)0.8-2 cm wide; L:W =10.7-1L7; higher order branch foliage leaf blades (3-)5-17.5 cm long, (0.5-)0.8-15 cm wide; ter- minal foliage leaf blade aaa unexpanded laterally, pasa: but persisting as a tail-like appendag 7-11.5cm long, 2-5 cm wide, determi- wo TRIPLETT ET AL., ANEW SPFEAILS UP 91 Fics. 4-5. Arundinaria appalachiana. 4. Habit, in Rhea Co., Tennessee. 5. Close-up of primary branch with compressed basal internodes, Dekalb Co., Alabama. (Photos by J. Triplett) nate, open-racemose, apparently borne on specialized nonleafy shoots; peduncle 10-17 cm long (3 measured), glabrous, terete; rachis glabrous; pedicels 4-25 mm long; 6-8 spikelets per synflorescence. Spikelets 3-5.5 cm long, laterally com- pressed, disarticulating above the glumes and between the florets, consisting of 1(-2) glumes, occasionally a basal sterile floret, 5-8 fertile florets and 1-3 progressively rudimentary apical sterile florets; rachilla internodes 3-4 mm long; glumes unequal, 5-nerved, attenuate, glabrous; glume I 3-6.5 mm long; glume II 5.5-9 mm long; fertile lemmas 11-16 mm long, 7-1l-nerved, apex acute or acuminate, abaxially glabrous, transverse veinlets barely perceptible or not at all manifest, usually somewhat reddish-purple; paleas 10-13 mm long, 8-10- nerved, broadly sulcate and 2-keeled dorsally; lodicules and ovary not seen. Sta- mens 3; anthers 5-7 mm long. Fruit not seen. Distribution and Ecology.—(Fig. 1). Endemic to the southern Appalachians and upper Piedmont of northeastern Alabama, northern Georgia, southwest- ern North Carolina, northwestern South Carolina, and southeastern Tennessee, United States; 300-800(-1065) m. In upland oak-hickory-pine forests on slopes, less typically in more mesic sites, seeps, or along small streams. Phenology.—Lack of specimens in flower or information on the extent of 92 BRIT.ORG/SIDA 22(1) blooming makes it impossible to determine flowering behavior in this species at present. Of the specimens cited below, only one confirmed flowering speci- men of this species was identified, suggesting that flowering may be an even rarer event in this species than in most woody bamboos. Etymology.—Arundinaria appalachiana is named for its distribution in the forests of the Appalachian Mountains. Common name.—hill cane. R ined: UNITE D STATES, ALABAMA. Dekalb Co.: Little River Canyon National peste plot PIEC 27, 2002, McDaniel s.n.(NCU); in steep wooded bank of West Fork, Little River, across stream from overlook shelter, DeSoto State neh 9 May 1959, Sherman & Carter 25747 (GH); Desoto State Park on trail by Laurel Creek (34° 30'N Lat; 85° 38’ W Long.), elev. ca. 515m, 25 Jul 2005, Triplett & Ozaki 100 USC, MO, US). GEORGIA. Bartow Co.: in woods on S side of Stamp Creek Rd. just E of Jones Mill Rd. (34° 14' N Lat; 84° 40' W Long.), elev. ca. 311 m, 26 Sep 2005, Triplett 166 CISC, MO, US). Rabun Co.: maple-birch-magnolia association, Laurel Creek Olivine Deposit, 1.5 mi E of Pine Mt., 21 Jun 1946, Radford s.n. (NCU ): + -hickory woods on Pine Mt., Bald Rd., 4 Jun 1952, Radford 6134 (NCU); Warwoman Road and Overflow Creek bridge, N side, 20 May 1996, Stancil 950 (GA); in swamp near top of Oakey Mountain, SE of Nacoochee Reservoir, 25 May 1941, Duncan 3283 (GA); on trail from 155 0.5 mi S of Kattie Place, elev. ca. 790 m, 22 Oct 1995, Milsted & Zhang 585 (GA); pine-oak woodland near roadside park, | mi N of Tallulah Falls River, 9 May 1967, Isely, Welsh, & Isely 10286 USC), 2 mi N of Warwoman Rd. along GA 28, scattered aan mature woods W of road (34° 57'N Lat; 83° LO’ W Long.), elev. ca. 687 m, 25 Oct 2003, Triplett & Clark 20 (SC, MO, US). Stephens Co.: Deep gorge, Cedar Creek, Camp Mikell Rd. off GA 184, just W of Camp Mikell and N of Toccoa (34 08' 06" N. lat; 83 20' 13” W. long), elev. ca. 300 m, 19 Jun 1975, Boufford & Wood 16501 (NCU), NW- facing ravines and ridges on the S side of Panther Creek, SW of Yonah Lake (Tugaloo River), N of Toccoa (34 43' 30” N. lat; 83 21' 13” W. long.), 25 Jun 1975, Boufford & Wood 16766 (NCU). NORTH CAROLINA. Clay Co.: Oak-hickory woods, 4 mi NW of Hayesville, 5Jun 1952, Radford & Wood 6162 (NCU). Gaston Co.: low woodland near the northern tip of Pasour Mt., about 3 mi SSW of High Shoals, 21 Jun 1956, Ahles & Leisner 15147 (NCU); oak-hickory woods on Crowder’s Mt, 4 Jun 1953, Radford 7084 (NCU). Graham Co.: small population on N side of Santeetlah Dam Rd. (SR 1146) just off Hwy 129 (35° 22'N Lat; 83° 5I’ W Long.), elev. ca. 537 m, 2 Oct aie eran MO, US). Jackson Co. on highway between Dillsboro & Park Entranc n e top of bank by roadside, 26 = Jul 1937, Coker s.n. (NCU); Cope Creek ae ne af NC 23 out ci ae growing among Polytrichum sp., Pine, and sedge (35° 23' N Lat; 83° Long.), elev. ca. 675 m, 25 Oct 2003, Triplett & Clark 21 CISC, MO, US). Lincoln Co.: rich oe forest and stream banks 0.4 mi W of Cat Square, 28 Apr 1957, Bell 6638 (NCU). Macon Co.: maple-birch-magnolia (cove) association, Corundum Hill Olivine Deposit, 1 1/2 mi NW Gneiss, 15 Jun 1946, Radford s.n.(NCU); Horse Cove near Highlands, 3 Sep 1948, Radford s.n. (NCU); in deep, shaded ravine at top of pasture at foot of Whiterock Mountain, 7 mi from Otto, NC on Tessentee Creek Rd., elev. ca. 1065 m, 7 Aug 1938, Stewart & Hechenbleikner s.n. (NCU) pine-broom-straw association, Corundum Hill Olivine Deposit, 1.5 mi NW Gneiss, 15 Aug 1946, Rad- ford sn. (NCU); Mulberry Rd. ca. 0.5 mi off 441, on steep hillside among oak, rhododendron, and maple (35° OLN Lat; 83° 23' W Long.), elev. ca. 647 m, 25 Oct 2003, Triplett, Clark, & Weakley 19 (SC, MC . McDowell Co.: near Marion, 25 Oct 1915, Ashe s.n. (NCU). Polk Co.: 4 mi W of Tyron, valley of Fall Creek in wet mez dows on eee in reservoir, 19 Jun 1942, Walker 3469 (US). Rutherford Co.: wet ditch on CR 1721 2 mi N of Sunshine, 24 Jun 1967, Smith 74 (NCU). Swain Co.: Bryson City, by stream, 1] Jul 1927, Hunne a sn. (GH); Bryson City, private property on W side of Wiggins Rd., 0.3 mi up from Betts Branch (SR 1343) (35° 26' N Lat; 83° 25' W Long.), elev. ca. 601 m, | Oct 2005, Triplett 184 (SC, MO, US). Transylvania Co.: rocks and cliffs on side of mountain, 100 to 200 feet ee the bank of Davidson River, Pisgah Forest, 28 Sep 1915, Ashe s.n. (NCU); Pisgah Forest, roac to TRIPLETT ET AL., A 93 Pink Beds, 23 Aug 1938, Stewart s.n. (NCU); Middle Bearcamp Creek area, roadside, Highlands, 2 Aug 1962, Rodgers & Shake 62165a (NCU), oak woods near Looking Glass Falls, 7 Jun 1952, Radford & Wood 6192 (NCU); Horsepasture Gorge, roadside about 1 mi W of crossing, elev. ca. 488 m, oyun 1961, Rodgers 6162a (NCU). SOUTH CAROLINA. Greenville Co.: Cedar Mountain, 0.4 mid road mi S of Caesar's Head SP Visitor Center (35° 05' N Lat; 82° 36’ W Long,), ae ca.515 m, 30 Sep 2005, Triplett 179 (SC, MO, US). Oconee Co.: oak-hickory woods ridge above Walhalla Fish Hatchery, ca. 11 mi N of junction of S.C. Routes 28 and 107 on Route 107, 9 Jun 1952, Wood 7879(A); Hill property, NW side Old Rocky Gap Road, W side West Village Creek near creek, Mountain Rest, Blue Ridge prov- ince, elev. ca. 520 m., 24 Aug 1991, Hill 22585 (GH). Pickens Co.: mixed deciduous forest, 3 mi N of pe Bottom near US 178, 22 Aug 1956, ta 16758 (NCU); Boggs’ Rock, granite-gneiss outcrop N of Liberty, 3 Jun 1974, Knox 407 (NCU) wooded slope, 2.4 mi S of NC line on US 178, 8 Jun 1956, small plant at summit of mountain, in hardwood forest. 29 Jun 1957, Pohl 7664A (ISC). Monroe Co.: White Cliff Springs, Jul 1890, Lamson-Scribner s.n. (US). Polk Co.: Boyd Gap Overlook above Ocoee River, 4.2 mi W of Hwy 68, on steep wooded hillside just beyond entrance to Boyd Gap Trail #331 (35° 02' N Lat; 84° 27' W Long.), elev. ca. 529 m, 26 Sep 2005, ae 165 (SC, MO, US). Rhea ve Firetower Rd just off Hwy 68 next to Grandview Community Center and below intersection wit Emergency Road (35° 44’ N Lat; 84° 50’ W Long.), elev. ca. 445 m, 3 Oct ee Triplett 188 (ISC, MO, ACKNOWLEDGMENTS Field work was supported by a Graduate Research Grant from the American Society of Plant Taxonomists and grants from the American Bamboo Society to JT as well as a National Geographic Society Grant (#7336-02) to LC. Support for JT and for the final preparation of this manuscript came from a National Science Foundation Grant (DEB-0515712) to LC. We are indebted to Angus Gholson (Chattahoochee, FL), Michael Hotchkiss (USDA, Byron, GA), Robert and Lauren Clark (Peachtree City, GA), Jim and Frankie Triplett (Winston-Sa- lem, NC), Gary Kauffman, Linda Lee Leslie, Adam and Sue Turtle (Earth Advo- cates Research Farm, Summertown, TN), and Julian Campbell (TNC, Lexing- ton, KY) for their hospitality or assistance in locating populations of Arundinaria. We also thank the curators of the herbaria that loaned specimens for this study. We are grateful to Gary Kauffman, Keith Langdon, Virginia McDowell, Tom Govus, and Mike Schafale for their interest and assistance in resolving the persistent taxonomic issue of the correct identity of the small upland mountain cane. Chris Stapleton (Royal Botanic Gardens, Kew) kindly pointed out to us the importance of compressed internodes in the taxonomy of this group and Emmet Judziewicz (Dept. of Biology, University of Wisconsin- Stevens Point) suggested the specific epithet. REFERENCES Beaote, C.D. 1914. Bamboo. In: Bailey, L.H. The standard cyclopedia of horticulture, Vol. 1. New York: The Macmillan Company. P. 446. Campsett, J.N. 1985. The land of cane and clover: presettlement vegetation in the so-called bluegrass region of Kentucky. Herbarium Report, University of Kentucky, Lexington. 94 BRIT.ORG/SIDA 22(1 Car, L.G. 1986. Systematics of Chusquea Section Chusquea, Section Swallenochloa, Sec- tion Verticillatae, Section Serpentes, and Section Longifoliae (Poaceae: Bambusoideae). Unpublished Ph.D. dissertation, lowa State University, Ames. CiarK, L.G.,S. DRANSFIELD, J. TRIPLETT, aNd J.G. SANCHEZ- ey (In eiese): Phylogenetic relationships among the one-flowered, determinat f Bambuseae (Poaceae: Bambusoideae). In:Columbus, J.T.,E.A. Friar, C.W. rion JM. es L.M.Prince,and M.G.Simpson,eds Monocots: comparative biology and evolution. 2 vols. Rancho Santa Ana Botanic Gar- den, Claremont, California, U.S.A. Eiuis, RP. 1976. A procedure for standardizing comparative leaf blade anatomy in the Poaceae. |. The leaf blade as viewed in transverse section. Bothalia 12:65-109. Eis, R.P. 1979. A procedure for standardizing comparative leaf blade anatomy in the Poaceae. Il. The epidermis as seen in surface view. Bothalia 12:641—-672 Gity, C.L. 1943. A preliminary investigation of the North American canes (Arundinaria). Bull. Torrey Bot. Club, 70:297-309. Guo, Q.and R.E. Rickters. 2000. Species richness in plant genera disjunct between temper- ate eastern Asia and North America. Bot.J. Linn. Soc. 134:401- Harper, R.M. 1928. Economic botany of Alabama.Part 2.Catalogue of the trees, shrubs, and vines of Alabama, with their economic properties and local distribution. Geol. Survey Alabama Monogr. 9:1-357. HitcHcock, A.S. 1951. Manual of grasses of the United States. Rev. by Agnes Chase. 2nd ed. USDA Misc. Pub. 200, U.S. Government Printing Office, Washington, DC. HitcHcock, A.S. 1971. Manual of the grasses of the United States. Dover Publ., New York. Hoimaren, PK.and N.H.Houmeren. 1998 onwards (continuously updated). Index herbariorum. New York Botanical Garden. http://sciweb.nybg.org/science2/IndexHerbariorum.asp Hucues, R.H. 1951. Observations of cane (Arundinaria) flowers, seed, and seedlings in the North Carolina coastal plain. Bull. Torrey Bot. Club 78:113-121. Hucues, RH. 1957.Response of cane to burning in the North Carolina coastal plain. North Carolina Agric. Exp. Stat. Bull. 402. Hucues, RH. 1966. Fire ecology of canebrakes. Proceedings of the Fifth Annual Tall Tim- bers Fire Ecology Conference, March 24-25, 1966. Mimeographed. uoziewicz, E.J., L.G. Ciark, X. LonDono, and MJ. Stern. 1999. American bamboos. Smithsonian Institute Press, Washington, DC. KeLCHNeR, S.A.and L.G.Cark. 1997.Molecular evolution and phylogenetic utility of the chlo- roplast rp/16 intron in Chusquea and the Bambuscideae (Poaceae). Molec. Phylogen. Evol. 8:385-397. Li,D.Z.1997.The flora of China Bambusoideae Project: Problems and current understand- ing of bamboo taxonomy in China. In: Chapman, G.P., ed. The bamboos. Linnean Soci- ety of London Symposium Series. Academic Press, London. McCture, F.A. 1963. A new feature in bamboo rhizome anatomy. Rhodora 65:134-136. McCiure, F.A. 1973. Genera of bamboos native to the New World (Gramineae: Bambusoideae). 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Handbook No. 193, Wash- ington, D.C. ZHANG, W.P. and L.G. Ciark. 2000. Phylogeny and classification of the Bambusoideae (Poaceae). In: Jacobs, $.W.L.,and J. Everetts, eds. Grasses: systematics and evolution. Pp. 35-42. CSIRO, Melbourne 96 BRIT.ORG/SIDA 22(1) ANNOUNCEMENT The 2006 Applications Delzie Demaree Travel Award Applications for the 2006 Delzie Demaree Travel Award should include a letter from the applicant telling how symposium attendance will benefit his/her graduate work and letter of recommendation sent by the major professor. Please send letters of application to: Dr Donna M.E. Ware, P.O. Box 8795, Herbarium, Biology Department, The College of William and Mary, Williamsburg, VA 23185- 8795, U.S.A. 1-757-221-2799, Email: ddmware@wm.edu. The period for receiv- ing applications will end three weeks prior to the date of the symposium if a sufficient number of applications are in hand at that time. Anyone wishing to apply after that date should inquire whether applications are still being ac- cepted before applying. The Systematics Symposium dates for 2006 are 13-14 October. The Delzie Demaree Travel Award was established in 1988 honoring Delzie Demaree who attended 35 out of a possible 36 symposia before he died in 1987. Delzie Demaree was a frontier botanist, explorer, discoverer, and teacher. His teaching career as a botanist began in Arkansas at Hendrix College in 1922. He also taught botany at the University of Arkansas, Navajo Indian School, Yale School of Forestry, Arkansas AQM, and Arkansas State University at Jonesboro where in retired as professor emeritus in 1953. One of the things he enjoyed most as a botanist was assisting students with their field botany research. — SIDA 22(1): 96. 2006 REVISION OF BAUHINIA SUBGENUS BAUHINIA SECTION AMARIA (CERCIDEAE: CAESALPINIOIDEAE: FABACEAE) Richard PR Wunderlin Department of Biolog University of South Florida Tampa, Florida 33620-5200 U.S.A rwunder@cas.usfedu ABSTRACT £R 1 ere | R . A A taxonomic treatment are recognized in series Decandrae andl one species is recognized in series Stenanthae. Bauhinia amatlana Wunderlin from southern Mexico, Bauhinia arborea Wunderlin from Ecuador, Bauhinia Fifteen species ayabacensis wie from Ecuador and Peru, and Bauhinia geniculata Wunderlin from Colombia are described and the combination Bauhinia aca eae ‘ Triana ex Hooker f. var. caudigera (S.F Blake) Wunderlin is proposed. Keys, descriptions data are provided. RESUMEN Se presenta un tratamiento taxonomico de Bauhinia subgénero Bauhinia secci6n Amaria. Se reconocen quince especies en la serie Decandrae y unaen oe serie ae nae Se edescPen Baubiiie d amatlana Wunderlin del Sur de México, Bauhinia arborea Wunderlin de Ecuador y Pert, y Bauhinia geniculata Wunderlin de Colombia, y se propone la combinacion Bauhinia petiolata (DC.) Triana ex Hooker f. var. caudigera (S.F Blake) Wunderlin. Se aportan claves, descripciones, y datos sobre la distribucion. The pantropical genus Bauhinia with approximately 300 species consists of four subgenera: Bauhinia, Piliostigma [=Elayunal, Barklya, and Phanera (Wunderlin et al. 1987). Bauhinia subgenus Bauhinia, as defined by Wunderlin et al., contains nine sections, of which three (Bauhinia, Pauletia, and Amaria) are New Wor This is the first of several proposed taxonomic treatments of the New World species of Bauhinia. Previous regional treatments in the last half-century such as those for Argentina (Fortunato 1986, 1997), Nicaragua (Wunderlin 2001), Panama (Wunderlin 1976), western Mexico (McVaugh 1987), Venezuelan Guayana (Wunderlin 1998), Middle America (Wunderlin 1983), and parts of the genus for Brazil (Vaz 1979; Vaz and Tozzi 2003) will be placed in the larger con- text of the entire genus for the Americas following Wunderlin et al. (1987). In the following revision, 16 species are recognized as comprising section Amaria. Fifteen of these are in series Decandrae and one in series Stenanthae. Series Triandrae, containing the single species Bauhinia pansamalana Donn. Sm. and placed in section Amaria by Wunderlin et al. (1987), is here transferred to section Bauhinia and combined in series Remotae with B. pinheiroi SIDA 22(1): 97- 122. 2006 = 98 BRIT.ORG/SIDA 22(1) Wunderlin with which it shares the characters of three fertile stamens and the same pollen type. The pollen sexine of B. pansamalanda is striate-reticulate and lacks the infratectal processes characteristic of section Amaria. Bauhinia section Amaria (S. Mutis ex Caldas) Endl. Gen. PI. 1318. 1840. Amarias. Mutis ex Caldas, Semanario Nueva Granada 2:25. 1810. TYPE DC. [=Bauhinia petiolata (S. Mutis ex DC.) Triana ex Hook. | ton and Killip (Ann. New York Acad. Sci. 36:164. 1936). : Amaria petiolata S. Mutis ex |. Lectotype designated by Brit- Trees or shrubs; branches lacking infrastipular spines. Leaves unlobed, bilobed, or rarely bifoliolate. Inflorescence terminal or axillary; floral bud ellipsoid to clavate. Hypanthium cupulate or turbinate to short-tubular; calyx spathaceous, sometimes irregularly splitting into several lobes in late aawhiesis petals 5, subequal; fertile stamens 10, the inner 5 shorter than the outer 5, or the inner 5 staminodial, the filaments connate at the base or free; pollen spheroidal, inaperturate or 3-colporoidate, the tectum reticulate or rarely striate-reticulate, with eT slunt spine-like or rarely low tectal processes continuous from the foot layer through the tectum (infratectal) or rarely supratectal; gynophore partly adnate to the abaxial wall of the hypanthium; stigma oblique-capitate. Fruit dehiscent. KEY TO THE SERIES OF BAUHINIA SECTION AMARIA 1. Fertile stamens 10; petals spreading | oO. | Series Decandrae . Fertile stamens 5; petals strongly imbricate, the corolla forming a pseudotube |. Series Stenanthae |. Bauhinia series Decandrae Wunderlin, K.Larsen & S.S.Larsen, Biol. Skr. 28:14. 1987. TypF: Bauhinia petiolata (S. Mutis ex DC.) Triana ex Hook. f KEY TO BAUHINIA SPECIES IN SERIES DECANDRAE 1, Leaves unlobed (rarely bilobate or bifoliolate in 8. beguinoti 2. Innermost pair of lateral nerves closer to the mid-nerve than to the adjacent lateral nerves 2. Innermost pair of lat than to the mid-n 3. Petals 3.5-4.5 cm Mae fruit 21-27 cm long 3. Petals 1-2 cm long; fruit 8-13 cm Pu 4, Leaves 12-30 cm long,6-13 cm wi inal s| lal tomentose; fruit elliptic-oblong, 3.5-4 cm wide 4, Leaves 7-11 cm long, 3-5.5 cm wide; staminal sheath pilose; gynoecium glabrous or glandular; fruit linear, 1.5-2.5 cm wide 1. Leaves bilobate (rarely bifoliolate or unlobed in 8. beguinotil). 5. Leaves with the innermost pair of lateral nerves closer to the mid-nerve than to the adjacent lateral nerves B. beguinotii 5. Leaves with the innermost pair of lateral nerves equidistant or closer to the ad- jacent lateral nerves than to the mid-nerve. Calyx prominently 5-winged 6, Calyx not winged or only very slightly winged (8. amatlana) B. beguinotii =quidistant or closer to the adjacent lateral nerves i B. petiolata ium B. arborea B. brachycalyx B. geniculata WUNDERLIN 7. Ovary glabrous or glabrat 8. ee yellow; yeaa in asciulat clusters, often appearing before e leaves; Mexico to Costa Ric 8. oe other than yellow; tower in racemes or panicles, appearing well after the leaves; Ecuador o 9, Leaves 1-4 cm long, piicbed to ca. 3/4 the blade length; fruit a 5 cm long; Peru B. cookii weberbaueri 9. Leaves 5-13. cm ae bilobed to ca. 1/2 the blade length; fruit . 25 cm long; Ecuado 10. Corolla white; cial sheath ca. 1/8 the stamen length B. seminarioi 10. Corolla red; staminal sheath 3/5 the stamen length B. haughtii 7, Ovary tomentose or pilose 11. Corolla pink, rose, pre with red or pink at the bas 12. Leaves ovate, 4-10 cm long, 4-7.5 cm wide; carole white with red or pink at the base; Colombia and Panama 12. Leaves suborbicular, 3-4 cm long, 3-4 cm wide; corolla deep pink or rose; Ecuador and Peru 11. Corolla white. 13. Inflorescence and the lower surface of the leaves brown- fe) B. picta . ayabacensis ntose B. seleriana 13. Inflorescence and the lower surface of the leaves other than brown-tomento 14. Leaf blade : ee 5.5 cm long; petals 2-2.7 cm long; Peru B. augusti 14. Leaf blade 0.8-3 cm long; petals ca. 1 cm long; Mexico. 15. Leaf blade 2-3 cm long, bilobed to 1/2 the length B. andrieuxii 15. Leaf blade 0.8-1.5 cm long, bilobed nearly to the base .amatlana Li ae amatlana Wunderlin, sp. nov. (Fig. 1). Type: MEXICO. Oaxaca: ca. L.5 km of San Cristobal Amatlan, ca. 80 km S of Oaxaca and 20 km E of (San Andrés) Miahuatlan ass 16°19.6'N, 96°22'W), ca. 1,500 m, 8 Jun 2000, J. Santiago Martinez s.n. (HOLOTYPE: USF: ISOTYPES: BRIT, F MO, NY). A Bauhinia andrieuxii Hemsley foliis minoribus bilobis prope basin differt. Shrub; branches brown-tomentose when young, soon glabrate. Leaves with the blade chartaceous, 5-7-nerved, broadly suborbicular, broader than long, bilo- bate nearly to the base, 0.8-1.5 cm long, 1.2-2.2 cm wide, the innermost pair of lateral nerves equidistant or closer to the adjacent lateral nerves than to the mid-nerve, the base truncate to slightly cordate, the apex of the lobes rounded, the upper surface glabrous, the lower surface sparsely subappressed, short-pu- bescent, the petiole 4-6 mm long, brown-tomentose; stipules ovate-lanceolate, ca. | mm long, caducous; adpetiolar intrastipular excrescence enlarged, form- ing a flattened subulate projection ca. 1 mm long, the others minute. Inflores- cence terminal, l- to 3-flowered, the peduncle or rachis strigose; floral bud lin- ear-lanceolate, 10-12 mm long, strigose, the apex usually with free calyx lobes BRIT.ORG/SIDA 22(1) UNIVERSITY OF SOUTH FLORIDA HERBARIUM PLANTS OF MEXICO HOLOTYPE Marti 8 UNIVERSITY OF SOUTH FLORIDA HERBARIUM ) Fic. 1. Bauhinia amatlana (holotype, USF WUNDERLIN, ACVIOIVIN UE 101 to 0.5 mm long, sometimes slightly winged and contorted, the pedicel 5-7 mm long; bract and bracteoles lanceolate, ca. 0.5 mm long. Hypanthium turbinate, 1-15 mm long; calyx spathaceous at anthesis; petals white, 10-12 mm long, slightly spreading, short-clawed, glabrous, the blade broadly elliptic, 5-6 mm wide, the margin entire or slightly crisped, the claw 1-2 mm long; fertile sta- mens 10, the outer five 7-8 mm long, the inner ones slightly shorter, the fila- ments short-connate at the base, forming a staminal sheath with laciniate pro- jections extending 1-2 mm above the sheath, the sheath short-pilose on the inner surface, glabrous or glabrate on the outer, the anthers linear-oblong, 1-2 long, sparsely pilose; gynoecium 5-7 mm long, the gynophore, ovary, and style subequal, brown-tomentose. Fruit not seen. Paratype. MEXICO. Oaxaca: ca. 1.5 km NE of San Cristobal Amatlan, ca. 80 km S$ of Oaxaca and 20 km E of (San Andrés) Miahuatlan (ca. 16°19.6'N, 96°22'W), ca. 1,500 m, 29 Aug 1995, J. Santiago Martinez s.n. (USF). Etymology.—The name is derived from the Spanish name of the village San Cristobal Amatlan, this in turn derived from Amatla, the Aztec name for the place of the tree (Ficus sp.) from which the bark is used to make paper. Distribution and ecology.—Mexico (Oaxaca). Known only from the type locality. Deciduous thorn scrub at ca. 1,500 m. Bauhinia amatlana is closely related to B. andrieuxii, also of Oaxaca, but differs by its leaf blade 0.8-1.5 cm long and bilobed nearly to the base (vs. 2-3 cm long and divided only to the middle in B. andrieuxii). 2. Bauhinia andrieuxii Hemsley, Diagn. Pl. Nov. Mexic. 48.1880; Biol. Cent-Amer, Bot. 1:336. 1880. Type: MEXICO. OAXACA: near Oaxaca, - 1834 (FD), G. Andrieux 411 (H¢ LOTYPE: K; IsoTYPES: F-fragment, G, K, P; photo ex G: F MO, US). Shrub or small tree; branches tomentose when young, soon glabrate. Leaves with the blade chartaceous, 7-9-nerved, suborbicular or broader than long, bilobate tol/2 the length, 2-3cm long, 2.5-3cm wide, the innermost pair of lateral nerves equidistant or closer to the adjacent lateral nerves than to the mid-nerve, the base cordate to rounded, the apex of the lobes rounded, the upper surface gla- brous, the lower surface tomentellous, the petiole 5-15 mm long, tomentellous; stipules ovate, ca. 1 mm long, caducous; adpetiolar intrastipular excrescence enlarged, forming a subulate projection ca. | mm long, the others minute. Inflo- rescence terminal, racemose, 3- to 8-flowered, the rachis strigose to tomentellous; floral bud linear-lanceolate, 12-18 mm long, strigose, the apex with free calyx lobes ca. 1.5mm long, the pedicel ca. 5 mm long; bract and bracteoles lanceolate, ca. 1.5mm long. Hypanthium cupulate, 2-3 mm long; calyx spathaceous at an- thesis; petals white, ca. 1 cm long, slightly spreading, subsessile, glabrous, the blade broadly elliptic, 4-6 mm wide, the apex rounded, the margin entire; fer- tile stamens 10, the outer five ca. 1 cm long, the inner ones ca. 5 mm long, the filaments unequally connate near the base for 1-3 mm, glabrous, the anthers 102 BRIT.ORG/SIDA 22(1) linear-oblong, ca. 2 mm long on the longer stamens, ca. 1.5 mm long on the shorter ones, glabrous; gynoecium ca. | cm long, the gynophore, ovary, and style subequal, tomentose. Fruit linear, 8-10 cm long, ca. 1.5 cm wide, apiculate with the persistent style, dark brown, tomentose, the gynophore 5-10 mm long; seeds not seen. Distribution and ecology.— Western Mexico (Morelos, Oaxaca, and Sinaloa). Deciduous thorn scrub at elevations from near sea level to ca. 1,700 m. Bauhinia andrieuxtii is closely related to B. amatlana, also of Oaxaca, but differs by its leaf blade 2-3 cm long, bilobed to the middle (vs. 0.8-1.5 cm long, bilobed sa to the base in B. amatlana). eae Specimens examined. MEXICO. Morelos: Canon de Lobos, 14 km on Cuernavaca-Yautepec road, Fryxell 764 (MEXU);, Canon de Lobos, Vazquez 1776 (MEXU). Oaxaca: without precise locality, Andrieux 371(G, P); Monte Alban, Rose & Hough 4590; Mitla, Smith 107 (US); ruins at Monte Alban, Sousa et dl. 6077 (kK, MEXU, MO, WIS); 3 km NE of La Ollaga, Téllez & Magallanes 233 (kK). Sinaloa: Cerro Llano Redondo, W of Caimanero, Gentry 7018 (NY). 3. Bauhinia arborea Wunderlin, sp. nov. (Fig. 2). ee ee BDOR Napo: Anangu, Parque Nacional Yasuni, 260-350 m, 30 May-21 Jun 1982 (fD, SE li lorean Forest ]8935 (HOLOTYPE: AAU; ISOTYPES: AAU, N A Bauhinia brachycalyx Ducke foliis majoribus, fructibus latioribus, staminum vagina glabra, nec ] LaF, 10N g\ Tree to 30 m; branches glabrous. Leaves with the blade coriaceous, (3-)5-7- nerved, elliptic-oblong, unlobed, 12-30 cm long, 6-13 cm wide, the innermost pair of lateral nerves equidistant or closer to the adjacent lateral nerves than to the mid-nerve, the base rounded to obtuse, the apex abruptly short-caudate, the upper surface glabrous, the lower surface minutely strigulose, slightly lighter in color than the upper surface, the petiole 2-4.5 cm long, glabrous; stipules broadly lanceolate, ca. 1 mm long, caducous; adpetiolar intrastipular excres- cence enlarged, forming a laterally flattened, obtuse projection to | mm long, the others minute. Inflorescence subterminal, short-racemose, 10- to 18-flow- ered, the flowers usually paired, the rachis strigose; floral bud elliptic-lanceolate, 1.5-2 cm long, strigose, the apex acute, the pedicel 1-3 mm long; bract and bracteoles triangular-ovate, ca. | mm long. Hypanthium short-tubular, 7-10 mm long; calyx spathaceous at anthesis; petals white, 12-15 mm long, slightly spreading, sessile, glabrous, the blade oblanceolate, 5-7 mm wide, the apex ob- ‘ive 12-14 mm —— tuse, the margin slightly crisped; fertile stamens 10, the outer long, the inner ones slightly shorter, the filaments free to the base or very short- connate, the anthers lanceolate, 4-6 mm long, sparsely short-pilose on the dor- sal surface; gynoecium ca. 8 mm long, the gynophore and style ca. 2 mm long, tomentose, the ovary ca. 5 mim long, tomentose. Fruit elliptic-oblong, ca. 13 ¢m long, 3-4 cm wide, apiculate with the persistent style base, brown, glabrous, C — — ae gynop hore 5-8 cm long; seeds not seen. WUNDERLIN, REVISION OF 103 A STUDIES OF ECUADOREAN FORESTS ; ORR, é IX Se Baul rea rhi mu ’ F * Richard P. Wu a University of South Florida = ie , = aan waded Co Duta The New ark Se L985 s by Crarina A nad, axi tow Botancal (nathyie, Unwerety of ABrhus Fic. 2. Bauhinia arborea (isotype, NY). BRIT.ORG/SIDA 22(1) Pararyers. ECUADOR. Morona-Santiago: 15 kim Nol Taisha(02°32'S.77°43W), Ortega 96 (USF). Napo: Reserva Etnica Huaorani, highway and pipeline “Maxus” under construction (O0°51'S, 76°26'W), Aulestia & Andi 938 (MO, USF); Parque Nacional Yasuni, trail toward Rio Daymi (00°52'S, 76°5'W), Ceron 3371 (MO, USF); ca. 30 km NNW of Coca, Rio Huashito, Pennington 10592 (kK); Anangu, Parque Nacional Y: i 31-32'S, 76°23'W), SEF 8762 (AAV), SEF 8834 (AAU), SEF 8867 (AAU), SEF 8922 _ago Agrio Canton, Reserva Cuyabeno, Tarapoa-Tipischa, Rio (Ay a SEF 10235( AAU). Sucumbios: | C ossing (76°15'W, 00°00'S), Palacios etal. 8917 (USF). } Etymology.—The name is derived from the tree habit remarkable for its size within section Amaria in Ecuador. Distribution and ecology.—Eastern Ecuador (Napo, Morona-Santiago, and Sucum bios). Rain forest on well-drained hilly ground at 250-450 m. The species is most closely related to B. brachycalyx but differs by having larger leaves, wider fruits, a glabrous staminal sheath, and a tomentose gyno- ecium. Bauhinia arborea is a tree to 30 m while B. brachycalyx is a shrub or small tree to 6 m, rarely to 15m. 4. Bauhinia augusti |larms, Repert. Spec. Nov. Regni Veg. 18:234. 1922. Type: PERU, RCA: near the confluence of the Rio Chinchipe and Rio Maranon, 700-800 m, | May CAJAMAR roTyPe: F here designated to replace destroyed B holotype: 1912 GD, A. Weberbauer 6224 (LEC ISOLECTOTYPES: F-fragment, G, NY, US; photo ex B: ER MO, NY, US). Shrub to 2m; branches pubescent when young, soon becoming glabrate. Leaves with the blade chartaceous, 7-9-nerved, broadly ovate, bilobed to ca. 1/2 the length, 3.5-5.5 cm long, 3-5 cm wide, the innermost pair of lateral nerves equi- distant or closer to the adjacent lateral nerves than to the mid-nerve, the base cordate, the apex of the lobes obtuse, the upper surface glabrous, the lower sur- face short-pilose, especially on the nerves, the petiole 0.5-1.5 cm long, short- detiolar pilose; stipules triangular-lanceolate, ca. 1.5 mm long, caducous; ad intrastipular excrescence occasionally enlarged, forming a subulate projection to 2 mm long, the others minute. Inflorescence terminal or subterminal and axillary, short-racemose or paniculate, 4+- to 12-flowered, the rachis densely short-pilose; floral bud narrowly ellipsoid, slightly arcuate, 2-2.5 cm long, short- pilose, the apex obtuse or acute, the pedicel 0.5-1 cm long; bract and bracteoles triangular-lanceolate, ca. | mm long. Hypanthium short-tubular, +-6 mm long; calyx spathaceous at anthesis; petals white, 2-2.7 cm long, slightly spreading, short-clawed, glabrous, the blade narrowly elliptic-oblanceolate, 0.5-0.7 cm wide, the apex obtuse, the margin slightly crisped, the claw 3-4 mm long; fer- tile stamens 10, the outer five 3-3.5cm long, the inner ones slightly shorter, the filaments short-connate at the base, sparsely pilose, especially toward the base, the anthers elliptic-oblong, ca. 5 mm long, glabrous; gynoecium 3-3.5 cm long, the gynophore, ovary, and style subequal, the ovary densely short-pilose, ne style and the gynophore sparsely mn Fruit not seen. Distribution and ne (Cajamarca). Known only collection. At 700-800 m. — — rom the type WUNDERLIN, REVISION OF BAUHINIA SUBGENUS BAUHINIA SECTION AMARIA ae Bauhinia augusti is related to the new species B. ayabacensis of Ecuador and Peru. Bauhinia augusti differs by its white corolla and lower leaf surface with erect trichomes in contrast to the pink corolla and lower leaf surface with appressed trichomes of B. ayabacensis. 5. Bauhinia ayabacensis Wunderlin, sp. nov. (Fig. 3). Typr: PERU. Piura: on road to Ayabaca, 18 km above Puente Tandopa (Rio Quiroz), 1,700 m, 24 Sep 1964 (fl), PC. Hutchinson & J.K. Wright 6689 (HOLOTYPE: F; IsSOTYPES: K, MO, NY, P. US). A Bauhinia augusti Harms corolla rosea lata, foliorum lamina inferiori trichomatibus brevibus adpressis obsita differt. Shrub or small tree to 4 m; branches pubescent when young, soon becoming glabrate. Leaves with the blade chartaceous, 7(-9)-nerved, suborbicular, bilobed to ca. 1/3 the length, 3-4 cm long, 3-4 cm wide, the innermost pair of lateral nerves equidistant or closer to the adjacent lateral nerves than to the mid-nerve, the base cordate, the apex of the lobes obtuse to rounded, the upper surface glabrous, the lower surface appressed short-puberulent, especially on the nerves, the petiole 1-15 cm long, pubescent; stipules triangular-lanceolate, ca. 1.5 mm long, caducous; adpetiolar intrastipular excrescence occasionally enlarged, forming a subulate projection to 2 mm long, the others minute. Inflorescence terminal or subterminal and axillary, short-racemose, 2- to 10-flowered, the rachis brown-tomentose; floral bud narrowly ellipsoid, slightly arcuate, 2-2.5 cm long, brown-tomentose, the apex acute, the pedicel 6-8 mm long; bract and bracteoles triangular-lanceolate, ca.1 mm long. Hypanthium short-tubular, 4- 6 mm long; calyx spathaceous at anthesis; petals deep pink or rose, 2.5-3 cm long, spreading, short-clawed, glabrous, the blade narrowly elliptic-obovate, l- L.5cm wide, the apex apiculate, the margins slightly crisped, the claw 3-4 mm long; fertile stamens 10, the outer five 2-2.5cm long, the inner ones 1-2 cm long, the filaments short-connate at the base, sparsely pilose, the anthers elliptic- oblong, ca. 5mm long, glabrous; gynoecium 2-2.5cm long, the gynophore, ovary, and style subequal, the gynophore and ovary tomentose, the style sparsely pi- lose. Fruit linear-elliptic, ca. 6 cm long, ca. 1.7 cm wide, apiculate with the per- sistent style base, brown, strigose to glabrate, the gynophore ca. lcm long; seeds ovate-reniform, ca. | cm long, 7-8 mm wide, the surface dark brown, dull, the funicular aril-lobe scars unequal, 1-3 mm long. Paratypes. ECUADOR. Loja: 5 km E of Celica on road to El Empalme, Harling & Andersson 18183 (GB, MO); El Empalme-Celica road, ca. 5 km above El Empalme, Harling & Andersson 18221 (MO). PERU. Cajamarca: on the crossing to San Luis, Cabanillas & Guevara 509 (F); El Camote, Carretera Chilete- Contumaza Sdagdstegui & Lopez 7636 (F MO, USF); between Rupe and Contumaza Sagdstegui et al. 9817 (MO); near San Pablo, Sagdstegui et al. 15380 (F); Caserio de Chilili, Sanchez 200 (F); between Chilete and Rupe, E of the Chilete-Contumaza highway, Sdnchez 4222 (F). Etymology.—The name derived from the town of Ayabaca (Province Ayabaca) near the type locality. BRIT.ORG/SIDA 22(1) UNIVERSITY OF CALIFORNIA Director, Paul C. Hutchison Additional field sores for fag and 3m. in diame : Tre r. Flowers deap pink. Irwin, 1966, comments ae description \ gives color as white." Bauhinia sages std Harm dupl. de STiadin; wes HOLOTYP Sie ayabacensis Wunderlin Dry open hillsides, scattered brash On road to Ayabaca, R4199r 18 km. : 1641236 rch. Wen ie en niversity of South Florida 1982 Biovioce of iArabaes: ficschae Hi ae FIELD MUSEUM OF Pau C. Hutcuison 7, NATURAL HISTORY Collection number 6689 of veh po on 1, UCy J. Kenneth Wr T 24 September 1964 determiners set, NY s Seaees USM, US, F, NY, MO, K, C MICH, M, P, LE, Fic. 3. Bauhinia ayabacensis (holotype, F) WUNDERLIN, REVISIUN UF 107 Distribution and ecology.—Ecuador (Loja) and Peru (Cajamarca and Piura). Dry, open hillsides in scattered brush or seasonal evergreen forest at 1,200-2,400 m. Bauhinia ayabacensis is closely related to B. augusti from which it differs by its wider deep pink- or rose-colored corolla (vs. white in B. augusti) and the lower leaf surface with appressed trichomes (vs. short pilose in B. augusti). The petals of B. ayabacensis are 1-15 cm wide, while those of B. dugusti are 0.5-0.7 cm wide. 6. Bauhinia beguinotii Cufod., Arch. Bot. Sist. 9:192. 1933. Type: COSTA RICA. LIMON: between Waldeck and Rio Barbilla, 40 m, 12 May 1930 (fD, G. Cufodontis 664 (HOLOTYPE: W, not seen; photo ex W; ISOTYPES: F MO, US). Shrub or small tree to 6(-10) m; branches brown-tomentose to -tomentellous when young, soon becoming glabrate. Leaves with the blade chartaceous, 7-9- nerved, oblong-ovate, unlobed or slightly bilobate (deeply bilobate or bifoliolate in var. gorgonae), 10-24 cm long, 5-16 cm wide, the innermost pair of lateral nerves closer to the mid-nerve than to the adjacent lateral nerves, the base rounded to truncate, the apex Gif the leaf unlobed) or the lobes (if the leaf bi- lobed or bifoliolate) acuminate to caudate, the upper surface glabrous, the lower surface brown-tomentellous to glabrate, the petiole 1.5-3.5 cm long, glabrate; stipules broadly ovate, 1-1.5 mm long, caducous; adpetiolar intrastipular ex- crescence occasionally enlarged, forming a subulate projection to 1 mm long, the apex glandular, the others minute. Inflorescence terminal or subterminal and axillary, racemose, 5-10-flowered, the rachis brown-tomentose to - tomentellous; floral bud elliptic-lanceolate, 1.5-2 cm long, brown-tomentose, the apex with minute free calyx lobes, the pedicel 2-5 mm long; bract and bracteoles ovate, ca. 1 mm long. Hypanthium short-tubular, 5-6 mm long; ca- lyx spathaceous at anthesis; petals white, 2.5-3 cm long, spreading, short-clawed, glabrous, the blade oblanceolate, 8-13 mm wide, rounded, the margin entire, the claw 3-5 mm long; fertile stamens 10, the outer five 2-3 cm long, the inner ones shorter, the filaments connate for ca. 1/2 the length of the shorter stamens, sparsely pilose near the base, the anthers triangular at anthesis, ca. 3 mm long on the longer stamens, ca. 2 mm long on the shorter ones, sparsely pilose to glabrate; gynoecium 2-3 cm long, the gynophore, ovary, and style subequal, brown-tomentose. Fruit linear, 12-15 cm long, ca. 2 cm wide, apiculate with the persistent style, brown, minutely strigose to glabrate, the gynophore 1-1.5 cm long; mature seeds not seen. KEY TO THE VARIETIES OF BAUHINIA BEGUINOTII 1. Leaves unlobed or only slightly bilobate; Costa Rica and Panama var. beguinotii 1. Leaves deeply bilobate or bifoliolate; Colombia var. gorgonae 108 BRIT.ORG/SIDA 22(1) 6a. Bauhinia beguinotii var. beguinotii Distribution and ecology.—Costa Rica (Limon) and Panama (Bocas del Toro, Panama, and San Blas). Tropical evergreen forest from near sea level to ca. 500 m. Specimens examined. COSTA RICA. Limon: drainage of the Rio Estrilla Valley, Shank & Molina 4466 (fF, US). PANAMA. Bocas del Toro: Cerro Bracha, NE of the encampment Changuinola | of Corriente Grande (IRHE), Correa et al. 3227 (PMA, USF); SE and NE of the encampment Changuinola | of the IREIE, Correa et al. 3316 (PMA, USF). Panama: 14.8 kin N of the Pan-American Highway, Folsom & Maas 5202 (MO); Carti road, 7 mi from turnoff at El Llano (9°15'N, 78°50'W), Hammel 13547 (MO. USF): 10-12 km N of Pan-American Highway, Mori & Kallunki 2847 (MO), El Llano-Carti-Tupile road, 16 km N of Pan-American Highway at El Llano, Nee 9362 (MO): ca. 10 km N of Pan-American Highway, Nee etal. 8881(MO). San Blas: FE] Llano-Carti road (09°20'N, 79°O'W), Hamilton & Stockwell COCK WELL 1089 (MO, USF): El Llano-Carti road, 19.1 km (09°19'N, 78°55'W ). de Nevers 5946 (MO, USF): ena along continental nes on E L L ke Carti sauce a“ LO'N, 78°15'W), de Nevers & Pérez 3701 ( JSF) ). Nusagandi | g the E aes Carti road (O9°19'N, 78°15'W), de — & Pérez 3582; Nusagandi g tl tal d El Llano-Carti road (09°1L9'N, 78°15'W), de Nevers & Pérez 3781 (MO, USP): Nusangandi ridge running NW from Punta Mama, (09°19'N, 78°15S'W), de Nevers et al. 3728 (MO, USF); El Llano-Carti road, 19.1 km from Interamerican | (O9°19'N, 78°55'W), de Nevers et al. 4957 (MO, USF); El Llano-Carti road, near Nusagandi, aor Sendero Nusagandi, W of buildings (09°15'N, 79°O'W), McPherson 11033 (MO, USF). 6b. Bauhinia beguinotii var. gorgonae (Killip ex R.S. Cowan) Wunderlin, Ann. Missouri Bot. Gard. 60:570. 1973. Bauhinia gorgonde Killip ex R.S. Cowan, Bol. Soc. Venez. Ci. Nat. 22:281. 196L. Type: COLOMBIA. NARINO: E side of Isla Gorgona, 50-100 m, 11 Feb 1939 ({D, EP. Killip & H. Garcia-Barriga 33170 (HOLOTYPE: US; ISOTYPES: COL, not seen, BM, US). Distribution and ecology.—Colombia (Narino). Tropical evergreen forest from near sea level up to 100 mm. The Colombian material is recognized at the varietal level and is distinquished from the Panamanian and Costa Rican material only in having the leaves deeply bilobate or bifoliolate rather than unlobed or only slightly bilobate. Since the degree of lobing of leaves in Bauhinia is often quite variable within species, further collections may prove var. gorgonde to be unworthy of recognition. Specimens examined. COLOMBIA. Narino: Isla Gorgona, Cheeseman [St. George Expedition] 380 (F K, US), Collenette [St. George Expedition] 628 (EK, US), Kelsall [St. George Expedition] 360 (K, US). 7. Bauhinia brachycalyx Ducke, Trop. Woods 90:12. 1947. Typr: BRAZIL. AMAZONAS: region of the Rio Purus near the mouth of the Rio Pauini [based on plant cultivated in the Horto Botanico do Museu Goeldi, Belém, Para, specimen made 30 Dec 1946 (I1)|, A. Ducke 2045 (HOLOTYPE: MG, not seen; IsOTYPES: NY, US). Shrub or small tree to 6(-15) m; branches glabrous. Leaves with the blade sub- coriaceous, 5-nerved, oblong-ovate, unlobed, 7-11 cm long, 3-5.5 cm wide, the innermost pair of lateral nerves equidistant or closer to the adjacent lateral nerves than to the mid-nerve, the base rounded to subtruncate, the apex witha caudate tip 1-2 cm long, the upper surface glabrous, the lower surface glabrous, — WUNDERLIN, REVISION Ur 109 the petiole 0.8-1.4 cm long, glabrous; stipules ovate, ca. | mm long, caducous; adpetiolar intrastipular excrescence often enlarged, forming a subulate projec- tion to 2 mm long, the others minute. Inflorescence terminal or subterminal and axillary, short-racemose, 6- to 12-flowered, the flowers usually paired, the rachis minutely puberulent; floral bud elliptic-oblanceolate, ca. 1.5 cm long, minutely puberulent, the apex apiculate, the pedicel 3-4 mm long; bract and bracteoles ovate-lanceolate, ca. | mm long. Hypanthium short-tubular, 4-6 mm long; calyx spathaceous at anthesis; petals white, ca. 1.5(-2) cm long, slightly spreading, subsessile, glabrous, the blade linear-oblanceolate, 4-5 mm wide, the apex acute, the margins slightly crisped; fertile stamens 10, the outer five 8-10 mm long, the inner ones ca. 1/2 as long, the filaments short-connate at the base, forming a staminal sheath with a laciniate projection extending 1-2 mm above the sheath, the sheath tomentose on the inner surface, glabrous on the outer, the anthers linear-oblong, ca. 3 mm long, sparsely pilose at the apex and the base; gynoecium 12-15 mm long, subequaling the androecium, arcuate, the ovary 5-6 mm long, the gynophore and the style 3-4 mm long, subequal, the style and the upper portion of the ovary with glandular trichomes in lines, the gynophore gla- brous, the stigma oblique. Fruit linear, 10-12 cm long, 1.5-2.5 cm wide, apiculate with the persistent style, brown, glabrous, the gynophore ca. 0.5 mm long; seeds suborbicular, 11-12 mm long, 9-10 mm wide, dark brown, the surface dull, the funicular aril-lobe scars subequal, extending ca. 1/2 way around the seed. Distribution and ecology.—Ecuador (Napo and Sucumbios), Peru (Amazonas, Loreto, and Pasco), and Brazil (Amazonas). In forest on well drained ie also often on inundated clay soil and along rivers at 200 m. pecimens examined. ECUADOR. Morona-Santiago: oil well “Garza” of Tenneco, ca. 35 km NE of Mere Ivo (01°49'S, 76°42'W), Zak & yee als 4618 (MO, USF) Napo: road construction on the Yuca Road, ca. 3 km from the Auca oil-field Road (ca. 00°31'S, 76°58'W,) Brandbyge et al. 30287 (AAU, USF); Parque Nacional Yasuni, oil well “bogi” of Conoco (00°43'S, 76°28W), Coello 225 (MO, USF); Anangu, N side of Rio Napo, trail from village to laguna (00°31'S, 76°23'W), Lawesson et al. 39571 (AAU, USF); Anangu, near the outlet of Rio Anangu into Be a in pus vues Nacional Yasuni, SEF 10133 (AAU); Anangu, near the outlet of Rio At | Yasuni, SEF 10389 (AAU). Sucumbios: Rio Gueppi, tributary ee Rio preys above Beauv border post of Puerto Pert, 8 km above mouth of river, Gentry et al. 21805 (MO, USF); Lago Agrio Canton Reserva Faunistica Cuyabeno, Rio Lagarto, on border with Peru (00°33'S, 75°16'W), Palacios et al. 8055 (MO, USF). PERU. Amazonas: Yamayakat, Quebrada Kusu-Chapi, Rio Maranon (00°55'S, 78°19W), Vasquez et al. 20104 (MO, USF). Loreto: Chambira, near Yurimaguas, Rio Huallaga Valley, Ferreyra 4952 (NY, US); Rio Gueppi, tributary of Rio Putumayo, northernmost tip of Peru on border with Ecuador, be- tween mouth of River and the border a of Puerto Pertica. 8 km upriver, Gentry et al. 21772 (MO, USF); trail from Indi iana on Rio Amazonas to Rio Napo, Gentry et al. 22188 (MO, USF); Rio Nanay on the road tothe M f Be ne rista, Rimachi 2800 (MO); San Miguel (Quebrada Tamshiyacu) ae a 72°40'W), Vasquez & Jaramillo 7153 (MO, USF); unos) saeaee a Suerte (Rio Itaya), Vasquez & 8418(MO, USF): Indiana, Yanamono, Rio A S,72°50'W), Vasquez & Jaramillo ‘11103 (MO, USP): Indiana, Yanamono, Explorama Lodge (03°28'S, 72°50 W), Vasquez et al. 12113 (MO USF). Pasco: Palcazu Valley, Iscozacin (10°12'S, 75°15'W), Foster 9463 (MO, USF). BRAZIL. Amazonas: Rio Javari, Miraflor, Lleras et al. [Prance] P16927 (K, MO, US) 110 BRIT.ORG/SIDA 22 1) 8. Bauhinia cookii Rose in Britton @ Rose, N. Amer. FL. 23:205. 1930. Type: COSTA RICA, ALAJUFLA: Cuesta de la Vieja, road to San Carlos, 300 m, 11 Apr 1903 (1D, O.F Cook & CB. Doyle 41 (HOLOTYPE: US; IsoTYPES: NY-fragment, US Casparia calderonii Rose in Britton & Rose, N. Amer. Fl. 23:217.1930. Ba ‘alderonii (Ros se) Lundell, pollen 1214. 1937. Type: EL SALVADOR, SONSONATE: San Julian, 1924 (Ir), S Calderén 2226 (HOLOTYPE: US; ISOTYPES: NY-fragment, US). Bauhinia gigas L oT Lyra 1:213. 1937. TYPE: ee Cayo: Valentin, Jun-Jul 1936 (st), CL. Lundell MICH; IsoTyPEs: NY, US). Tree to 15(-40) m; branches strigose or puberulent when young, soon becom- ing glabrate. Leaves with the blade submembranaceous or chartaceous, 7-9- nerved, broadly ovate, bilobate to ca. 1/2 the length, 5-10(-15) cm long, 7-10(- 12) cm wide, the innermost pair of lateral nerves equidistant or closer to the adjacent lateral nerves than to the mid-nerve, the base cordate to truncate, the apex of the lobes acute to obtuse, the upper surface sent the lower surface strigose, at least on the nerves, the petiole 2-3(-5) cm long, slightly strigose; stipules lanceolate, ca. 1.5 mm long, caducous; adpetiolar intrastipular excres- cence enlarged, forming a subulate projection to 1.5 mm long, the others minute. Inflorescence paniculate, congested into a fasciculate cluster, sometimes pro- duced before the leaves and appearing cauliflorous or ramiflorous, 15- to 60- flowered, the rachis strigose; floral bud linear-elliptic, 8-10 mm long, strigose, the apex with minute free calyx lobes, the pedicel 5-7 mm long, strigose; bract and bracteoles ovate-lanceolate, ca. | mm long. Hypanthium cupulate, 2-3 mm long; calyx spathaceous at anthesis; petals light sulfur-yellow, 10-15 mm long, spreading, sessile or short-clawed, glabrate, the blade elliptic to oblanceolate, 5-7 mm wide, the apex obtuse, the margin entire; fertile stamens 10, the outer five LO-15 mm long, the inner ones 5-10 mm long, the filaments connate at the base linear-oblong, ca. 2 mm long on the longer stamens, ca. 1.5 mm long on the or ca. L mm, with a laciniate rim projecting above, glabrate, the anthers shorter ones; gynoecium 5-10 mm long, the gynophore, ovary, and style subequal. Fruit linear, 5-7 cm long, ca. 1.5 cm wide, apiculate with the persis- tent style, light brown, See the gynophore 5-10 mm long; seeds obovate, 7-10mm ee 5-8 mm wide, the surface dull, dark brown, the funicular branch scars equal, ca. 4/5 the length of the seed. Distribution and ecology.—Western Mexico in Sinaloa southeast through Mexico and Central America to Costa Rica. Not known from Nicaragua, but to be expected there. Tropical deciduous or semideciduous forests at elevations from near sea level to ca. 4,200 m, but more commonly 300-1,000 m. It appar- ently flowers throughout the year, but usually from March to July. Flowers are sometimes produced before the leaves appear or are much developed. The species is unusual not only in section Amaria by having yellow flow- — —_ ers, but in fact, no other New World species of subgenus Bauhinia has flowers of this color. WUNDERLIN, REVISION Ur 111 Specimens examined. MEXICO. Chiapas: vicinity of La Chacona, Miranda 6485 (US). Colima: vicin- ity of Manzanillo, Ferris 6022 (A, F US); ca. 15 mi SSW of Colima on Manzanillo road (3 mi above Terolapa), Mc Vaugh 15541 (MICH); ca. 14 mi WNW of Santiago on road to Cihuatlan, oe isco, Mc a 20776 (G, MICH, US). Guerrero: Canon de la Mano Negra, near Iguala, Rose et al. 9344 (F GH, NY, US). Jali ween Bahia Navidad and La Manzanilla on Bahia Tenacatita, 3 mi W of ee hae road, McVaugh a 007 (MICH); near Playa de Cuastecomate, 8 km by road NW of Navidad, McVaugh & Koelz 1674 (MICH). Nayarit: 7-8 mi W of Compostela along road to Mazatlan, Gentry & Gilly 10848 (DUKE, MEXU, TEX, WIS); 6-7 mi S of Compostela, McVaugh 18753 (G, MICH, US). Oaxaca: Ejido Benito Juarez, Sebastopol, Tuxtepec, Dioscorea Brigade 2741 (MEXU). Sinaloa: Mazatlan, Reko 4520 (MEXU, US). Veracruz: Barranca de ae Purpus 8797 (UC), Purpus 8896 (GH, MO, NY, UC, Uo) near Rancho Remundadero, Purpus 11068 (NY, US), Purpus 11069 (A), Purpus 11170 (MO); Municipio Puente Nacional, La Ceiba, Ventura 12760 (MEXU). GUATEMALA. Escuintla: Hacienda ae Salas 1406 (F); Rio Guncalate Standley 60187 (F, US). Guatemala: without precise locality, Aquilar 610 (F). Retathuleu: Rio Talculan, 5 mi W of Retalhuleu, Standley 87377 (F). Suchitepéquez: along Rio Madre Vieja Joa oe ae 62207 (F). HONDURAS. Comayagua: Pito Solo, Lake Yojoa, Edwards P-404 (Fk, US). EL DOR. Ahuachapan: Finca San Benito, near Rio Guayapa (13°49'N, 89°56'W), aoe et al. oon U sea Finca San Bene alone Rio Nacimiento del Guayapa (13°5 EN, 89°55'W), Witsberger Nts COSTA RICA. Al etween Canas and Upala, 10 km N of Pyaepe Croat 36478 (MO, US nee of the Rio Sue aes Pittier 16695 (US). San José: Cerro Bijaqualito-Tarcoles, Poveda 1097 es 9. Bauhinia geniculata Wunderlin, sp. nov. (Fig. 4). Type COLOMBIA. RISARALDA: Haci- enda Alejandria, 6 km on highway between La Virginia and Cerrito, low hills of the extreme north of the wide part of the Valle del Rio Cauca, ca. 940 m, 27 May 1989 (f1), PA. Silverstone- Sopkin, J. Giraldo, & M. Cabrera 5270 (HOLOTYPE: CUVC; ISOTYPE: USF) A Bauhinia petiolata (DC.) Triana ex Hooker f. foliis lobatis, calyce alto, nec non androecii forma singulari differt. Tree to 5 m; branches glabrous. Leaves with the blade coriaceous, 7-9-nerved, oblong-ovate, bilobed to ca. 2/3 the length, 14-26 cm long, 9.5-17 cm wide, the innermost pair of lateral nerves equidistant or closer to the adjacent lateral nerves than to the mid-nerve, the base shallowly cordate to subtruncate, the apex of the lobes acuminate, the upper surface inconspicuously reticulate, gla- brous, the lower surface evidently reticulate, with inconspicuous, colorless, medially attached, biradiate, glandular trichomes, minutely strigose, at least on the veins near the base, the petiole 2-3(-4) cm long, glabrous; stipules trian- gular-ovate to lanceolate, 1-1.5 mm long, caducous; adpetiolar intrastipular ex- crescence sometimes enlarged and forming a laterally flattened, blunt projec- tion to 2 mm long, others minute. Inflorescence terminal or subterminal and axillary, racemose, 2- to 10-flowered, the flowers solitary or paired, the rachis reddish brown-tomentose; floral bud linear-clavate, 4-4.5 cm long, sparsely brown-tomentose, evidently 10-nerved, becoming prominently 5-winged above the hypanthium with the wings gradually enlarging toward the apex to 4mm, the pedicel ca. 1.5cm long; bract and bracteoles triangular-ovate, ca. 1 mm long. Hypanthium short-tubular, L.5-2.2 cm long; calyx green, spathaceous at anthe- sis, each sepal with a thickened nectiferous area ca. 5-7 mm long and wide near the base; petals white, 3.5-4.5 cm long, slightly spreading, subsessile, glabrous, 112 BRIT.ORG/SIDA 22(1) UNIVERSITY OF SOUTH FLORIDA HERBARIUM k Fic. 4. Bauhinia geniculata (isotype, USF). WUNDERLIN, ACVIOIUIN UE 113 the blade oblanceolate, 15-18 mm wide, gradually tapering to the thickened midrib at the base, the apex rounded to slightly g : fertile stamens 10, the outer five 4-4.5 cm long, the inner ones shorter, the filaments connate at the base for 2-3 mm and forming a staminal sheath, the uppermost stamen of the inner whorl the shortest and strongly geniculate near the point of attach- ment near the apex of the shield-like portion of the staminal sheath, the stami- nal sheath with a ligulate projection ca. 1mm long behind the lower 3 stamens of the outer whorl and with a minute projection behind the lower 2 stamens of the inner whorl, the sheath reddish tomentose on the inner surface, the anthers linear, 7-8 mm long, glabrous; gynoecium 3.5-4.5 cm, the gynophore ca. 1.5 cm long, reddish tomentose, the ovary ca. 2 cm long, reddish tomentose, the style ca. 15cm long, reddish tomentellous to glabrate near the apex. Fruit linear, 29- 41 cm long, 2.8-3.3 cm wide, apiculate with the persistent style base, brown, puberulent, the gynophore 2.5-4 cm long; seeds suborbicular to oblong, strongly compressed, 17-18 mm long, 12-15 mm wide, the surface dull brown, the fu- nicular scars subequal, extending 2/3 to nearly the length of the seed. ParatyPe. COLOMBIA. Risaralda: Hacienda Alejandria, 6 km on highway between La Virginia and Cerrito, low hills of the extreme north of the wide part of the Valle del Rio Cauca, Silverstone-Sopkin et al. 5809 (CUVC, USF). Etymology—The name derived from the bent uppermost stamen of the inner staminal whor Distribution and ecology—Colombia (Risaralda). Primary forest at 900-940 m. Bauhinia geniculata is most closely related to B. petiolata, but differs by its lobed leaves (vs. unlobed in B. petiolata), winged calyx (vs. not winged in B. petiolata), and unique androecial morphology. 10. Bauhinia haughtii Wunderlin, Brittonia 35:337. 1983. Type: ECUADOR. MANaBi: Bella Vista, SE of Santa Ana, 100 m, 5 Sep 1942 (f1, fr), O. Haught 348] (HOLOTYPE: US; ISOTYPES: Shrub or tree to 10 m; branches glabrous. Leaves with the blade submembrana- ceous, 9(-11)-nerved, suborbicular, bilobed to ca. 1/2 the length, 5-8 cm long, 6- 10 cm wide, the innermost pair of lateral nerves equidistant or closer to the adjacent lateral nerves than to the mid-nerve, the base deeply cordate, the apex of the lobes obtuse to rounded, the upper surface aac the lower surface minutely strigose, the petiole 3-5 cm long, glal iangular-lanceolate, ca. l mm long, caducous; intrastipular ¢ excrescences ‘minute. Inflorescence ter- minal or subterminal and axillary, racemose, 6- to 10-flowered, the flowers usually paired, the rachis minutely strigulose-tomentose; floral bud narrowly elliptic, 5-6 cm long, minutely strigulose-tomentose, red, the apex attenuate, the pedicel 2-5 mm long: bract and bracteoles triangular-lanceolate, ca. 1 mm long. Hy- panthium tubular, 2-2.5cm long, slightly arcuate; calyx spathaceous at anthe- sis; petals red, 4.5-5cm long, slightly spreading, short-clawed, glabrous, the blade 114 BRIT.ORG/SIDA 22(1) oblanceolate, 1-1.5 cm wide, the apex rounded, the margin entire, the claw 3-4 mim long; fertile stamens LO, red, the outer five 4.5-5.5 cm long, the inner ones slightly shorter, 9 connate into a staminal tube for ca. 3/5 their length, the other one free to ca. 0.5 cm lower and slightly longer, the staminal sheath sparsely short pilose on the inner surface, the anthers elliptic-oblong, ca. 5 mm long, glabrous; gynoecium 6.5-8.5 cm long, the gynophore, ovary, and style subequal, glabrous. Fruit linear, 16 cm long, 1.7 em wide, apiculate with the persistent style, light brown, glabrous, the gynophore 4.5 cm long: seeds not seen. Distribution and ecology.—Ecuador (Manabi and Pichincha). In tropical wet forest at L1OO-300 m. Bauhinia haughtii is a distinctive species most closely related to B. seminarioi, also endemic to Manabi, Ecuador. It differs from that species by its red flowers (vs. white in B. seminarioi) and a much longer staminal sheath (ca. 3.5.cm vs. ca.6 mm in B. seminarioi). The only other red-flowered species in Ecuador, B. stenantha, is readily distinguished by its five fertile stamens (vs. ten) and smaller flowers (2.5-3.0 cm long) and smaller fruits (6-10 cm long). Specimen examined. ECUADOR. Manabi: 170-175 km on road between Santa Domingo and Quininde, Acosta-Solis 13653 (F). 11. Bauhinia petiolata (DC.) Hook. f., Bot. Mag. 103: tab. 6277. 1877. Amaria petiolata DC., Prodr, 2: 519. 1825. Type: COLOMBIA: without precise locality, 1760-1808, 5. Mutis 2398 (HOLOTYPE: MA, not seen; ISOTYPE: US). Amaria sessilifolia DC, Prodr. 2:519, 1825. Bauhinia sessilifolia (DC.) Quinones, Ann. Missouri Bot. Gard. 75:1156. 1988. TyPE: COLOMBIA: without precise locality, 1760-1808, S. Mutis 2724 (HOLOTYPE: MA, not seen; Isotype: US-fragment). ane Shrub or tree to 15 m; branches glabrous. Leaves with the blade chartaceous to coriaceous, 5-nerved, ovate, unlobed, 8-14 cm long, 4-8 cm wide, the innermost pair of lateral nerves equidistant or closer to the adjacent lateral nerves than to the mid-nerve, the base rounded to deeply cordate, the apex acute, sometimes with a caudate tip to 3. cm long, the upper surface glabrous, the lower surface glabrous, often glaucous, conspicuously reticulate-nerved, the petiole 0.5-3.5 cm long, glabrous; stipules triangular, ca. 1 mm long, caducous; adpetiolar intrastipular excrescence often enlarged and forming a blunt subulate projec- tion ca. | mm long, the others minute. Inflorescence terminal or subterminal and axillary, short-racemose, 4- to 8-flowered, the rachis sparsely strigose- tomentellous to glabrate; floral bud elliptic-ovate, 2-3 cm long, strigulose-to- mentose, the apex attenuate, glabrous, the pedicel to 1 cm long; bract and bracteoles triangular-ovate, ca. 1 mm long. Hypanthium short-tubular, ca. 1.5 cm long; calyx spathaceous at anthesis; petals white, 3.5-4.5 cm long, slightly spreading, short-clawed, glabrous, the blade narrowly oblanceolate, 1.2-1.5 cm wide, the apex rounded, the margin slightly crisped, the claw 2-3 mm long: tertile stamens 10, the outer five 3-5(-7.5) cm long, the inner ones slightly shorter, —" WUNDERLIN, AEVISIUN UF 115 the filaments short-connate at the base, glabrous, the anthers linear-oblong, 8- 12 mm long, glabrous; gynoecium 4-5 cm long, the gynophore ca. 1 cm long, glabrous, the ovary ca. 3 cm long, tomentose (glabrous or glabrate in var. caudigera), the style ca. 1.5cm long, glabrous. Fruit linear, 21-27 cm long, 2-2.5 cm wide, apiculate with the persistent style, brown, sparsely pubescent, gla- brescent at maturity, the gynophore ca. 2.5 cm long; seeds not seen. Bauhinia petiolata is a highly distinctive though poorly known species represented by two varieties. The species is most closely related to B. geniculata which is also from the upper Magdalena Valley (Risaralda Prov.) in Colombia but is readily distinguished by its unlobed leaves and flower buds not winged. Italso has affinities with B. arborea (Ecuador) and B. brachycalyx (Ecuador, Peru, and Brazil), two other species with unlobed leaves in section Amaria. Quinones (1988) argues that B. sessilifolia is a distinct species and differs from B. petiolata by its sessile leaves, the leaf base deeply cordate, the inflores- cence of axillary umbels, the petals emarginate, the stamens of five long and five short and included, and the fruit 2 cm wide. Bauhinia petiolata on the other hand has leaves with the petiole 0.5-3 cm long, the leaf base truncate or rounded, the inflorescence a terminal or axillary in a short raceme, the petals acute, the stamens subequal and excluded, and the fruit 1.5 cm wide. These characters, with the exception of the sessile leaves, do not hold up in the material exam- ined. Even though I have not seen the MA material cited by Quinones, I believe that B. sessilifolia is based on aberrant, sessile-leaved specimens of B. petiolata. The sessile-leaved condition is approached in some material seen for B. petiolata. For example, André 1810 has petioles only 5 mm long. KEY TO THE VARIETIES OF BAUHINIA PETIOLATA 1. Ovary tomentose; central Colombia var. petiolata 1. Ovary glabrous or glabrate; Panama and Venezuela var. caudigera lla. Bauhinia petiolata var. petiolata Distribution and ecology.—Upper Magdalena Valley of central Colombia (Cundinamarca and Tolima). Wet forest at ca. 500 m. Specimens examined. COLOMBIA. Cundinamarca: Quebrada Cachimbulo near Tocaima, Rio Funza valley, André 1810 (kK); Rio Magdalena valley, Goudot s.n. (P); Hacienda El Cucharo, between Tocaima and Pubenza, Killip et al. 38314 (COL, FE NY, US), Killip et al. 38350 (COL, K, US); between Nilo an Puebla Nuevo, Murillo & Jaramillo 260 (COL, NY); Tocaima, Triana 4412 (BM, COL, G, NY); La Parada Limba and the Rio Magdalena valley, Triana s.n. (P); Tocaima, Triana s.n.(K); Tocaima, Rio Magdalena, Triana s.n. (kX). Tolima: Chicoral, Haught 6306 (COL, NY, US). State unknown: without locality, Mutis 2377 (fragment ex MA: US); Mutis 2549 (fragment ex MA: US); Mutis 4146 (fragment ex MA: US). Cultivated at Kew from material provided by Linden from Colombia (K). 11b. ue petiolata var. caudigera (S.F Blake) vi peeiete comb. nov. BASIONYM: a caudigera S.F. Blake, Contr. U.S. Natl. Herb. 20:522. 1924. TYPE: VENEZUELA. Baie ales, alone road from Puerto Cabello to os Felipe, 10-100 m, 20 May-10 BRIT.ORG/SIDA 22(1) Jun, 1920 ({D, H. Pittier 8851 (HOLOTYPE: US; photo ex US: US, NY, ISOTYPE: P). Distribution and ecology.—Panama (Colén) and Venezuela (Carabobo and Falcon). Rainforest and wet forest at 10-600 m. The variety was probably more widely distributed from Panama through northern Colombia to Venezuela in the past. Its present disjunct distribution is relictual, apparently now confined to wet river valley refugia. This taxon was placed in synonymy of B. petiolata by Wunderlin (1983). Since then, additional material has become available and it is here considered as a distinct variety on the basis of its glabrous or glabrate ovary (vs. tomentose in var. petiolata). } Specimens examined. PANAMA. Colon: ca. 2-3 mi up the Ri mouth, Kennedy & Fos- ter 2127 (MO, USF). VENEZUELA. Carabobo: upper Ciaremales ve from Puerto Cabello to San Fe- lipe, Pittier 8914 (NY, US). Faleon: Parque Nacional Quebrada de la Cueva E1 Toro (L0°50'N, 69°07'W), Liesner etal. 7727 (MO. USF), Liesner et al. 7736 (MO), Wingfield 7501 (USF). 12. Bauhinia picta (Kunth) DC,, Prodr. 2: O15. 1825. ale etia picta RUE, in a Eisibolie! al., Nov. Gen. Sp. 6:316. 1824. Type: COLOMBIA. ! A g the Ri near the confluence with Rio Opon, between Bojorque and Isla de etree May 1801 (fD), RWH.A. Humboldt & AJ.A. Bonpland 1604 (HOLOTYPE: P, not seen, microfiche IDC 156/C6; ISOTYPEs: P, 5). BCW) not seen; photo ex BCW): F G, MO, NY Bauhinia ligulata Pittier, Contr. U.S. Natl. Herb. 20:112. 1918. Type: PANAMA. SAN BLAS: near Pu- erto Obaldia, 0-50 m, Aug 1911 ({L), H. Pittier 4334 (HOLOTYPE: US; IsoTYPES: BM, F NY, US). Seana kalbreyeri Harms, Repert. Spec. Nov. Regni Veg. 19:65. 1923. TYPE: COLOMBIA. ANTIOQUIA: Murri, 1,000-1,100 m, Jun 1880 (£D, W. ees 1802 (LECTOTYPE: K, here ae nated to replace paves B holotype; ISOLECTOTYPES: F-fragment, US; photo ex K: EF NY photo ex B: EK, S Tree to 20(-40) m; branches ferruginous-tomentose when young, soon becom- ing glabrate. Leaves with the blade chartaceous, 9-13-nerved, ovate, bilobate to ca. 1/4 the length, 4-10(-21) cm long, 4-7.5(-15) cm wide, the innermost pair of lateral nerves equidistant or closer to the adjacent lateral nerves than to the mid-nerve, the base cordate to rounded, the apex of the lobes apiculate to obtuse or rounded, the upper surface glabrous, the lower surface finely ferruginous- tomentellous (at least on the nerves), the petiole 2-3.5 cm long, ferruginous- tomentellous to glabrate; stipules ovate, ca. 1 mm long, caducous; adpetiolar intrastipular excrescence enlarged, forming a subulate projection to 2 mm long, the apex with a lateral glandular area, the others minute. Inflorescence axil- lary, subterminal, short-racemose, 10- to 20-flowered, the flowers solitary or paired, the rachis ferruginous-tomentose; floral bud linear-clavate, 2-2.5 cm long, ferruginous-tomentose, the apex with minute free calyx lobes, the pedicel 2-6 mm long; bract and bracteoles lanceolate, ca. 1mm long. Hypanthium short- tubular, 6-8 mm long; calyx spathaceous at anthesis, soon splitting partly or fully to the hypanthium into 2 to 5 lobes; petals white with pink or roseate base, sometimes lavender-pink or rose, 2-3 cm long, spreading, short-clawed, WUNDERLIN, ACVIOIVUN UE 117 glabrous, the blade ovate-elliptic, 6-13(-20) mm wide, the apex rounded, the margin entire, the claw 2-4 mm long; fertile stamens 10, the outer five 2-3 cm long, the inner ones slightly shorter, the filaments free to base or short-connate, short-hirsute or with only a few short trichomes on the lower 1/2, the upper stamen attached near the apex or the middle of the ligulate part of the staminal sheath, short-hirsute (rarely glabrate); gynoecium 2-3 cm long, the gynophore, ovary, and style subequal, the gynophore sparsely tomentose below the ovary on the lower surface or glabrate, the ovary tomentose, the style sparsely tomentellous to glabrate. Fruit linear, 15-25 cm long, ca. 2 cm wide, apiculate with the persistent style, brown, tomentellous to glabrate, the gynophore 1-2 cm long; seeds elliptic-ovate, ca. 15 mm long, ca. 12 mm wide, dark brown, the surface dull, the funicular aril-lobes 6-7 mm long. Distribution and ecology.—Colombia (Antioquia, Bolivar, Boyaca, Choco, Cundinamarca, Risaralda, and Santander) and Panama (San Blas). In open to closed forest from near sea level to about 1,500 m. The species is apparently most closely related to B. petiolata, but differs in having bilobed leaves and petals white with a pink or roseate base. The type of B. ligulata from Panama isa large tree (40 m) with slightly smaller flowers and the adaxial stamen attached lower on the ligulate portion of the staminate sheath. It agrees in all other respects with the Colombia material. The material from Choco Province, Colombia differs by its slightly larger leaves (16-21 cm long compared with the usual 4-10 cm long), the glabrate staminal sheath and filament bases (these usally short-hirsute), and the less tomentose ovary. Bauhinia picta is occasionally cultivated in Colombia for its showy flowers. 1. COLOMBIA. Antioquia: 14-17 km N of Remedios on the way to Zaragoza, Cerro Cakes aa along the Rio Tucupé (07°20N, 74°30'W), Callejas et al. 5177 (USF); Vereda Alicante, Finca Penjamo, Quebrada Penjamo, on the San Juan de Bedout-La Cabana road (06°39'N, 74°31 W), Callejaset a 9388 (HUA nee Vereda La Pava, 14 km by Andes highway toward La Vereda El Libano, Fonnegra et al. 2441 (USF); Segovia, Sandeman 5575 (K). eee : near the confluence of the Rio Ité and Rio Tamar into the Rio Cimitarra, ca. 38 km W of mn eee ja (06°55 'N, 74°15'W), - Br a 1554 (MO, NY, US). Boyaca: fl Umbo region, ie rence 492 (K). ae Rio Tagachi, ca. 12 of Rio Atrato (O6°LSN, 76°50'W), Gentry et al. 37068 (MO, USF); Mecana, Juncosa 1923 (MO, = we Cundinamarca: 16 km NW of Alban along highway to Villeta, Barclay et al. 3666 (US). Risaralda: Hacienda ee ndria, 6 km on La Virginia-Cerritos highway, extreme northern part of Valle del Rio Cauca, Silverstone-Sopkin 5895 (CUVC, USF). Santander: Puerto Berrio, between Rio Carare and Rio Magdalena, ee 1772 (P); Bucaramanga and vicinity, Killip & ain 14978 (NY). State unknown: Rio Carare, Dawe 456 (K); Cordillera Orientale, Goudot s.n. (P). 13. Bauhinia seleriana Harms in Loes., Bull. Herb. Boissier 7:549. 1899. Type: GUA- AL A rae JETENANGO: near Quen Santo, ca. 1,100 m, C. Seler & E. Seler 2797 (LECTOTYPE: K,h stroyed B holotype; ISOLECTOTYPE: F-fragment, NY, US; photo ex K: E NY, US: cae EG, MO, NY US:). Bauhinia paradisi Standley & L.O. Williams, Ceiba 1:80. 1950. TyPpE: HONDURAS. EL PARAISO: Rio Lizapa floresta de Robles area, between Galeras and Lizapa Grande, 1,000 m, 14 May 1948 (fl, A. Molina 860 (HOLOTYPE: US; photo ex US: NY, US; ISOTYPES: BM, F). 118 BRIT.ORG/SIDA 22(1) Shrub or small tree to 12 m; branches short-hirsute when young, soon becom- ing glabrate. Leaves with the blade chartaceous, 9-11-nerved, broadly ovate, bi- lobate to ca. 1/3 the length, 6-10 cm long, 6-10 cm wide, the innermost pair of lateral nerves equidistant or closer to the adjacent lateral nerves than to the mid-nerve, the base cordate to rounded, the apex of the lobes obtuse to rounded, the upper surface glabrous, the lower surface short hirsute or tomentellous, the petiole (I-)2-4(-5) cm long, hirsute or tomentellous; stipules triangular-ovate, ca. 4mm long, caducous; adpetiolar intrastipular excrescence often enlarged and forming a subulate projection to 1.5 mm long, the others minute. Inflores- cence terminal or subterminal and axillary, racemose, 5- to 30-flowered, the rachis short-hirsute or tomentellous; floral bud elliptic, 8-15 mm long, short- hirsute or tomentellous, the apex with free lobes to 2 mm long, the pedicel 4-5 mm long; bract and bracteoles lanceolate to linear, 1-4 mm long. Hypanthium cupulate, 1-2 mm long; calyx spathaceous at anthesis; petals white, 11-18 mm ong, spreading, subsessile or short-clawed, glabrous except for the blade apex, the blade elliptic, 4-8 mm wide, the apex apiculate, the margin entire, the claw ca. 1 mm long, glabrous; fertile stamens 10, the outer five 11-18 mm long, the inner ones 1/2 the length of the outer, the filaments unequally connate at the base for 1-3 mm, with a laciniate rim extending above the point of adnation, the inner surface tomentose, the anthers linear-oblong, ca. 2 mm long on the longer stamens, ca. 1.5 mm long on the shorter; gynoecium 11-18 mm long, the gynophore, ovary, and style subequal, tomentose. Fruit linear, 8-10 cm long, ca. 15cm wide, apiculate with the persistent style, dark brown, tomentose, the gy- nophore 0.5-1 cm long; seeds not seen. Distribution and ecology.—Southern Mexico, Guatemala, and Honduras. Deciduous forest at 700-1,500 m. The species is closely related to B.andrieuxii of western Mexico from which itis Gennes by its larger leaves 6-10 cm long (vs. 2-3 cm in B. andrieuxii) and conspicuously hirsute parts. Bauhinia seleriana is a forest species while B. andrieuxti occurs in open deciduous scrub. —_— Representative specimens examined: MEXICO. Chiapas: along Mexican Highway 190 3 mi S of La Trinitaria, Breedlove 14485 (NY). México: gan ancho, Hinton etal. a One ca. paces EF of Laollaga, Téllez & Magallanes 233 (MEXU, MO). GUATEMALA. I a7 Las Palmas, ne eee 51582 (F US). ane Loma El Picacho, above Santa Rosalia Steyermark 42730 (F NY, US). H RAS. Choluteca: 6 km SW of Panamerican Highway, Harmon & Fuentes 5962 (UMO). El Paraiso: ie Rio California, Sierra de la Villa Santa, Williams & Molina 10479 (F, MO, UC, US, VT). Francisco Morazan: along Quebrada Suyapa, near Suyapa, Molina 553 (F& MO, US). 14. Bauhinia seminarioi Harms ex Eggers, Bot. Centralbl. 73:69. 1898. Type: ECUA- DOR. MANABI near mecienda El pec 16 mae ee (fl, fr), HEA. von Eggers 14925 (LECTO- YPE: F-fragment, h d B holotype; photo ex B: ISOLECTOTYPES: FE a US). Shrub or small tree 2-3(-5) m; branches sparsely tomentellous when young, WUNDERLIN, REVISION OF 119 soon becoming glabrate. Leaves with the blade submembranaceous to chartaceous, (5-)7-9-nerved, suborbicular, bilobed to ca. 1/2 the length, 5-13 cm long, 6-12 cm wide, the innermost pair of lateral nerves equidistant or closer to the adjacent lateral nerves than to the mid-nerve, the base cordate, the apex of the lobes obtuse to rounded, the. upper surface glabrous, the lower surface minutely strigulose, short-pilose along the veins, especially near the base, the petiole 2-4.5 cm long, tomentellous, soon becoming glabrate; stipules triangu- lar-lanceolate, ca. 1 mm long, caducous; adpetiolar intrastipular excrescence occasionally enlarged and forming a subulate projection to 2mm long, the oth- ers minute. Inflorescence terminal or subterminal and axillary, racemose, 8- to 18-flowered, the flowers usually paired, the rachis tomentellous to glabrate; flo- ral bud linear-clavate, 4-6 cm long, tomentellous to glabrate, the apex rounded, the pedicel 0.5-1.8 cm long; bract and bracteoles triangular-lanceolate, 0.5 mm long. Hypanthium short-tubular, 1.5-2 cm long; calyx spathaceous at anthesis, petals white, ca. 5cm long, slightly spreading, short-clawed, glabrous, the blade linear-oblanceolate, 5-6 mm wide, the apex acute to obtuse, the margin slightly crisped, the claw ca. 0.5cm long; fertile stamens 10, the outer five ca. 4 cm long, the inner ones slightly shorter, the filaments short-connate at the base to ca. 6 mim, the staminal sheath pilose on the inner surface near the apex and slightly above on the filaments, the anthers elliptic-oblong, ca. 5 mm long, glabrous; gynoecium 5.5-6.5cm long, the gynophore, ovary, and style subequal, glabrous. Fruit linear, 15-25 cm long, 2-2.3 cm wide, apiculate with the persistent style, dehiscent, brown, glabrous, the gynophore 2-3 cm long; seeds not seen. Distribution and ecology.—Ecuador (Guayas and Manabi). Tropical moist forest on inner coastal hills at 100-150 m. The type collection (von Eggers 14925) is apparently a mixture of this spe- cies and Bauhinia aculeata subsp. grandiflora Jussieu ex Poiret) Wunderlin of subg. Bauhinia section Bauhinia. Specimens of B. aculeata subsp. grandiflora with various collection dates were distributed as the type of B. seminarioi by C to K, F and undoubtedly elsewhere. The description, except for the statement that the species is aculeatous, clearly fits what is known to be B. seminarioi as represented by the photograph of the holotype (now destroyed) and not B. aculeata subsp. grandiflora. The adpetiolar intrastipular excrescences in B. seminarioi sometimes enlarge to 2 mm, especially in the inflorescence, but the older branches can not be described as aculeatous. Since the holotype is of young floriferous material with evident enlarged excrescences, Harms apparently in- correctly interpreted the materia. The Iltis & Iltis E-221 specimen bears only small blunt excrescences. mens ined. ECUADOR. eal Bosque Protector Cerro Blanco, road to Salinas, 15 km (02° 10'S, 79°58'W), Rubio 2020 (MO, USF). Manabi: Hacienda El Recreo, von Eggers 14886 (F, K, P, US); 4 km E of Hacienda El Recreo (2 as S along coast from Canoa, 18 km due N of Bahia de Caraquez; 00°27'S, 80°26'W) and the Pacific Ocean, Iltis & Iltis E-221 (USF WIS) 120 BRIT.ORG/SIDA 22(1) 15. Bauhinia webert i Harms, Bot. Jahrb. Syst. 42:91. 1908. Type: PERU. CAJAMARCA: near eas 2 sia 2,100 m, Jan 1905 (fD, A. Weberbauer 4802 (LECTOTYPE: G, here desig- nated t stroyed B holotype; ISOLECTOTYPE: F-fragment; photo ex B: F MO). Bauhinia ruiziana Harms, Repert. Spec. at Regni Veg. 18:234. 1922. Type: PERU. HUANUCO: erie. io ee ay H. Rune aoe 1795 [H. Ruiz Lopez & J.A. Pav6én 1300] (LECTOTYPE: K, erate ISOLECTOTYPES: F, MA, not seen, P; photo ex K, F US; Tes MA F nets ex B. Fi G MO, U Shrub to 1.5m; branches strigulose when young, soon becoming glabrous. Leaves with the blade chartaceous, 7-nerved, suborbicular, bilobed to ca. 3/4 the length, 15-4 cm long, 15-4 cm wide, the innermost pair of lateral nerves equidistant or closer to the adjacent lateral nerves than to the mid-nerve, the base cordate, the apex of the lobes obtuse to rounded, the upper surface glabrous, the lower surface strigulose, the petiole 0.5-1 cm long, minutely strigulose or glabrous; stipules triangular-lanceolate, |-1.5 mm long, caducous; adpetiolar intrastipular excrescence occasionally enlarged and forming a subulate pone to2mm long, the others minute. Inflorescence terminal or subterminal, the flowers soli- tary, the rachis strigulose; floral bud elliptic-lanceolate, ca. 2 cm long, red, strigulose, the pedicel ca. 1 cm long; bracts and bracteoles triangular, ca. | mm long. Hypanthium short-tubular, 4-6 mm long; calyx spathaceous at anthesis: petals bright pink or rose-red, 3-4 cm long, spreading, short-clawed, glabrous, the blade oblanceolate, 1-1.8 cm wide, the apex rounded, the margin entire, the claw 3-5 mm long; fertile stamens 10, red, the outer five 2.5-3.cm long, the inner ones slightly shorter, the filaments very short-connate, sparsely pilose toward the base, the anthers elliptic-oblong, 3-4 mm long, glabrous; gynoecium 3-4 cm long, the gynophore, ovary, and style subequal, glabrous. Fruit narrowly el- liptic, 4.5-5 cm long, ca. 1.5 em wide, apiculate with the persistent syle, dehis- cent, light brown, glabrous, the gynophore 4.5-5 cm long; seeds hee tees y el- liptic-ovate, dark brown, the surface dull, the funicular aril-lobe scars subequal, 2-3 mm long. Distribution and ecology.—Peru (Cajamarca, La Libertad,and Huanuco). On steep rocky slopes above rivers at 2,100-2,400 m. Bauhinia weberbaueri is related to B. ayabacensis and B. dugusti, but is eas- ily distinguished from both by its glabrous ovary (vs. tomentose in B. augusti and B. ayabacensis) and from B. augusti by its pink to rose petals (vs. white in B. d soci fo PERU. La Libertad: between Los Alisos and Pataz, Alayo 18 (USF). Huanuco: above eiasenda bute, between Ambo and Huanuco, Ferreyra 6553 (F); Ambo, Macbride & Featherstone 2411 (F), Macbride 3164 (FG), Pavon s.n. (BM), Sawada P-103 (F); Huanuco, Machride 3503 (FG), Matthews 927 (kK), without precise locality, Vita, McLean s.n. (K). Il. Series Stenanthae Wunderlin, K.Larsen & S.S.Larsen. Biol. Skr. 28: 14. 1987. Type: Bauhinia stenantha Diels Petals strongly imbricate, the corolla forming a pseudotube; fertile stamens 5. WUNDERLIN, REVISION OF 121 1. Bauhinia stenantha Diels, Biblioth. Bot. 29(116):93. 1937. Type: ECUADOR, CHIMBORAZO: canyon of the Boo panciae near Paes each ] ie m, 7-14 May 1945 (£1, fr), W.H. Camp E-3041 (NEOTYPE: Fh B holotype; ISONEOTYPE: K, P). [ECUADOR. CHIMBORAZO: en of the Rio c shanchan near Huigra, 1,260 m, 21 Sep 1933 (fl, fr), EL.E. Diels 1130 (HOLOTYPE: B, destroyed)]. Shrub or small tree to 3 m; branches puberulent when young, soon becoming glabrate. Leaves with the blade submembranaceous or chartaceous, 7-9-nerved, suborbicular, bilobed to ca. 3/4 the length, 2.5-5.5 cm long, 3-5.5 cm wide, the innermost pair of lateral nerves equidistant or closer to the adjacent lateral nerves than to the mid-nerve, the base cordate, the apex of the lobes obtuse to rounded, the upper surface glabrous, subglaucous, ns LOWE sue sparsely to moderately tomentose, the petiole 1-3cm long g te; stipules triangular-ovate, ca. 1mm long, caducous; adpetiolar intrastipular excrescence occasionally enlarged and forming a subulate projection to 2 mm long, the oth- ers minute. Inflorescence terminal or subterminal, short-racemose, 2- to 1O- flowered, the rachis tomentellous or strigulose; floral bud elliptic-oblanceolate, ca. 2.5 cm long, rose-red, tomentellous or strigulose, the apex apiculate, the pedicel 1-1.5cm long; bract and bracteoles triangular-lanceolate, ca. 1 mm long. Hypanthium short-tubular, subgibbous at the base, 6-8 mm long; calyx spathaceous at anthesis, soon splitting to the hypanthium into 2 to 5 lobes; pet- als rose-red, 2.5-3 cm long, erect and pseudotubular, short-clawed, glabrous, the blade oblanceolate, 7-8 mm wide, the apex rounded, the margin slightly crisped, the claw 8-9 mm long; fertile stamens 5, red, 2.5-3 mm long, the filaments very short-connate, the staminal sheath with a small laciniate rim, glabrous, the anthers oblong, ca. 5mm long, glabrous, the staminodes 5, rudimentary or to 1/ 2 the length of the fertile stamens, the aborted anthers globose; gynoecium 3.5- 45cm long, the gynophore, ovary, the style subequal, glabrous. Fruit linear, 6- 10 cm long, ca. 1.5 cm wide, apiculate with the persistent style base, dehiscent, light brown, glabrous, the gynophore ca. 1.5 cm long; mature seeds not seen. Distribution and ecology.—Ecuador (Azuay and Chimborazo). Dry scrub hillsides at (600-)1,200-1,800 m. |. ECUADOR. Azuay: along Pasaje-Santa Isabel-Giron highway, ee of the Rio Tenens Harling & Andersson 14404 (GB, USF); Cuenca-Machala road, between Girén and Santa Isa- bel, 17 km from Girdén (03°14'S, 79°13'W), Lojtnant & Molau 14176 (AAU); 17 om SW of Giron, MacBryde 451 (MO). Chimborazo: Huigra, Asplund 7736 (S), Asplund 15507 (K, S); vicinity of Huigra Rose & Rose 22119 (photo ex NY: US); Huigra, White s.n. (BM). State unknown: without definite local- ity, Acosta-Solis 11675 (F). ACKNOWLEDGMENTS | gratefully acknowledge the curators of A, AAU, BM, COL, CUVC, DUKE, EG, GB, GH, HUA, K, MEXU, MICH, MO, NY, P PMA, S, TEX, UC, UMO, US, VT, WIS for making specimens available for study. I thank Philip A. Silverstone-Sopkin, (CUVC) for information on Bauhinia geniculata, Gerald G. Robinson (USF), for 122 BRIT.ORG/SIDA 22(1) information on Bauhinia amatlana, Bruce F. Hansen (USF) for his helpful sug- gestions on the manuscript, and Kathleen Hotchkiss (USF) for assistance with graphics. | also thank the three reviewers of the manuscript for their helpful suggestions. REFERENCES ForTUNATO, R.H. 1986. Revision del genero Bauhinia (Cercideae, Caesalpinioideae, Fabaceae) para la Argentina. Darwiniana 27:527-557. Fortunato, RH. 1997. Bauhinia. Flora Fanerogamica Argentina. 33:4—9. Programa Proflora (CONICET), Cordoba, Argentina. McVaucu, R. 1987. Leguminosae. In:W.R. Anderson, ed. Flora Novo-Galiciana. 5:1—786. Uni- versity of Michigan Press, Ann Arbor. Quinones, L.M. 1988. Una nueva combinacion en Bauhinia (Fabaceae—Caesalpinicideae). Ann. Missouri Bot. Gard. 75:1155-1156. Vaz, A.M.S.DAF.1979.Consideragdes sobre a taxonomia do género Bauhinia L.sect. Tyloteae Vogel (Leguminosae-Caesalpinioideae) do Brasil. Rodriguésia 31: Vaz, A.M.S. ba F. and A.M.G. pe A. Tozz!. 2003. Bauhinia ser. Cansenia (Leguminosae: Caesalpinioideae) no Brasil. Rodriguésia 54:55—143. Wuwoeruin, R.P. 1976. The Panamanian species of Bauhinia (Leguminosae). Ann. Missouri Bot. Gard. 63:346-354. WUNDERLIN, R.P. 1983. Revision of the arborescent Bauhinias (Fabaceae: Caesalpinioideae: Cercideae) native to Middle America. Ann. Missouri Bot Gard. 70:95-127. WuNbeatin, R.P. 1998. Bauhinia. In:J.A. Steyermark, PE. Berry, and B.K. Holst, eds. Flora of Ven- ezuelan Guayana. 4:5-13. Missouri Botanical Garden Press, St. Louis. Wunberun, R.P. 2001. Bauhinia. In: W.D. Stevens, C.U. Ulloa, A. Poole, and O.M. Montiel, eds. Flora de Nicaragua. 1:522-526. Missouri Botanical Garden Press, St. Louis. WUNDERLIN, R.P., K. LARSEN, and S.S. Larsen. 1987, Reorganization of the Cercideae (Fabaceae: Caesalpinioideae). Biol. Skr. 28:1—40. NO ™ DENDROPHORBIUM RESTINGAE (ASTERACEAE: SENECIONEAE), A NEW SPECIES FROM SAO PAULO, BRAZIL Aristonio M. Teles Jimi N. Nakajima Universidade Federal de Minas Gerais Universidade Federal de Uberlandia Instituto de Bichel ias iB Qeaiea? Rua Ceara s/n - Bloco 2D - campus Umuarama Departam Uberlandia, Minas Gerais Av. Anténio ete 6627, Pampulha, 31.270-110 Belo Horizonte, Minas Gerais, BRASIL aristonio@hotmail.com Joao R.Stehmann Universidade Federal de Minas Gerais ie tuto ae as éncias Ries Ay Antonio ane 6627 Pampulha, 31.270-110 Belo Horizonte, Minas Gerais, BRASIL ABS RACT Dendrophorbium restingae (Asteraceae: Senecioneae), a new species from Sao Paulo state, Brazil, is described, illustrated, and compared with the related species D. paranense (Malme) Matzenb. & Baptista RESUMEN Dendrophorbium restingae eeu ee Seceipncae) una nueva epens ae Bae Ee, Brasil, es 2. | descrita, ilustrada y i i aptista L Dendrophorbium (Cuatrec.) C. Jeffrey is a predominantly Andean genus with about 50 species ranging from the Antilles to southern Brazil (Jeffrey 1992). Hind (1993) recorded nine species for Brazil; some years later Matzenbacher and Baptista (1997) proposed one more new combination in the genus, for a total of ten Brazilian species. Dendrophorbium was segregated from the large genus Senecio (Nordenstam 1996), and is recognized by its large and generally dentate leaves, corymbiform to paniculiform capitulescences, radiate heads with 8-13 glabrous phyllaries, style branches with a truncate, obtuse, or conical apex bearing a tuft of trichomes, and 5-8-ribbed cypselae with an annuliform carpopodium (Nordenstam 1996). Dendrophorbium restingae A. Teles J. N. .Nakaj. & Stehmann, 2 nov. (Figs. la, b, TYPE: BRAZII P gica Juréia-Itatins, restinga da praia do Arpoador, 22 Jun 1994, I. Cae M.M.R. Fiuza de Melo, RJ. Oliveira & M. Barros 1531 (HOLOTYPE: SP; ISOTYPE: K). SIDA 22(1): 123 - 128. 2006 BRIT.ORG/SIDA 22(1) 124 lotype, SP). a ' right (HUIULY Pe, TELES ET AL., A NEW SPECIES OF DENDROPHORBIUM FROM BRAZIL 125 INSTITUTO DE BOTANICA sp SSo Paulo — Brasil Fic. 1b. Basal leaves (holotype, SP). BRIT.ORG/SIDA 22(1) 10 mm Fic.2.Dend bi j A. Head. B. Phyllaries.C. Ray floret. D. Disk floret. E. Cypselae of disk floret. F. Stamen. All from the holoiyne (SP). A enone horbium paranense affinis, sed foliis majoribus vald ‘datis ( nulatisq ecules aan pevols non auriculatis, floribus dian 30- 35 (vs beac 25) et cypsel is 5-8-costatis (vs. 10 Ca a Robust herbs 1-1.5 m high. Stems fistulose, multisulcate and lanate. Leaves al- ternate, coriaceous, discolorous, glabrous and brownish adaxially and greyish- tomentose abaxially when dry, penninervate with 9-16 secondary veins promi- nent abaxially, margins crenulate; basal leaves non-auriculate, petiolate, the petioles 5.5-12 cm long, winged, clasping the stem, lanate, the blades ovate-cor- date, 7-35 cm long, 6.5-26 cm wide, with apex acute, base cordate; distal leaves sessile and invaginating, the blades oblong, 2-14 cm long, 0.7-3.5 cm wide, with acute apex and obtuse base. Capitulescences corymbiform. Heads heterogamous, TELES ET AL., A NEW SPECIES OF DENDROPHORBIUM FROM BRAZIL 127 radiate, peduncle 1-4 cm long, lanate, bracteole lanceolate. Involucres |-1.3 cm long, 1-1.2 cm wide, campanulate, calyculate, calyculus 5-bracteolate, bracteole 5-6 mm long, linear, glabrous; phyllaries ca. 13, uniseriate, lanceolate, 10-11 mm long, 1.5-2 mm wide, the apex acute and ciliate, the margins glabrous and scari- ous. Receptacles plane, alveolate, epaleaceous. Ray florets 4-8, pistillate; corolla liguliform to 13 mm long, bidentate to tridentate apically; style to 10 mm long, branches straight, ca. 1 mm long, the apex with a tuft of pennicilate trichomes. Cypselae 3-4 mm long, cylindrical, 5-ribbed, glabrate, with inconspicuous carpopodium; pappus ca. 7 mm long, deciduous. Disk florets perfect, 30-35; corolla tubulose, 7-9 mm long, 5-lobate, the lobes 2.5-3 mm long, the midvein visible along the distal half of the tube; stamens with anthers to 3 mm long, exserted, caudate at base to 0.2 mm long, the filaments with anther collar di- lated, the connective with an oblong conspicuous appendage to | mm long; style 7-12 mm long, branches 1-2.5 mm long, divergent, apex truncate crowned by simple trichomes. Oypeclac 3-5 mm long, cylindrical, 5-8ribbed, glabrate, carpopodium annuliform; pappus 6-7 mm long, deciduous. Dendrophorbium restingae mor phological) resembles Dend rophorbium paranense (Malme) Matzenb. & Baptista, but is disti ble by its crenulate and larger leaves and more ray florets, as well as its distinct geographical range and habitat. The new species so far has been collected only in coastal forests on sandy soils of the state of Sao Paulo. This area belongs to the edaphic vegeta- tional complex called Restinga that occupies a narrow belt along the Brazilian coast (Veloso et al. 1991; Rizzini 1997). Other endemic species of Asteraceae grow- ing in this environment, such as species of Noticastrum from Santa Catarina and Rio Grande do Sul (Zardini 1985) and Litothamnus from Bahia (Holmes 1996). Additionally, D. paranense occurs in humid soils of riverine habitats in the cloud forests of Parana, Santa Catarina, and Rio Grande do Sul. Both species can be distinguished by the characters set in the following key: Leaves coriaceous with crenulate margins; petioles not auriculate; phyllaries apically ciliate; disk florets 30-35, corollas 7-9 mm long; cypselae of disk florets 5-8-ribbed D. restingae eee pee earn ql Fee | remot 1 ni- alee dik florets 22-25, corollas 3.5- 5n mm long; ees of disk florets . ae D. paranense Etymology.—The epithet alludes to the habitat of the species, the Brazilian coastal vegetation on sandy soils named restinga. Until the present, no species of this genus had been collected in this vegetation. Geographical distribution and habitat.—This species so far has been col- lected only in restinga vegetation at the Juréia-Itatins Ecological Station, in the municipality of Peruibe in the state of Sao Paulo, where it grows in rocky places along the shore line. Phenology.—Flowers and fruits were collected from June to October. 128 BRIT.ORG/SIDA 22(1) PARATYPE: BRAZIL. Sao Paulo: Estacao Ecoldgica de Juréia-Itatins, Municipio de Peruibe, nucleo, praia do Arpoador, vegetacao de restinga arbustiva, 18 Oct 2004, Moreno et al. 100 (SP). ACKNOWLEDGMENTS We thank Inés Cordeiro, curator of the herbarium SP and collector of the type material, Marcos Sobral for suggestions and comments on the manuscript, John Pruski and Guy Nesom for reviewing the manuscript, Rosa Alves for the line drawings, CAPES and CNPq for the grants to the first and last author respec- tively. REFERENCES Hino, DJ.N. 1993. A checklist of the Brazilian Senecioneae (Compositae). Kew Bull. 48: 279-295. Hoimes, W.C. 1996. Litothamnus nitidus (Compositae: Eupatorieae), a new combination based on Mikania nitida (DC.) R. King & H. Robinson. Phytologia 81:385-390. Jerrrey, C. 1992. The tribe Senecioneae (Compositae) in the Mascarene Islands with an annotated world check-list of the genera of the tribe. Notes on Compositae: VI. Kew Bull. 47:49-109. Martzensacuer, N.I. and L.R.M. Baptista. 1997. Uma nova combinagdao no género Dendrophorbium (Cuatrec.) C.Jeffrey (Asteraceae - Senecioneae). Bol. Mus. Bot. Prefeitura Municipal Curitiba, MBM. 65(65):1-3. Norbenstam, B. 1996. Recent revision of Senecione and Calenduleae systematics. In DJ.N. Hind & HJ. Beentje, eds. Compositae: Systematics. Proceedings of the International Compositae Conference. Vol. 1. Kew: Royal Botanic Gardens. Pp. 591-596. Rizzini, C.T. 1997. Tratado de fitogeografia do Brasil: aspectos ecoldgicos, socioldgicos e floristicos. Ambito Cultural Edicdes Ltda, Rio de Janeiro. Vetoso, H.P, A.L.R. Ranaet FitHo, and J.C.A. Lima. 1991. Classificagao da vegetagao brasileira, adaptada a um sistema universal. IBGE, Rio de Janeiro. ZARDINI, EM. 1985. Revision del genero Noticastrum (Compositae — Astereae). Revista Mus. La Plata, Secc. Bot. 13(83):313-424. NOVEDADES EN GENTIANACEAE PARA AMERICA DEL SUR Eva M. Filippa Gloria E. Barboza Instituto Multidisciplinario de Instituto Multidisciplinario de Biol ogia Vegetal (CONI CET- UNC) y Biologia | vegeldl Vaan T- UNC) y Facultad de Ciencias Quimicas (UNC) Facultad de Ciencias fees (UNC) Casilla de Correo 495, 5000, Cordoba, Casilla de Correo 495, 5000, Cérdoba, ARGENTINA. efilippa@imbiv.unc.edu.ar ARGENTINA. gbarboza@imbiv.unc.edu.ar ABSTRACT Gentianella fabrisii (Gentianaceae) is proposed as a new species. It is known only from the type collection from the northwestern of Argentina (Prov. Tucuman); a key to distinguish this species from. its closer eae is provided: ne syuonyas for Cicendia quadrangularis, Curtia tenuifolia, 4 spe cies o een are proposed. The ocurrence of Gietenanthas Vernier 1 Schultesia gui is confirmed. These re ] t . Pal : fo) oO cor d and illustrated in detail. RESUMEN Se describe e ilustra una nueva especie de Gentianella a fa crs) que see en el noroeste de la Argentina (Prov. Tucu man), y se provee UNa Clave | Se dan a conocer nuevos sinénimos para Cicendia adams Girea tenuifolia, Gentiana prostrata, Zygostigma australe y para 4 especies de G nella. Se confirma la pre a de Chelonanthus viridiflorus y Schultesia guianensis para : ae y aoe las que se pe e ilustran detalladamente. INTRODUCCION Li tesdl el ti t tartico, Gentianaceae es una familia con ca. 1690 especies que habitan, por lo comun, en regiones templadas, subtropicales y tropicales (Struwe et al. 2002). Una moderna clasificacion en tribus y subtribus basada, en gran parte, en estudios filogenéticos derivados de datos moleculares (trnL, matK e ITS), acepta 6 tribus y 87 géneros (Struwe et al. 2002). Esta familia se caracteriza por presentar algunas entidades bien conocidas por sus principios amarogénicos y colorantes, de uso tanto en la in- dustria alimenticia como en la farmacéutica Jensen & Schripsema 2002). En el Nuevo Mundo, la mayor diversidad de géneros (de 47 nativos, 36 son endémicos) se encuentra en la region tropical de América Central y América del Sur (Struwe et al. 2002). Con motivo de estar realizando la revision de Gentianaceae para el Catalogo del Cono Sur y para la Flora Fanerogamica de Argentina, surgieron algunas novedades, que ya se dieron a conocer en parte (Filippa & Barboza 2001; 2003), a las que se suman ahora las de la presente contribucion. SIDA 22(1): 129 — 143. 2006 130 BRIT.ORG/SIDA 22(1) UNA NUEVA ESPECIE DE GENTIANELLA Gentianella fabrisii Filippa et Barboza, sp. nov. (Fig. 1). Tivo: ARGENTINA. TUCUMAN, Dpto. Tafi: Quebrada del Baron, Los Planchones, 3300 m, 7 Feb 1958, H.A. Fabris 1417 a (HOLOTIPO: BM; ISOTIPO: H.A. Fabris 1417 a pp., LP). A Gentianella bromifolia foliis ovatis vel late ovatis, floribus hermaphroditus et femineis, calycis lobis anguste ovatis, corollae lobis tubo 3-5 plo longioribus, g ssili differt. Hierba anual, ginomonoica, erecta, hasta ca. 40 cm alt, glabrescente. Tallo solitario, laxamente ramificado desde la base o mas arriba, poco hojoso. Hojas sésiles, las basales escasas, con lamina espatulada, base atenuada en pseudopeciolo y apice agudo, hojas caulinares sésiles, ovadas a anchamente ovadas, agudas, con 3-5-nervios sobresalientes en el hipofilo, de 10-44 mm long. x 5-15 mm lat.,con tricomas glandulares no capitados en la base foliar adaxial. Flores terminales solitarias 0 en cimas 3-5-floras, raro 8-floras y flores axilares solitarias o geminadas; pedicelos de 20-60 mm long.; caliz acrescente y persistente, de 10-16 mm long., lobulos angosto-ovados, agudos, desiguales, hasta 3 veces mas largos que el tubo, tubo con anillo de tricomas glandulares pluricelulares en su borde inferior interno, de 3-5 mm long. corola lila, violacea oscura en la base de cada lobulo, de 17-26 mm long., tubo breve (5-7 mm long.), glabrescente, con tricomas glandulares pluricelulares por encima de la union de los estambres, Lobulos elipticos a espatulados, agudos, de 10-19 mm long., 3- 5 veces mas largos que el tubo, nectarios epipétalos en U; filamentos estaminales planos, mas largos que la altura media de la corola e insertos cerca de la mitad inferior del tubo en las flores perfectas, anteras versatiles (primero introrsas, y extrorsas en la dehiscencia), elipsoides, inclusas, de + 3 mm long, tecas libres en su tercio inferior; en las flores pistiladas, estambres reducidos a estaminodios (filamentos breves con anteras atrofiadas, vacias); gineceo angosto-elipsoide, sésil, lobulos estigmaticos ovados. Capsula de color marron claro, cuando madura excede la corola persistente. Semillas numerosas, castaho oscuras, subesféricas, menores de | mm de diam., con tegumento levemente reticulado. Etimologia.—El epiteto se debe a Humberto A. Fabris 1924-1976), destacado docente e investigador botanico argentino, quien colecciono6 el material tipo y dedico una buena parte de su vida al estudio de las Gentianaceae, en especial Gentianella, de América del Sur. Observacion.—En la coleccién tipo de G. fabrisii se encontraron semillas maduras provenientes de frutos derivados tanto de las flores pistiladas como de las perfectas (Fig. 1 F-H). Habitat y distribucién geografica.—Hasta el momento, solo se conoce esta especie por el material tipo, proveniente de las montanas del oeste de la Prov. de Tucuman, por arriba de los 3000 m. Al igual que sus congéneres, habita en praderas humedas de altura donde convive con Gentianella bromifolia (Griseb.) T.N. Ho et S.W. Liu. FILIPPA AND RARRNZA AMERICA DEI Sl JR Fic.1 tianella fabrisii.A. Habito. B. Corolad 1. do fl re | I E. Caliz, vista externa. F. Embrion.G. Semilla, vista erie ore sr. Lb la . ime 1 1 y lateral. H.T Py I PY -l K. Flor, vista lateral. L. Botén floral despl 10, ot fi I ' E, I,K, L. De Fabris 14174. BRIT.ORG/SIDA 22(1) Gentianella fabrisii es afin a Gentianella bromifolia por compartir la ramificacion de los tallos, la longitud de los peciolos, la disposicion de las flores, y la forma y el color de los lobulos corolinos; se diferencian por su sexualidad (planta ginomonoica vs. planta monoico-monoclina), la forma de las hojas caulinares (ovadas a anchamente ovadas vs. linear-elipticas), la forma de los lobulos calicinos (angosto-ovados, agudos vs. linear-triangulares, aleznados), la relacion longitud tubo/Id6bulos de la corola (1/3-1/5 vs. 1/2-1/3) y por el gineceo (sésil vs. estipitado). En los tratamientos de Gilg (1906, 1916), ya se hace mencion de la presencia de poligamia entre las especies de Gentianella (sub nom. Gentiana). Si bien la mayoria de las especies poseen flores monoclinas, se pudo comprobar que de las 29 especies que crecen en la Argentina, en 5 de ellas ocurre ginomonoecia, entre las que se encuentra G. fabrisii,en tanto que solamente G. florida presenta ginodioecia Juarez de Varela 1997; Filippa & Barboza, inédito). Un caso muy particular es el de G. thiosphaera debido a la ocurrencia de trimonoecia (segun Font Quer 1979), es decir flores perfectas en su mayoria, y pocas estaminadas y pistiladas en un mismo individuo (Filippa & Barboza 2001) LAS 7 ESPECIES POLIGAMAS DE LA ARGENTINA SE DIFERENCIAN EN LA SIGUIENTE CLAVE pe cee terminales densas. Anteras hasta de 2 mm long. Pedicelos cortos, en oe o blanco-amarillenta, con tubo notablemente mas corto que los lObulos. Anteras completamente hee Plantas aes G. florida Corola amarillao lila cina,cont {ny j {| it { Anteras inclusas 0 apenas exertas, Plantas no pines ee 37 L I: 4 eet | IAtaral es ae mm. ). Plantas trimonoicas G. thiosphaera 1 | 3. 4s largo que los ldbulos. Gineceo sésil. Plantas ginomonoicas. 4, ils amarilla. Flores axilares en verticilos. Hojas opuestas o en verticilos G. multiflora 4 ae ere Flores axilares no verticiladas. Hojas siempre opuestas G.imberbis 1. Infl i inales | bien fl lit Anteras de 2-4 mm.Pedicelas comparativamente largos, de 15-90 mm 5. Tallo solitario. Hojas ovadas a anchamente ovadas. Lobulos calicinos angasto- vados G. fabrisii 5. Tallos numerosos. Hojas elfpticas 0 angosto- a linear-elipticas. Lobulos calicinos ara a linear-triangulares. 6. Hojas caulinares elipticas, siem} puestas. Nectarios en V. Semillas de 1-1,1 mm X 0,7-0,9 mm G. kurtzii . Hojas caulinares angosto- a linear-elipticas, opuestas o en verticilos trimeros. Rlactarias en U. Semillas de 0,5-0,9 mm X 0,3-0,4 m G. myriantha CILIDDA AND RARRNZA AMERICA NEI SUR 133 NUEVOS SINONIMOS PARA ESPECIES DE GENTIANACEAE CICENDIA Cicendia quadrangularis (Lam.) Griseb., Gen. sp. Gent. 157. 1839. Basonimo: Genti- uadrangularis Lam., Encycl. 2:645. 1788. Tipo: PERU: “Environs of Lima,” Dombey s.n. (HOLOTIPO: P!) Gentiana flava Larranaga, Escritos de Don D.A. Larranaga 2:100. 1923. Tipo: No citado. Syn. nov. Si bien el herbario de Larranaga fue destruido por accion de los insectos (Alonso Paz, com. pers.), gran parte de su obra escrita pudo rescatarse y fue publicada en varias etapas; en 1923, se dieron a conocer las descripciones de las especies y cuatro afios mas tarde (1927) - en el Atlas (Parte I. Botanica)-, las magnificas ilustraciones de referencia. Cuando Larranaga describe Gentiana flava no precisa ningun material tipo, solo sefiala que se encuentra “..en todos los prados; pero es tan pequena que se confunde muy facilmente con el pasto tierno entre que se oculta.” La diagnosis es breve (planta muy pequena, tallo poco ramificado y unifloro, hojas lineares, corola tetramera, glabra, amarilla, hipocrateriforme); sin embargo, los caracteres que utiliza, refrendados por la ilustracion (Lam. LXXV1J), son lo suficientemente precisos para decidir su sinonimia con Cicendia quadrangularis. CURTIA Curtia tenuifolia Ce oe Bot. Zentralbl. 60:357. 1894. (Fig. 2). Basonimo: Hist - Guiane 1:70, .26, f.2. 1775. Tipo: “Habitant in pratis humidis Ca z & Guian ae’ ‘ | lensis Ar , Anales Mus. Nac id 2, 1:65, f.3. 1911. Tipo: [URUGUAY] “Montevideo, arenales hamedos de Carrasco,” Abr 1874, J. Arechavaleta 163 (HOLOTIPO: MVM)). Syn. nov. Arechavaleta (1911) describe C. montevidensis sin mencionar los caracteres que la diferencian de sus congé El habito y la disposicion, forma y longitud de las hojas (opuesta, linear-subulada y ca. 4 mm long.) y la longitud del caliz y corola (ca.4 mm _y 5mm long. respectivamente) observados en el material tipo, son coincidentes con los de C. tenuifolia. Por otro parte, cuando Fabris (1973) cita por primera vez la existencia de Curtia malmeana en la Argentina, destaca su afinidad y posible sinonimia con C. tenuifolia (esta situacion esta propuesta en una preliminar Checklist de Bo- livia virtual: www.efloras.org). Mediante el analisis del material tipo de C. malmeana (Brasil: Malme 424b, S!), corroboramos esta sinonimia. De esta manera, Curtia tenuifolia (Fig. 2), es la Gnica especie que crece en la Argentina (Prov. Corrientes) y sur de Uruguay (Dpto. Montevideo y Canelones). GENTIANA Gentiana prostrata Haenke, Collectanea 2:66, pl.17, f.2. 1789. Tipo: ‘Rara habitat in jugis nivosis alpium Kartal et Frosnitz, principatus Salisburgensis...”. (n.v.). 134 BRIT.ORG/SIDA 22(1) Fic. 2. Curtia tenuifolia. A. Habito. B. Corola despelgada. C. Gineceo. D—-F. Anteras en vista ventral, dorsal y lateral respectivamente G. Semill lado. H. Flor. I. F CILIDDA AND RADRNZTA AMERICA NEI SUR 135 Gentiana minima Phil. Linnaea 29:13. 1858. Tipo: CHILE: “In Andibus interioribus prov. Santiago,” Philippi s.n. (HOLOTIPO: SGO 053950). Syn. nov. Fabris (1953) hace referencia al gran polimorfismo que presenta G. prostrata, lo que ha motivado una profusa sinonimia para esta especie (Fabris 1983). Genti- ana minima, descripta para Chile, ha pasado inadvertida en la literatura de los ultimos anos. El analisis del holotipo depositado en SGO, puso en evidencia que se trata de un sindnimo mas de G. prostrata. GENTIANELLA Gentianella cosmantha (Griseb.) J.S. Pringle, Phytologia 48:281. 1981. Basonimo: Gentiana cosmantha Griseb., Pl. lorentz.: 161. 1874. Tipo: ARGENTINA. TUCUMAN: Cuesta de la Casilla, Apr 1872, Lorentz 774 (HOLOTIPO: Bl; ISOTIPO: CORD)). Gentianella cosmantha f. violacea Fabris, en A.L. Cabrera (dir), Flora prov. Jujuy 8:66. 1983. Tipo: ARGENTINA. Jujuy: Dpto. Capital, Laguna de Yala, 26 Mar 1979, A.L. Cabrera et al. 30678 (HOLOTIPO: SI). Syn. nov. Fabris (1983) dif ia laf. violacea de la f. cosmantha por el tamano y color de la corola (mas pequena y de color violaceo en vez de rojo 0 rojo-anaranajado). Las observaciones realizadas a campo y de ejemplares de herbarios provenientes del area de distribucion de Gentianella cosmantha, revelaron que poblaciones con corolas rojizas poseen tamano dentro del rango de variacion senalado por Fabris para la f. violacea. Con respecto al color, es frecuente que al envejecer la flor, la corola vire de rojo a rojo-vinoso. Gentianella magellanica (Gaudich.) Fabris ex D. M. Moore, Sci. Rep. Brit. Antarc. Surv. 60:103. 1968. BAsonimo: Gentiana magellanica Gaudich., Ann. Sci. Nat. (Paris) seas 102. 1825. Tipo: [ARGENTINA| “Iles Malouines,” Gaudichaud s.n. (HOLOTIPO: B+; ISOTIPOS: Gentiana patagonica & darwinii Griseb., Gen. sp. Gent. 237. 1839. Tipo: CHILE. “Ad fretum Magellanicum, Elizabeth Island,” Darwin s.n. (HOLOTIPO: K!). Syn. ti Phil., Linnaea 29:109. 1858. Tipo: [CHILE]: “In ae Bee dictis ad altit., ca. ed.s.m.,3 ie 1837,” F Fonck 130 SGO 0539206!). Syn. nov. eee lactea Phil., Anales ce eae 90: 206, 1895, Tipo: [CHILE]. VALDIVIA: “Cordillera Pelada, 1889,” F Philippi s.n. (HOLOTIPO: SGO aon ISOTIPO: BM!). Syn. nov. Gentiana patagonica var. gracilis v, Revista Mus. La Plata, Secc. Bot. 7:348. 1896. TIPO: [ARGENTINA “Vallee de POlivaia, cr ea 1896,” N. Albov 661 y 662 (SINTIPOS: LP!). Syn. nov. Gentiana pearcei f. nana Hauman, Anales Soc. Ci. Argent. 86(3-4):299. 1918. SinTIPOs: [ARGEN- ROV. MENDOZA|: “Las Cuevas, Sep-Oct 1909,” M.S. Pennington 20 (BAF); “Cordillére de Tunuyan, 4 2700 metres,” leg. Sanzin 1301 (Herb. desconocido). Syn. nov. Los caracteres utilizados por Grisebach para Gentiana patagonica { darwinii (numero de piezas de la corola) asi como los empleados por Philippi para Gen- tiana lactea (grado de ramificacion de los tallos, forma de las hojas e inflorescencias paucifloras) y los usados por Philippi, Albov y Hauman para Gentiana modesta, Gentiana patagonica var. gracilis f. pumila uniflora y Genti- 136 BRIT.ORG/SIDA 22(1) ana pearcei f. nana respectivamente (altura de la planta y numero de flores), coinciden en un todo con los que definen a Gentianella magellanica Por otro lado, Spegazzini (1902) describio algunos taxones infravarietales de Gentiana magellanica(G. magellanica var. darwinti f. cyanescens, G. magellanica var. darwinii f. albiflora; G. magellanica var. typica f. cyanescens; G. magellanica var. typica f. albiflora) que coinciden con el rango de variacion del color de la corola de Gentianella magellanica. Estos nombres subespecificos carecen de toda validez taxonomica, ya que no se mencionan especimenes tipos ni ilustraciones. Gentianella multicaulis (Gillies ex Griseb.) Fabris, Bol. Soc. Argent. Bot. 7:89. 1958. BASONIMO: Gentiana multicaulis Gillies ex Griseb., Gen. sp. Gent. 225. 1839. Tipo: [ARGEN- TINA. MENDOZAl “Habitat in humidis pr. Uspallata et pr. Manantiales del Portillo in Andibus Mendozianis, Fl. Jun.,” J. Gillies s.n. GHOLOTIPO: K n.v.). Gentianella achalensis (Gilg) T. N. Ho et S. W. Liu, Bull. Brit. Mus. (Nat. Hist.), Bot. 23(2):61. 1993. BASONIMO: Gentiana achalensis Hieron, ex Gilg, Bot. Jahrb. Syst. 22:322. 1896. Tipo: ARGEN- TINA, CORDOBA: San Miguel, en la Sierra Grande (Achala), 27 Mar 1875, J.H.E.W. Hieronymus 215 (LECTOTIPO: CORD!)). Syn. nov. Gentianella mendocina (Gilg) J. S. Pringle, Sida 1368. 1986. BASONIMO: Gentiana mendocina Gilg, Bot. Jahrb. Syst. 54, 2 (Beibl. 118):36. 1916. Tipo: ARGENTINA. MENDOZA: 1876, R.A. Philippi s.n. (HOLOTIPO: B+). Syn. noy OC. ( Al igual que G. magellanica, G. multicaulis es una especie muy variable tanto en sus caracteres vegetativos como florales. El rango de variacion se observa en el numero, grosor y altura de los tallos, en la presencia y cantidad de hojas basales, en la cantidad de oe Bas ya ae cencia y en el color de la corola. El lectotipo hal de Gentianad ac Gentianella multicaulis. De igual manera, la diagnosis de G. mendocina se ajusta a dicha especie. Gentianella ottonis (Phil.) Munoz, Sinopsis Fl. Chile: 134. 1959. Basonimo: Gentiana ottonis Phil. Linnaea 33:177. 1805. SINTIPOS: ARGENTINA. MENDOZA: “Portillo de los Puiquenes,” R.A. Philippi s.n. (B+); CHILE: “Andes de Santiago,” R.A. Philippi s.n.(SGO 053960). iana ottonis var. hexamera (Phil.) Reiche, Fl. Chile 5:131.1907. BASONIMO: Gent hexamera Phil, Anales Univ. Chile 43:510. 1873. Tipo: CHILE: “Valle del Yeso de la Cordillera de San- tiago, Ene 1861,” R.A. Philippi s.n. (HOLOTIPO: SGO 053939!, 041974! ISOTIPO: BAF!). Syn. nov. Gen Cuando Reiche (1907) subordina G. hexamera a nivel varietal de G. ottonis, senala que los caracteres de esta especie (tallos unifloros y flores 6-7-meras) podrian corresponder a un hecho tanto anémalo como frecuente. Si bien la mayoria de los ejemplares analizados de G. ottonis presentan flores pentameras, las plantas de las dos cartulinas del holotipo y la del isotipo de G. hexamera, no dejan dudas de esta sinonimia por su habito rizomatoso, el tamano de las flores y la forma de los l6bulos corolinos. ZYGOSTIGMA Zygostigma australe (Cham. & Schltdl.) Griseb., Gen. sp. Gent. 150. 1839. Basonimo: Sabatia australe Cham. & Schltdl, Linnaea 1:194. 1826. Tipo: BRASIL: “Prov. Cisplatina, Braxuelo, Novembri mense floret.,” Sellow s.n. (W?). Gentiana rubra Larrafaga, Escritos de Don D.A. Larranaga 2:100. 1923. Tivo: No citado. Syn. nov. FILIPPA AND RARRO7ZA AMERICA DEL SUR 137 Tal como en el caso de Gentiana flava, Larranaga describe e ilustra esta especie con bastante precision, sin mencionar ejemplar de referencia, salvo su nombre vulgar y fecha de coleccion: “Canchalagua, Ener. 1811.” En base al protdlogo, no quedan dudas que se trata de Zygostigma austra © Sobre la ampliacion del area de distribucion de Chelonanthus viridiflorus en América del sur. Chelonanthus (Griseb.) Gilg es el género neotropical con mayor area de distribucion de la tribu Helieae Gilg, extendiéndose desde el norte de México hasta el sur de Brasil y Bolivia; hasta el t eptanca.8 (Struwe et al. 2002). Su taxonomia es muy confusa; en efecto, muchas ae sus especies han sido descriptas o incluidas en distintos géneros tales como Lisyanthus Aublet (Don 1838; Grisebach 1839, 1845; Progel 1865, Bentham 1876), Helia Mart. (Kuntze 1891) e Irlbachia Mart. (Maas 1985; Pringle 1995). En la actualidad, algunas especies transferidas a Irlbachia (Maas 1985), han sido rehabilitadas a Chelonanthus (Struwe et al. 1998, 1999) por lo que su circunscripcion no resulta facil. Por otro lado, al no haber un tratamiento monografico del género atn permanecen sin resolverse muchos problemas nomenclaturales y de circunscripcion de taxones (Struwe et al. 2002). En la Argentina, y como consecuencia de los cambios en la taxonomia de Chelonanthus, el registro de sus especies ha sido variable. En 1953, Fabris identifico como C. grandiflorus (Aubl.) Hassl., un Gnico ejemplar proveniente del litoral (Prov. Misiones: Molfino s.n.). Hunziker 1984) puso en duda la existencia de esta especie en ese pais debido a su escasa representatividad en los herbarios argentinos. Trabajos recientes registran la ocurrencia de Irlbachia alata (Aubl.) Maas en estudios p de especies del litoral fluvial (Pire et al. 2004) e L alata subsp. viridiflora (Mart.) Pers. @ Maas en estudios floristicos de los esteros del Ibera, en Corrientes (Arbo & Tressens 2000). Al analizar los ejemplares de referencia y otras colecciones del nordeste de la Argentina, se pudo verificar que solo Chelonanthus viridiflorus habita en este pais; asi, su area de distribucion se extiende hacia el sur del continente. En esta contribucion se proporciona una detallada descripcion y se ilustra C. viridiflorus a la par que se adicionan otras colecciones que reafirman la distribucion de esta especie en territorio argentino y paraguayo. 1c 1. Chelonanthus viridiflorus (Mart.) Gilg, en A. Engler u. K. ee Nat. Pflanzenfam. 4(2):98. 1895. (Fig. 3). Lisianthus viridiflorus Mart., Nov. 1. 2(2):94, t.173. 1827. Helia viridifl Mart.) Kuntze, Revis. gen. pl. 2:428. 1891. hie oe subsp. eel (Mart.) oe & Maas, Proc. Kon. Ned. Akad. Wetensch., C 88(4):410. 1985. Tipo: “Crescit in subalpinis campis prope Villa Rica, Sabara, et alibi in Provincia Minarum” Martius 72 (SINTIPO: MI). Hierba anual. Tallo erecto, simple, terete en la base y subcuadrangular superiormente, de 65-100 cm alt. Hojas enteras, pinnatinervias, las inferiores 138 BRIT.ORG/SIDA 22(1 hel fiflorus. A. Habito. B. Semilla. C. Corola desplegada con lusos. D. Transcorte de ovario. E. Antera. F. Frut aliz y estilo persistentes. G. Frut dehi ia. H. Flor. 1. Tri glandular del caliz.J Gineceo. K. Caliz, vist 1o las glandulas.L. pical de infl A, E-G de Schinini et al.11248; B de Arbo etal. 8492; C, D, H—L de Molfino s.n. La escala de 12 mm vale para C,J,L;lad FH. FG K lad CIIDDA AND RAPRNTA AMERICA NEI SUR 139 brevemente pecioladas, subconnadas (las bases confluyen en una vaina membranosa), las superiores sésiles, lamina eliptica, aguda, base atenuada, de (1,5-)2,5-9 cm de long. x (0,6-)1,5-4,5 cm de lat., hipofilo con 2 6 3 pares de nervios secundarios, opuestos y sobresalientes, peciolo de 4+,5-6 mm de long. Cimas dicotomicas terminales, por lo general paucifloras, cada rama con 2-6 flores, bracteas elipticas, agudas. Flores pentameras, perfectas, levemente zigomorfas, horizontales, reflexas en post-antesis, cortamente pediceladas (5-7 mm); bractéolas escuamiformes, hasta ca.7 mm de long, caliz campanulado, persistente y poco acrescente, de 6-9 mm de long., tubo muy breve, con tricomas glandulares pluricelulares en la base, de 2-3 mm, l6bulos crasos, ovados, obtusos, por fuera y en el centro un area glandular elipsoidea sobresaliente; margen translucido; corola verde palida, de (2-)2,5-5 cm de long,, tubo angosto en la base, ensanchado en campana ventricosa por arriba del caliz, de (1,6-)2-4,4 cm de long., lobulos ovados, rotundos, pentanervados, hasta unas 7 veces mas cortos que el tubo, apice grueso y oscuro, recurvado; estambres insertos debajo del 1/3 inferior del tubo, filamentos laminares subiguales, anteras ovoides 0 sagitadas, libres en su 1/3 basal, con un apéndice estéril en el apice, polen en tétrades, blanco, abundante; gineceo bicarpelar, ovario claviforme, placentas introflexas, estilo filiforme, de igual longitud que los estambres, en post-antesis igual o mas largo que la corola, estigma en dos lamelas ovadas. Capsula septicida, reflexa, ovoide-cénica, rostrada (estilo persistente), dehiscente por el medio, en dos valvas al madurar. Semillas numerosas por fruto, anguladas, menores de 1 mm de diam. Habitat y distribucion geogrdfica.—Especie caracteristica de zonas pantanosas, sabanas htimedas, borde de rios y embalsados. Habita en Venezu- ela, Colombia, Guyana, este de Bolivia (Dpto. Santa Cruz), centro y sudeste de Brasil (Edos. Mato Grosso, Goias, Sao Paulo), sudeste de Paraguay (Dptos. Alto Parana, Caaguazu, Canendiyt, Itapua y San Pedro) y noreste de Argentina (Provs. Corrientes y Misiones). Material examinado: ARGENTINA. Prov. Corrientes. Dpto. Ituzaingo: Desembocadura del Arroyo Garapé en el Rio Parana, 45 km al E de Ituzaing6, en pantano, 24 Abr 1975, Schinini et al. 11248 (CTES), Dpto. Santo Tomé: Reserva Natural Provincial del Ibera, Laguna Galarza, 28° 05'S, 56° 40' W, 0-100 m,embalsado al borde del canal de a ala Laguna, 25 Abr 1995, Arbo et al. 6553 (CTES), Laguna de Luna, 28° 04'S, 56° 49’ W, costa occidental, en embalsado, 24 Nov 1999, Arbo et al. 8492 (CTES); Laguna Galarza y Luna, 03 Abr 1998, Honfi & Davina 00895 (MNES); Laguna Galarza, embalsado, 9 Dic 1974, Quarin et al. 2509 (CTES). Prov. Misiones. Dpto. San Javier: San Javier, Feb 1922, Molfinos.n. ae PARAGUAY. Dpto. Caaguazu: Yhu, 24 Ene 1951, Sparre et Vervoorst 2060 (CTES, LIL); 10 km al e Caaguazu, camino a Yhu, orilla de arroyo en terreno pan 0, 8 Feb 1966, Krapovickas et i 9H ok Dpto. Canendiyu: Car pa-cué, pastizal sobre slo sees se Mar 1996, Jimenez & Marin 110 (CTES, PY); Ballinoti cué, cerrado, eas al b turad Feb 1988, Jimenez & eae BJ (CTES). Dpto. Itapua: Isla Yacyret, f I que rod ee de Arary, 22 Abr 1993, Quintana et al. 265 (CTES, PY). 140 BRIT.ORG/SIDA 22(1) Sobre la presencia de Schultesia guianensis en Argentina y Paraguay La cita de Fabris (1953) para Schultesia guidnensis en la Argentina estaba documentada solo por un ejemplar del Prof. J. Molfino s.n., procedente de Misiones. Se pudo localizar este ejemplar en el herbario BAF y corroborar que, efectivamente se trata de esta especie al compararlo con el material tipo depositado en Paris. En este trabajo, se describe e ilustra a esta especie y se la documenta ademas, para Paraguay. Schultesia guianensis (Aubl.) Malme, Ark. Bot. 30.2):9. 1904. (Fig. 4). Exacum eguianensis Aubl, Hist. pl. Guiane 1:68, t.26, f.1. 1775. Tipo: “Guiane” (PD). Hierbas anuales, pequenas, glabras, de 4-15 cm alt. Tallo erecto, tetragono, con alas reducidas en los angulos, poco ramificado superiormente. Hojas enteras, opuestas, las inferiores pseudopecioladas, espatuladas, obtusas, de 7-10 mm de long. x 5mm de lat.; las superiores sésiles, elipticas, agudas, subconnadas en la base, de 10-25 mm de long. X 5-8 mm de lat. Flores solitarias 0 en cimas dicotomicas en el apice de las ramas; bracteas elipticas, agudas. Flores tetrameras, perfectas, sésiles o con pedicelos breves (hasta de 5 mm); caliz urceolado, persistente, poco acrescente, de 12-19 mm de long., 4-alado, alas marcadamente retinervadas (1,5-2 mm de ancho), tubo de 8-11 mm, l6bulos subtriangulares largamente aleznados; corola rosada a purptrea, marcescente, e 15-22 mm de long., tubo angosto en la garganta, de 9-12 mm, lobulos obovados, rotundos, a veces apiculados, pentanervados, un poco mas cortos que el tubo; estambres insertos en la 1/2 basal del tubo corolino, inclusos 0 apenas exertos, subiguales, sector libre de los filamentos alado hasta su altura media, alas irregularmente partidas en el apice; anteras ovoides 0 sagitadas, libres en su 1/3 basal; polen en tétrades; gineceo bicarpelar, ovario elipsoide, un poco mas corto que el tubo corolino, placentas introflexas, estilo filiforme, deciduo, estigma con dos lamelas trans-ovadas a subcirculares. Capsula septicida, elipsoide. Semillas numerosas por fruto, subesféricas, ca | mm de didm.; episperma reticulado, células poligonales con paredes ligeramente onduladas. Observacion.—Acorde con Struwe et al. (1999) y Pringle (com. pers.), los especimenes procedentes de otras areas de América muestran algunas diferencias con respecto a nuestros datos, sobre todo en la altura de la planta y la longitud del caliz y la corola. Asi, Schultesia guianensis en su conjunto, presenta un rango de variacidn mas amplio en cuanto a los caracteres mencionados, extendiéndose la altura de la planta desde 3-45 cm, la longitud del caliz de 8-22 mm y, la de la corola de 8-26 mm Habitat y distribucion geogrdfica.—Especie palustre de amplia distribucion tropical y subtropical, que se extiende desde México hasta el noroeste de Ar- gentina y Paraguay. En el Cono Sur habita en campos bajos y anegados en la ribera de cursos de agua y lagunas. FILIPPA AND RARRNZA AMERICA DEL SUR 141 & Fic. 4. Schultesi j is. A. Habito. B. Caliz abierto.C. Corola { y gi D. Semilla | i f ticul De Molfino s.n. (BA 52976). lade5 le para B, C 142 BRIT.ORG/SIDA 22(1) Material examinado: ARGENTINA. Prov. Misiones. Dpto. Capital: Posadas a San José, Feb 1922, Molfino s.n(BA 52976); Posadas, hacia San José, en campo bajo, Molfino s.n., Feb 1922, (BAF). PARAGUAY. Dpto. Cordillera: In regione lacus rte: 1913, Hassler 12596 (BAF, LP); Camino de Altos a Loma Grande 25° 17'S, 55° 15' O, 20 Mar 1989, N. Soria 3438 (FCQ); Cordillera, 25.08° S 57.12' W, 25 Febr 1990, E. Zardini y R. Velazquez awe (AS). BRASIL. Edo. Rio de Janeiro: Cabo Frio, Oct 1899, E. Ule 4747 (CORD). BOLIVIA. Dpto. Santa Cruz: Fortin Ra-velo-Cerro San Miguel, Mar 1989, F. Mereles 2744 (PY). AGRADECIMIENTOS Las autoras expresan su gratitud a los Dres. JS. ae y E.F Guimaraes por la lectura critica de este trabajo y sus acertadas observaciones para mejorar el manuscrito. A los curadores y asistentes curatoriales de los herbarios AS, BA, BAP BM € CORD CLES FCO. KLE LP VMN ES MYNME Pi 5, 5OU. SL yy, Z, quienes pusieron a nuestra disposicion el material de herbario estudiado. Al Dr. P. Maas por habernos facilitado la obtencién de bibliografia especifica. A M. Ehn (S) y a los Dres. M. Mufioz (SGO) y H.S. Osorio (MVM) por el envio de imagenes digitales de materiales tipo. Al Dr. E. Alonso Paz (MVFQ) por la informacion brindada sobre el herbario de Don D. Larranaga. Al Dr. L. Ariza Espinar por suconstante estimulo. A los dibujantes L. Ribulgo y P. Peralta por la confeccion de las ilustraciones. Al Consejo Nacional de Investigaciones Cientificas y Técnicas (CONICET Argentina), Agencia Cordoba Ciencia (ACC- Argentina), Secretaria de Ciencia y Tecnologia (UNC-Argentina) y a la Myndel Botanica Foundation (Argentina) por el apoyo financiero otorgado. REFERENCIAS Arso, M.M. & S.G. Tressens (eds.) 2000. Flora del Ibera. EUDENE, Corrientes. Pp. 1-613. ARECHAVALETA, J. 1911. Vegetacion Uruguaya. Varias especies nuevas. Anales Mus. Nac. Montevideo, ser. 2, 1:59-83. BentHAM, G. 1876.Gentianeae. En:G. Bentham & J.D. Hooker,eds.Genera plantarum 2. Lovell Reeve & Co.,Williams & Norgate, London. Pp. 799-820. Don, G. 1838. A general history of the dichlamydeous plants 4. J.G.& F. Rivington et al. London. Pp. 173-214 Fasris, H.A. 1953. Sinopsis preliminar de Gentiandceas argentinas. Bol. Soc. Argent. Bot. 4:233-259. Fasris, H.A.1973.Un nuevo género de Genciandceas para la Flora Argentina, Curtia Cham. & Schidtl. Bol. Soc. Argent. Bot. 15:130-131. Fasris, H.A. 1983. Gentianaceae. En: A.L. Cabrera, ed., Fl. Prov. Jujuy 8. Coleccion Cientifica Instituto Nacional de Tecnologia Agropecuaria, Buenos Aires. Pp. 55-85 Fivippa, E. & G.E. Barsoza. 2001. Novedades en Gentianaceae de Argentina. Kurtziana 29: 79-83. Fiuippa, EM & G.E. Barsoza. 2003. Novedades en Gentianaceae de Argentina. Il. Kurtziana 65-67 Fiippa, E.M & G.E. Barsoza. Gentianaceae, Flora Fanerogamica Argentina. PROFLORA- FILIPPA AND RARRNTA AMERICA NEI SUR 143 CONICET. Inédito. Font Quer, P. 1979. Diccionario de Botanica. Ed. Labor, S.A. Barcelona. Pp. 1-1244. Gi, E. 1906. Beitrage zur Kenntnis der Gentianaceae Ill. Gentianaceae andinae. Repert. Spec. Nov. Regni Veg. 16-17:33-56. Gitc, E. 1916. Gentianaceae andinae. Monographischem Zusammenstellung der Genti- ana-Arten Stid-Amerikas. Bot. Jahrb. Syst. 54-2. Beibl.118:80-85. GRrISEBACH, H.R. 1839. Genera et species Gentianearum.J.G. Cotta, Stuttgart & Tubingen. Pp. 364 GriseBACH, H.R. 1845. Gentianaceae. En: A. De Candolle, ed. Prodromus systematis naturalis regni vegetabilis 9. Fortin, Masson et Sociorum, Paris. Pp. 38-141. Hunziker, A.T. 1984.Los géneros de Fanerdgamas de Argentina.Claves para su identificacion. Bol. Soc. Argent. Bot. 23:158 JENSEN, S.R. & J. ScHripsema. 2002. Ct t and pharmacology of Gentianaceae. En: L. Struwe and V.A. Albert, eds. Gentianaceae - Systematics and Natural History. Cam- bridge University Press, Cambridge. Pp. 573-631. JUAREZ DE VARELA, F. 1997. Gentianaceae. En: Flora del Valle de Lerma, Aportes Botanicos de Salta, Ser. Flora 4:1-19. Kuntze, C.E.O. 1891.Revisio generum plantarum 2. Arthur Félix, Leipzig. Pp. 426-432. Larraniaca, D.A. 1923. Escritos de Don Damaso Antonio Larrafaga. Il. Botanica-Zoologia. Instituto Histdrico y Geografico del Uruguay, Montevideo. Pp. 1-512. LARRANAGA, D.A. 1927. Escritos de Don Damaso Antonio Larranaga. Atlas, Parte |. Botanica, Instituto Histérico y Geografico del Uruguay, Montevideo. Lam I-CXXxXI. Maas, PJ.M.1985.Nomenclatural notes on Neotropical Lisyantheae (Gentianaceae). Proc. Kon. Ned. Akad. Wetensch C 88:405-41 2. Prinate, J.S.1995.Gentianaceae. En:G. Harling and L. Anderson, eds. Flora of Ecuador 53.Pp. Sie Procet, A. 1865. Gentianaceae. En: C.F.P von Martius, ed. Flora Brasiliensis 6 (1). Leipzig. Pp. 197-248. Pire, S.M., LM. AnzoteGu! & G.A. Cuanrano. 2004. Estudios palinoldgicos en el Litoral fluvial argentino. Instituto Superior de Correlaci6n Geoldgica, Ser. Misc. 12:139-146. ReicHe, C. 1907. Jencianaceas, Fl. Chile 5 [Gentianaceae: 120-134]. Specazzini, C. 1902. Nova addenda ad Floram Patagonicam Il. Anales Soc. Ci. Argent. 53 [Gentianaceae: 73-75]. Stauwe, L.& V.A. Aceert. 1998. Lisianthus (Gentianaceae), its probable homonym Lisyanthus, and the priority of Helia over Iribachia as its substitute. Harvard Pap. Bot. 3:63-71. Struwe, L., PJ.M. Maas, O. PIHLAR & V.A. AcperT. 1999. Gentianaceae. En: PE. Berry, K. Yatskievych & B.K. Holst, eds. Fl. Venezuelan Guyana 5:474-542. Struwe, L., J.W. Kavereit, J. KLACKENBERG, S. Nisson, M. THiv, K.B. Von HAGEN & V.A. AvBerT. 2002. Systematics, character evolution, and biogeography of Gentianaceae, including a new tribal and subtribal classification. En:L. Struwe and V.A. Albert, eds. Gentianaceae - Sys- tematics and Natural History. Cambridge University Press, Cambridge. Pp. 21-309. BRIT.ORG/SIDA 22(1) Book REVIEW Oi Lers. 2005?. Coffee, cappuccino, espresso and much more. (ISBN 3-921799- 48-1, hbk.). Kochbuch-Verlag?, Munich. $30.00, 159 pp, 14 color illus., 6 3/4" x 8 3/8". Robusta, arabica, peaberry, mocha...? “Perfectly home made by Olli Leeb,” follows the title of this charming homespun publication. A loosely edited translation from the German results in a quaint posualys across-the-table Bo of } ur i€ if narrative. Add to that the ty pevnile: styled, unjustified, co fun book! And asan added anomaly, dered only slightly UOUL f | tha Ca VEE Yy tui except for a few color graphics, the book i is printe di in an apropos light sepia tone. This is haan a coffee cookbook which the author makes no bones about. It does, eee begin with several one o He skims through such topics as cultivation, different variet- two page “chapters” on various to ies, processing, opus - movement oa the globe, roasting, and even the origins of (gasp!) in- drepat ation of the different styles of coffee drinks tor stant coffee. From tt (cappuccino, Turkish, café latte etc.) ia the use of coffee in other drinks. Finally, there are several chapters on food prepared with coffee including: cakes, ice cream, pastries, and other desserts. All is mments on the role of coffee in history, politics, culture, and social custom. interspersed with ¢ » way of example here is one of my favorite quotes from the book: “The small coffee ae met at its greatest success during the industrialization when beer, which until then had b work and even at breakfast, was gradually being pushed aside by coffee. This brought forth a more productive labor force, which was better protected when handling machinery and worked more ef - ficiently.” Regardless of depth of knowledge anyone interested in the bean will enjoy this delightful Research Institute of Texas, Fort Worth, TX 76102-4060, U.S.A. bli Robert]. George, Botanical publication, reeorge@britorg. SIDA 22(1): 144. 2006 AMELICHLOA: A NEW GENUS IN THE STIPEAE (POACEAE) Mirta O. Arriaga Mary E. Barkworth Herbario Intermorntain Herharilsm Museo Argentino de ie: (as aaIIGIES Department of Biology ernar dino R Utah State University C1405DJR Buenos Aires, ARGENTINA Logan, Utah 84322-5305, U.S.A. ABSTRACT A new genus of Sti} P Amelichloa, is described. It differs f ther genera in the tribe in the woody, sharp tips of it basal leaves, the presence of smooth longitudinal ribs on its caryopses, its persistent stylar bases gnc in the edpent pie of cleistogamous panicles in the axils of its basal leaf sheaths. TI ambigua, A. brachychaeta, A. brevipes, A. caudata, and A. clandestina. The fie four sneciee are native to South America; A. clandestina is native to northern Mexico. A key to the ee is ia ided. Three of them, A. brachychaeta, A. caudata, and A. clandestina, have becom ished in disturbed areas on other continents. The name honors Dr. Maria Amelia Torres. RESUMEN Se do Ameliehloe: un nuevo genero pan la ube onpeae (Poaceae). El cu ono difiere a otros presentar a muy longitudinales e en la supe del cariopse y fee ere persistentes, asi como en la frecuente presencia de nos en las axilas de las hojas basales. El género se encuentra constituido por cinco Spo A. ambigua, A. Oe ae on Prey e! A. Eauace y A. sain esas Las cuatro r . te Tres de | ies A. brachychaeta, A.caudata y A.clandesti | blecid A li bad | i Se dedica este géneroa la Dra. Maria Amelia Torres. This paper arises from the need to determine the appropriate generic treatment of three grass species introduced in the United States. In traditional generic treat- ments they are known as Stipa brachychaeta, S. caudata, and S. clandestina. All three have names in Achnatherum (Barkworth 1993; Jacobs & Everett 1996). The first two have, in addition, names in Jarava (Penailillo 2002). Soreng et al. (2003) treated S. brachychaeta and S. caudata as species of Jarava and S. clandestina as a species of Achnatherum. We reviewed fresh material, herbarium samples and type specimens of all three species and became convinced that, not only do they all belong to the same genus, but that they do not belong to any currently recognized genus. In this paper, we present the new genus, explain how it differs from other Ameri- can genera, list the species that it includes, and provide a key to the species. Etymology.—We are naming the new genus Amelichloa in honor of Dr. Maria Amelia Torres who has generously shared with us her extensive knowledge of South American Stipede. SIDA 22(1): 145 — 149. 2006 146 BRIT.ORG/SIDA 22(1) Amelichloa Arriaga & Barkworth, gen. nov. Type: Amelichloa ambigua (Speg,) Arriaga & Barkworth (Stipa ambigua Speg.). Species Amelichl liff | li Stipearum laminis apicibus rigidisimis a ae | i pag dorsali et duobus costis 1:} ] : 1; 1G] . : Pec SPs eee We fal; oO et acutissimis ) 121 * basalium quam inter species alias in triby pro ai te one Plants perennial, cespitose. Culms erect, with 2-3 nodes. Leaves mostly basal; sheaths open, smooth, glabrous; auricles absent; ligules scarious, rounded to acute; blades stiff, involute, apices very stiff, sharply pointed. Inflorescences paniculate, main panicle terminal, apparently wholly chasmogamous, reduced cleistogamous panicles often present axillary to the basal leaf sheaths. Spike- lets of terminal panicles with | floret, disarticulating above the glumes and be- low the floret. Glumes exceeding the floret, acute to acuminate, 1-5-veined; anthoecia fusiform, terete; calluses antrorsely strigose, blunt; lemmas pubescent, often more densely and/or more persistently so over the midvein and lateral veins, hairs on the lower portion about 0.7-2 mm long, hairs on the distal por- tion often longer; crown not developed; awns 1-2-geniculate, scabrous, persis- tent; lodicules 3; stamens 3, anthers sometimes all of equal size and more than 2 mm long, sometimes 1 more than 2 mm long and 2 much shorter, sometimes all short; ovaries glabrous, style with 2 branches, united at the base, stigmas plu- mose. Caryopses fusiform, with 3 smooth, longitudinal ribs at maturity, stylar bases 1-2 mm long, persistent, sometimes eccentric; hila linear, about as long as the caryopses. Spikelets of axillary panicles 0.5-1 mm long, glumes thin, shorter than the florets, florets unawned or with reduced awns. x = 1]? Amelichloa includes five species, four of which are South American. The fifth species, A. clandestina, grows in northern Mexico. Some species have also become established in other parts of the world. The short anthers are probably associated with cleistogamy. A similar variation in anther length is also found in the primarily South American genus Nassella. In N. leucotricha (Trin. & Rupr.) R.W. Pohl, the proportion of florets with the different combinations varies in response to environmental conditions. Cattle avoid the species of Amelichloa because of the pointed leaves. The frequent presence of cleistogenes also favors establishment of the species in ar- eas that are grazed or mown. The combination of the sharp leaves and cleistogenes means that members of the genus could become serious problems in rangelands. The only chromosome count that has been made for the genus is of 2n = 44 for Amelichloa brachychaeta (Bowden and Senn 1962). None of the species has been included in any molecular studies. We shall be initiating a study that in- cludes some species of Amelichloa, plus many other South American taxa, in summer, 20006. ADDIACA ANN RADVINODTU AMECLICUINA 147 1. Hairs at the top of the lemmas 4-5 mm long, numerous A.ambigua 1. Hairs at the top of the lemmas up to 2.5 mm, long, sparse. 2. Awns 27-30 mm; florets 7-9.8 mm A. brevipes 2. Awns 10-25 mm; florets 4-8 mm. 3, Mature caryopses with inclined, eccentric stylar bases; lemmas glabrous be- tween the midvein and the lateral vein, even at the base, usually also gla- brous between the lateral vein a the marginal vein A. caudata . Mature caryopses with erect, usually centric stylar bases; lemmas pubescent between the lateral and marginal veins on the lower 1/2, usually also be- tween the midvein and lateral veins. 4. Florets 4—-5.5 mm long; awns usually 1-geniculate A. brachychaeta 4, Florets 5.5-8 mm long; awns usually 2-geniculate A. clandestina (eS) Amelichloa ambigua (Speg.) Arriaga & Barkworth, comb. nov. Basionym: Stipa ambigua Speg,, April 1925. Revista Argent. Bot. 1:27. Jarava ambigua (Speg.) Penailillo, Gayana, Bot. 59:30. 2002. TypE: ARGENTINA: PROV. BUENOS AIRES: Pdo. Gral. Pinto, “in pratis circa Estacion Iriate,” 24 Nov 1905, Spegazzini s.n. (HOLO- YPE: LP ex LPS-2412!; isotype: BAA-2946 (col. Typus!)). Pee dusenii Hitchc., Contr. U.S. Natl. Herb. 24:271.1925. Type: ARGENTINA: BUENOS AIRES: plains Sierra de la Ventana., 29 Nov 1904, P Duséns.n. (HOLOTYPE: US-1161165; PARATYPE: BAA-3038 ex Herb. Parodi 5054 (col. Typus!)). Distribution.—Argentina and Uruguay (Rosengurtt et al. 1970; Zuloaga et al. 1994; Torres 1993, 1997), introduced to France (Verloove 2005). Amelichloa brachychaeta (Godr.) Arriaga & Barkworth, comb. nov. BasionyM: Stipa brachychaeta Godr., Mém. Sect. Sci. Acad. Sci. Montpellier 1:450. 1853. Nassella brachychaeta (Godr.) Barkworth, Taxon 39:609. 990) Achnatherum brachychaetum r.) Barkworth, Phytologia 74:6. apes Jarava brac! (Godr.) Penailillo, Gayana, Bot. 59: 5 2002. TyPE: FRANCE: du Por l ‘au Jardin Botanique, Patria ignota, “plantes etrangerse-an, Port Juvenal. Anon. igh 22 May pla inane PE (probable): MPU; OM: BAA- 2979, hojas a & b! (col. typus, fragm. d tipo). ce eminens fo. viridis Kuntze, Revis. =e PL 3:37). 888 TYPE: ARGENTINA. SANTA FE: Ceres, 92, Kuntze s.n. (HOLOTYPE: NY; ISOTYPE: COR - vanes Griseb., Symb. Fl. Argent. 298- 290, a TYE: ARGENTINA: Cordoba, PG. Lorentz 46,187? (HOLOTYPE: GOET, ISOTYPE: CORD, US 866090! (fragm. ex GOET)). Distribution.—Argentina, Chile, Pert and Uruguay (Matthei 1965; Rosengurtt et al. 1970; Zuloaga et al. 1994; Torres 1993, 1997). Although Tovar (1993) and Brako and Zarucchi (1993) do not mention it, the presence of this species in Perti is cited in Caro and Sanchez (1971), Torres (1997), and Verloove (2005). In- troduced to the U.S.A., Australia Jacobs & Everett 1993), France, and Spain (Verloove 2005). Amelichloa brevipes (E. Desv.) Arriaga & Barkworth, comb. nov. BasionyM: Stipa brevipes E. Desv, Fl. Chil. 6:282. 1854. as brevipes (E.Desv.) Penailillo, Gayana, Bot. 59:30. 2002. Type: CHILE: en las provincias ntrales de la Republica, C. Gay 1106. s.d. (HOLOTYPE: P; ISOTYPE: BAA! (col. typus fragm ues (fragm.)). 148 BRIT.ORG/SIDA 22(1) Stipa hirtiflora Hack. in Dusén, Ark. Bot. 7:5, t.3, (f.7), 7, CL. 4-5). 1908. TYPE: ARGENTINA. SANTA Cruz: Arroyo Pelque, Jan 1905, P. Dusén 5640 (HOLOTYPE: W, ISOTYPE: US 1161168, 3168627 (fragm.)). Distribution.—Argentina (Roig in Correa 1978; Torres 1993; Zuloaga et al. 1994), Chile (Matthei 1965). Amelichloa caudata (Trin.) Arriaga & Barkworth, comb. nov. BasionyM: Stipa caudata Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 1:75. 1830 A] th tum (Trin.) S.\W.L. Jacobs & J. Everett, Telopea 6:582. 1996. Jarava caudata (Trin.) Penailillo, Gayana, Bot. 59:30-31. 2002. TYPE: Specimena Chilensia, J. Lindley s.n. (HOLOTYPE: LE-TRIN 1392.01 (& fig); IsoTYPE: BAA 3001! (col. typus fragm), US 2489479 (fragm)). Stipa bertrandii Phil. Linnaea 33(3-4):283. 1864. Type: CHILE: Andes de Santiago, Bertrand s.n (HOLOTYPE: SGO-PHIL-102; IsoTYPE: BAA 2972! (col. typus frag. ex holotype), SGO 62787, 63156; US 866140). Stipa amphicarpa Phil. Anales Mus Nac. Santiago de Chile 11, lam.3, f.2. 1892. Type: CHILE: prope Angol, Dec 1887, R.A. Philippi (HOLOTYPE: SGO-PHIL 99, BAA 2952! (col.typus fragmento ex SGO); ISOTYPE: SGO 37393, 62793, US A866145). Stipa litoralis Phil. Anales Univ. Chile 93:717. 1896. Type: CHILE: Isla Quiriquina, Mar 1880, R.A Philippi s.n. (HOLOTYPE: SGO 62798; US 825120 (fragm. ex SGO)) Distribution.—Argentina, Chile and Uruguay (Matthei 1965; Rosengurtt et al. 1970; Zuloaga et al. 1994; Torres 1993, 1997); introduced to Australia Jacobs & Everett 1993), England, Italy, Spain (Verloove 2005), and U.S.A. Amelichloa clandestina (Hack.) Arriaga & ees comb. nov. BASIONYM: Stipa clandestina Hack., Repert. Spec. Nov. Regni Veg. 8:516. 1 Achnath ‘land (Hack.) Bark worth, Phytologia 74:6. 1993. TYPE: MEXICO: COAHUILA: saltillo, Mar 1908, Arséne aan ECTOTYPE: W, ISOLECTOTYPE: MO 845904t US 1030718, 3168626 (ragm. ex W). Lectotype designated by Hitchcock, Contr. U.S. Natl. Herb. 24:238. 192 Distribution.—México; introduced to U.S.A. Hitchcock (1925) included a pittieri in S. clandestina and, as a result, included Colombia in the range of S. clandestina. The type of S. pittieri (US 531634!), however, belongs in Nassella, as is reflected in the combination N. pittieri (Hitche.) Penailillo (1998). Thus, it appears that A. clandestina is restricted to northern México. ACKNOWLEDGMENTS We thank Maria Amelia Torres for her ongoing interest in the tribe, her will- ingness to share her knowledge, and her well documented publications. We also thank the curators of BAA, CDA, LP, MO, and US for making their specimens available to us for this and our other studies in the Stipeae, J. Landers and J. Valdés Reyna for bringing the presence of A. clandestina to Barkworth’s atten- tion, RJ. Soreng for clarifying Hitchcock's attribution of Stipa clandestina to Colombia, and G. Davidse and S.WL. Jacobs for their careful review and thought- ful comments. ARDIAGA AND RARPKUIWOARTH AMELICHINA 149 REFERENCES BarkworthH, M. 1993. North American Stipeae (Graminae): taxonomic changes and other comments. Phytologia 74:1-25. Bowoen, W.M. and H.S. Senn. 1962. Chromosome numbers in 28 grass genera from South America. Canad. J. Bot.40:1115-1124. Brako, L.and J.L. ZaruccHi. 1993. Catalogue of the flowering plants and Gymnosperms of Peru. Monogr. Syst. Bot., Missouri Bot. Gard. 45. Brown, W.V. 1952.The relation of soil moisture to cleistogamy in Stipa leucotricha. Bot. Gaz. 113:438-444. HitcHcock. A.S. 1925. North American species of Stipa. Contr. U.S. Natl. Herb. 24:215-262. Jacoss, S.W.L.and J. Everett. 1996. Austrostipa,a new genus, and new names for Australasian species formerly included in Stipa (Gramineae). Telopea 6:579-595. MattHel, O.R. 1965. Estudio critico de las gramineas del género Stipa en Chile. Gayana, Bot. 13:1-137. PENAILILLO, P. 1998. Nuevas combinacidnes en el género Nassella E. Desv.emend.Barkworth (1990) (Poaceae, Stipeae). Gayana, Bot. 55:85-88. PENAILILLO, P. 2002. El género Jarava Ruiz et Pavon (Stipeae-Poaeae): delimitacion y nuevas combinaciones. Gayana, Bot. 59:30. Roic, FA. 1978. Género Stipa L. In: Correa, Flora Patagénica, Coleccidn Cientifica del INTA, 7(3):288-327. Rosencurrt, B., B. ARRILLAGA DE Marrel, and P. Izacuirre De Artucio. 1970. Gramineas Uruguayas. Univers. de la Republica, Depto. Publicacidnes. Montevideo, Uruguay. SoRENG, R.J., BM. Peterson, G. Davipse, E.J. Jupziewicz, F.O. ZULOAGA, T.S. FILGUEIRAS, and O. Morrone. 2003. Catalogue of New World grasses (Poaceae): IV. Subfamily Pocideae. Contr. U.S. Natl. Herb. 48:1-730. Torres, M.A. 1993.Revision del género Stipa (Poaceae) en la Prov.de Buenos Aires. Comision de Investigaciones Cientificas, Prov. Buenos Aires, Monografia 12:1-62. Torres, M.A. 1997. Stipa (Gramineae) del noroeste de la Argentina. Comisién de Investigaciones Cientificas, Prov. Buenos Aires, Monografia 13:42-65. Tovar, O. 1993.Las Gramineas (Poaceae) del Peru. Ruizia 13:1-480. Vertoove, F. 2005.A synopsis of Jarava Ruiz & Pav.and Nassella E. Desv. (Stipa L.,s./.) (Poaceae: Stipeae) in southwestern Europe. Candollea 60:97-117. ULOAGA, F.O., E.G. Nicora, Z.E. RUGOLO DE AGRASAR, O. Morrone, J. PENSIERO, aNd A.M. CIALDELLA. 1994. Catalogo de la Familia Poaceae en la Republica Argentina. Monogr. Syst. Bot., Missouri Bot. Gard. 47:1-178. 150 BRIT.ORG/SIDA 22(1) Book NOTICES Flora of North America (Asteraceae) Fiora oF Nort AMERICA EpitoriAL Committee (Eds.) 2006. Flora of North America North of Mexico. Vol. 19. Magnoliophyta: Asteridae, part 6: Asteraceae, part 1. (ISBN 0-19-530563-9: 978-0-19-530563-0, hbk.). Oxford Univ. Press, 198 Madison Avenue, New York, NY 10016, U.S.A. (Orders: http:// www.oup.com). $95.00, 579 pp., illustrations, maps, 8 1/2" x 11". FLora or NortH AMeriIca EpttortAL Committee (Eds.) 2006. Flora of North America North of Mexico. Vol. 20. Magnoliophyta: Asteridae, part 7: Asteraceae, part 2. ISBN 0-19-530564-7; 978-0-19-530564-7, hbk.). Oxford Univ. Press, 198 Madison Avenue, New York, NY 1OO16, U.S.A. (Orders: http:// www.oup.com). $95.00, 666 pp., illustrations, maps, 8 1/2" x 11" Frora oF NortH America Epiroriat Committee (Eds.) 2006. Flora of North America North of Mexico. Vol. 21. Magnoliophyta: Asteridae, part 8: Asteraceae, part 3. ISBN 0-19-530564-4; 978-0-19-530565-4, hbk.). Oxford Univ. Press, 198 Madison Avenue, New York, NY 10016, U.S.A. (Orders: http:// www.oup.com). $95.00, 616 pp., illustrations, maps, 81/2" x 11" From FNA website.—Covering the Asteraceae, or Composite, family, the largest family in the Flora of North America area, these three volumes contain ne cae 420 species classified in +18 genera and 14 tribes. More than 60% of the s > the flora region and almost 10% are known as introductions that have miatanaticedl Members of ‘his tamily of higher plants can be found to flour- ish in many habitats. Sunflowers, daisies, goldenrods, sagebrush, asters, zinnias, arnicas, chamomile, cory, lettuce, coneflowers, ragweeds, tickseeds, thistles, dandelions, > chrysanthemums, marigolds, ch tarweeds, cockle dee ereen and many other familiar plants are members of the Asteraceae. Each of the genera within these volumes has one or more representative species illustrated with a line drawing; more than 660 species are illustrated. Occurrence maps for nearly 3,300 species anc accompany the identification keys, descriptions, and discussions about these subspecies, b plants. SIDA 22(1): 150. 2006 TAXONOMY OF THE CUSCUTA PENTAGONA COMPLEX (CONVOLVULACEAE) IN NORTH AMERICA Mihai Costea (corresponding author Guy L.Nesom Department of Biology Botanical Research Institute of Texas fa a pallies! University can Stre 5 | West Fort Worth, Texas 76102- He U.S.A. Waterloo, Ontario esom@brit.org N2L 3C5, CANADA mceostea@wlu.ca Sasa Stefanovic Department of Biology University of Toronto at : ississauga 359 Mississauga Road Mississauga, cakes L5L 106, CANADA ABSTRACT Cuscutd pentagona, C. campestris, and C. glabrior are closely similar taxa recognized here at specific a 8 y P rank. Closely related to these are C. runyonii, C. harperi, C. plattensis, and C. sandwichiana, as well as G: pu UoNG var. cle an and C. polygonorum. Cuscuta runyonii, C. harperi, ane Ss ea aan L are relatively narrow € : pla ttensisand C.¢labrior have wider but ay parently while C. obtusiflora var. ¢ Pagid C. polygonorum, and especially C. pentagona ae Ee Peierls are widespread and br oadly sympatric in North America, at least in some regions. Cuscuta campestris is one of the most widespread species in the world, being additionally recorded from South America, Europe, Asia, Africa, and eee Comparisons among these nine species are presented as a key and as comparative d M images are provided for flowers, fruits, seeds, and pollen. RESUMEN Cuscutd ponapane: . canon y GC. ais son taxa fuertemente semejantes, reconocidos aqui como especies as estan C. runyonii, C. harperi, y C. Ss como tambien G; gps ite var, eagles yiG: pono Cuscuta runyonii, C. ee C. plattensis y C.glabrior tienen ran I 14 di 8 eee Cn ca tusiflora var.glandu losa, a y especialmente G pemieepng y.G. eee son comunes y eae iamente Sue en d Norteameéric siendo mencionada a Sudamérica, Europa, Asia, Arica y Australia Las comparaciones entre estos seis Se ofrecen imagenes de SEM de las I E f ores Hates semillas y polen. Engelmann (1859) distinguished three varieties of Cuscuta pentagona Engelm. in addition to the typical form: var. calycina, var. verrucosa, and var. microcalyx. Yuncker (1932, 1965) treated var. calycina and var. verrucosa at specific rank but provided a new name for each: C. campestris Yuncker (nom. nov.) and C. SIDA 22(1 = > 151-175. 2006 152 BRIT.ORG/SIDA 22(1) glabrior (Engelm.) Yuncker (comb. nov.), respectively. He treated var. microcalyx as a synonym of typical C. pentagona. Recent studies have not recognized C. campestrisat any rank (e.g, Beliz 1986; Gandhi et al. 1987), while C. glabrior has been regarded asa variety of C. pentagona (Gandhi et al. 1987) or as nomencla- tural synonym of C. indecora Choisy var. neuropetala (Engelm.) Hitche. (Beliz 1986). Three other species from North America, C. harperi Small, C. plattensis A.Nels.,and C. runyonii Yuncker, were included by Yuncker in the same group (subsect. Arvenses Yuncker) with C. pentagona and C. campestris (Yuncker 1965). Although C. runyonii has been considered a good species by all authors, C. harperi and C. plattensis were treated by Beliz in her Ph.D. dissertation (1986) as nomenclatural synonyms of “C. pentagona var. polygonorum’ and C. indecora, respectively, The current paper is the second of several to present taxonomic analyses of Cuscutad species groups and complexes. The first (Costea et al. 2005) gave an assessment of Cuscuta nevadensis and Cuscuta veatchii; three others follow in this issue (Costea et al 2006a,b,c). Other two papers that are now in preparation will deal with the phylogeny of Cuscuta and its infrageneric taxonomy, includ- ing the formal delimitation of the groups of species treated in the four papers published in this issue. Evaluations of the status of all these taxa are necessary for the accounts of Cuscuta in the Flora of North America and the forthcoming revision of The Jepson Manual (California). The comments below on reproduc- tive biology and taxonomy are in perspective of the whole genus. pes) — REPRODUCTIVE BIOLOGY AND TAXONOMY OF CUSCUTA Scarcity of information regarding the reproductive biology of Cuscuta species limits progress toward understanding the taxonomy and evolution of this ge- nus. For example, a detailed study of floral biology is available only for C. attenuata Waterfall (Prather & Tyrl 1993), which is very closely related to to C. indecora. Although they may be sympatric over an extensive geographical range, species such as C. campestris and C. pentagona have never been found “growing together at the same site” (Musselman 1986). Apparently a single her- baceous host cannot support more than one parasite (often a single Cuscuta plant simultaneously parasitizes several different host plants), although it does not seem impossible that colonies of different Cuscuta species might exist side by side. Furthermore, although some species (e.g., C. epithymum Murray, C. rostrata Shuttlw. ex Engelm. &@ A.Gray) have been reported to be pollinated by insects (Beliz 1986; Musselman 1986; Holm et al. 1997), others such as C. pentagona and its relatives apparently are autogamous (Verdcourt 1948; Beliz 1986; Musselman 1986; Prather & Tyrl 1993; Dawson et al. 1994). In the C. pentagona complex, we observed that anthers dehisce longitudinally before {lowers open, further substantiating previous observations of autogamy. The low pollen/ovule ratio (= 65) found by Beliz 1986) in C. pentagona Gncluding C. COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 153 campestris) supports the idea that autogamy may be common (see Cruden 1977). The occurrence of inbreeding may help explain the relatively high num- ber of taxa (species and varieties) that are recognizable as morphological enti- ties in Cuscuta. The existence of intermediate-like plants observed rarely in populations of the C. pentagona complex probably reflects populational vari- ability in closely related taxa. Hybridization also is a possible explanation, but experimental attempts to produce hybrids (e.g. between C. indecora and C. attenuata) have failed (Prather & Tyrl 1993). Seed dispersal of many Cuscuta species seems unspecialized (Kuijt 1969). “Natural” means of seed dispersal are unknown. Lyshede (1984) suggested that wind may play a significant role in dispersal because of the pits present on the seed coat of dry seeds. However, Cuscuta seeds do not possess “classical” adap- tations for wind dispersal, and the alveolate/papillate seed coat seems more an adaptation related to the germination process (Costea & Tardif 2006). Kuijt (1969) mentioned that seeds remain viable while they pass through the diges- tive system of sheep, but Gdmez (1994) in Sierra Nevada, Spain, reported that hosts infested with C. epithymum were avoided by the Spanish ibex, the main herbivore in the area. Herbivores may avoid plants infested with Cuscuta be- cause of the yellowish color (Costea & Tardif 2006). Capsules and seeds of Cuscuta float, and although water dispersal has never clearly been documented, it may occur only in the species that prefer the vicinity of water, such as C. gronovii Willd.ex J. A. Schult. Dispersal over long distances of weed species such as C. campestris has been connected with anthropomorphic activities. The prin- cipal means of world-wide dispersal of Cuscuta weeds has been through con- taminated seeds of forage legumes (alfalfa, clover, and lespedeza, especially Les- pedeza cuneata (Dumont) G. Don; reviewed by Dawson et al. 1994; Costea & Tardif 2006). Furthermore, when wetted, the seed coat in Cuscuta species be- comes papillose-gelatinous and adheres easily to soil particles, and seeds may thus be carried by farm machinery (Costea & Tardif 2006). METHODS Descriptions of morphology and micromorphology are based mostly on speci- mens from herbarium NY, which includes Yuncker’s herbarium (Appendix 1). Measurements of floral parts except anthers were done on rehydrated her- barium material. Length of flowers was measured from the base of calyx to the tip of straightened corolla lobes. Pollen, anthers, seeds, and capsules were mea- sured on SEM pictures taken with a scanning electron microscope Hitachi S- 570 at 15 KV. Samples were coated with 30 nm gold using an Emitech K 550 sputter coater. Standard terminology for cell types and surface sculpturing pat- terns follows Barthlott et al. 1998); pollen terminology follows Hoen (1999). Conservation status was assessed using NatureServe (2005) ranks and criteria. Identification of Cuscuta often isa lengthy process because rehydration of 154 BRIT.ORG/SIDA 22(1) flowers, dissection, and examination under a microscope are usually necessary. Variation may be expected on the same individual (e.g, Gandhi @ Thomas 1983; Severova 1991). Furthermore, because flowers do not mature simultaneously, the size and ratios between different floral parts on the same plant may vary to some extent. A range of flowers, fruits, and seeds belonging to the same plant should be examined. The angled calyx of C. pentagona and C. harperi is best ower and during early fructification; full development of capsules — —_— observed inf may obscure the character. MICROMORPHOLOGY Flowers and capsules.—Laticifers, papillae, and multicellular protuberances may occur on the floral parts and capsules and distinctions between these features can be important in recognizing members of the C. pentagona complex (mag- nification of LOOX or more recommended). Laticifers are visible in the calyx and sometimes in the corolla and capsules of all taxa. Yuncker usually referred to them as “pellucid glandular-appearing cells” (Yuncker 1921, 1932, 1965). La- ticifers may appear isolated, rotund, ovoidal or tangentially elongated, or they can be organized in rows (Fig. 1 c,e). Laticifers from the stems are perivascular, multinucleate, and articulated (Lyshede 1985). Papillae are more or less radi- ally elongated cells, and they are present in the calyces, corollas, and capsules of C. harperi, C.glabrior, and C. runyonii (Fig. Le,f; 2. a,b,c). Multicellular protu- berances are larger and they may occur in C. runyonii along the midveins of the calyx lobes (Fig. 2c). Epicuticular wax on the perianth, when present, is repre- sented by longitudinally reticulated rodlets (Fig. Lf). Pollen.—Pollen is relatively uniform among species. Grains are 3(-4)- zonocolpate (rarely +-6 loxocolporate in aberrant grains),and they may be poly- morphic in the same anther or flower, from spheroidal to prolate (Fig. 3). Orna- mentation varies from tectum imperforatum or witha few puncta in C. glabrior, C.runyonii,and C. plattensis, to tectum perforatum in C. pentagona, C. campestris, C. harperi, and C. obtusiflora var. glandulosa (Fig. 3). The latter three species have sexine often persistent in punctum apertures (Fig. 3). Cuscuta pentagona and C. obtusiflora var. glandulosa have the largest puncta, up to 0.7 fm in diam- —_— = eter, sometimes approaching a microreticulate ornamentation (Fig. 3). Seeds.—A striking feature of the seed coat epidermis is that when seeds dry, cells of the outer cell wall invaginate, which causes the seed surface to be- come alveolate (Fig. 4; see also Lyshede 1984; Knepper et al. 1990; Costea & Tardif 2006). Water uptake induces bulging of the invaginated epidermal walls, and epidermis cells become swollen and papillose (Fig. 4c). The diameter of seed coat epidermal cells is 17-50 pm in C. pentagona, C. campestris, C. glabrior, C. runyonii, and C. harperi and 15-20 um in C. plattensis, C. obtusiflora var. glandulosa, and C. polygonorum. The seeds of C. runyonii and C. polygonorum have epicuticular wax organized as longitudinal rodlets (Fig. 4d), a feature not COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX Fic. 1.a. Calyx of C. campestris (scale bar = 1 mm);b. Calyx of C. p (scale t 0.5 ey | 1 capsule of C. ); d. Flower of C. (scale | 0.75 mm); e. Flower of C. bar = 0.5 mm); f. P pill th lla lob fC harperi (scale bar = 0.43 um). with ithma arperi (scale 156 BRIT.ORG/SIDA 22(1 = Fic. 2. a. Flower with maturing capsule of C. gigenes (scale ae 1 mm); b. panies on the capsule of C. glabrior (scale bar = 0.75 Lim); c. Flower of C. runyonii projections; d. Flower of C. plattenis (scale bars = 1mm). observed in the other species. The hilum is round to broadly elliptic; the cells are smaller than in the rest of the seed coat and they are radially oriented around the vascular scar of the funiculum (Fig. 4 b,d). The vascular scar is linear, ob- lique to vertical (on broadest diameter of the seeds), 0.025-0.09 mm in C. pentagona, C.campestris, C.glabrior, C. pe C. harperi,and C.sandwichiana and 0.125-0.25 mm in C. plattensis, C. obtusiflora var. glandulosa, and C. polygonorum (Fig. 4 ae,f). TAXONOMY This g er oup of species is characterized by globose or mostly depressed-glo capsules with relatively large interstylar apertures, which sometimes are thick- COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 157 eee rectum Sanu but sexine he S Saat of pollen a-b. bua bast grains oe nee «Cp is (scale bar = 8.6 um); d. C. BRIT.ORG/SIDA 22(1) Fig. 4, Malahat f Is. a. C. campestris, ventral view (scale ba ranvoti teal bar = 120 j1m); e, See (scale bar = 0.5 mm). = 0.5 mm); b. Hilum and alveolate seed coat of C. = 150 um); ¢. Ppl on yaa jaa co tof gna Gale a = a yale d. Hilum of ¢. dof © harp C. plattensis, ventral COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 159 ened but not raised to form a stylopodium. The corolla is persistent at the base of the capsules or surrounding them. Cuscuta sand wichiana, a Hawaiian endemic, was placed by Yuncker (1932) in subsect. Californicae because it exhibits a reduction of the infrastaminal scales similar to the other species (C. sandwichiana was omitted in his 1965 treat- ment). This species, however, does not appear to be evolutionarily allied to C. californica complex. Beliz 1986) hypothesized that C. sandwichiana has evolved from C. cephalanthi Engelm., which is a member of the C. gronovii Willd. ex Roem. & Schult. complex. However, the stems of C. sandwichiana grow and branch as in species of the C. pentagona complex: the main and secondary stems grow continuously and never twine around the host. Instead, tendril-like branches produced from axillary buds of the rudimentary leaf-scales fix the parasite to the host. In C.gronovii and its relatives (e.g.,C. cephalanthi, C. umbrosa Beyr. ex Hook., and C. rostrata Shuttlw. ex Engelm. & A. Gray), haustoria are formed as the main stem twines around the host, and no tendril-like axillary branches are generated (as first observed by Dawson 1984 in C. gronovii). The calyx and corolla of C. sandwichiana are relatively similar in morphology to those of C. campestris, although flowers are larger in C. sandwichiana. Seeds of C.sandwichiana are commonly angled, like those of C. campestris, and not dor- soventrally compressed as in C. cephalanthi. Contrary tosome relatively recent treatments (e.g. Austin 1986; Musselman 1986), C. campestris has not been accepted as a distinct taxon in modern North American overviews of the genus (Beliz 1986; Gandhi et al. 1987; Kartesz et al. 1999; USDA NRCS 2004). This judgment has been based on the allegedly vari- able morphology of the calyx, which was the main character used by Yuncker to differentiate this taxon from C. pentagona s. str. Beliz (1986) wrote: “The angled calyx (of C. pentagonas. str.) is not a constant feature, it is present in some flow- ers and absent on others in an individual, ... this feature may be an artifact cre- ated when flowers dry out.” In contrast, we have found the angled calyx of C. pentagona to bea remarkably constant feature (Fig. 1b). Because the calyx lobes of C. campestris are overlapping, the calyx sometimes may appear angled in pressed flowers, but if the flowers are hydrated, the calyx usually appears “rounded” (Fig. la). The prominent angles of the calyx in C. pentagona are the result of a different morphology of the calyx lobes (Fig.1b). Furthermore, small but consistent differences in the size of floral parts, pollen, capsules, and seeds (see key below) usually separate the two taxa. Size overlap between the two taxa does occur, as well as intermediate-like plants, although the latter are uncom- mon. Austin (1979) discussed methods of pressing to avoid obscuring calyx traits in all species, but particularly in C. pentagona and C. campestris. Four years before Yuncker formally recognized Cuscuta campestris as a separate species from C. pentagona, he was aware that his taxonomic solution might be received with reluctance. Ina letter addressed to WC. Ferguson, accom- 160 BRIT.ORG/SIDA 22(1) panying a collection of C.campestrissent to him for identification (Ferguson 7795, NY), Yuncker (1929) wrote: “I suspect I will be accused of various bad tendencies in the way of species making, but I think I have clarified the ‘arvensis’ situation somewhat.” The assessment presented here also finds that C. campestris is con- sistently distinct from C. pentagona, although differences may seem subtle, and we believe their recognition as species best reflects the biological situation. Similarly, Cuscuta campestris may be sometimes difficult to distinguish from C. glabrior, but the two taxa can be usually differentiated using a combi- nation of features indicated in the key and descriptions. Cuscuta runyonii is most similar to C. glabrior, and C. harperi to C. pentagona (see descriptions; Yuncker 1932; 1942). Cuscuta plattensis does not appear to be evolutionarily al- lied to C. indecora, in contrast to the suggestion by Beliz (1986). Yuncker (1932, p. 140) considered C. plattensis to be “very closely related” to C. campestris. The morphology of seeds of C. indecora is different from the species of C. pentagona complex and more similar to that of C. gronovii and its relatives. Two North American taxa previously treated within Cuscuta subsect. Platycarpae (Yuncker 1932), C. obtusiflora var. glandulosa Engelm. and C. polygonorum Engelm. are included here in subsect. Arvenses on the basis of their overall morphological similarity with the other species of this subsection. This close relationship was also emphasized by Beliz (1986), who referred to C. polygonorum as “C. pentagona var. polygonorum,” within which C. harperi was included asa synonym. In contrast, we found that although C. polygonorumand C. obtusiflora var. glandulosa apparently are related to the species of subsect. Arvenses, each is clearly delimited morphologically. Cuscuta australis R. Br from Australia and Asia, whichis known only from one collection in North America Jefferson Co., New York, 1926, Muenscher s.n., NY), probably belongs to this group of species as well. This latter species has not been included in this taxo- nomic treatment limited in scope to North America. In the Cuscuta pentagona c th ively narrow endemics (C. runyonii, C. harperi, and C. sandwichiana), two with wider but apparently discrete ranges (C. plattensis and C. glabrior), and four species (C. pentagona, C. campestris, C. polygonorum, and C. obtusiflora var. glandulosa) widely distrib- uted and broadly sympatric, at least in some regions. The exact North Ameri- can distributions of C. campestris and C. pentagona, particularly, require addi- tional floristic study. For additional synonymy see Yuncker (1932, 1965). — 1 KEY TO SPECIES OF THE CUSCUTA PENTAGONA COMPLEX IN NORTH AMERICA 1. Infrastaminal scales well-developed, densely fringed or fimbriate. 2. Corolla lobes obtuse, straight 8.C. obtusiflora var. glandulosa 2, Corolla lobes acute, inflexed at apices. 3. Calyx angled, loose around the corolla tube, 0.3-1.1 mm long; lobes broadly- ovate rhombic, + auriculate, overlapping and forming prominent angles at COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 161 sinuses; flowers 0.9-2.5 mm one: corolla tube 0.5-1.2 mm; anthers 0.2-0.35 mm long; capsules 1.9-2.4 Xx 1 4. Flowers commonly 5-merous, 1.4—2.1(—2.5) mm long; calyx lobes not cari- nate and without rows of prominent laticifers along midveins (Fig. 1 b, d); seeds usually 4 per capsule, angled, subrotund to broadly elliptic, longitu- dinal groove absent 1.C. pentagona 4. Flowers commonly 4-merous, 0.9-1.1(-1.5) mm long; calyx lobes carinate or with rows of prominent laticifers along midveins (Fig. le); seeds usually 1 per capsule, almost spherical, with a longitudinal groove on the ventral face (Fig. 4e) 2.C. harperi . Calyx rounded, closely enclosing the corolla tube, 1-2.1 mm long;lobes ovate- triangular to triangular, not auriculate, overlapping or not and not forming prominent angles at sinuses; flowers 2.1-4.6(-5) mm long, corolla tube 1.1- 2.5 mm; anthers 0.4-0.7 mm long; capsules 1.3-2.8 X 1.9-3.8 mm 5. Calyx lobes about as wide as long, evidently overlapping (Fig.1a); persis- tent corolla enveloping 1/3 or less of the capsule (the basal portion) (Fig. 1 Ww C). 3.C. campestris 5. Calyx lobes longer than wide, not rei or only slightly so; persis- tent corolla enveloping 1/2 or more of the capsule. 6, Flowers (whole flowers or sometimes ee mae lyx) and re papillate; reddish-brown when dried, corolla erpandiic. alehere cate between the lines of stamen attachments; infrastaminal scales ing corolla tube; seeds an 7. Calyx without Sane Bee (Fig. 2a) 4.C. glabrior 7. Base of calyx corresponding to each lobe with a reflexed spur-like multicellular projection 0.1-0.6 mm long, and often with smaller mul- ticellular projections on the midveins of calyx lobes (Fig. 2c) 5.C. runyonii 6. Flowers (calyx, corolla, ovary) and capsules not papillate, creamy white when dried; corolla tube narrowly campanulate to cylindric-campanu- late, not saccate between the lines of stamen attachments (Fig. 2d); infrastaminal scales shorter than corolla tube; seeds dorsoventrally compressed (Fig. 4f) 6.C. plattensis 1. Infrastaminal scales absent or reduced, bifid, or with a few distal teeth or fimbriae. 8. Flowers (4-)5-merous, 3-4(—5) mm;infrastaminal scales absent or reduced, trun- cate or triangular, bifid or distally with a few teeth or fimbriae; endemic to Hawaii 7.C, sandwichiana 8, Flowers (3-)4-merous, 2—2.7 mm; infrastaminal scales oblong, mostly shallowly bifid with 1-3 fimbriae on each side of the filament attachment or a few irregu- lar fimbriae at the apex; widespread distribution in N America 9.C. polygonorum |. Cuscuta pentagona Engelm., Amer. J. Sci. Arts 43:340. 1842. Cuscuta arvensis Beyr. ?ngelm. var. pentagona (Engelm.) Engelm., Trans. Acad. Sci. St. Louis 1:494. 1859. LECTO- TYPE (Yuncker 1932): U.S.A. VIRGINIA: Norfolk, “On Euphorbia or Tragia,” 1849, Rugel s.n. (MO). Cuscuta pentagona Engelm. var. microcalyx Engelm., Amer J. Sci. 45:76.1843. LECTOTYPE (Yuncker 1921): U.S.A. ILLINOIS. [Cass Co.| Beardstown, 1842, Geyer s.n. (MO). Engelmann cited only “Illinois.” Cuscuta arvensis Beyr. ex Engelm. in A. Gray, Man. Bot. (ed. 2) 336. 1856. Epithymum arvense (Beyr. ex Engelm.) Niewl. & Lunell, Amer. Midl. Naturalist 4:51. 1916. Type: Engelmann cited only “(in herb. Berlin),” perhaps referring to a Beyrich collection. As noted by Yuncker (1921, p.51),a MO collection by Beyrich in 1845 from North Carolina may be “the type?” (an isotype). Evidently 162 BRIT.ORG/SIDA 22(1) Engelmann intended C. arvensisasa broad concept variants might be iden- tified as C. pentagona, C. pentagona var. microcalyx, C. pentagona var. calycina, or C. verrucosa. Stems |.4-2.1(-2.5) mm in diameter, yellow to orange. Inflorescences dense, corymbiform to glomerulate of 3-15(-20) flowers; pedicels 0.5-3(-4.5) mm; bracts | at the base of clusters and 0-1 at the base of pedicels, 0.4-2.2 x 0.2-13 mm, membranous, ovate, ovate-triangular to lanceolate, margins entire, apex acute. Flowers (4-)5-merous, 1.4-2.1(-2.5) mm (Fig. 1d), membranous, whitish when fresh, yellowish to brown when dried; papillae sometimes present on the corolla lobes; laticifers evident in the calyx and less obvious in the corolla and ovary/capsule, isolated or arranged in rows, ovoid or elongated; calyx yellow to brown, * reticulate, shiny or not, angled, cupulate, ca. as long as the corolla tube, divided 1/2-2/3 the length, lobes overlapping and forming prominent angles at sinuses, broadly-ovate to rhombic, auriculate, not carinate, margins entire, apex rounded (Fig. 1b); corolla persistent; tube campanulate, 0.7-1.2 mm, not saccate between the lines of stamen attachments; lobes spreading, 0.65-1.1 mm long, triangular-lanceolate, margins entire, apex acute to acuminate, in- flexed; stamens exserted, shorter than corolla lobes, anthers subrotund to broadly elliptic, 0.25-0.30 x 0.20-0.28 mm, filaments 0.3-0.4 mm; pollen grains 18-25.2 um long; tectum perforatum; maximum diameter of puncta 0.6 pm; supieicelal Sea eae to conical + uniformly distributed (Fig. 3e); rolla tube, oblong-ovate, rounded, + uniformly, dense fimbriate; styles everily filif orm, O./-1.1 mm, ca.as long as the ovary; stig- mas capitate, globose. Capsules indehiscent to irregularly dehiscent, globose- depressed to ovoid, 1.9-2.4 x 1.6-2.5 mm, not thickened or raised around the medium to large interstylar aperture, translucent or not, persistent corolla at the capsule base. Seeds 4 per capsule, angled, subrotund to broadly elliptic, 0.9- Ll x 0.8-1 mm, seed coat cells reticulate to alveolate/papillate, 18-35 tm in diameter, epicuticular wax absent; hilum region round 0.4-0.5 mm in diam- eter, vascular scar 0.07-0.09 mm long, vertical to slightly oblique. 2n = 56 (Fogelberg 1938), ca. 44 (Pazy & Plitmann 1995). Distribution and ecology.—CANADA: Manitoba (based on only 2 collections from DAO and MTMG respectively). U.S.A.: Alabama, Arkansas, D.C., Delaware, Florida, Georgia, Kansas, Illinois, Indiana, Maryland, Massachusetts, Michigan, Minnesota, Mississippi, Missouri, Montana, New Jersey, New York, North Caro- lina, North Dakota, Oklahoma, Pennsylvania, South Carolina, South Dakota, Tennessee, Texas, Virginia, Washington. We have seen no evidence that C. pentagona has spread outside of North America, where it is less common than C.ca mpestris. Isolated European records identified as this taxon (e.g, Buia 1938) are instead C. campestris. Flowering Jun-Nov. Hosts: numerous hosts (Gaertner 1950) growing in a wide variety of climates and soil types. Conservation status.—G4G5 (apparently secure to secure) (G5, NatureServe 2005). COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 163 2. Cuscuta harperi Small, Fl. S.E. US. ed. 2, 1361, 1375. 1913. Type: US.A. Georcia: Altamaha Grit, between Peacocks and Harrison, on Chond rophora virgata, 20 Jul 1906, Harper s.n. (HOLOTYPE: NY, see Yuncker 1943; ISOTYPE: NY). Stems 0.15-0.3 mm in diameter, orange-yellow. Inflorescences loose, corymbiform of 2-10(-15) subsessile or short pedicellate flowers; pedicels 0.5- 2.5(-3) mm; bracts | at the base of clusters and absent (rarely present) at the base of pedicels, like those of C. pentagona. Flowers 4(-5)-merous, 0.9-1.1(-L5) mm (Fig. le), fleshy, creamy-white when fresh, yellow to brownish when dried; papillae present on the corolla lobes (Fig. Lf); laticifers evident along midveins of calyx and less obvious in the corolla and ovary/capsule, isolated or in rows, elongated; calyx yellow-brownish, + reticulate, rarely shiny, about as long as the corolla tube, angled, cupulate, lobes basally overlapping and forming promi- nent angles at sinuses, broadly-ovate rhombic, + auriculate, + carinate or with laticifers along midveins, margins entire, apex rounded; corolla persistent; tube campanulate, not saccate, 0.5-0.7 mm; lobes erect to reflexed, 0.4-0.7 mm, tri- angular-ovate, margins entire, apex subacute to acute, inflexed; stamens ex- serted, slightly shorter than corolla lobes, anthers subrotund to broadly ellip- tic, 0.2-0.25 x 0.15-0.18 mm, filaments 0.16-0.25 mm; pollen grains 17.5-25 um long; tectum perforatum but sexine often persistent in puncta aperture; diam- eter of puncta 0.4 um; supratectal process granular granular to conical + uni- formly distributed (Fig. 3 a,b); infrastaminal scales as long as the corolla tube, narrowly oblong, rounded, shortly and irregularly-dense fringed in the distal half; styles uniformly narrow terete to filiform, 0.5-0.9 mm, shorter than the ovary; stigmas capitate, globose. Capsules indehiscent to irregularly dehiscent, globose to ovoid, 1.2-2.3 x 1.2-1.6 mm; not thickened around the small to mod- erately large interstylar aperture, almost translucent, with persistent corolla enveloping the 1/4-1/3 of the capsule bases. Seeds usually 1-2 per capsule, subglobose, with a longitudinal groove on the ventral face, 0.92-1.11 x 0.85-L1 mm, seed coat cells alveolate/papillate, 35-50 um in diameter; epicuticular wax absent; hilum region terminal, round, 0.14-0.16 mm in diameter, vascular scar 0.025-0.03 mm long, vertical to slightly oblique. 2n = ? Distribution and ecology.—U.S.A.: Alabama and Georgia. Flowering Sep- Nov. Hosts: Bigelowia nuttallii, Croton willdenowii, Helianthus longifolius, Hy- pericum gentianoides, Liatris microcephala, and other species of sandstone outcrops (see also Yuncker 1943). onservation status.—G1G2 (critically imperiled to imperiled) (G2, NatureServe 2005). 3. Cuscuta campestris Yuncker [nom. nov], Mem. Torrey Bot. Club 18:138. 1932. Based on Cuscuta pentagona Engelm. var. calycina Engelm., Amer. J. Sci. 45:76. 1843 [1845]. Cuscuta arvensis Beyr. ex Engelm. var. calycina (Engelm.) Engelm., Trans. Acad. Sci. St. Louis 1:495. 1859. Lectotype (Yuncker 1921): U.S.A. TEXAS: wet prairies, [no date], Lindheimer 126 (MO). Stems 0.3-0.5 mm in diameter, yellow to orange. Inflorescences dense, 164 BRIT.ORG/SIDA 22(1) corymbiform or glomerulate of (3-)6-25(-30) subsessile to short pedicellate flowers; pedicels 0.3-2.5(-3.5) mm; bracts | at the base of clusters and O-1 at the base of pedicels, like those of C. pentagona. Flowers (4-)5-merous, (1.9-)2.1-3.6 mm (Fig. Ic), membranous, white-creamy when fresh, creamy or golden-yel- low when dried; papillae absent; laticifers evident in the calyx and less obvious in the corolla and ovary/capsule, isolated or arranged in rows, rotund, ovoid or elongated; calyx yellow, reticulate, shiny, rarely obscurely angled, cupulate, about as long as corolla tube, divided 2/5-3/5 the length; lobes overlapping but not forming or rarely forming obscure angles at sinuses; ovate triangular, not cari- nate, margins entire, apex obtuse to rounded (Fig. la); corolla persistent; tube campanulate, not saccate, (1.1-)1.5-1.9 mm; lobes spreading, triangular-lan- ceolate, (1.1-)1.4-175 mm long, margins entire, apex acute to acuminate, inflexed; stamens exserted, shorter than corolla lobes, anthers broadly elliptic, (0.3-)0.4- 0.5 x 0.25-0.3 mm, filaments 0.4-0.7 mm long; pollen grains 18.5-28 um long; tectum perforatum but sexine often persistent in puncta aperture; maximum diameter of puncta 0.4 um; supratectal process granular, often in groups of 2-4 (Fig. 3f); infrastaminal scales equaling or slightly exceeding corolla tube, ob- long-ovate to spathulate, rounded, uniformly dense fimbriate; styles evenly fili- form, 0.8-16 mm, ca. as long as the ovary; stigmas capitate, globose. Capsules indehiscent to irregularly dehiscent, globose-depressed to depressed, 1.3-2.8 x 1.9-3.8 mm; not thickened or raised around the large interstylar aperture, some- times translucent, persistent corolla enveloping 1/3 or less of the capsule base. Seeds 4 per capsule, angled, subrotund to broadly elliptic, 112-154 x 0.9-L1 mm, seed coat cells alveolate/papillate, 32-40 um in diameter; epicuticular wax absent; hilum region subterminal, round, 0.45-0.5 mm in diameter, vascular scar, 0.085-0.01 mm long, vertical or slightly oblique (Fig. 4 a,b). 2n = 56 (Fogelberg 1938; Ward 1984). Distribution and ecology.—CANADA: Alberta, British Columbia, Saskatchewan, Manitoba, Nova Scotia, Ontario and Québec. Darbyshire (2003) also mentioned it from NewFoundland, New Brunswick and Prince Edward Island. Although we found no herbarium collections, its presence in these prov- inces is possible. U.S.A.: Arizona, Arkansas, California, Colorado, Florida, Geor- gia, Idaho, Illinois, Indiana, lowa?, Kansas, Kentucky, Louisiana, Massachusetts, Maryland, Michigan, Minnesota, Mississippi, Missouri, Montana?, Nebraska, Nevada, New Mexico, New Jersey, New York, North Carolina, North Dakota, Ohio, Oklahoma, Oregon, Pennsylvania, South Dakota, Tennessee, Texas, Utah, Vir- ginia, Washington, West Virginia, Wisconsin, Wyoming. Cuscutd campestris is the second most common species in North America, after C. gronovii (Yuncker 1932), and perhaps the most successful and widespread Cuscuta weed world- wide, recorded from South America, Europe, Asia, Africa, and Australia (Holm et al. 1997). Flowering Jun-Nov. Hosts: numerous species from hundreds of gen- era in various families, including Hydrophyllaceae, Chenopodiaceae, COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 165 Verbenaceae, Convolvulaceae, Acanthaceae, Brassi Euvhorbi Care aceae, Urticaceae, Polygonaceae, Fabaceae, Asteraceae, and others (Gaertner 1950), growing in a wide variety of climates and soil types; reported not to sur- vive on Amaranthus retroflexus, Arctium lappa, Atriplex spp., Brassica nigra, Equisetum arvense, Glycine max, selected cultivars of Lycopersicon esculentum Portulaca oleracea, Tanacetum vulgare, Vicia villosa, Cyperaceae and Poaceae (Gaertner 1950; Parker & Riches 1993; Dawsonet al. 1994; Costea & Tardif 2006). Conservation status.—G5 (common) (the same assessment in NatureServe 2005). — 4. Cuscuta glabrior (Engelm.) Yuncker, Mem. Torrey Bot. Club 18:140. 1932. Cuscuta pentagona Engelm. var. ae brior (Engelm.) Gandhi, Thomas, & Hatch, Sida 12:372. 1987. Cuscuta verrucosa Engelm. var. glabrior Engelm., Amer. J. Sci. 43:341. 1842. LECTOTYPE (Yucker 1932): U.S.A. TEXAS. [Harris Co.?|: dry sterile prairies west of Houston, Lindheimer s.n. ( The protologue noted that var. glabrior [was growing] “with the preceding variety [ie., C verrucosa var. hispidulal, F Lindheimer: on Petalostemon, Drummond (Grd collection, No. 247).” Choice of the Drummond collection (III, 247) as lectotype of Cuscuta verrucosa Engelm. var. glabrior has been associated with Yuncker (1921), but there Yuncker instead chose the DEdnond type for Cpe ntagona Var. VErrucosd, citing C. verrucosa var. glabr iorasa synonym. Cuscuta verrucosa Engelm., Amer. J. Sci. 43:341. 1842 oo. invalid, non Sweet 1823). Cuscuta rvensis var. verrucosa Engelm., Trans. Acad. Sci. St. s 1:495. 1859. Cuscuta pentagona Engelm. var. verrucosa (Engelm.) Yuncker, Illinois it Monogr. 6:142. 1921. LECTOTYPE (Yuncker 1921): U.S.A. TEXAS: no other locality, Drummond III 247 (MO; ISOLECTOTYPE: TEX). Engelmann described C. verrucosa and two varieties of it: var. hispidula (treated by us as C. indecora var. indecora) and var. glabrior. He cited calectoe associated with both varieties but none for C. verrucosa in its typical form. In 1859, in valid use of the epithet as C. arvensis var. verrucosa, he cited “Texas, Drummond! III. 247; Lindheimer! 127” and others as syntypes. Yuncker (1921, p. 52), under the combination C. pentagona var. verrucosa, indicated that “Drummond III 247” at MO was the type (lectotype). Cuscuta arvensis Beyr. ex Engelm. var. pubescens Engelm., Trans. Acad. Sci. St. Louis 1:495. 1859. Cuscuta pentagona Engelm. var. pubescens (Engelm.) Yuncker, Illinois Biol. Monogr. 6:142.1921. Cuscuta glabrior var. pubescens (Engelm.) Yuncker, Mem. Torrey Bot. Club 18:141. 1932. LEc- TOTYPE (Yuncker 1921): “Texas (... L ica in ee )’? (MO). Engelmann’s protologue cited “Western Texas, Lindheimer! Wright! 135 (57 Stems 0.35-0.5 mm in diameter, creamy. Inflorescences loose to compact, glomerulate or corymbiform of 3-25(-30) subsessile to pedicellate flowers; pedicels 0.8-4(-5) mm; bracts like in C. campestris. Flowers 5-merous, |.4-2.1(- 2.5) mm (Fig. 2a), membranous, white or commonly reddish when fresh, yel- low to reddish-brown when dried; papillae present on the perianth, ovaries/ capsules and sometimes on the pedicels (Fig. 2b); laticifers obvious in the calyx and less evident in the corolla and ovary /capsule, isolated or arranged in rows, rotund, ovoid or elongated; calyx yellow to reddish-brown, shiny, + reticulate, cupulate, about as long as corolla tube; divided 1/2-2/3, lobes not overlapping, ovate triangular, not carinate, margins entire, apex obtuse to subacute; corolla persistent; tube campanulate-globose, saccate between the lines of stamen at- tachments, (1.1-)1.5-2.1 mm long; lobes spreading to reflexed, lobes 1.2-1.75mm 166 BRIT.ORG/SIDA 22(1) long, triangular to sublanceolate, margins entire, apex acute to acuminate, in- flexed; stamens exserted, shorter than corolla lobes, anthers broadly elliptic to elliptic-oblong, 0.44-0.67 X 0.22-0.3 mm, filaments 0.4-0.7 mm; pollen grains 18-22.6 um long; tectum imperforatum or witha few puncta; maximum diam- eter of puncta 0.2 1m; supratectal process granular to conical + uniformly dis- tributed; infrastaminal scales reaching the filament bases, ovate to spatulate, rounded, uniformly dense fimibriate; styles evenly filiform, 0.9-1.6 mm, as long as or longer than the ovary; stigmas capitate, globose. Capsules indehiscent to irregularly dehiscent, globose-depressed to depressed, 1.5-2.8 x 2.1-3.5mm, not thickened or raised around the medium to large interstylar aperture, not trans- lucent, persistent corolla enveloping 1/2-2/3 of the capsule. Seeds 4 per cap- sule, angled, subrotund to broadly elliptic, 0.95-L15 x 0.85-1 mm, seed coat cells alveolate/papillate, 35-50 um in diameter (Fig. 4c), epicuticular wax absent; hilum region subterminal, round, 0.13-0.2 mm in diameter, vascular scar 0.03- 0.05 mm long, oblique. 2n = ? Distribution and ecology.—U.S.A.: Louisiana, New Mexico, Oklahoma, Texas, Utah; Mexico. Flowering Jun-Sep. Hosts: wide variety of herbaceous spe- cies, including Amaranthus, Ambrosia, Amphiachyris, Asclepias, Convolvulus, Coreopsis, Croton, Dalea, Dyschoriste, Evolvulus, Gaura, Gilia, Hedeoma, Helenium, Lespedeza, Liatris, Machaeranthera, Medicago, Mimosa, Plantago, Prosopis, Tragia, Oenothera, Justicia, Polygonum, Ruellia, Solanum, Symphyotrichum, Thelesperma, Verbena. Conservation status.-G4G5 (the same assessment in NatureServe 2005). According to Yuncker (1965, p. 6), Cuscuta pentagona var. pubescens (Engelm.) Yuncker “represents the extreme degree of papillation which covers the flower and extends down onto the pedicels. In var. glabrior the ovary and capsules are almost always more or less papillate while the corolla shows con- siderable variation in this character.” We find continuous variation, and two varieties cannot be separated. Occasional plants that are hardly papillate can be distinguished from C. campestris by their calyx lobes, which are broadly triangular-lanceolate and not overlapping at the base and by the corolla envel- oping the mature capsule (Fig. 2a). ~— 5. Cuscuta runyonii Yuncker, Bull. Torrey Bot. Club 69:541. 1942. Type: U.S.A. TEXAS. Hidalgo Co.: La Joya, occasional, on dry hill tops, clay, 45 m, flowers cream white, hosts: Coldenia canescens and Nama hispida, 8 Jun 1941, Runyon 2732 (HOLOTYPE: US; ISOTYPES: F, GH, MO, NY, TEX). Stems 0.35-0.5 mm in diameter, yellow-orange. Inflorescences loose, + umbel- late of 2-15(-25) subsessile to pedicellate flowers; pedicels 0.8-4(-5) mm; bracts as in C. campestris. Flowers 5-merous, 2.5-3.5(-4) mm (Fig. 2c), white to creamy when fresh, reddish-brown when dried, membranous, papillae present on the perianth, ovary/capsules and sometimes on the pedicels; laticifers evident in —_— COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 167 the perianth and less obvious in the ovary/capsule, isolated or arranged in rows, rotund, ovoid or tangentially elongated; calyx reddish-brown, + reticulate, shiny, cupulate, shorter than corolla tube or about equaling it, divided 1/2-2/3 the length, lobes not basally overlapping, triangular, sometimes carinate, base of calyx corresponding to each lobe with a reflexed spur-like projection, 0.1-0.6 mim long (Fig. 2c), and often with smaller protuberances on the midveins of calyx lobes, margins entire, apex obtuse to subacute; corolla persistent; tube campanulate-globose, saccate between the lines of stamen attachments, (1.2- )1.6-2.3 mm, lobes reflexed, 1.2-2 mm, triangular-ovate to lanceolate, margins entire, apex acute to acuminate, inflexed; stamens exerted, shorter than corolla lobes anthers broadly elliptic to elliptic-oblong, 0.4-0.6 x 0.25-0.3 mm, fila- ments 0.4-0.6 mm long; pollen grains 18-23.8 ,1m long; tectum imperforatum or with a few puncta; maximum diameter of puncta = 0.3 um; supratectal pro- cess granular to conical + uniformly distributed; infrastaminal scales reach- ing the filament bases, oblong to spatulate, rounded, uniformly dense fimbri- ate; styles evenly filiform, 0.9-1.8 mm, equaling to longer than the ovary; stigmas capitate, globose. Capsules indehiscent to irregularly dehiscent, globose-de- pressed, 1.6-3 X 2-3.4 mm; not thickened or raised around the large interstylar aperture large, not translucent, persistent corolla enveloping 1/2-1/lof the cap- sule. Seeds 4 per capsule, angled, ovate to broadly elliptic, 0.95-1.40 x 0.85-L1 mm, seed coat cells alveolate/papillate, 30-50 um in diameter; epicuticular wax present with longitudinal; rodlets hilum region subterminal round to subrotund, 0.30-0.40 mm, vascular scar, 0.08-0.09 mm long, oblique (Fig. +d). 2n =? Distribution and ecology.—U.S.A.: Texas. Flowering Mar-Apr(-May, -July, - Oct). Hosts: species of Dalea, Dyschoriste, Erigeron, Gutierrezia, Hymenoxys, Justicia, Linum, Melampodium, Oenothera, Nama, Spermolepis, Tetraneuris, Tiquilia, Thamnosma, Thelesperma. Conservation status.—G3 (vulnerable) (G4, NatureServe 2005). 6. Cuscuta plattensis A. Nels. Bull. Torrey Bot. Club 26:131. 1899. Type: U.S.A. WYOMING. [Carbon Co.?| Platte Canyon, 27 Aug 1896, A. Nelson 2768 (HOLOTYPE: RM; IsOTYPEs: NY, US). Stems 0.15-0.3 mm in diameter, yellow to pale orange. Inflorescences loose, paniculiform of 2-10(-15) subsessile to pedicellate flowers; pedicels 0.5-2.5(-3) mm.; bracts asin C. campestris. Flowers 5-merous, 3-4.6(-5) mm (Fig. 2d), mem- branous, white-creamy when fresh, whitish-yellow when dried; papillae ab- sent; laticifers present in the calyx and less obvious in the corolla and capsule, isolated, ovoid to elongated; calyx yellow, reticulate, not shiny, cylindric-cupu- late, ca. equaling the corolla tube, divided 1/2-2/3, lobes slightly basally over- lapping, ovate triangular, not carinate, margins entire, obtuse to subacute; co- rolla persistent; tube narrowly campanulate to cylindric campanulate, not saccate, 1.9-2.5 mm, lobes spreading to reflexed, 1.4-2.2 mm long, triangular, 168 BRIT.ORG/SIDA 22(1) margins entire, apex narrowly acute, inflexed; stamens exserted, shorter than corolla lobes anthers subrotund, broadly elliptic to elliptic-oblong 0.4-0.7 x 0.3-0.4 mm, filaments 0.3-0.4 mm; pollen grains 18-26.3 um long: tectum imperforatum or with a few puncta; diameter of puncta 0.2 pm; supratectal process granular + uniformly distributed (Fig. 3c); infrastaminal scales 3/4- 4/5 of the corolla tube, oblong-spathulate, rounded uniformly dense fringed to fimbriate; styles uniformly filiform, 1.3-L6 mm, about as long as the ovary; stig- mas capitate, globose. Capsules indehiscent to irregularly dehiscent, globose to globose-depressed, 1.8-3.2 x 2.2-3.6 mm, not thickened or raised around the large intersylar aperture; not translucent, withered corolla persistent envelop- ing 1/2 or more of the capsule base. Seeds 1-4 per capsule, dorsoventrally com- pressed, broadly elliptic to obovate, 1.07-1.42 x 0.9-1.21 mm, seed coat cells al- veolate/papillate, 15-20 1m in diameter; epicuticular wax absent; hilum region subterminal, subrotund, 0.30-0.35 X 0.28-0.30 mm. vascular scar 0.20-0.25 mm long, vertical to slightly oblique (Fig. 4f). 2n = ? Distribution and ecology.—U.S.A.: Nebraska, Washington, and Wyoming. Flowering Aug. Hosts: species of Grindelia, Helianthus, Humulus Psoralea, Ru- bus, Solidago. Conservation status.—Gl (critically imperiled) (the same assessment in NatureServe 2005). 7. Cuscuta sandwichiana Choisy, Mem. Soc. Phys. Hist. Nat. Geneve 9:280. 1841. Tyre: U.S.A. HAWAII Sandwich Islands, 1830, Gaudichaud-Beaupré I (HOLOTYPE: G-DC, frag- ment NY; ISOTYPE: P, fragment NY). Stems, 0.2-0.5 mm, yellow-orange. Inflorescence loose, paniculiform of 3-7(-1D pedicellate flowers; pedicels 2-5 mm; bracts | at the base of clusters, usually lacking at the base of pedicels, membranous, narrow-triangular, I-1.6 x 0.7-1 mm, margins entire, apex acute. Flowers (4-)5-merous, 3-4(-5) mm long (Fig. 5a), fleshy, white when fresh, yellow-brownish when dried; papillae absent; la- ticifers conspicuous in the corolla lobes, sometimes in the calyx and ovary/ capsule as well, isolated or in rows, ovoid to elongated; calyx yellow-brown, not reticulate or shiny, cupulate, almost as long as corolla tube, divided 1/2-1/3 the length, lobes not basally overlapping, triangular-ovate, in some flowers cari- nate, margins entire, apex obtuse to subacute; corolla persistent; tube campanu- late to globular, not saccate, 2-3 mm long, lobes erect to slightly spreading, 0.8- L5 mm long, ovate-triangular, margins entire, apex obtuse to subacute with inflexed tips; stamens mostly included, shorter than corolla lobes, anthers broadly-elliptic 0.5-0.7 x 0.4-0.5 mm, filaments 0.2-0.7 mm; pollen grains 22- 27.5 um long; tectum imperforatum or with a few puncta; maximum diameter of puncta 0.3 um; supratectal process granular to conical, + uniformly distrib- uted; infrastaminal scales absent or reduced, bifid, truncated or triangular, dis- tally with a few teeth; styles uniformly stoutish, 0.5-1L1 mm, ca. as long as the COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 169 ovary; stigmas capitate, globose. Capsules indehiscent to irregularly dehiscent, globose to globose-depressed, 2.5-3.5 mm X 33-4 mm, + thickened but not raised around the large interstylar aperture, not translucent, withered corolla surrounding the capsule. Seeds usually 2 per capsule, angled, elliptic to broadly- elliptic, 11-2 x 1-17 mm, seed coat cells alveolate/papillate, 35-50 um in diam- eter, epicuticular wax absent; hilum region subterminal, round, 0.30-0.45 mm in diameter, vascular scar 0.19-0.25 mm long, vertical or slightly oblique (Fig. 5b). 2n =? Distribution and ecology.—U.S.A.: Hawaii. Flowering summer-fall JQun- Oct). Hosts: plants in seashores and arid habitats. Conservation status.—G2 (imperiled) (the same assessment in NatureServe 2005). 8. Cuscuta obtusiflora Kunth var. glandulosa Engelm., Trans. Acad. Sci. St. Louis Q?. 1859. Cuscuta gla ndulosad (Engelm.) Small, Fl. S.E. U.S. 969, 1337. 1903. LECTOTYPE (Yuncker 1921): U.S.A. GEORGIA. [Muscogee Co.]|: Columbus, 1838, Boykin s.n. (MO). The protologue cited numerous collections in addition to the Boykin one. Stems 0.25-0.4 mm in diameter, orange. Inflorescences dense, glomerulate of 5-18 sessile or subsessile flowers; pedicels absent to | mm; bracts 1 at the base of clusters and 0-1 at the base of pedicels/flowers, membranous, ovate, 0.5-1.3 x 0.2-1.1 mm, margins entire, apex acute to obtuse. Flowers 5-merous, 1.8-2.5 mm, membranous, creamy white when fresh, yellow-brown when dried; papillae absent; laticifers numerous, evident in the perianth and ovary/capsule, isolated, round to ovoid; ealyx yellow-brownish not reticulate or shiny, shallowly cupu- late, about equaling the corolla tube, divided ca. 1/2 the length, lobes barely basally overlapping, ovate, not carinate, margins entire, apex obtuse; corolla persistent; tube campanulate, not saccate, 1-15 mm, lobes erect to spreading, 0.8-1.5 mm long, ovate to ovate-oblong, margins entire, apex obtuse, straight; stamens exserted, shorter than corolla lobes, anthers ovate to broadly elliptic, 0.3-0.4 x 0.2-0.3 mm, filaments 0.4-0.6 mm; pollen grains (18-)20-24(-27) um, tectum perforatum; maximum diameter of puncta 0.6 um; supratectal process granular to conical + uniformly distributed (Fig. 5c); infrastaminal scales reach- ing the filament bases, oblong, rounded, with a few basal fimbriae, densely fringed in the distal 3/4; styles uniformly stoutish, 0.4-1.1 mm, shorter than or equaling the ovary, stigmas capitate, globose. Capsules indehiscent to irregu- larly dehiscent, depressed-globose, 1.5-3 x 2.5-4 mm, not thickened or raised around the large interstylar aperture, not translucent, withered corolla persis- tent at the base. Seeds usually 4 per capsule, dorsoventrally compressed, broadly- ovate to broadly elliptic, 14-156 x 1.23-1.35 mm, seed coat cells alveolate/papil- late, 20-30 pm in diameter, epicuticular wax usually present (Fig. 5d). 2n = ? Distribution and ecology.—U.S.A.: Alabama, Arkansas, Connecticut, Florida, Georgia, Kentucky, Louisiana, Minnesota, Mississippi, New York., Oklahoma., 170 BRIT.ORG/SIDA 22(1) howe =& "es A as pk ft a eet Ch Bh AS Sey Denes ‘3 ‘ et ee) ie rere sf — Fic. 5. Cuscuta sandwichiana. a. Flower (scale bar = 1 mm);b. seed (scale bar = 0.6 mm). Cuscuta obtusiflora var. glandulosa. c. Pollen (scale bar = 6 im); d. seed (scale bar = 0.5 mm). Cuscuta polygonorum.e. Seed—general morphology (scale bar = 0.60 mm); f. seed—surface micromorphology (scale bar = 100 Lm). COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 171 Pennsylvania, Texas; Mexico, West Indies (Puerto Rico, Cuba). Flowering Jul- Sep, ~Oct. Hosts: species of Alternanthera, Dalea, Hygrophila, Lythrum, Polygonum, Xanthium. Conservation status.—G3G4 (vulnerable to apparently secure) (G4, NatureServe 2005). 9, Cuscuta polygonorum Engelm., Amer. J. Sci. Arts 43:342. 1842. Type US.A. Mis- souRE W of St. Louis, “bottoms of sink-holes, or margins of ponds”], on Polygonum, Aug 1839, Lindheimer s.n. (HOLOTYPE: MO). d ,glomerulate Stems 0.2-0.4 mm in diameter, yellow-orange. Infl of 4-18 sessile to subsessile flowers; bracts 1 at the base of clusters and 0-1 at the base of pedicels, membranous, ovate-triangular to lanceolate, 0.3-1 x 0.2- 0.9 mm, margins entire, apex acute; pedicels absent to | mm. Flowers (3-)4- merous, 2-2.7 mm, membranous, white when fresh, yellow-brown when dried; papillae absent; laticifers evident in the corolla and the ovary/capsule, isolated, round to ovoid; calyx yellow-brownish, not shiny or reticulate, cupulate, ca. equaling the corolla tube, divided 1/2-2/3 to the base, lobes not basally over- lapping, triangular-ovate, not carinate, margins entire, apex obtuse to rounded; corolla persistent, tube cupulate to shallowly campanulate, hot saccate, l-14 mm, lobes erect, l.1-1L6 mm long, triangular, margin ntire, apex acute, straight; stamens exserted, shorter to nearly equaling corel lobes, anthers ovate to broadly elliptic, 0.3-0.4 x 0.2-0.3 mm, filaments 0.4-0.7 mm; pollen grains (18-) 21-24.5 (-26) um long, tectum imperforatum or with a few puncta; maximum diameter of puncta 0.3 um; supratectal process granular to conical + uniformly distributed; infrastaminal scales usually reaching the filament bases, oblong, mostly shallowly bifid with 1-3 fimbriae on each side of the filament attach- ment or with a few irregular fimbriae at the apex; styles evenly subulate, 0.4- 0.9 mm, shorter than the ovary; stigmas capitate, globose. Capsules indehis- cent to irregularly dehiscent, depressed-globose, often appearing angled, 1.6-3 < 2.5-5 mm, not thickened or raised around the large interstylar aperture, not translucent, withered corolla persistent the capsule base. Seeds usually 4 per capsule, dorsoventrally compressed, subrotund, broadly ovate to broadly el- liptic, 145-16 x 1.25-1.39 mm, seed coat cells alveolate/papillate or poliedric, 20-30 um in diameter, epicuticular wax usually present; hilum region subter- minal scar area slightly raised, elliptic, 0.45-0.6 x 0.3-0.35 mm; vascular scar 0.15-0.2 mm, vertical or slightly oblique (Fig. 5e,f). 2n = Distribution and ecology.—CANADA: Ontario, Québec. U.S.A.: Arkansas, Connecticut, District of Columbia, Delaware, Iowa, Illinois, Indiana, Kansas, Kentucky, Louisiana, Massachusetts, Maryland, Maine, Michigan, Minnesota, Missouri, North Dakota, Nebraska, New Jersey, New York, Ohio, Oklahoma, Pennsylvania, Rhode Island, Tennessee, Texas, Virginia, Wisconsin. Flowering rt —" 172 BRIT.ORG/SIDA 22(1) Jul-Oct. Hosts: usually Polygonum but occasionally also on Impatiens, Ipomoea, Lycopus, Penthorum, Xanthium, and others. Conservation status.—G3 (vulnerable) (G5, NatureServe 2005). APPENDIX lL. VOUCHERS FOR THE SEM STUDY Vouchers are from NY unless otherwise indicated. 1. Cuscuta pentagona (16 collections examined).—U.S.A. DISTRICT OF COLUMBIA. Eastern sae near Mt. Hamilton, 2 Aug 1919, Killip 6338. FLORIDA. Lake Co.: 3 mi S of Sorrento, Welch GEORGIA. Whitfield Co.: field E of Dicks Ridge, 1000 ft, 27 Jul 1900, Wilson s.n. INDIANA eee Co.: 2 mi W of Bedford, flood plain of Salt Creek, 23 Aug 1934, Kriebel 2518. KANSA S. Trego Co.: 19 mi S and 2 mi W of Collier, 6 Aug 1952, David & Harr 4136. MASSACHUSETTS. Middlesex Co.: Winchester, Winter Pond, 22 Sep 1908, Fernald & Weatherby s.n. MICHIGAN. Kalamazoo Co.: Fort Custer, 12 Aug 1945, Hanes 4541.MISSOURI. Phelps Co.: Jerome, 27 Jul 1931, Kellog s.n. NEW JERSEY. Ocean Co.: Bay Head, 31 Jul 1910, Mackenzie 4742.NEW YORK. (no county given).Long Island, 8 Aug 1903, Bicknell s.n. NORTH CAROLINA. Franklin Co.: Franklin, 23 Jul 1888, Johnson s.n. NORTH DAKOTA. Barnes Co.: Eckelson, 15 Sep 1937, Stevens s.n. Benson Co.: Cran- berry Lake, 8 Sep 1930, Stevens s.n. TENNESSEE. Wayne Co.: 4 mi W of Waynesboro, 10 Aug 1934, Welch 1382. VIRGINIA. Arlington Co.: 1/4 mi SE Hatfield,9 Jul 1939, Hermann 10391;W of Williamsburg, 15 Aug 1921, Weatherby 4230 2.Cuscuta campestris (23 Akai alunt onaiecinalac QUEBEC. paiies te 940, Cartier s.n. oNTAR 10, Waterloo Co.: Galt, 26 Sep 1908, Herriot s.n. Wellington Co.: 3 Aug 1943, Howitt s.n. I rt cel ae Jonesboro, 260 ft,6 Jun 1948, ens 26643, Garland Co.: Hot Springs, 500 ft, 17 Jul 1938, Demaree 17932. CALIFORNIA. Los Angeles Co.: Los Angeles, 10 Jul 1933, et 10533. IDAHO. Ada Co.: Harrison dee on NE side of Boise, T4N R2E Sec 34 SENW, 2710 ft, 1 g 1980, Ertter & Strachon 3951. FLORIDA. Dade Co.: at 14000 SW 8th Road, Miami, 12 Nov cen Y).GEORGIA. Clarke Co.: He at 285 Hillcrest, 800 ft, 11 Nov 1946, Cronquist 4209. INDIANA. Harrison Co.: 4 mi SE of Central, 26 Aug 1945 Deams.n.. Wells Co.: 5 mi NE of Bluffton, 26 Aug 1940, Deam 59866. KENTUCKY. Fayette Co.: Gentry-Thompson stockyards, Lexington, 2 Sep 1944, McFarland s.n. MARYLAND. Wicomico Co.: Willards,4 Sep 1942, Moldenke 13847. NEVADA. Douglas Co.: eae River, ca. 7 mi N of Minden, 4700 ft, 14 Sep 1969, gla Howell ioe alae EXICO. Grant Co.: | 881, Rusby 295. NEW YORK. Onondaga Co:: Syra- cuse, shore of Onondaga a5 cere One GON. Marion Co.: Salem, Willamette River, 13 Jul 1922, ieee TENNESSE N of Knox lle Wallace’s Orchard, 6 Aug 1943, Wilson 3007. TEXAS. Childress Co.: 3.4 mi : a ress,S of Hwy. 287,6 Sep 1945, Whitehouse 10715.Tom Green Co.: South Caucho River, a mi N 2 E of Christoval, 29 Aug 1943, Cory s.n. UTAH. Sanpete Co.: 1145, R2E,SEC 22,4 mi NE of F a non U-132,9000 ft, 19 Jul 1977, Neese & White 3682. VIRGINIA. Nansemond Co.: Suffolk, 24. jul 1893, Heller s.n. ta glabrior (15 collections examined).—U.S.A. OKLAHOMA. Murray Co.: Sulphur, Platt an Park, 28 May 1 bee Merril 522.Pontotoc Co.: 2 mi SW of Ada,near State Hwy. 12,26 Jun 1947, Robbins 2607.NEW MEXICO.(no locality given) 1851-1852, Wright 1635. TEXAS. Bell Co.: Near Little River, 11 June 1930, ee Killen, 14 Aug 1931, Wolff 3270. Bexar Co.: 16 mi E of San Anto- nio, 27 Apr 1921, Schulz 453;San Antonio, 17 Apr 191 oo s.n. Burnet Co.: along State Hwy. 29, mi E of Burnet, Webster & Rogers 6476.Coleman Co.: 7 mi E of ee Anna, be Jun 1958, Correll & Johnston 19019. Dallas Co.: Dallas, 30 Jun 1872, Hall s.n. Edwards Co.: 29 mi NW of Rocksprings, 27 Jun 1943, Cory s.n. Ellie Co.: above Red Oak Creek off Waco road, | ; ite Deni 9353. Neueces Co.: Corpus Christi, 0-40 ft, 9-12 Apr 1894, Heller s.n. Randall Co.: Buffalo Lake National COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 173 Wildlife Refuge, 3 Aug 1975, Higgins 9567. elias a low hills in upper big Canyon, ca. 30 mi N of Sanderson toward Sheffield, 2800 ft, Warnock 14 Cuscuta runyonii (5 collections So eee Duval Co.: of Hyw. 44 NE of San Diego, 20 Apr 1949, Lundell 14906. Hidalgo Co.: 2 mi N of La Joy,45 m, 13 Jul 1941, Runyon 2825. Starr Co.: Brownsville, 50 m,27 Apr 1941, Runyon 2622. Webb Co.: ca. 21 mi from junction of Hwys. 81 & 83, along Hwy. 83, 1 Apr 1949, Tharp & York 52-163; 46 mi NE of Laredo, route #59, 17 Jul 1958, Correll & Johnston 14906 5. Cuscuta harperi (5 collections examined).—U.S.A. ALABAMA. Cherokee Co.: Lookout Mt. at the edge of gorge of May's Gulf, Harper s.n. De Kalb Co.: Desoto State Park, above Fort Payne, in Little River Canyon, 8 Oct 1960, Sherman et al. 27757; Glades, 1200 ft, 13 Oct 1962, Demaree 46295; Lookout Mt, Jul 1898, Ruth 473. Etowah Co.: Black Creek just above Noccalula Falls, Lookout Mt., 31 Aug 1911, Harper 147 scuta plattensis (2 collections examined).—U.S.A.WYOMING. Goshen Co.: 3 mi NE of Torrington, T25N R6QW Sec 31 S1/2 SW 1/4, 4200 ft, 15 Aug 1993, Dorn 5470. Niobrara Co.: near Newcastle, 24 Jul 1942, Degener & Peiler 16242. 7. Cuscuta sandwichiana (3 collections examined).—U.S.A. HAWAII. Hawaii Co.: East Maui, Makawao, Kaunauhae, La Perouse Bay, 30 Apr 1978, Sy/ aaa Ka’ena Pt.,29 Nov 1969, Char 40; O'ahu, Makapu’u Beach Park, 4 Oct 1960, Fujiwara 8.Cuscuta obtusiflora var. glandulosa (5 eee examine), —U.S.A.ARKANSAS. Ash- ley Co.: Lone Prairie, Prairies, 15 Sep 1940. Demaree 21543. va ae White Co.: in the Tippecanoe River, ca. 1 mi below Monticello, 14 Aug 1925, Deam 41990. AHOMA. Haskell Co.: stream ca.5 mi S of the intersection of Hwy 2 with Hwy 31,22 Jul 36 cha , 3387. TEXAS. Dallas Co.: Bachmans Dam, bordering lake, wet area, 1 Sep 1942, Lundell & Lund 717,24 Aug 1942, Lundell 11626. scuta polygonorum (7 collections eee ana ARKANSAS. St. Francise Co.: 5 mi S of Forrest City, 19 Sep 1959, McDaniel 1419. INDIANA. Putnam Co.: 2 mi E of Bainbridge, 20 Oct 1941, Yuncker 10836. NEBRASKA. Otoe Co.: extreme S part of SE1/4,NE1/4, Sec 25, T8N-R14E, 11 Sep 1974, Sutherland 4096. NEW JERSEY, Somerset Co.: Millstone, plaine de débordement de la riviére Millstone, 4 Aug 1956, Lavoie 452 (QUE). NEW YORK. [county not indicated; Nassau Co.?] Long Island, Plattsdale, 19 Sep 1925, Ferguson 4608. OHIO. Ottawa Co.: Duck Club,Winous Pt.,4 Aug 1949, Core & Anderson s.n. WISCONSIN. Winnebago Co.: swampy area along the shore of lake Winnebago, N be- yond the end of Hazel St., N of Murdock, in Gin 25 Jul 1965, Arriman 598 (UNB) ACKNOWLEDGMENTS We thank directors and/or f ACAD, ALTA, ARIZ, ASU, BRIT, DAO, F GH, HAM, MEXU, MICH, MT, MTMG, NFLD, NSPM, OAC, QFA, QUE, RBG, RSA, SASK SES, TEX GLE TUR UBG UGG | BPS -UNB UNM, US-USAS UWO, UWPG, WAT, WIN, WIS, WTU, and XAL for loans to Costea. Dan Austin, Lytton Musselman and Alan Prather provided useful comments and suggestions for an earlier version of the manuscript. Thierry Deroin helped us with informa- tion about the type of Cuscuta sandwichianaand kindly donated all the Cuscuta fragments annotated by Yuncker (held in P) to WLU. Special appreciation goes to NY staff for approving and preparing the numerous voluminous loans. Al- exandra Smith assisted us with the scanning electron microscope. REFERENCES Austin, D.F.1979.Comments on Cuscuta — for collectors and curators. Bull. Torrey Bot.Club 106:227-228. 174 BRIT.ORG/SIDA 22(1) Austin, D.F. 1986. Cuscutaceae. In:T. Barkley, ed. Flora of the Great Plains. University Press of Kansas, Lawrence. Pp. 661-666. BartHtort, W,, C. Neinuis, D.Cutcer, F. Dirscy, |. MEUSEL, |. THEISEN, and H.WiLHeLmI. 1998. Classifica- tion and terminology of plant epicuticular waxes. Bot. J. Linn. Soc. 126:237-260. Beuz, T. 1986. A revision of Cuscuta sect. Cleistogrammica using phenetic and cladistic analyses with a comparison of reproductive mechanisms and host preferences in spe- cies from California, Mexico, and Central America. Ph.D. diss., Univ. of California, Berkeley Buia, A. 1938. Cuscutele Romaniei. Bull. Fac. Agron. Cluj 7:1-147. Costes, M.and FJ. Taroir. 2006. Biology of Canadian weeds. Cuscuta campestris Yuncker, C. gronovii Willd. ex Schult.,C. umbrosa Beyr.ex Hook.,C. epithymum (L.) Land C.epilinum Weihe. Canad. J. Pl. Sci. 86:293-316. Costea, M.,G.L.Nesom, and F.Taroir.2005. Taxonomic status of Cuscuta nevadensis and Cuscuta veatchii (Convolvulaceae). Brittonia 57:264—272. Costea, M., G.L. Nesom, and S. Steranovic, 2006a. Taxonomy of the Cuscuta salina-californica complex (Convolvulaceae). Sida 22:177-195. Costea, M., G.L. Nesom, and S. Steranovic. 2006b. Taxonomy of Cuscuta gronovii and Cuscuta umbrosa (Convolvulaceae). Sida 22:197-20/7. Costes, M., G.L. Nesom, and S. Steranovic. 2006c. Taxonomy of the Cuscuta indecora (Convolvulaceae) complex in North America. Sida 22:209-225. Crupen, RW. 1977. Pollen-ovule ratios: a conservative indicator of breeding systems in flowering plants. Evolution 31:32-46. Dawson, J.H. 1984.A vegetative character that separates species of Cuscuta. Proc. 3rd Interntl. Symposium on Parasitic Weeds. Pp. 184-187. Dawson, J.H., L.J.Mussetman, P.Wotswinket, and |. Dorr. 1994. Biology and control of Cuscuta. Rev.Weed Sci. 6:265-317. Dareysuire, S.J. 2003. Inventory of Canadian agricultural weeds. Agriculture and Agri-Food Canada, Research Branch, Ottawa, Ontario. 396 pp. Electronic Publication: ENceLMANN, G. 1859. Systematic arrangement of the species of the genus Cuscuta with critical remarks on old species and descriptions of new ones. Trans. Acad. Sci. St. Louis 1:453-523 Foceteers, 5.0. 1938. The cytology of Cuscuta. Bull. Torrey Bot. Club. 65:631-645. GaerINeR, E.E. 1950. Studies of seed germination, seed identification, and host relation- ships in Dodders, Cuscuta spp. Mem. Cornell Agric. Exp. Sta. 294:1—56. GANDHI, K.N.and R.D.THomas. 1983.Variations in the floral structure of Cuscuta L.Phytologia 53:184-186. GANDHI, K.N., R.D. THomas, and S.L. Hatcu. 1987. Cuscutaceae of Louisiana. Sida 12:361-379. Gomez, JM. 1994. Importance of direct and indirect effects in the interaction between a parasitic angiosperm (Cuscuta epithymum) and its host (Hormatophylla spinosa).Oikos 71:97-106 Hoen, P. 1999. Glossary of pollen and spore terminology. Laboratory of Palaeobotany and Att / | Palynology, Utrecht. COSTEA ET AL., TAXONOMY OF CUSCUTA PENTAGONA COMPLEX 175 Howm, L., J. Dott., E. Houm, J.PANCHO and J. Hersercer. 1997.World weeds:natural histories and distribution. John Wiley & Sons Inc., Toronto, ON. Kaatesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland. First edition. In: Kartesz, J.T. and C.A.Meacham. 1999. Synthesis of the North American flora, Version 1.0.North Caro- lina Botanical Garden, Chapel Hill. Knepper, D.A., R.A. Creacer and L.J. Mussetman. 1990. Identifying dodder seed as contami- nants in seed shipments. Seed Sci. Technol. 18:731-741 Kuur, J. 1969. The biology of parasitic flowering plants. Univ. of California Press, Berkeley. LysHebe, B.O. 1984. Seed structure and germination in Cuscuta pedicellata with some notes on Cuscuta campestris. Nordic J. Bot. 4:669-664. LysHebe, B.O. 1985. Morphological and anatomical features of Cuscuta pedicellata and C. campestris. Nordic J.Bot.5:65-77. Mussetman, LJ. 1986. The genus Cuscuta in Virginia. Castanea. 51:188-196. NatureServe. 2005. NatureServe Explorer: An online encyclopedia of life [web applicatian) Version 4.0. NatureServe, Arlington, Virginia. (Accessed: November 16, 2005). Parker, C.and C.R.RicHes. 1993.Parasitic weeds of the world. Biology and control.CAB Inter- national, Wallingford, UK. Pazy, B. and U. PLiTMANN 1995. Chromosome divergence in the genus Cuscuta and its sys- tematic implications. Caryologia 48:1 73-180. Prater, L.A.and R.J.Tyrt. 1993.The biology of Cuscuta attenuata Waterfall. Proc. Okla. Acad. SCL 73-13 Severova, E.E. 1991. Morphology and taxonomy of dodder (Cuscuta L.) of the European part ofthe USSR. Byull. Mosk. Obshch. Ispyt, Prir,, Biol. 96:59-63. USDA, NRCS. 2004. The PLANTS database. National Plant Data Center, Baton Rouge, LA. Electronic Publication,Version 3.5: Verocourt, B. 1948. Biological flora of the British Isles. Cuscuta L. J. Ecol. 36:356-365. Warp, D.E. 1984. Chromosome counts from New Mexico and Mexico. Phytologia 56:55-60. Yuncker, 1.G. 1921. Revision of the North American and West Indian species of Cuscuta. Illinois Biol. Monogr. 6:91-231. Reprinted 1970, Johnson Reprint Company, N.Y. Yuncker, 1.G. 1932. The genus Cuscuta. Mem. Torrey Bot. Club 18:113-331. Yuncker, 1.G. 1942. Three new Cuscutas from western North America. Bull. Torrey Bot. Club 69:541-543. Yuncker, T.G. 1943. Nomenclatural changes in the genus Cuscuta and notes on some American species. Bull. Torrey Bot. Club 70:61-67. Yuncker, T.G. 1965. Cuscuta. North American flora, ser. 2,4:1-51. 176 BRIT.ORG/SIDA 22(1) Book REVIEW DieTRICH FROHNE and Haws J. Pranper. Patrick McKinney, Kirk Cumpston (Con- sult. Eds.). (Translated by Inge Alford). 2005. Poisonous Plants: A Hand- book for Doctors, Pharmacists, Biologists and Veterinarians, Second Edi- tion. (ISBN 008192-750-3, hbk.). Timber Press Inc, 133 S.W. Second Avenue, Suite +50, Portland, OR 97204-3527, U.S.A.(Orders: wwwtimbet press.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax.). $150.00, 480 Pp. 224 color photos, 131 b/w illustrations, 20 tables, 115 dia- grams, 91/2" x 10 1/2". Many seemingly innocuous plants harbor toxins that can cause serious illness and even n. This authoritative text is intended for pharmacists, poison control center personnel, doctors, veterinar- ians, and other sores who deal with poisoning cases, although it is useful to those interested in wild foods and wi iderat Hone medicinal herbs. It epens! with a concise introduction to poisonous plants, including hov ning cai yee doi pected cases [ poisoning and descrip- k contains clear descripti - poisonous plants, tions of toxic ees als. The rest of fig arranged by family from Aceraceae to ee ‘The text is aoe with descriptions of macroscopic and microscopic plant morphology, symptoms, treatments, and references. Excellent color photographs illustrate sae section a met English cauion of a sie was ne 20 gli -} years ago. This second E dix with notes on Eaiconetie ee found in Nor th America. Not intended to ae a field ¢ muitde hae book is still an excellent reference book.—Marissa Oppel, MS, Herbarium Technician, Botanical Re- search Institute of Texas, Fort Worth, Texas 76102-4060, U.S.A. Book Notice CARMEN ULLOA ULLoA and D.A. Neil. 2005. Cinco aitos de adiciones a la flora del Ecuador: 1999-2004. (ISBN 9978-09-578-0, pbk.). Universidad Técnica Particula de Loja, Missouri Botanical Garden, Funbotanica. Editorial, UTPL, Loja. (Orders: Available at Missouri Botanical Garden Press, http:// www.nbgpress.info/). $16.00, [Reduced price availabl« only at UTPL, Loja]. ToD o1/ 2 oC AL Summary.—Of the 1246 new additions to the Ecuadorian flora, 820 are new taxa described from Ecuadorian material, 337 names are new records for Ecuador, and 89 are taxonomic changes. The new total for the flora is of 17,058 species of vascular plants. Some 242 new bibliographic references Qu are liste SIDA 22(1):17 176. 2006 TAXONOMY OF THE CUSCUTA SALINA-CALIFORNICA COMPLEX (CONVOIVULACEAE) M ihal Coster (corresponding author) Guy |. Nesom Department of Biology Botanical Research Institute of Texas Wilfrid Laurier University Pecan Stree 75 University A West Fort Worth, Texas 76102-4060, U.S.A. Waterloo, Ontario gnesom@brit.org N2L 3C5, CANADA meostea@wlu.ca Sasa Stefanovic Department of Biology vaivelsigye oF Toronto at Mississauga I UUYU hodad Missis issauga, Ontario LSL 106, CANADA ABSTRACT The Cuscuta califor plex (Cuscuta subsect. Californicae) is recircumscribed to include the species from aieceee PS ioineticae The species include C. salina, C. susksdorfii, C. californica, C. occidentalis, C. subinclusa, and C. howelliana. Asummary of relevant taxonomic information is pro- vided, along with a key to species and varieties, synonymy, distribution, host range, and conserva- tion status. Morphology and micromorphology of flowers, seeds, and capsules are analyzed and il- lustrated. RESUMEN uscuta subsect. Californicae) Aci i incluir las especies El complejo Cuscuta c lifornica ( de subsect. Subinclusae. Las especies ee C. salina, C. susksdorfii, C. californica, C. occidentalis, C. subinclusa, y C. howelliana. En este trabajo se presenta un resumen de la informacion taxonémica junto con una clave para identificar las especies y variedades, sinonimos, distribucion, variabilidad de huéspedes y estato de conservacion. Se analizan e ilustran la morfologia y micromorfologia de las flores, de y de las capsulas. Yuncker (1932) described and defined Cuscuta subsect. Californicae and subsect. Subinclusae mainly on the basis of absence or presence, respectively, of infrastaminal scales. He suggested that both sections were derived indepen- dently but in parallel from subsect. Arvenses. In Yuncker’s view, subsect. Californicae comprised five species: C. californica Hook. & Arn., C.occidentalis Millsp., C. brachycalyx Yuncker, C.jepsonii Yuncker,and C. sandwichiana Choisy, while subsect. Subinclusae included three species: C. salina Engelm., C. suksdorfii Yuncker, and C. subinclusa Durand & Hilgard (Yuncker 1932, 1965). Cuscuta howelliana Rubtzoff was recently described (Rubtzoff 1966) and noted by its author to have similarities to C. suksdorfii of subsect. Subinclusae. SIDA 22(1): 177 — 195. 2006 178 BRIT.ORG/SIDA 22(1) Definitions and interrelationships among these taxa are complex. Beliz (1993) included Cuscuta brachycalyx, C. occidentalis, and C. suksdorfii as syn- onyms of C. californica var. breviflora Engelm. Cuscuta subinclusa and C. howelliana were hypothesized to be sister species in a separate clade (Beliz 1986). Some taxa were described in one subsection and later were reclassified in the other. For example, Engelmann (1859) initially described C. salina as “°C. californica var. squamigera” and as C. subinclusa var. abbreviata (see also the example below of C. suksdorfii). Cuscuta sandwichiana (Beliz 1986, Costea et al. 2006a) and C. jepsonii (Beliz 1986; Costea et al. 2006c) do not appear to be evo- lutionarily related to this group. Circumscriptions of these taxa and an assessment of their we are provided here on the basis of the morphology and micromorphology of flow- ers, capsules, seeds, and pollen. Conservation status is assessed for all taxa. METHODS Descriptions of morphology (see Costea et al. 2006a) are based on samples from specimens of the NY, JEPS, and UC herbaria (Appendix 1). Measurements and pictures were taken with a scanning electron microscope Hitachi S-570 at 15 KV.Samples were coated with 30 nm gold using an Emitech K 550 sputter coater. Conservation status was determined using NatureServe (2005) ranks and Criteria. TAXONOMY Delimitation of the Cuscuta salina-californica complex Our observations suggest that the species of subsections Californicae and Subinclusae form a single phylogenetic group. We hypothesize that infrastaminal scales have undergone a gradual reduction from fimbriate scales or ridges in C. salina, to dentate wings in C. suksdorfii, to complete reduction in C. californica and C. occidentalis. A similar reduction of infrastaminal scales has occurred in C. indecora complex (subsect. Indecorae Yuncker) (Costea et al. 2006c). Cuscuta subinclusa and C. howelliana have well-developed infrastaminal scales and were suggested by Beliz (1986) to form a distinct clade in which a cylindric-campanulate corolla has evolved as a specialization to butterfly pol- lination. Although C. subinclusa and C. howelliana are distinct in their microreticulate pollen (see descriptions below), close similarities in morphol- ogy and micromorphology of calyx, corolla lobes, seeds, and capsules (see be- low) indicate that they probably belong to the same phylogenetic group as C californica, C. salina, and C. suksdorfiti. A cylindric-campanulate corolla some- times occurs in C. californica, and Engelmann (1876) noted that “in many re- spects [C. salina] is intermediate between the preceding [C. californica] and the following species [C. subinclusal.” Yuncker (1932, 1965) observed that the corolla of C. subinclusa is “usually showing horizontal ridges between the stamen at- — COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 179 tachments” and we find this characteristic in some flowers of C. californica as well. The original description of C. howelliana (Rubtzoff 1966) noted that it is similar to C. suksdorfii var. subpedicellata. Cuscuta jepsonii and C. sandwichiana were included by Yuncker (1932) in subsect. Californicae (the latter species omitted in the treatment from 1965) because they exhibit a similar reduction of the infrastaminal scales. Cuscuta jepsonii, however, may be evolutionarily related to taxa of Cuscuta indecora com- plex (Costea et al. 2006c). Cuscuta sandwichiana is a Hawaiian endemic and was hypothesized by Costea et al. (2006a) to belong to the C. pentagona com- plex. In conclusion, the Cuscuta salina-californica complex in our view includes C. salina, C. susksdorfit, C. californica, C. occidentalis, C. subinclusa, and C. howelliana, which are recognized here as Cuscuta subsect. Californicae Yuncker (including subsect. Subinclusae). Taxonomically and nomenclaturally problem- atic species are discussed below. Cuscuta subinclusa.—Curran (1885) noted that “from the description, [C. ceanothi is]evidently C. subinclusa,’ and the former name has been used by most authors (e.g. Yuncker 1965) because it has priority. From Curran’s observation it can be inferred that the type collection of C. ceanothi was not available even at 1885. This collection may have been destroyed during the fire following the San Francisco earthquake from 1906 or even at an earlier date. As observed by Beliz 986), it is possible that the name C. ceanothi refers to a species distinct from C. subinclusa. The protologue of C. ceanothi states that flowers are urceolate with obtuse calyx lobes, features not encountered in C. subinclusa or in any species hypothesized to be closely related. In a letter addressed to Engelmann on 10 Nov 1860 (Ertter 2003), Behr wrote: “Of Cuscuta I know two kinds, the the latter I gave once a diagnosis in the transactions (Proc. Calif.) under the p pres liminary name Cuscuta ceanothi. This diagnosis is by the nature of a search very incomplete, as for comparison I had only the just-mentioned parasitic kind on the Salicornia and none of the more closely related exotics.” In a summary of the San Francisco flora (1888), approximately three decades after describing C. ceanothi, Behr mentioned C. subinclusa but not C. ceanothi. Without neotypification, there apparently is no way to establish the identity of C. ceanothi, and until more certainty might exist that C. ceanothi is not a distinct and evidently rare species, we use the later name C. subinclusa for the known species. Cuscuta suksdorfii.— Yuncker (1921) described C. salina var.acuminata Yuncker, which he later (1932) treated at specific rank as C. suksdorfii. This species is “closely allied with C. salina” from which it differs in the morphology of calyx and corolla, infrastaminal scales reduced to dentate wings, and multiseeded 180 BRIT.ORG/SIDA 22(1) capsules. Beliz (1986) originally treated C. suksdorfii as a variety of C. salina, but later (1993) she included it as a synonym of C. californica var. breviflora, together with C. occidentalis and C. brachycalyx. Indeed, C. suksdorfii shares some characteristics with both C. salina (e.g., presence of infrastaminal scales as well as seed and pollen morphology—see descriptions) and C. californica/C. occidentalis (e.g., multiseeded capsules), but we consider that treating it at spe- cific rank better represents the differences in morphology and biology as well as the evolutionary relationships between all the taxa involved (see below). Cuscuta suksdorfii var. subpedicellata Yuncker with flowers sessile or Peis shorter calyx (16-2 mm long) and capsule globose to depressed globose treated formally at varietal rank. Cuscuta californica and C. occidentalis —By describing C. brachycalyx, Yuncker (1932, 1965) obscured the already tenuous distinction between C. californica and C. occidentalis. Although homotypic, C. californica var. brachycalyx Yuncker and C. brachycalyx have slightly different protologues. Cuscuta californica var. brachycalyx has “corolla campanulate, lobes ... shorter than the tube...” (Yuncker 1921, p. 62); C. brachycalyx has “corolla campanulate-globose, saccate between the stamen attachments, lobes ... shorter than or about equal- ing the tube” (Yuncker 1932, p. 159) and “is closely related to C. californica, but differs by its very short calyx and more obtuse perianth lobes.” Yuncker did not mention the saccate corolla that would separate C. brachycalyx from C. californica but that would bring it close to C. occidentalis. Not surprising]y, Beliz (1986) concluded that “all names proposed for the numerous perianth and an- ther size variants within C. californica do not warrant taxonomic recognition . and she included both C. occidentalis and C. brachycalyx as synonyms of C. californica var. breviflora (Beliz 1986, 1993). We find that indeed two different major entities, corresponding to C. californica and C. occidentalis, can be dis- tinguished in most cases, based on a combination of characters. Plants called C. brachycalyx by Yuncker, with a short calyx and a long (ca. 2.5 mm) campanu- late corolla tube that may become somewhat saccate in fruit, possess all the characteristics of C. californica (see below). Although such plants are occasion- ally distinct, at other times the calyx/corolla tube ratio may vary even on the same plant, from flowers with calyx ca. equaling the corolla tube to flowers with calyx ca. 1/2 of the corolla tube length (Fig. 1 a, b,c). Apparently the co- rolla tube may continue to grow from the beginning of anthesis until fructifi- cation, significantly altering the ratio between calyx and corolla tube lengths. Similar infraspecific variation of the ratio between calyx and corolla tube lengths may be encountered in C. gronovii (between var. gronovii and var. latiflora Engelm.), but here the entities are relatively discrete (Costea et al. 2006b). For these reasons, C. brachycalyx is here considered conspecific with C. californica and not recognized at any rank. ~— COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX flowers of the same individual (scale ale m).d talis—flower (scale bar = (scale bar = 0.75 mm) ill f C. howelli 7um).g-h ae aS by corolla:g C cee h.C. penta (scale bar = 1mm). Fic. 1. Morphology of flowers and capsules. a—c. Cuscuta aig var. californica. ae of corolla tube length in mm).e. C. suksdorfii — calyx 182 BRIT.ORG/SIDA 22(1) Some plants of this complex produce a short (1.5-2 mm), campanulate-glo- bose corolla tube (as in Cuscuta occidentalis) and long anthers/styles (as in C. californica), or plants may show reversed character states (corolla tube long and campanulate and anthers/styles short). Such intermediate-like plants oc- cur at a frequency of ca. 1-2% of specimens examined. It is unclear if they are result of hybridization, which has never been clearly documented in Cuscuta, or evidence of variation in closely related and incompletely differentiated taxa. Because such possibly intermediate plants are relatively few, and to avoid no- menclatural changes, we maintain C. californica and C. occidentalis as distinct species. Several varieties within C. californica usually can be distinguished, based on single characters. Cuscuta occidentalis is less variable; a few collec- tions were found to possess papillose flowers like C. californica var. papillosa, but the origin of this variation is unclear. KEY TO SPECIES OF THE CUSCUTA SALINA~CALIFORNICA COMPLEX 1. Infrastaminal scales present (Sometimes reduced to ridges), fimbriate 2. Flowers 5—7(-—9) mm long; calyx ca. 1/2 of the corolla tube, with lobes overlap- ping at base; corolla lobes 1/4-1/3 as long as the tube; anthers 0.8-2 mm long .C. subinclusa 2. Flowers 2.8-5(-6.) mm long; calyx ca.equaling or somewhat longer than corolla ube, with non-overlapping lobes; corolla lobes + equaling the tube; anthers 0.3-0.7 mm lon 3. Flowers 5-merous; calyx and corolla lobes acute to acuminate; capsules ellip- tical-ovate, + thickened around the interstylar aperture, with 1 seed 1.C. salina 3. Flowers 4- and 5-merous; at least some calyx and corolla lobes in the same flower long-attenuate; capsules globose to slightly ice apically, with 1-4 seeds C. howelliana . Infrastaminal scales SOMPISIEY Ansel or represented by lateral, iemiate en 4. Flowers 4-5 llalobes long ac represented by lateral, dentate wings; withered corolla surrounding ae in the lower half 4 e suksdorfii 4. Flowers 5 | lla lot but not ac inf! scales completely absent; withered corolla completely enveloping the capsule or leaving only its top visi 5. Flowers short- medic taee corolla not saccate between the stamen attach- ments (if slightly saccate then with all the following characteristics); styles 1.2-3 mm long; anthers oblong to linear,0.7—1.1 mm long; capsule completely enclosed by corolla (top not visible); seeds not gg through the corolla and pericarp, which are thicker, not semitransparen 5.C. californica 5. Fl il | il II te bet the st ttachments; styles 0.5-1(-1.5) mm,anthers broadly-elliptic,0.25—0.5 mm long; capsule not completely enclosed by corolla (at least the top of capsule is visible); seeds visible through the thin and semitransparent corolla and pericarp 6.C. occidentalis 1. Cuscuta salina Engelm. in W.H. Brewer, S. Watson, & A. Gray, Bot. Calif. 1:536. 1876. Grammica salina (Engelm.) Taylor & MacBryde, Canad. J. Bot. 56:186. 1978. LECTOTYPE COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 183 (Yuncker 1932): U.S.A. UTAH: Rio Virgen, on Suaeda, saline soil, Nov 1885, Remy s.n. (MO, frag- ment NY). This was essentially a renaming at specific rank (as “C. salina, pogo n. sp.”) of what Engelmann had earlier published as C. subinclusa var. abbreviata and C. californica var. squamigera—both varieties were ated in ue y. Yuncker (1932) speci fically referred to the MO es Remy, in the herbariur f the Missouri B | Garden”) as the type of C.salin TInfl eA ae Taser Stems ae orange-yellow. : pedicels 0.5-2.5 mm long; bracts 1(-0), ovate-lanceolate to lanceolate. Flowers 5-merous, white, 2.5-5(-6.2) mm long; papillae present or absent in the corolla; laticifers conspicuous in the perianth, ovary and capsule. Calyx campanulate to narrow- campanulate, about as long as the corolla tube, divided ca. 1/2 the length, glossy yellow or brownish when dried, lobes ovate-lanceolate, acute to acuminate, + unequal, not basally overlapping or slightly so. Corolla 2.4-4.8(-6) mm long; tube cylindric-campanulate to campanulate, 1.2-2.5 mm long, lobes ovate-lan- ceolate, acute to acuminate, ca. as long as the tube, erect to spreading, some- times basally overlapping. Stamens included or exerted, anthers elliptical, 0.3- 0.7 mm long, filaments equaling to longer than anthers. Pollen 3(-4)-zonocolpate 19-22(-26) um, polymorphic, subsphaerical to subprolate, rounded at poles, tectum perforatum, puncta, 0.3-0.5 um in diameter, granulate (Fig. 2a). Infrastaminal scales oblong, short-fringed, rarely reduced to sparingly fringed ridges ca. 1/2-2/3 the corolla tube length. Styles distinct, 0.4-1 mm long; stigmas capitate, globose. Capsules elliptical-ovate, 2-3.6 x 1.4-2.1 mm, + thickened around the interstylar aperture, indehiscent or irregularly dehiscent, surrounded or capped by the withered corolla. Seeds | per capsule, not visible through the persistent corolla and pericarp, 1.35-1.57 x 1.25-1.43 mm, + dorsoventrally com- pressed, broadly elliptic to subround, hilum subterminal, subround, 0.11-0.14 x 0.7-0.11 mm, vascular scar linear, 0.02-0.05 mm, oblique; surface of seed coat epidermis alveolate when dried and papillate when hydrated, cells 30-40 um in diameter. n = 14 (Beliz 1986); 2n = 30 (Pazy & Plitmann 1995). KEY TO VARIETIES OF CUSCUTA SALINA 1. Papillae present on pedicels, calyx and/or corolla;infrastaminal scales represented by narrow, sparingly fringed ridges 1c. C. salina var. papillata 1, Papillae absent; infrastaminal scales narrowly oblong, few-toothed. 2. Corolla 2-3.5 mm; inland salt flats 1a. C. salina var. salina 2. Corolla 3-4.8(-6) mm; coastal salt marshes, tidal flats 1b. C. salina var. major la. Cuscuta salina var. salina. Cuscuta subinclusa var. abbreviata Engelm., Trans. Acad. Sci. St. Louis 1:500. 1859. Type: U.S.A. CALIFORNIA. [Solano Co.]: Mare Island in San Francisco Bay, on Arthrocnemum, Wright s.n. HOLOTYPE: MO). Cuscuta i Hook. & Arn. var. squamigera Engelm., Trans. Acad. Sci. St. Louis 1499. 1859. a salina var. sq uaMugen (Engelm.) Yuncker, Illinois Biol. Monogr. 6, pt. 2-3:71, fig. 126. 192] Im.) Piper, Contr. U.S. Natl. Herb. 11:455. 1906. Type: U.S.A. UTAH: Rio ne ea Sudeds line ol Nov 1855, J. Remy s.n. (HOLOTYPE: P; ISOTYPE: MO, fragment NY). Engelmann’s protologue cere noted “J. Remy! in Hb. Mus. Paris.” Yuncker (1921) described the type of var. squamigera as “Rio Virgen, (Remy in 1855, ...a fragment in the En- BRIT.ORG/SIDA 22(1) 184 10 m).b. C. occidentalis (scale bar = 10 m).c.—-d. C. howelliana (scale bar = 10 and 3 tm, respectively). e—f. C. subinclusa (scale bar = 10 and 3 |im, respectively). { MPs r P Fic.2.M COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 185 gelmann Herb.).” Distribution and ecology —CANADA: British Columbia. U.S.A.: Arizona, Cali- fornia, Nevada, New Mexico, Oregon, Utah, Washing MEXICO: Baja Califor- nia. Flowering Apr-Nov. Hosts: Atriplex, Cressa, Centromadia, Jaumea, Plan- tago, Salicornia, Salsola, Suaeda, Trichostema, Wislizenia, in inland salt flats, marshes, ponds. Conservation status.-T4 (apparently secure) (not yet assessed by NatureServe 2005). 1b. Cuscuta salina var. major Yuncker, Illinois Biol. Monogr. 6:161. 1921. Type: U.S.A. CALIFORNIA. Santa Clara Co. Palo Alto, frequent on Salicornia in the marshes, 14 Sep 1901, Baker 41 (HOLOTYPE: NY; ISOTYPES: CAS, GH, RSA). Distribution and ecology.—CANADA: British Columbia. U.S.A.: California, Or- egon, Washington. Flowering Jul-Oct. Hosts: various species, especially Salicornia, in coastal salt marshes, tidal flats. Conservation status.—T3T4 (vulnerable to apparently secure) (not yet as- sessed by NatureServe 2005). 1c. Cuscuta salina var. papillata Yuncker, Bull. Torrey Bot. Club 69: 543. 1942. YPE: U.S.A. CALIFORNIA. Mendocino Co.: Fort Bragg, 8-16 Aug 1912, Eastwood 1593 (HOLOTYPE: GH, fragment NY). Distribution and ecology. —U.S.A.: Arizona, California, Utah. Flowering Jun-Oct. Hosts: various species in salt marshes, flats, ponds. Conservation status.—T3T4 (vulnerable to apparently secure) (not yet as- sessed by NatureServe 2005). 2. Cuscuta howelliana Rubtzoff, Leafl. W. Bot. 10:335. 1966. Type: U.S.A. CALIFORNIA. ke Co. Boggs Lake, open dry margin of the lake, inundated in winter, 20 Aug 1966, Rubtzoff & Arnaud 5792 (HOLOTYPE: CAS, ISOTYPES: CAS, GH, OSC, RSA, UC). Stems slender, yellow to orange. Inflorescences few- to many-flowered + glomerulate cymes, pedicels 0-0.6 mm long; bracts 1-0, lanceolate. Flowers embedded in the inflorescence of the host, 4-5-merous, 3-4.5 mm long, whitish, papillate-glandular. Papillae present in the calyx and corolla; elongated latici- fers present in the calyx and corolla but not obvious (Fig. lf). Calyx campanu- late, ca. equaling corolla tube or exceeding it, divided 1/2-2/3 to the base, lobes unequal, triangular-ovate, acute, acuminate to long-attenuate and recurved at the apex, non-overlapping. Corolla tube cylindrical-campanulate, later more or less urceolate, ca. 1.5-2.2 mm long, lobes unequal, ca. equaling the tube, tri- eeu as ovate, way pee to nea Aung recurved tips, suberect to spread- ing ant 1,0.4-0.7 mm long; filaments 0.1-0.4 mm long. Pollen 3(-4)-zonocolpate (15- )17-22(-24) um, polymorph, sphaerical to subprolate (subsphaerical ), rounded at poles, tectum perforatum to microreticulate, puncta, 0.4-0.8 4m in diameter, granulate (Fig. 2 c, d). 186 BRIT.ORG/SIDA 22 = 1) Infrastaminal scales oblong-ovate, fringed, reaching to about the middle of the corolla tube; Styles distinct, evenly filiform, 0.4-11 mm; stigmas capitate, glo- bose. Capsules globose to slightly depressed, 1.2-1.5 « 0.8-1.2 mm, not thick- ened apically, indehiscent or irregularly dehiscent, completely enclosed by the withered corolla and latter capped by it. Seeds 1-4 per capsule visible through the semi-transparent corolla and pericarp, 0.9-1.43 x 0.8-1.3 mm, dorsoventrally compressed to slightly angled, subround to broadly-elliptic, hilum subtermi- nal, scar area clearly differentiated from the rest of the seed, broadly elliptic 0.21-0.25 x 0.12-0.15 mm, hilum linear 0.05-0.06 mm, vertical; heterogeneous; some areas are alveolate with cells 35-50 um in diameter; some areas irregu- larly wrinkled (Fig. 3c, e, f, g). 2n = 26 (Beliz 1986). Distribution and ecology.—U.S.A.: Endemic in California. Flowering Aug- Sep. Hosts: mostly Eryngium aristulatum, E. vaseyi, E.castrense, E.alismaefolium, Navarretia leucocephala, and N. minima, but also on Polygonum kelloggii, Epilobium pygmaeum, margins of vernal pools. Conservation status.—G2G3 (imperiled to vulnerable) (G3, NatureServe 2005). The biology of this species deserves future study. The parasite becomes strictly localized to the inflorescence region of the host. Flowers of C. howelliana develop inside the dense host inflorescences and apparently synchronize their anthesis with that of the host's flowers. The parasite achieves both protection and proximity to the flux of assimilates intended for the development of host’s reproductive structures. — 3. Cuscuta subinclusa Durand & Hilgard, J. Acad. Nat. Sci. Philad. ser. 2, 3:42. Type: U.S.A. CALIFORNIA. [Los Angeles Co]: Tejon Pass, on a willow, Heermann s.n. CHO- LOTYPE: PH presumably, fragment MO). Cuscuta eda Behr, Proc. Cali i ae (ed. 2) 1:16. 1854. The protologue he no ormariOn anda probably will be required to firmly establish th tit = of this name (see comments ca) Cuscuta ceanothi is the earlier name but is tenta- tively placed here with C. subinclusa. Stems medium, sometimes fleshy, creamy. Inflorescences few- to several-[low- ered, in scattered to densely aggregated clusters, pedicels 0-1 mm long; bracts 1-0, ovate to lanceolate. Flowers 5-merous, 5-7(-9 mm) long, white. Papillae present in the corolla lobes; laticifers isolated or in rows obvious in the calyx, corolla and fruit. Calyx campanulate, ca. 1/2 as long as the corolla tube, divided 3/5-2/3 the length, lobes broadly ovate to lanceolate, acute, sometimes cuspi- date, basally overlapping. Corolla tube cylindric, 2.5-3.5(-4.5) mm long, usu- ally showing horizontal ridges between the stamen attachments when dry, lobes ovate-triangular, acute and often slightly acuminate, 1/4-1/3 as long as the tube, widely spreading to reflexed. Stamens subincluded; anthers linear 0.8-2 mm; filaments 0-O.1 mim. Pollen 3(-4)-zonocolpate (15-)17-22(-24) um long, poly- morph, subsphaerical to subprolate, rounded at poles, tectum microreticulate, COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 187 Fic. 3. Morphology of is.a. C. californica (var. californica), lateral view. b. C. occidentalis, lateral view. c. C howelliana lateral view (scale bar = 0.38 mm). d. Hilum of C. californica. e—g. C. howelliana: e. hilum, f-g. variation of seed coat micromorphology (scale bar = 100 um). 188 BRIT.ORG/SIDA 22(1) puncta, 0.4-0.9 tum in diameter, granulate (Fig. 2 ef). Infrastaminal scales ob- long to spatulate, irregularly short-fimbriate, ca. 1/2 as long as the corolla tube; styles 1-1.5 mm long; stigmas capitate, globose. Capsules ovate to elliptical, 1.5- 3 x 1.2-2.5mm, pointed, thickened in the form of a collar about the interstylar aperture, indehiscent or irregularly dehiscent, capped by the withered corolla. Seeds mostly | per capsule, not visible through the capsule, subglobose to broadly ovoid, rarely slightly dorsoventrally compressed, 1.3-1.7 x 1.2-1.5 mm; hilum terminal; seed coat cells alveolate/papillate when dried and papillate when hydrated, cells 30-40 um in diameter. n = 14 (Beliz 1986). Distribution and ecology—U.S.A.: California, Oregon. MEXICO: Baja Cali- fornia. Flowering Apr-Oct. Hosts: wide variety of woody and sometimes her- baceous plants growing along river banks and canyon bottoms, sometimes in salt marshes and deserts, including species of Adenostoma, Amelanchier, Arc- tostaphylos, Artemisia, Asclepias, Ceanothus, Cercis, Citrus, Clematis, Erigeron, Grindelia, Heteromeles, Monardella, Rhus, Schinus, Populus, Rhododendron, Rosa, Salix, Solidago, Vitis, particularly common on Rhus and Eriogonum. Conservation status.—G4 (apparently secure) (not yet assessed by NatureServe 2005). 4. Cuscuta suksdorfii Yuncker, Mem. Torrey Bot. Club 18:167. 1932. Type: US.A / ASHINGTON. Skamania Co. on an island of a mountain lake, on Aster, 24 Sep 1891, Suksdorf 1487 (HOLOTYPE: US, fragment NY, IsoTypEs: GH, NY-2 sheets). This was essentially a renam- ing at varietal rank of Cuscuta salina var. acuminata (see below), but apparently Yunc — Ker intended for it to be taken as a homotypic new species. Stems slender, yellow. Inflorescences few-flowered umbellate clusters: pedicels 0-2 mm long; bracts 0-1, ovate lanceolate. Flowers 4-5-merous, white, 2.8-3.3 mm long; papillae absent; laticifers isolated in the calyx, corolla and ovary/ capsule. Calyx broadly campanulate often zygomorphic, reaching ca 1/2 tothe corolla lobes tips, divided 1/2-3/5, lobes ovate more or less unequal, with long- attenuate tips, not basally overlapping (Fig. le). Corolla 2.9-3.2 mm long: tube campanulate, 1.2-1.5 mm, lobes triangular-ovate, with lanceolate-attenuate tips, longer than the tube, suberect. Stamens included or barely visible through the corolla sinuses, anthers broadly elliptic, 0.2-0.4 mm long, filaments longer than anthers. Pollen 3(-4)-zonocolpate 18-22(-24) ttm, polymorph, subsphaerical to subprolate, rounded at poles, tectum perforatum, puncta, 0.3-0.5 1m in diam- eter, granulate. Infrastaminal scales oblong, represented by shallowly toothed wings, |/2-3/4 as longas the corolla tube; styles distinct, terete to slightly subu- late, 0.3-0.7 mm long; stigmas capitate, globose. Capsules elliptical-ovoid, ovoid- conic, globose to depressed globose, 2-3.2 x 2-3.6 mm, irregularly dehiscent, with withered corolla surrounding lower half. Seeds 2-4 per capsule, 0.80-1.1 x 0.8-1.02 mm, dorsoventrally compressed, subround, hilum subterminal, 0.25- 0.3 x 0.2-0.28 mm, vascular scar, 0.04-0.07 mm, oblique; alveolate when dry and papillose when hydrated, cells 30-50 um in diameter. n = 14 (Beliz 1986). — —_ COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 189 REY: 1O VARIETIES OF CUSCUIA SURSDOREL . Flowers on pedicels 0.5-2 mm; calyx 2.2-2.6 mm long; capsule ae to ovoid- conic 4a. C. suksdorfii var. suksdorfii . Flowers sessile or subsessile; calyx 1.6-2 mm long; capsule me to depressed glo eo 4b. C. suksdorfii var. Serenata 4a. Cuscuta suksdorfii var. suksdorfii. Cuscuta salina var. acuminata Yuncker, Illinois Biol. Monogr. 6, pt. 2, 3:72, fig. 32,89. 1921 (non C. acuminata Nutt. ex Engelm. 1859, in adnot.). TYPE: U.S.A. WASHINGTON. Skamania Co: on an island of a mountain lake, 24 Sep 1891, Suksdorf 1487 (HOLOTYPE: US, fragment NY; IsOTYPES: CAS, GH, MO, NY-2 sheets). Distribution and ecology.—U.S.A.: California, Oregon, Washington. Flowering Jul-Sep. Hosts: herbaceous species, mostly Asteraceae, Calyptridium, Trifolium. Mountain meadows, 5000-8500 ft Conservation status.—G2G3 (imperiled to vulnerable) (t] in NatureServe 2005). 4b. Cuscuta suksdorfii var. and Aare C uscuta suksdorfii var. sulpedieallcia Yuncker, Bull. Torrey Bot. Club 62:512. 1935. TyPE: U.S.A. CALIFORNIA. Siskiyou Co. Siskiyou Mts., Head E Fork Horse Creek, 6500 ft, 59 T47 N, R1O W, a Calyptridium umbellatum, 21 Aug 1934, L.C. Wheeler 3192 (HOLOTYPE: NY). Distribution and ecology.—U.S.A.: California, Flowering Jul-Sep. Hosts: Calyptridium. Mountain meadows, 6500 ft. Conservation status.—G2G3 (imperiled to vulnerable) (the same assessment in NatureServe 2005). 5. Cuscuta californica Hook. & Arn., Bot. Beechey Voy., 364. 1839 (non Choisy 1842). Type: Presumably in K, not designated by Hooker and Arnott. Engelmann (1859) com- mented that “Both [Choisy and Hooker & Arnott] described this plant from Douglas's speci- mens under the same name and in the same year ( )” De Candolle (Prodr. vol. 9, 1845, p. jes cited “hab. Nov. Californiam ubi rep. Douglas! h. soc. hort. Lond. Hook. bot. Beech. suppl ie 364.” Yuncker (1921, 1932) apparently repeated de Candolle’s citation, noting for the type v. Californiam’ (Douglas). Not seen.’ Stems —— yellow to orange. Inflorescences loose, cymose-paniculate clus- ters; pedicels (0.5)1-2.5(-3 mm); bracts lanceolate to ovate. Flowers 5-merous, white, 3-5(-6.5) mm long; papillae sometimes present on the pedicels, receptacle, calyx and corolla; laticifers isolated, elongated present in the corolla and capsules. Calyx turbinate-campanulate, sometimes together with the receptacle, fleshy at the base, 1/2 to ca. as long as the corolla tube, divided 1/2-1/3 to the base, lobes triangular to lanceolate, acute to acuminate, basally overlapping. Corolla persis- tent; tube campanulate-cylindric, 1.9-2.7 mm, sometimes showing horizontal ridges between the stamen attachments when dry, rarely somewhat saccate, lobes narrowly lanceolate, acute, shorter than, equaling or longer than the tube, ini- tially erect, later reflexed (Fig. 1a, b,c). Stamens + exerted when flowers are com- pletely open, anthers oblong to linear, 0.7-1.1 mm long, filaments 0.1-1.1 mm long. Pollen 3(-4)-zonocolpate (16-)17.5-22.6(-26) um, polymorph, sphaerical to pro- 190 BRIT.ORG/SIDA 22(1) late (the latter more common), rounded at poles, tectum imperforatum or witha few puncta, 0.2-0.3 um in diameter, granulate. Infrastaminal scales completely lacking or reduced, 0.1 mm long at the base of the corolla tube. Styles distinct, evenly filiform, 12-3 mm long, stigmas capitate, globose. Capsules globose or ovoid-conic, 1.6-2.3 x 1.8-2.5 mm, sometimes apically pointed, indehiscent or ir- regularly dehiscent, completely hidden by the withered corolla (Fig. 1g); pericarp and persistent corolla not semi-transparent (thicker than in C occidentalis). Seeds 1-4 per capsule, not visible through the persistent corolla and pericarp, 0.90-1.4 x 0.85-1.2 mm, dorsoventrally compressed, broadly-elliptic to obovate, hilum lateral, inconspicuous, subround 0.10-0.15 mm, vascular scar 0.05-0,06 mm, ver- tical; entirely alveolate (hilum area included) when dry and papillose when hy- drated, cells 25-50 um in diameter (Fig. 3a, d).n = 14 (Beliz 1986). In some plants of Cuscuta californica (particularly of vars. apodanthera and apiculata), the receptacle and perianth cells located at the base of capsules and seeds are fleshy and become mucilaginous when brought in contact with water. A similar phenomenon occurs in C. nevadensis LM. Johnston and to a lesser extent in C. veatchii Brandegee; Costea et al. 2005 suggest these features may be related to vivipary observed in C. nevadensis. The fleshy receptacle and perianth base in C. californica may be homologous with those of C. nevadensis, but in contrast to C. nevadensis, C. californica does not have seeds with a globu- lar-enlarged embryo at the hilum end. Apart from scattered observations such as these, the reproductive biology of Cuscuta species is poorly known (reviewed by Dawson et al. 1994; Costea & Tardif 2006). KEY TO VARIETIES OF CUSCUTA CALIFORNICA . Ovary and fruit conic, apically pointed; 1-seeded 5b. C. californica var. apiculata eee and fruit globose to globose depressed; (1-)2-4 seeded. 2. Flowers and pedicels densely papillate___ 5c. C. californica var. papillosa 2. Flowers and pedicels not papillate. 3. Anthers sessile or subsessile 5d. C. californica var. apodanther 3. Anthers not sessile, filaments shorter to equaling anthers 5a.C. Seni var. californica 5a. Cuscuta californica var. californica. Cuscuta californica Hook. & Arn. var. graciliflora Engelm., Trans. Acad. Sci. St. Louis 1499. 1859, LECTOTYPE (designated noe : oe CALIFOR- NIA: “Almost without flowers, on Monarda,” datal, 1846 (MO). Engelmann cited “California, Dotigiast Fremont! 506: ae Cuscutad ca omnes Hook. & Arn. var. li loba Engelm., Trans. Acad. Sci. St. Louis 1:499. 1859. LECTOTYPE (designated here): U.S.A. CALIFORNIA: San Felipe, on Eriogonum polifolium, Jan 1852, Thurber 633 (MO). Engelmann pee ‘Sta. Barbara, Nuttall! San Diego, Thurber! 570 & 633; Newberry!” Engelmann’s material (at MO) of the Nuttall collection apparently is a fragment of the son ee specimen at PH, judging from notes inside the packet Cus tlifornica Hook. & Arn, var. brachycalyx Yuncker, ite Biol. Monogr. 6, nos. 2, 3:62, fig. 45, 75, 51921 Type: U'S.A. CALIFORNIA: Near Hanford, dry soil on Centromadia pungens, 21 Jun L901, Kearney 52 (HOLOTYPE: NY; ISOTYPE: US). COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 191 Cuscuta brachycalyx Yuncker, Mem. Torrey Bot. Club 18:159. 1932. Type: U.S.A. Cam IFORNIA: Near Hanford, dry soil on Centromadia pungens, 21 Jun 1901, Kearney 52 ( ). Yuncker treated this as a new species, even though he cited C. californica var. br ee in synonymy. Distribution and ecology.—U.S.A.: Arizona, California, Nevada, Oregon, Utah, Washington; Mexico (Baja California). Flowering Mar-Aug(-Sep). Hosts: Abronia, Adenostoma, Agastache, Ambrosia, Asclepias, Convolvulus, Corethrogyne, Croton, Eriodictyon, Eriogonum, Holocarpha, Iva, Lupinus, Salvia, and others. 200-22650 feet. Conservation status.—T5 (common) (the same assessment in NatureServe 2005) 5b. Cuscuta californica var. apiculata Engelm., Trans. Acad. Sci. St. Louis 1:499. TYPE: U.S.A. CALIFORNIA: “Dry arroyos, on Dalea spinosa, [on the] Colorado,” 22 Feb 1884, J.M. Bigelow s.n. (HOLOTYPE: MO; ISOTYPES: GH, NY). Distribution and ecology—U.S.A.: California, Nevada. MEXICO: Baja California. Flowering Mar-Aug. Hosts: various species in sandy desert areas. Conservation status.—T1T2 (critically imperiled to imperiled), (T3?, Natureserve 2005). 5c. Cuscuta californica var. papillosa Yuncker, Illinois Biol. Monogr. 6:152. 1921. TYPE: U.S.A. CALIFORNIA. San Bernardino Co. San Bernardino Valley, Parish 5524 (HOLOTYPE: el Distribution and ecology.—U.S.A.: California (endemic): Imperial, Riverside, San Bernadino, and San Diego cos. Flowering Mar-Aug, -Sep. Hosts: various species in sandy desert areas. 300-4600 feet. Conservation status.—T3 (vulnerable) (the same assessment in NatureServe 2005) 5d. Cuscuta californica var. apodanthera Yuncker, Illinois Biol. Monogr. 6:152. 1921. Cuscuta brachycalyx var. apodanthera (Yuncker) Yuncker, Mem. Torrey Bot. Club 18:159. 1932. Type: U.S.A. CALIFORNIA: Yosemite Valley, 7-12 Jul 1896, Jepson 80a (HOLOTYPE: JEPS, frag- ment NY). Distribution and ecology.—U.S.A.: Endemic in California. Flowering Aug-Sep. Hosts: Eriogonum and other herbs. Conservation status.—T1T2 (critically imperiled to imperiled) (T2, NatureServe 2005). 6. Cuscuta occidentalis Millsp. nom. nov. Bee Field Columbian Mus., Bot. Ser. 204.1923. Based on C Arn. var. breviflora Engelm., Trans. Acad. Sci. St. Louis 1:499. 1859. Grammnicd Bei (Millsp.) cca & Chrtek, Folia Geobot. Phytotax. 8:220. (Aug) 1973. Grammica occidentalis (Millsp.) W.A. Weber, Southw. Naturalist 18:319. (Oct) 1973. TYPE: U.S.A. CALIFORNIA. Monterey Co.: rae in fields, [no date], Hartweg 1863 (HOLOTYPE: MO; ISOTYPES: GH, NY) Stems medium, yellowish to orange. Inflorescences compact glomerulate clus- ters; pedicels 0-0.5(-1.5 mm); bracts lanceolate to ovate. Flowers 5-merous, white 192 BRIT.ORG/SIDA 22(1) to creamy; 2.7-3.4 mm long; papillae rarely present; laticifers not conspicuous (Fig. 1d). Calyx campanulate, somewhat shorter to ca. as long as the corolla tube, divided 2/5-1/2 the length, rarely fleshy at base, lobes narrowly ovate to lan- ceolate, acuminate, not basally overlapping; corolla tube cylindric-campanu- late, 1.4-2.1 mm long, saccate between the stamen attachments, lobes lanceolate, acuminate, shorter than the tube, initially erect, later usually spreading (some- times reflexed) giving flowers a star-shaped appearance. Stamens + exserted; anthers broadly-elliptic, 0.25-0.5 mm long, filaments 0.3-1 mm long. Pollen as in C. californica, 17-)19-24(-26) um long. Infrastaminal scales as in C. californica; styles distinct, evenly filiform, styles 0.5-1(-L5) mm long; stigmas capitate, globose. Capsules globose to globose-depressed, 1.8-2.2 x 2-2.6 mm, indehiscent or irregularly dehiscent, not completely enclosed by the corolla (the top of capsule visible) (Fig. 1h); corolla and pericarp thin, membranous and semi-transparent. Seeds 2-4 per capsule, visible through both corolla and peri- carp, 0.85-L3 « 0.8-1.1, dorsoventrally compressed, subround to broadly-ellip- tic, hilum subterminal, poorly differentiated from the rest of the seed, subround 0.10-0.15 mm, vascular scar 0.05-0.06 mm, vertical, surface of seed coat alveo- ate when dry (including hilum area), cells 25-50 um in diameter (Fig. 3b). Distribution and ecology.—U.S.A.: California, Colorado, Idaho, Nevada, Or- egon, Utah, Washington, Wyoming. MEXICO: Baja California. Flowering Mar- Aug(-Sep). Hosts: Artemisia, Boisduvalia, Calyptridium, Corethrogyne, Ericameria, Eriogonum, Hemizonia, lva, Lotus, Lupinus, Monardella, Oenothera, Polygonum, Salvia, Sisymbrium, and others. 25-6500 feet. Conservation status.—G4G5 (apparently secure to secure) (not assessed yet by NatureServe 2005). APPENDIX 1.~VOUCHERS FOR THE SEM STUDY Vouchers are from NY unless otherwise indicated. 1. Cuscuta salina (see Costea et al. 2005). 2.Cuscuta howelliana (7 collection d).— U.S.A. CALIFORNIA. Butte Co.: ca 7 mi N of Oroville, Table Mountain, E side of Cherokee Rd., 1325 ft, 18 Jul 1998, Ahart 6044 (JEPS). Sacra- mento Co.: ca 1 mi S of White Rock Rd. and Scott Rd. intersection, 27 May 1985, Bowcutt 337 (UC). Shasta Co.: between Goose Valley and Burney Valley, ca. 3.5 mi NNW from Burney, 3200 ft, 8 Aug 1988, Taylor 10026 (JEPS). Siskiyou Co.: 16 mi N of Fall River Mills (Shasta Co.) and 6 mi W of Day (Modoc Co.), 1.2 mi E of Spring Creek Rd., 3600 ft, 19 Jul 1989, Powell 3533 EPS). Tehama Co.:ca 10 mi SE of Corning, S Fork Hall Creek, 460 ft, 12 May 1995, Taylor 14897 (JJEPS); BLM parcel along Hwy 36 NE of Red Bluff, 870 ft, 8 Jul 1996, Oswald & Ahart 7978 (JEPS): Inks Creek, ca. 1.5 mi N of Dales Lake, 612 ft, 27 May 1992, Taylor 12661 (UC). 3. Cuscuta subinclusa (11 collections examined). oe S.A. CALIFORNIA. [no county given, Los Angeles or San Bernardino Co.?] Swartout, 1 Sep 1939, Grace s.n. Los Angeles Co.: Liebre Mts., entrance to Ruby Canyon of Forest ae Rd., ca. : 0 mi from junction with Lake Hughes Rd., 2000 ft, 23 Oct 1996, Raz & Boyd 015; Lower Spunky Canyon, stretch draining SSE exiting into Bouquet Reservoir, 2995-3065 ft, 20 Sep 1994, Ross & Boyd 8260; San Gabriel Mts., Tujunga Creek, COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 193 near mouth of Canyon, 23 Aug 1937, Ewan 11049. Marin Co.: mouth of San Antonio Creek, 3 Sep 1939, Howell 15355; Mouth of San Gabriel Canyon, 17 Dec 1935, Hastings s.n.;San Antonio eo N of Novato, 5 ft, 3 Sep 1939, Rose 39363. eOner ey Co.: Banks of San Antonio River, Santa Lucia Mts., 25 Oct 1930, Mason 5766.Nevada Co.: Banner Mt.,ca.4 mi E of Nevada City, 3900 ft, 15 Jul er True & Howell 2315. Plumas Co.: Mill erealiun 1877, Austin s.n. San Joaquin Co.: San Joaquin Bridge near Lathrop, 9 Sep 1892, Brandegee s.n 4.Cuscuta suksdorfii (5 collections examined).—U.S.A. CALIFORNIA. Humboldt Co:: Trin- ity Summit, near Box Camp, 5000 ft, 4 Aug 1949, Tracy 18430 (UC). Siskiyou Co.: Siskiyou Mts., M Diablo, 1 mi E Dry Lake Lookout, 6000 ft, 31 Jul 1934, Wheeler 3011 UJEPS);.[Co. inepecieal ae Canyon National Park, Woods Creek, near base of Castle Dome, 8500 ft, 27 Jul 1948, Bailey & Bailey 2672a (UC). [Co. unspecified] ca 1/4 mi E of Yuba Gap and Hwy 80, E side of the paved road to Lake Valley Reservoir, 5840 m, 22 Jul 2002, Ahart 9885 (JEPS). [No location, no date, no collector] (UC 23290). 5. Cuscuta uae 5a. uta californica var. californica (13 collections examined).—U.S.A. CALIFORNIA. [locality nee 21 Jun 1931, Yuncker & Welch 3748. Kern Co.: “brachycalyx,” 1.5 mi N of Kernville, ep 1962, Howell 38877. Las Angeles Co.:Liebre Mts., Knapp Ranch at the ee end of Castaic Creek drainage in pio alluvial valley at head of Cienaga Canyon, S of Liebre 2840- 3080 ft, 28 May 1997, Boyd 9839; [some flowers “brachycalyx"-like] Whittier Hills, S- aan see of Turnbull Canyon, 980-1000 ft, 15 Jun 1992, Ross 6575; Mount Wilson, 3000 ft, 17 Jul 1931, Yuncker & Welch 3704, 3707. Orange Co.: Santa Ana Mts., Silverado Canyon, 21 Jun 1931, Fosberg 5143. River- side Co.: 8 mi SE of Corona, along road to Elsinore, 17 Mar 1964, Hitchcock & Muhlick 23076. a Bernardino a foot hills of San Bernardino Mountains, 15 Jun 1898, Parish s.n. San Diego Co. Jamul Valley, Jun 1875, Palmer 439; San Diego, Mission Hills, 14(?) May 1903, Abrams 3462; along re 78E 5.3 Mi W of Ramona, 15 Jun 1973, LeDoux et al. 114; [some flowers “brachycalyx”-like] Coronado Beach, 24 Jun 1932, Moldenke 7037. San Luis Obispo Co.: Pismo, 25 Aug 1932, Demaree 9281.NE- VADA. Clark Co.: Hells Kitchen area near Gold Butte, Devils Cove Rd., 2.9 mi S of Connoly Spring, 2400 ft, 14 May 1986, Pinal 72 5b. Cuscuta californica var. iepapillosa (5 collections examined).—U.S.A. CALIFORNIA. [no a Aug 1901, Grant 4449. Riverside Co.: Cathedral City, 23 Apr 1945, Lillian s.n.; ca. 500 ft, 27 1945, Rose 45331; Cathedral Canyon, 23 Dec 1946; Rose 46300. San Bernardino Co.: 1/2 a E of ae Ranch, Horsethief Canyon, Mohave River tributary, 3300 ft, 17 Sep 1933, Wheeler 271 Cuscuta californica. var. apiculata (3 collections examined).—U.S.A. PAIEORAIA Riv- erside Co.: Cathedral City,no date (probably 1945), Blake s.n. NEVADA. Clarke Co.: 4 mi SE of Muddy Peak at head of Callville Wash, 2500 ft, 28 May 1937, LaRivers & Hancock 261; 29 May 1937,LaRivers & Hancock 231 ta identalis (14 collections examined).—U.S.A. Tee Fresno Co.: Cop- per Creek Canyon Creek Trail, 7200 ft, 2 Aug 1958, Howell 34198.Mono Co.: 1 mi Eof Mammoth Post Office, 29 Jul 1951, Raven 3720.Monterey Co.: Point Lobos State Park, 16 Jun ees. 9219. Siskiyou Co.: along Klamath River Hwy.W of Pacific Hwy., 2000 ft, 20 Jun 1940, Yuncker 15144. Trin- ity Co.: Cold Springs Area, South Fork Mt.,5400 ft, 28 Aug 1941, Parks & Tracy 11517. Washoe Co. 1 mi ENE of Grass Valley Reservoir, 6000 ft, 14 Aug 1991, Schoolcraft et al. 2220. COLORADO. Duis oe Paonia, 23 Jul 1911, Osterhout 4602.IDAHO. Adams Co.: 2 Mi S of Council, 26 Aug 1937, Christ & ard 8787.Gooding Co.: Little City of the Rocks in the Mt. Bennett Hills N of Gooding, 28 Jul 1976, eae NEVADA. Douglas Co.: Antelope Valley at the E edge of Red Hill on the E side of Gray Hills, 4990 ft, 14 Jul 1997, Tiehm 12257. Eureka Co.: Roberts Mts., 1.7 mi N of Three Bars Ranch on the main road to Tonkin Summit, 6600 ft, 22 Aug 2002, Tiehm 14108. Mineral Co.: E slope of Wassuk Range, base of Big Indian Mt., 9500 ft, 11 Sep 1938, Archer 7032. Washoe Co.: dry bed of Ice Pond, Whittell-Audubon Area, Carson Range foothills, 5480 ft, 12 Aug 1974, Howell et al. 50745.UTAH. Salt Lake Co.: [no locality], 16 Aug 1907, Garrett 2170. 194 BRIT.ORG/SIDA 22(1) ACKNOWLEDGMENTS We thank directors and/or curators from ACAD, ALTA, ARIZ, ASU, BRIT, DAO, F GH, HAM, MEXU, MICH, MT, MTMG, NFLD, NSPM, OAC, QFA, QUE, RBG, Roa opoK ols, (EA Oo LE.TUP UBC, UC EPS. UNE, UNM, US USis.U WO: UWPG, WAT, WIN, WIS, WTU, and XAL for loans to Costea. Special thanks are due to NY staff for approving and preparing the four voluminous loans con- taining the herbarium of T.G. Yuncker. Dan Austin and an: mous reviewer provided useful comments and suggestions for an earlier version of the manu- script. Thierry Deroin kindly donated all the Cuscuta fragments annotated by Yuncker (held in P) to WLU. Barbara Ertter (UC) sent us copies of the relevant publications of Behr and Currant. Alexandra Smith assisted us with the scan- ning electron microscope and Gabriel Moreno-Hagelsieb translated the Span- ish abstract. REFERENCES BeHr, H. 1888. Flora of the vicinity of San Francisco. San Francisco, California (privately published). Beuz, T. 1986.A revision of Cuscuta sect. Cleistogrammica using phenetic and cladistic analy- ses with a comparison of reproductive mechanisms and host preferences in species from California, Mexico, and Central America. Ph.D. diss., Univ. of California, Berkeley. Betiz, T. 1993. Cuscuta.|n: J. Hickman, ed. Jepson’s manual: higher plants of California. Univ. of California Press, Berkeley. Pp. 538—539. Costea, M. and FJ. Taroir. 2006. Biology of Canadian weeds. Cuscuta campestris Yuncker, C. gronovii Willd. ex Schult., C. umbrosa Beyr.ex Hook., C. epithymum (L.) L.and C.epilinum Weihe. Can. J. Pl. Sci. 86:293-316 Costea, M.,G.L.Nesom, and FE. Tarpir.2005. Taxonomic status of Cuscuta nevadensis and Cuscuta veatchii (Convolvulaceae). Brittonia 57:264-272. Costea, M., G.L. Nesom, and S. Steranovic. 2006a. Taxonomy of the Cuscuta pentagona com- plex (subsect. Arvenses: Convolvulaceae) in North America. Sida 22:151-175. Costea, M., G.L. Nesom, and S. Steranovic. 2006b. Taxonomy of Cuscuta gronovii and Cuscuta umbrosa (Convolvulaceae). Sida 22:197-207. Costea, M., G.L. Nesom, and S. Steranovic. 2006c. Taxonomy of the Cuscuta indecora (Convolvulaceae) complex in North America. Sida 22:209-225. Curran, M.K. 1885. List of the plants described in California, principally in the Proc. of the Calif. Acad. of Sciences, by Dr. Albert Kellogg, Dr. H.H. Behr, and Mr. H.N. Bolander: with an attempt at their identification. Bull. Calif. Acad. Sci. 1:128-151. Dawson, J.H., LJ. Musseuman, P.WotswinkeL, and |. Dorr. 1994. Biology and control of Cuscuta. Rev.Weed Sci. 6:265-317. ENGELMANN, G. 1859. Systematic arrangement of the species of the genus Cuscuta with critical remarks on old species and descriptions of new ones. Trans. Acad. Sci. St. Louis 1:453-523. COSTEA ET AL., TAXONOMY OF CUSCUTA SALINA-CALIFORNICA COMPLEX 195 ENceLMANN, G. 1876. Cuscuta salina. In: W.H. Brewer, S. Watson, and A. Gray. The botany of California, Univ. Press, Cambridge. P. 536 Ertter, B. 2003. Mein lieber herr College: letters from Behr to Engelmann. Fremontia 31: 18-22. NatureServe. 2005. NatureServe Explorer:An online Sneyclopediar of life [web pplication! Version 4.0, NatureServe, Arlington, Virginia. . Dawson, J.H. 1984.A vegetative character that separates species of Cuscuta.Proc. 3rd Interntl. Symposium on Parasitic Weeds. Pp.184-187 COSTEA ET AL., TAXONOMY OF CUSCUTA GRONOVII AND C. UMBROSA 207 ENGELMANN, G. 1842. A monograph of the North American Cuscutineae. Amer. J. Sci. 43: 333-346 ENGELMANN, G. 1859. Systematic arrangement of the species of the genus Cuscuta with critical remarks on old species and descriptions of new ones. Trans. Acad. Sci. St. Louis 1:453-523. Feinsrun, N. 1972. Cuscuta L.In:T.G.Tutin, V.H. Heywood, N.A. Burges, D.M. Moore, D.H.Valen- tine, S.M.Walters, D.A.Webb, eds. Flora Europaea, Vol. 3, Diapensiaceae to Myoporaceae. Cambridge University Press. Pp. 74-77. FOGELBERG, S.O. 1938. The cytology of Cuscuta. Bull. Torrey Bot. Club. 65:631-645. Gaertner, E.E. 1950. Studies of seed germination, seed identification, and host relation- ships in Dodders, Cuscuta spp. Mem. Cornell Agric. Exp. Sta. 294:1-56. Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland. First edition. In: Kartesz, J.T. and C.A. Meacham. 1999. Synthesis of the North American Flora, Version 1.0. North Carolina Botanical Garden, Chapel Hill, NC. MitcHett, R.S.and G.C. Tucker. 1997.A checklist of New York state plants (rev. ed.). New York State Bull. 490. New York State Museum, Albany. NatureServe. 2005. NatureServe Explorer: An online encyclopedia of life [web application, Version 4.0. NatureServe, Arlington, Virginia. K++ Version 4.0. NatureServe, Arlington, Virginia. tion, the auth northern one “Amen but aie it isan ecosystem a extends from the higher mountains of thern Peru, thus embracing these countries as well as Colombia, Ecua- ou Costa Rica anc dor, and Venezuela. It is an ecosystem rich in biodiversity and endemism and one of ecological ex- tremes, to which the flora has adapted. Plant characteristics include acaulescent rosettes, shrubs with minute leaves, prostrate dwarf shrubs, and geophytes. The purpose of the book, however, is not to introduce these plants but to serve students a professionals as a guide to the g of vascular plants, ultimately as an incentive for the protection of this unique anc luabl 5 With this objective in mind, the authors provide many keys: a basic one to each of e on owering plants, with especial attention to leaf structures, which ¢ groups from pteridophytes to [1 count for five of the eight general keys. Within each of the 12 families esa aanees eee are keys to the 540 genera and keys to tribes. The text ful ly eae ibes the genus and and altitudes at which the plants grow. Accompa he text is a picture, in most cases a crawls e species in the genus or a repre eseniative aes almost all of these were previously pub- — of the sing lished and are so acknowledged in an appendix. References follow each entr The five authors are at eee meena institutions. The leading ae Petr Sklenaf is associ- aches at Charles University, Prague. Nine other pro- fessionals are cited a rie ee TCU Library), Botanical Research Insti- tute of Texas, Fort ate TX 76102- 4060, US. :ditor’s note.—See page 724. ated with the New York B 4 SIDA 22(1): 242. 2006 BIG-FRUITED BUCKTHORN, SIDEROXYLON MACROCARPUM (SAPOTACEAE), A LONG-FORGOTTEN GEORGIA ENDEMIC James R. Allison PO. Box 511 edge, Georgia 30663-0511, U.S.A. jallison@mindspring.com ae Rut ABSTRACT Bumelia macrocarpa Nutt. languished in obscurity for more than 150 years. Thomas Nuttall (1786- 1859) collected uae low shrub in 1830 and described it in 1849. Asa Gray relegated it to the syn onymy of B. lanuginosa (Michx.) Pers. in 1886, apparently due to the lack of subsequent collections and without seeing any material of Nuttall’s plant. In 1940 Robert Clark identified type material of B. macrocarpa as B. reclinata (Michx.) Vent, sete to notice the fact that more than one species had been mounted on the sheet, with only ot all fragment actually representing B. macrocarpa. Since Bumelia Swartz is now considered to asynonym of Sideroxylon me the restoration of this distinc- tive species, endemic to southeastern Georgia (U.S.A.), requires a new combination, S. macrocarpum (Nutt.) J.R. Allison. RESUMEN Bumelia macrocarpa Nutt. languidecio en la oscuridad durante mas de 150 anos. Thomas Nuttall (1786-1859) coleccion6 este arbusto bajo en 1830 y lo describid en 1849. Asa Gray - aesee ala 1 | sinonimia de B. lanuginosa (Michx.) Pers. en 1886, aparentemente debido subsiguientes y sin ver ningtin material de la planta de Nuttall. En oe Robert Clark identifico el material tipo de B. macrocarpa como B. reclinata cas Vent., al no notar el hecho que se nab montado en la lamina mas de una especie, con B. macrocarpa. Puesto que Bumelia Swartz s la restauracion de esta especie distintiv: lémica al UU), requiere una nueva sidera actu: sienenis un sinonimo de Sideroxylon L., combinacion, S. macrocarpum (Nutt.) J.R. Allison. INTRODUCTION Among the genera of woody plants listed in the work that launched the mod- ern system of binomial nomenclature, Linnaeus’ Species Plantarum of 1753)! was Sideroxylon L. ( 17 um diam), and / or micrograins. Conversely, a pollen sample was identified as being produced by a sexual plant if it generally included a low proportion of aborted grains, was uniformly small (ca. 12-15 um diam), and lacked micrograins. Measurements of grains, as required, were performed on pollen images captured and analyzed using AnalySIS (v3.1; Soft Imaging Sys- tem, GmbH 1989-2001). The longitude and latitude of each collection location was determined from the data provided on the herbarium specimen label. This was accomplished initially using the mapping software TOPO! Texas (Copyright 2004, National Geographic Maps, Evergreen, CO) and later using TOPO USA (version 5.0, DeLorme, Yarmouth, ME), facilitated on occasion with text web inquiries ae Google (Google, Inc, Mountain View, CA). Locality data were compiled in Excel (version 10.0.4, Microsoft Corp.). If not explicitly stated, distances and direc- tions from towns given on herbarium specimen labels were assumed to lie along existing roads. In most cases, the collection data were sufficient to estimate co- 268 BRIT.ORG/SIDA 22(1) ordinates with a high degree of confidence. For a few older specimens, only a county or town was provided, in which case coordinates were selected at the geographic center of the county or town. To evaluate differences in the phenol- ogy of apomictic vs. sexual plants, collection dates were compared statistically using Excel. This test assumed that the herbarium specimen collection dates, taken as a whole, accurately reflected the duration of flowering in the species. Localities were mapped using Geographical Information System (GIS) soft- ware ESRI ArcInfo (v. 9.0, copyright 1999-2004; http://wwwesri.com). Dupli- cate speci within or between herbaria were removed from the database so that subsequent analyses were performed only on unique collections. Asa check on locality coding, county names for data plotted in ArcInfo were extracted and compared with the original data set. In addition, ca. 20% of the herbarium speci- mens were selected at random and plotted localities were evaluated for accu- racy. Geographic distributions of sexual and apomictic plants were analyzed visually and statistically with respect to rainfall, natural areas, and soils using GIS data layers for Texas available from the Texas General Land Office (http:// www.glostate.tx.us/gisdata), the Texas Park and Wildlife Department (http:// www.tpwd.state.tx.us/landwater/land/maps/gis/map_downloads), and the United States Department of Agriculture (http://www.ncgce.nrcs.usda.gov/ products/datasets/statsgo/data/tx.html), respectively. The soil classification employed followed the taxonomy proposed by the United States Department of Agriculture that recognizes 12 soil orders and 64 soil suborders (Soil Survey Staff 1999). Specific environmental point data for collections were obtained using the Intersect Point Tool (Beyer 2004), and statistical analyses were con- ducted using Excel. RESULTS A total of 311 Texas collections were studied. These included 184 specimens from BRIT and 127 specimens from TEX. In addition, floral material from the holo- type of Erigeron traversii Shinners (C.L. Lundell 11,093, 11 Apr 1942, BRIT) was obtained via special arrangement with curators (185 BRIT Texas collections analyzed, 312 collections total). Twenty collections were duplicated between BRIT and TEX, one collection was represented by two separate sheets at TEX, and three collections were represented by two separate sheets at BRIT. In total, there were 288 unique collections between the two institutions. Three collec- tions (1%), all from TEX, were excluded from subsequent analyses: G.L. Fisher 38,018 (1938) and W.R. Carr 11,924 (1992) lacked capitula at a suitable age from which pollen could be obtained, and the locality (even to county) could not be determined for A. Gieseuschlag s.n. 1928). The earliest collection (FE Lindheimer 6978) was made near Austin in 1844, and the largest portion of specimens (113, 40%) was collected in the 1940s (Fig. 1). Although there were specimens pre- pared by 95 principal collectors, the majority (52%) was collected by just seven NOYES ET AL., ERIGERON STRIGOSUS IN TEXAS 269 120 100 a “” O ss 80 iS) 2 © 60 O fe) = 40 ji EL | 0 +=; T T T T Tt .. T T T LLL, — —_ — —_ = —_ oh — —_ — =k ah — hk = = ie) o7on7n ow Mm oOo mo 6 6 6 6 © © 6 © © © 6 khaonntowoeosz NY © FB aAON © © GS a — = — 0 fF DH DH HF HH HF HHH HG H Fic. 1. Erigeron strigosus collections by decade in Texas. Solid b ictic (n = 245) b I (n = 40). r 7 es r c : : s < < botanists: L. Shinners, 40 (14%); E. Whitehouse, 29 (10%), V.L. Cory, 27 (9.5%), DS. Correll, 13 (5%); B.C. Tharp, 13 (5%), and CL. Lundell, 11 (4%), who were prolific in the middle decades of the 20th century, and WR. Carr, 14 (5%), whose collections were prepared in the 1990s. The 285 specimens were collected in 87 of 254 Texas counties (Fig. 2A), which are concentrated in north central and east Texas. Most of the collections (87%) were restricted to Pineywoods (88, 31%), Post Oak Savannah, (87, 31%) and Blackland Prairie (73, 26%) (Fig. 2B). The remaining 37 collections (13%) occurred in adjacent regions to the west (Edwards Plateau, 12; Rolling Plains, 14; Llano Uplift, 4) or to the south (Gulf Coast Prairies and Marshes, 5; South Texas Brush Country, 1). Additionally, one specimen occurred in the extreme north Panhandle in Lipscomb County. It was the only specimen to occur in the High Plains region of the state. No specimens were collected in the Trans Pecos region or Coastal Sand Plain of southern Texas. The western boundary of Eri- geron strigosus in Texas occurs abruptly at ca. 99°W in the central part of the state. This corresponds approximately to the 28 in. precipitation isohyet (Fig. 2C). The Lipscomb county outlier is exceptional in longitude and rainfall, oc- curring at LOO’W in a region with approximately 23 in. annual rainfall. Of the 285 specimens, 40 (14%) were determined to possess pollen charac- teristic of sexual diploids, and 245 (86%) were determined to possess pollen characteristic of polyploid apomicts (Fig. 3). Polyploid apomicts through- out the range of the species in Texas. Sexual diploid plants, however, were re- stricted to Pineywoods (34 / 40, 85%) and Post Oak Savannah (6 / 40, 15%). 270 BRIT.ORG/SIDA 22(1) oO sexual A apomictic precipitation. Isohyet values in inches. Scale bars = 100 km. Sexual plants were collected in 17 counties within these two regions. In six of these counties, only sexual plants have been collected. However, in the other 11 counties, both sexual and apomictic plants have been collected. The oldest sexual specimen was collected by CLL. York in 1928 (s.n., Gregg Co.). There is no significant trend in the proportion of sexual plants collected by decade in the 20th century (r = 0.19, P = 0.58). This indicates that both sexual and apomictic NOYES ET AL., ERIGERON STRIGOSUS IN TEXAS 271 Fic. 3. Pollen from herbarium specimens of Erigeron strigosus. A. The holotype of E. traversii (CL. Lundell 11,093, Nacogdoches Co., BRIT), sexual. B. L.H. Shinners 32,543 (Morris Co., BRIT), apomictic. Arrow indicates a micrograin. Scale bar = 20 pm. plants have coexisted in Texas since at least the 1920s, and that their relative proportions have not changed dramatically. The holotype of Erigeron traversii was determined to have pollen characteristic of sexual diploids, as did the speci- mens L. H. Shinners 7618 (BRIT), 18,387 (BRIT), 18,402 (BRIT), 18,450 (BRIT), 18,503 (BRIT), and B.L. Turner 4377 (TEX), which were cited in the original de- scription of EF. traversii (Shinners, 1964). E. Whitehouse 20,861 (BRIT), the eighth specimen cited by Shinners for Texas, was not present among the loan materials. Precipitation at sites where sexual plants were collected ranged from 41 to 57 in. / year (mean = 48.2 in., SD = 4.9), while values at apomictic sites ranged from 23 to 57 in. / year (mean 39.1 in., SD = 6.3). Precipitation values for apomic- tic plants, though overlapping, were significantly lower than those for sexual plants (Student t, P = 6.04E-17). Sexual plants predominantly occurred on leached forest soils with subsurface clay (alfisols, 18 collections; ultisols, 18 col- lections). Four sexual collections occured on poorly developed entisols. Apomic- tic plants in Oak Savannah and Pineywoods regions also occurred on alfisols (72 collections) and ultisols (28 collections), but also were found on clay-rich vertisols (11 collections), and dark grassland mollisols (12 collections). Six apomictic collections corresponded to entisols. Collections dates of specimens of Erigeron strigosus (Fig. 4) indicate that flowering commences in March, peaks in April and May, and continues spo- radically at reduced levels until October or early November. One exceptional apomictic specimen (R,J. Fleet wood 10,050, Nacogdoches Co., TEX) was collected 5 Jan 1972 (the collector noted the presence of ice on the plant). The average collection date for sexual plants (for 39 collections made from March to June) was 22 April (SD = 23.3 days). The average collection date for 210 apomictic plants made during the same time period was significantly later (10 May, SD = 272 BRIT.ORG/SIDA 22(1) | q | | q . . | ‘| apomictic = } C] sexual Relative Proportion T | 1 T | t I T I qT I rn T t | A + | t | qT | t | 1 T I JtrF>m{ta};{m{si fila Fig. 4. Erig eu? W * oo | JIi+,f £ } } + + 7b 22.5 days, Student t, P = 2.5E-06). The same tendency was observed for asexual plants occurring in only those counties where sexual plants also occurred (n = 22, avg. = 9 May, SD = 20.4 days; Student t, P = 0.006). DISCUSSION Sexual and apomictic Erigeron strigosus in Texas The greater range of habitat, precipitation, and tolerated by apomictic Eri- geron strigosus in Texas is consistent with the hypothesis (Baker 1965; Lynch 1984) that asexual lineages, through polyploidy, hybridization, and genome level selection, evolve general-purpose genotypes that are capable of exploiting di- verse habitats. However, this pattern is al nsistent with the hypothesis that apomictic reproduction may be favored in fragmented marginal habitats be- cause it circumvents the deleterious effects of inbreeding depression and mate scarcity (Haag & Ebert 2004). Detailed, fine-scale analysis of populations, es- pecially in regard to population size, habitat, and disturbance, will be required to differentiate between these two hypotheses. Superficially, the geographic pattern for E. strigosus is similar to that for Taraxacum officinale in Europe. Sexual diploid populations of that taxon are restricted to low-elevation habi- tats in south central and southeastern Europe, while apomictic populations are found throughout northern Europe (Den Nijs @ Menken 1994). The historical record indicates that sexual and apomictic Erigeron strigosus have co-existed in Texas at least since the early 20th century, and that there- ay | NOYES ET AL., ERIGERON STRIGOSUS IN TEXAS 273 1:1 fore, the two different forms might be in stable eq ium. Investigations of E. strigosus in Georgia document that gene flow from apomicts to sexual diploids may occur, such that progeny of the latter may include a significant proportion of apomicts (Noyes unpublished). Whether or not sexual and apomictic plants occur simultaneously at the same sites in Texas, and hybridize, is unknown. However, because the phenology of the two forms overlaps, hybridization is likely. Hybridization between sexual and apomictic plants has been thought to be important as a source of novel apomictic genotypes (Stebbins 1950) and has been hypothesized to be an important part of the dynamics between apomictic and sexual Taraxacum in Europe (Menken, Smit, @ Den Nijs 1995; Verduijn, van Dijk, & van Damme 2004). However, in theory, apomictic plants should ultimately replace sexual populations through reproductive advantage (Maynard Smith 1978) or by unidirectional gene flow via pollen from apomic- tic to sexual plants (Mogie 1992). To determine if apomictic and sexual Erigeron strigosus are in dynamic equilibrium in Texas, or if replacement is occurring, detailed demographic study will be required. Status of E. traversii and a new combination Although not known by Shinners at the time he published the species, pollen evidence indicates that Erigeron traversii is a sexual diploid taxon. It is there- fore distinct from other Erigeron strigosus in Texas, which, based on pollen, is apomictic and polyploid. The analysis of collection dates supports Shinners (1964) observation that E. traversii, on average, flowers earlier than E. strigosus. Although Shinners documented E. traversii in Pineywoods habitats, our data also document the existence of sexual diploids in Post Oak Savannah. Accord- ing to Shinners, E. traversii was distinct in the field from E. strigosus in produc- ing showier plants. Whether that difference is due to features of the capitulum (colors, shapes, presentation) or shape and structure of the inflorescence has not been evaluated. These traits, as well as geographic variation among sexual plants, require further study. The question of whether or not E. traversii should be resurrected based on the available evidence is not without controversy. Erigeron traversii is distinct from E. strigosus in ploidal level and mode of reproduction but the two taxa are sympatric and likely hybridize. However, the sexual taxon appears to be a con- sistent component of forested habitats of east Texas. To recognize the distinc- tiveness of E. traversii yet acknowledge the close relationship between E. strigosus and E. traversii, the following combination is proposed: Erigeron strigosus Muhl. ex Willd. var. traversii Soda roe soul & stat nov. Erigeron traversii Shinners, Sida 1:376. 1964. TyPE: U.S.A. T Cc US 59, ca. 8 mi S of Nacogdoches, 1] Apr 1942, CL. Lundelland A.AL ae eee BRIT). The holotype of E. strigosus var. traversii shows strong morphological similar- ity tosexual diploid plants from South Carolina, Georgia, Florida,and Alabama 274 BRIT.ORG/SIDA 22(1) (Noyes & Allison 2005). In particular, these plants are distinguished by nar- rowly spathulate rosette and lower stem leaves that bear small, alternate, mu- cronate lobes along the petiole margin. Thus it is possible that E. strigosus var. traversii extends continuously from east Texas to South Carolina on the coastal plain and piedmont. Molecular phylogenetic analysis indicates that sexual dip- loid southeastern populations of Erigeron strigosus are distinct from sexual dip- loid edaphic endemic populations of E. strigosus var. calcicola and E. strigosus var. dolomiticola (Noyes 2006a). On the other hand, ongoing geographic map- ping for E. strigosus indicate that sexual diploid populations occur in western Louisiana, where they are contiguous with those in East Texas, and also in scat- tered localities in the Ozarks of western Arkansas and adjacent Oklahoma (Noyes 2006b). Detailed biosystematic analyses will be required to determine if these geographically disparate sexual populations constitute a single taxon, or several distinct taxa. The precise leaf morphology observed in sexual plants of Erigeron strigosus has not been observed for apomictic polyploid E. strigosus, which may entirely lack petiole lobes, or exhibit larger, more irregular lobing. Variability in apomic- tic E. strigosus is complex, extending from E. strigosus var. septentrionalis (Fernald & Wiegand) Fernald, which includes plants approaching E. annuus (L.) Pers., to apomictic Erigeron strigosus var. strigosus, which can be very simi- lar to E. strigosus var. traversii. In addition, the closely related species E. tenuis Torr. & A. Gray, also occurs in Texas and adjacent states. The likelihood of hy- bridization and morphological similarity between early flowering E. strigosus (likely corresponding to E. strigosus var. traversti) and E. tenuis was discussed by Van Vleet (1951). Further, based on consideration of published chromosome counts, F. tenuis also likely includes sexual and apomictic plants, but the geo- graphic distribution of mode of reproduction in the species is unknown. Thus, elucidation of evolutionary relationships for Erigeron strigosus must also in- clude detailed study of E. tenuis. The recognition of E. strigosus var. traversii brings to three the number of recognized sexual taxa related to the widespread apomictic taxa EF. strigosus, E. annuus, and E. tenuis and contributes to our understanding of the evolution and dynamics within the species complex. The following key is provided to aid in distinguishing Erigeron strigosus var. traversii from its close relatives in Texas: 1. Pappus of ray and disc florets of numerous conspicuous capillary pappus bristles; the species including both sexual and apomictic plants and in need of taxonomic study E. tenuis . Pappus of ray florets a low crown-like series of scales, lacking Conspicuous capillary pappus bristles, disc florets with capillary bristles 2. Plants relatively tall (to 1.5 m) with ace heal pliable leaves; dense, long, flat- tened trichomes present along the main stem and on the involucre; apomictic and usually triploid; relatively uncommon and restricted to eastern Texas E.annuus NOYES ET AL., ERIGERON STRIGOSUS IN TEXAS 275 2. Plants relatively shorter (to 0.7 m) with narrower, stiffer | flatt | trichomes if present, restricted to the base of the plant or relatively sparse or absent; ere or apomic tic. 3, Plants generally lacking flattened trichomes throughout; plants sexual, dip- loid; pollen of high quality, relatively uniform in size, grains < 15 um diam; restricted to Pineywoods habitat in East Texas and adjacent Louisiana ___ E. strigosus var. traversii 3, Plant stems and involucres sometimes possessing spreading flattened tri- chomes; plants polyploid, apomictic, highly variable in morphology; pollen of poor quality, highly variable in size, largest viable grains > 17 um diam; rela- tively widesprea 4. Hairs cae flattened, mostly 0.5—1.2 mm; hairs of stems 0.5-1 mm appressed to spreadin E. stri cae var. rseptentrionali 4, Hairs of involucres terete, mostly 0.1—-0.5 mm;hairs of stems mostly mm, appressed to ascending E. aa var. strigosus ACKNOWLEDGMENTS We thank the curators at BRIT and TEX for specimen loans, staff at COLO for providing assistance managing specimens, and Kiyoshi Yamashita, Christof Bigler, and Jason Knouft, of the University of Colorado, for help implementing GIS analyses. Guy Nesom (BRIT) and Tim Lowrey (UNM) provided valuable comments in review. REFERENCES Asker, S.E and L. Jertina. 1992. Apomixis in plants. CRC Press, Boca Raton, FL. Baker, H.G. 1965. Characteristics and modes of origin of weeds. In: G.L. Stebbins, ed. The genetics of colonizing species. Academic Press, New York. Pp. 147-172. Bark.ey, T.M.(ed.). 1986. Flora of the Great Plains. University Press of Kansas, Lawrence, Kansas. Beyer, H.L. 2004. Hawth’s analysis tools for ArcGIS (http://ww tialecology.com/htools). BierzycHubek, P. 1987. Patterns in plant parthenogenesis. In: S.C. Stearns, ed. The evolution of sex and its consequences. Birkhauser Verlag, Basel, Boston. Pp. 197-217 Corrett, D.S. and M.C. JoHNsTon. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner, TX. Cronauist, A. 1947. Revision of the North American species of Erigeron, north of Mexico. Brittonia 6:121-302. Den Nus, J.C.M.and S.B.J.Menken. 1994. Breeding systems and evolution in Taraxacum. Evol. Trends PI. 8:11-20. Grant, V. 1982. Plant speciation, 2nd. Columbia University Press, New York. Gustarsson, A. 1946-1947. Apomixis in higher plants. Lunds Univ. Arsskr.42—43:1-370. Haac, C.R. and D. Esert. 2004. A new hypothesis to explain geographic parthenogenesis. Ann. Zool. Fennici.41:539-544. Hatcu, S.L., KN. GANDHI, and L.E. Brown. 2001. A checklist of the vascular plants of Texas, version 2. (http://www.csdl.tamu.edu/FLORA/taes/tracy). Texas A&M University, Col- lege Station. 276 BRIT.ORG/SIDA 22 = 1) Hickman, J.C. (ed.). 1993. The Jepson manual: higher plants of California. University of Cali- fornia Press, Berkeley. Hoimeren, |. 1919. Zytologische Studien uber die Fortpflzung bei den Gattung Erigeron und Eupatorium Kong. Svenska Vetenskapsakad. Handl. 59:1-118. Joxnston, M.C.1990.The vascular plants of Texas: a list up-dating the manual of the vascu- lar plants of Texas, 2nd.M.C. Johnston, Austin, TX. Kartesz,J.1.1992.A synonymized checklist of the vascular flora of the United States, Canada, and Greenland, 2nd. Timber Press, Portland, OR. LYNCH, ue 984. Destabilizing Pee general-purpose genotypes and geographic is. Quart. Rev. Biol. 59:257-290. re Smith, J. 1978. The evolution of ae University Press, Cambridge. Menken, $.B.J.,E.Smit,and J.C.M. Den Nus.1995.Genetic population structure in plants:gene flow between diploid sexual and triploid asexual dandelions (Taraxacum section Ruderalia). Evolution 49:1108-1118. Mosie,M. 1992.The evolution of asexual reproduction in plants. Chapman and Hall, London. Nesom, G.L. 1978.Chromosome numbers in species of Erigeron and Conyza (Compositae). Sida 7:375-381. Noyes, R.D.2006a. Intraspecific nuclear ribosomal DNA divergence and reticulation in sexual diploid Erigeron strigosus (Asteraceae). Amer. J. Bot. 93:470-479. Noyes, R.D.2006b. Reticulation and the evolution of apomixis in Erigeron sect.Phaiacroloma (Asteraceae) In: E. Hérandl, U. Grossniklaus, P Van Dijk, and T. Sharbel, eds. Apomixis: evolution, mechanisms and perspectives. Regnum Veg. 147. (Gantner Verlag, Ruggell, Liechtenstein) (Accepted for publication) Noyes, R.D.and J.R. ALuson.2005.Cytology, ovule development and pollen quality in sexual Erigeron strigosus (Asteraceae). Int. J. Pl. Sci. 166:149-159. SHinNeRS, L.H. 1964. New names and records for Texas Compositae. Sida 1:373-379. Soi Suavey Starr. 1999. Soil taxonomy:a basic system of soil classification for making and interpreting soil surveys. USDA Natural Resource Conservation Service Agriculture Handbook # 436, U.S. Government Printing Office, Washington DC. STANLEY, R.G.and H.F. Linskens. 1974. Viability tests. In: Pollen: biology, biochemistry,and man- agement. Springer, Berlin. Pp.67-86. Steesins. G.L. 1950. Variation and evolution in plants. Columbia University Press, New York. USDA, NRCS. 2005. The PLANTS Database, version 3.5 (http://plants.usda.gov). Data com- piled from various sources by Mark W. Skinner. National Plant Data Center, Baton Rouge, LA. Van VieeT,R.L.1951.Phenotypic variations of Erigeron strigosus Muhl.(Compositae) in east- ern Texas. Field & Lab. 19:161-162. VerRDUUN, M.H., PJ.vAN Duk, and J.M.M.van Damme. 2004. Distribution, phenology and demog- raphy of sympatric sexual and asexual dandelions (Taraxacum officinale s..): geographic parthenogenesis on a small scale. Biol. J. Linn. Soc. 82:205-218. MORE AMERICAN BLACK SAPOTES: NEW DIOSPYROS (EBENACEAE) FOR MEXICO AND CENTRAL AMERICA Mitchell C. Provance Andrew C. Sanders Depainens orp raya! and ea : ences UCR Herbarium College of Natural ciences Department ee and Plant Sciences Boies ity of Cal ifornia na University of California, Riverside Riverside, ee ig 92571-0124, U.S.A Riverside, ie 92521-0124, U.S.A. chtell@prodigy.net andrew.sanders@ucr.edu ABSTRACT During the preparation of a monograph of the Ebenaceae for Mexico, numerous llections previ- ously determined to be Diospyros conzattii Standley and Diospyros riojae Gomez-Pompa could not be reconciled with their original descriptions, type specimens and illustrations. It was determined that collections of putative D. conzattii from Costa Rica represent a new species, D. costaricensis, described here. All but one collection of putative D. riojae from the Mexican states of Queretaro, Hidalgo, Tamaulipas, and San Luis Potosi, represent a new species, D. gomeziorum, described here. Putative collections of D. fies from Veracruz additionally include specimens of D. conzattii, a spe- cies previously unreported for the state, and a new species from the Sierra de Los Tuxtlas, D. tuxtlensis described here. We provide emended descriptions of D. riojae and D. conzattii, and describe three id distribution, ecology new species of Diospyros for Tropical America. We also conservation, ethnobotany, and provide illustrations for peach of these taxa. Keyworps: black sapote, Costa Rica, Diospyros gomeziorum, D. conzattii, D. costaricensts, pergamentacea, D. riojae, ee rosei Complex, D. tuxtlensis, Ebenaceae, granadilla, ae Mexico, new species, persimmon, za gro silvestre, zapotillo RESUMEN pieen - Bispace de la ee aaa de ehaeaues para México, numerosas colecciones tii Standley y Diospyros I iojae Gome Pompa no pudieon ser reconciliadas con las Ree eons originales, el espécimen tipo y las ilustraciones. Se determino que la coleccion putativa D. conzattii de Costa Rica, representa una especie nueva, D. costaricensis descrita aqui. Todas, excepto una de las eolecone putativas de D. riojae de los estados mexicanos de Querétaro, Hidalgo, Tamaulipas y San Luis Poto n una especie nueva de D. gomeziorum, descrita aqui. Adicionalmente, las eee mietie de D. riojae de Veracruz, ie especimenes de D. conzattii, una especie no citada previamente para ese estado y de una especie nueva de la Sierra de los Tuxtlas D. tuxtlensis, descrita aqui. Proveemos descripciones corregidas de D. riojae y D. conzattii y describimos tres especies nuevas de Diospyros para América tropical. También hacemos comentarios sobre la distribucion, ecologia, conservacion, etnobotanica y aportamos ilustraciones de todos los taxa. PaLaBras CLavE: Complejo de Diospyros rosei, Costa Rica, sate panes D. conzattii, D. costaricensis, D. pergamentacea, D. riojae, D. tuxtlensis, Ebenaceae, especie nueva, granadilla, guacalillo, México, persimo, zapote negro, zapote negro silvestre 6 SIDA 22(1): 277 - 304. 2006 278 BRIT.ORG/SIDA 22(1) INTRODUCTION The pantropical genus Diospyros (Ebenaceae) consists of about 500 species of trees, shrubs, and suffrutices (White 1983). A treatment of the Mexican species of Diospyros in full has not appeared since Standley (1924). Modern treatments have dealt with smaller areas, such as Veracruz (Pacheco 1981), northeastern Michoacan, Guanajuato, and Queretaro (Carranza 2000), the Yucatan Penin- sula (Lundell 1942), and the Tehuacan-Cuicatlan Valley, Oaxaca (Kelly 2001), or covered portions of southern Mexico within a larger flora (Whitefoord & Knapp 2001). Circumscription of the Mexican and Central America taxa is not entirely agreed upon. However, based on our preliminary work, a reasonable estimate of the number of native species occurring in Mexico is around 25. In 1918, C. Conzatti collected a new black zapote from the coastal moun- tains northeast of San Pedro Pochutla, Oaxaca. He recorded some characteris- tics of the tree, especially those pertaining to the edible fruit, the taste of which he enjoyed. He believed that the fruit of the ‘zapote negro silvestre’ (Conzatti 3167) had potential as a new tree crop (Standley 1922). This zapote was formally described a few years later as Diospyros conzattii Standl. (Standley 1922). Ac- cording to Trabut (1926), D. conzattii was introduced to California and Florida. However, he did not provide any specific details, and we are unaware of any other information that might corroborate the putative introduction of this spe- cies to the United States. Diospyros riojae Gomez-Pompa was described from one fruiting collection made in the Misantla region of central Veracruz, Mexico (Gomez-Pompa 1964). Since then, collections of putative D. riojae have been made in several more lo- cations in Veracruz, Iamaulipas, San Luis Potosi, Hidalgo, and numerous places in northeastern Queretaro. Diospyros riojae is considered an endangered spe- cies (Sanchez-Coello 2002 During the preparation of a monograph of the Ebenaceae for Mexico, we noticed a large amount of morphological variation among collections of puta- tive D. riojae and D.conzattii. While we had intended to examine the taxonomy of both species at a later time, in view of the gered status of D. riojae,and asense of some urgency regarding its protection (e.g., Sosa et al. 1998), we thought it would be prudent to proceed with the treatment of the following taxa now, lest the designation of protected areas, or expensive and time-consuming eco- logical research, be confounded by the unrecognized presence of similar look- ing taxa. About eighty-five specimens from seventeen herbaria were examined for this study. With the exception of one collection from San Luis Potosi, specimens of putative D. riojae collected outside of Veracruz represent a new species, D. gomeziorum, described here. Much of the material collected in Veracruz and attributed to D. riojae has been found to represent D. conzattii, and a species PROVANCE AND SANDERS, NEW SPECIES OF DIOSF AMERICA 279 new to science, D. tuxtlensis, described here. Costa Rican material, previously ascribed to D. conzattii, clearly represents a third new species, D. costaricensis, which is described here. These species are best considered members of what we have informally recognized as the ‘Diospyros rosei Complex’ (Provance & Sanders 2005), a group that additionally includes: Diospyros rosei StandL., Diospyrosoaxacana StandL., Diospyros palmeri Eastwood, Diospyros californica LM. Johnston, Diospyros sonorae Standl., Diospyros texana Scheele, Diospyros riojae Gomez-Pompa, Diospyros conzattii Standl., Diospyros rekoi Standl., Diospyros torresii M.C. Provance & A.C. Sanders, Diospyros morenoi A. Pool and Diospyros xolocotzii Madrigal & Rzedowski. Previously (Provance & Sanders 2005), we called into question the distinctness of D. morenoi. At that time, we only had the holotype to examine. Very recent collections from Chiapas, Mexico, have inclined us to reevaluate our position on the status of this species. We now believe that it is a valid taxon, and we plan to address this issue further in the near future. In this treatment we use ‘lanceolate’ in the sense of Jackson (1916), being broadest near the lower third of the leaf, not at mid-leaf in the sense of Stearn (2000). ‘Scintillant’ refers to the sparkling appearance of the epidermis of some plant structures when viewed under a dissecting microscope using bright light. Lengths for winged petioles are unavoidably approximate. Female inflores- cences are inferred from fruiting specimens and persistent pedicels, where they are otherwise unknown for these taxa. The leaf venation for most taxa is ‘arcolanguid, defined here as an intermediate state between eucamptodromous and brochidodromous, in which major lateral veins nearly form distinct loops with superadjacent lateral veins, but tardily wane, becoming difficult to dis- tinguish as loops. Tne term is coblas for Simtel ley and out of necessity, since the term ‘ lod ’ of Todzia and Keating (1991) refers toa different type of pone Geographical coordinates, elevations, and similar estimations when made by us are presented in brackets. TAXONOMIC TREATMENT 1. Diospyros riojae Gomez-Pompa, J. Arnold Arbor. 45:465. 1964. (Fig. 1). Type: MEXICO. VERACRUZ: between Chiconquiaco and Misantla, in Liquidambar-Quercus forest, with Magnolia, Meliosma, Juglans, and Tu ae nia, 1350 m, 13 Aug 1962 (fr), A.Gémez-Pompa 789 (HOLOTYPE: A; ISOTYPES: MEXU, US on fiche from UCSBI). Original material examined: (TYP. 1C.: A.Géomez-Pompa 7891, J. Arnold hee 45:467. 1964.). Trees, rarely shrubs, probably facultatively deciduous, to 25m tall,and to 2.5m in diameter. (Pacheco 1981); trunk channeled, rimose, bark dark-brown (Pacheco 1981); aged stems subterete to terete, bark shallow-fissured with dark ridges, gray and black, stemwood pale yellow; 2nd-3rd year stems short-fissured, gla- brous to furfuraceous pulver ulent, punctic ulate, lenticellate, half-netted, gray above, tawny below; current year’s stems angular, smooth to sulcate, glabrous BRIT.ORG/SIDA 22(1) Fic. 1. A-G. Diospyros riojae Gomez-Pompa. A. Abaxial surface (left) and adaxial surf ight) surface of staminat petal lobe. B. Adaxi Mt ft) and | ralifig) ) f staminate infl bract. C Tpicaltanitier D. Staminate flower. E. Abaxial f f sh il of F. Fruiting stem. G. Fruit. A—D. Based on F Ventura A. 15078. E-F. Based on J. RecorraiZ: 83.G. Based ane Pacheco & J.I. Calzada 18 PROVANCE AND SANDERS, NEW SPECIES OF DIOSPYROS FROM MEXICO AND CENTRAL AMERICA 281 to sparingly subappressed puberulent, hirtellous at the base and near the apex, tan to reddish-brown, glandular. Petioles stout, thicker than width, (2-)4-6(-7) mm long, winged, rugose below, slightly pruinose, deeply concave to V-grooved above, ciliolate, glandular. Leaves alternate, simple, entire, subcoriaceous to co- riaceous, (5-)7-13(-14.2) cm long, (2.5-)3-6(-7.5) cm wide, elliptic, widely el- liptic, oblong, widely obovate, sub-oval, apex acutely to obtusely rounded, or short-acuminate to a rounded point, margin slightly recurved, + thickened, sometimes ciliolate, base obtusely rounded, cuneate, or short-acuminate; lower leaf surface glabrous, puncticulate, sometimes slightly pruinose or scintillant, shiny, olive to bronze to brown; upper leaf surface glabrous, shiny, pale light- green to gray-green or copper-green, epidermal cells enlarged, the anticlinal walls thickened. Venation arcolanguid; midrib prominent below, chartreuse to reddish-brown, shallowly concave above, smooth to clavate glandular hairy, shiny, olive to yellow-green; 2°-3° veins reticulated, raised below, fine with sharp relief, sometimes pruinose-scintillant, granular-papillose, shiny, usually promi- nent above, lateral veins 8-12 pairs. Laminar extrafloral nectaries abaxial, near base and midrib, round to narrowly elliptic, sometimes on lateral veins. Male inflorescence fascicle, 1-4 flowers from leaf axils of the previous year’s growth, or from the base of new stems, sometimes a pair of solitary flowers from the new stem just above the fascicle; pedicels +-7(-10) mm long, sparingly brown to claret velutinous, epidermis green; pedicel bracteoles 2, | mm long, oblong, + flat to slightly concave, ascending wavy hairy below, the hairs thick, glisten- ing. Male flowers 5-merous; calyx infundibuliform; calyx tube 1.5 mm long, gla- brous or scantly minute subulate hairy; sepals triangular to ovate, 1.3-1.8 mm long, 0.7-0.9 mm wide, ciliolate, scantly minute appressed subulate hairy ba- sally, becoming dark glandular wavy hairy apically; corolla urceolate-campanu- late, tan to brown, reportedly white in living material; corolla tube constricted below the lobes, 4 mm long, 3 mm wide, interior grading from glabrous dis- tally to minutely deltoid scaly at mid-tube, again becoming glabrous near the base, except near filament bases, exterior puberulent with hairs erratically di- rected, some glandular; corolla lobes oblong-squarish, emarginate, 2 mm long, 2mm wide, interior pubescent, exterior minutely sericeous-puberulent, exmedially appressed, several black hairs present near the apex. Stamens 18, subexserted, adnate to corolla at mid-tube or lower, some inserted on the recep- tacle, some lower stamens extrorse, abruptly geniculate, and connected to introrse upper stamens by a superdermal vascular trace; filaments | mm long, hirtellous, especially on margin; anthers basifixed, lance-ovate, + 2.5 mm long, laterally concave, apically constricted, opening by short apical slits; pistillode + wheel- shaped, lobes 5, bifurcate, glabrous, dark gray. Female inflorescence flowers soli- tary, from caducous bract scar axils near the base of young stems. Fruiting pedicels 7-12mm long, stout, very minutely hirtellous. Fruiting calyx 5-lobed; fruiting calyx tube 3.5-4 mm long, explanate to ref lexed; fruiting sepals accrescent, coriaceous, 282 BRIT.ORG/SIDA 22 — ) arcuate-reflexed to spreading, 7-25 mm long, 7-13 mm wide, oblong, narrowly triangular or ovate, apex usually ascending to incurved, sometimes glaucous- pruinose to scintillant. Fruit berry, 3-5.5cm long, up to 4. 7cm mace, SHE Opes, locules8-10; mesocarp [leshy, yellow-orange in dried and I epidermis smooth to bullate, shiny in living s material, oreen to brown or black, herbarium material reddish-brown to dark-brown, immature fruits sometimes atropurpureous. Seeds not seen in available material (seeds orangish in photograph in Sanchez-Coello 2002 from Conde 2000), described by Pacheco (1981) as brownish to grayish, shiny, very rugose, and with a prominent raphe. Additional specimens examined. MEXICO. oa ae Potosi. Mpio. Tamazunchale: Tamazunchale, [21° 16'N, 98° 47'W, 153 ml, 24 Jul 1937, M.T.E ), US). Veracruz. Mpio. Alto Lucero: Cruz Blanca, 19° 46'N, 96° 40'W, 900 m, 30 Apr 1980, L. Pacheco & J.1. Calzada 23 & 24(XAL); same locality 19 Sep 1979, L. Pacheco & J.I. Calzada 18 (X AL); Cruz Blanca, 19° 40'N, 96° 50'W, [900 ml], 25 May 1998, Tono Vazquez 98-045 (UCR). Mpio. Chiconquiaco: Canada del Huerfano, 19° 49'N, 96° 48'W, [900- 1200 ml, 10 Jul 1966, A. Gomez- ie 1602 (XAL). Mpio. Martinez de la Torre: Malaupan [Santa Ana Maloapan, 20° 04'N, 97° 04'W], LOO m, 15 Mar 1978, F. Ve He 15078 (EB, XAL). Mpio. Tantima: Sierra de Tantima, 21° 17'N, 97° 51 7: 740 m, 23 Aug 1979, .1. Calzada 5550 (XAL). ae Tepetzintla: Sierra de la Pena Blanca, Sierra de Tantima, [21° 13'N, 97° 55'W, = 700 ml, 21 Sep 1989, P Zamora C. et al. aes ), San José de Copaltitla al NE de Tepetzintla, 21° 12'N, 97° 52'W, 350 m, 28 Aug 1981, G. Castillo C.& A. Benavides M. 2265(XAL ; de Mixtepec, Sierra de Tantima, 21° 18'N, 97° 50'W, LOOO m, ; va 1988, J. Becerra Z.83(XA Distribution and Ecology.—The a known collection of this taxon made out- side of the state of Veracruz is from Tamazunchale, San Luis Potosi. This 1937 collection (M.T. Edwards 670) is the earliest one of the species known to us. Tamazunchale is at 153 m, a low elevation based on the literature (e.g. Pacheco 1981). In northern Veracruz, this species occurs in the Sierra de Tantima. In cen- tral Veracruz, occurrences are in the Sierra de Chiconquiaco, the lowlands of its northern base, and the Misantla region. Populations in northern Veracruz are in isolated cloud forest patches. In central Veracruz, collection sites have been characterized as forested escarpments near pasture (Sosa et al. 1998), cloud for- est, and deciduous forest (Pacheco 1981) at 750-1000 m. It seems not to be widely realized that this taxon also occurs at lower elevations at 100-300 m. Low el- evation occurrences are associated with tropical evergreen forest with Quercus. In this community, they are known to flower during March. This species is con- sidered to be in danger of extinction (Sanchez-Coello 2002). Ethnobotany.—The tree is called ‘granadilla’ in the Sierra de Tantima re- gion of northern Veracruz where it is considered edible (G. Castillo C. & A. Benavides M. 2265). The name '‘sapote prieto’, is indicated on the collection from Tamazunchale, San Luis Potosi. 2. Diospyros gomeziorum M.C. Provance & A.C. Sanders, sp. nov. (Fig. 2e). Typr: MEXICO. QUERETARO: Mpio. Jalpan de Serra, 5-6 km al NW de San Juan de ee Duran, El Ar- royo, 21.48°N, 99.12°W, escaso, bosque de pino-encino, cedro blanco, orilla de arroyo, canada 1500-1600 m, 1 Aug 1991 (fr), Benito Servin 1274 (HOLOTYPE: IEB-150586). PROVANCE AND SANDERS, NEW SPECIES OF DIOSPYROS FROM MEXICO AND CENTRAL AMERICA Fic. 2.A—-E. Di ji M.C. Provance & A.C. Sanders, sp.nov. A.5 fl B. Typical anther. C. Adaxial ¢ L surface of staminate petal lobe. D. Lateral v E Fruiting pene A-C. Based on R. Ferndndez N. 2829. D. Based on Hiram Rubio 1520. E. Based on Benito Servin 1274. 284 BRIT.ORG/SIDA 22(1) Arbor [rutexve usque ad 20 m altus, D. riojae Gomez-Pompa similis sed differt margine foliia pagina je adaxiali usque ad paginam abaxialem exter obliquantibus; cellulis epidermalibus paginae laminae = ed oe 16 tie cl 7 ; ; i} Vaal peed il 1 Ee Be . LS [ ] : : ] ie sf; igs sai fb ~ ] | ot \; 14 [ C rT i t t 7 frouctiteris 7 ] Trees or shrubs to 20 m tall, facultatively deciduous; trunk reported to 12 cm diameter for a 5-6 m tree (Servin 1274) and 20 cm in diameter for a 4+ m tree (R. Fernandez N. 2459), erect, longitudinally fissured, bark gray, reportedly shiny; aged stems angular, bark verrucose, scarred, short-fissured, cinereous, stem wood yellow to yellow-orange; 2nd-3rd year stems angular to subterete, sometimes hirtellous, + half-netted, the outer epidermal layer gray, tawny or reddish- brown, the lower layer cream to orangish; lenticels protruding, sometimes 2-3 mm long; current year’s stems subterete, smooth to striate, densely hirtellous, reddish-brown, puncticulate. Petioles stout, often thicker than wide, 2-5(-8) mm long, glabrous to minutely erect hairy to deltoid scaly, rounded to rumi- nate below, usually glistening, dark brown to olive green, narrowly channeled, flat or barely raised, furfuraceous-glandular, ascending ciliolate on margin, yel- low-green to olive above. Leaves alternate, simple, entire, pergamentaceous to chartaceous, sometimes subcoriaceous, lanceolate to ovate to elliptic, 6-11(-1L5) cm long, 2-4.8 cm wide, apex acute, obtuse or acuminate to a rounded point, margin beveled outward from upper surface at + 45° to the lower surface, some- what sharp, shiny, sparsely to moderately ciliolate, base acute to obtuse, decur- rent onto petiole; lower leaf surface glabrous to sparsely subappressed pubes- cent, hairs black or reddish, most common near the base, midrib, and apex, surface shiny, green to olive, puncticulate; upper leaf surface glabrous, glau- cous to pruinose-scintillant, localized or covering most of surface, but not on the beveled margin, granular-papillose, olive or pale blue-olive to dark-olive or dark blue-green, usually darker above than below, young leaves tend to be lighter above. Venation arcolanguid, granular-papillose, glaucous to pruinose- scintillant; midrib prominent below, glabrous to subappressed pubescent, hairs — — dlack, reddish-brown, or white, narrowly canaliculate above, sparsely to densely hirtellous to deltoid scaly, rarely glabrous, yellow-green; 2° venation subprominent below, lateral veins (8-)11-13(-14) pairs; 3° venation reticulate, obscure to prominent below, usually prominent above. Laminar extrafloral nec- taries abaxial, minute, circular. Male inflorescence fascicles of 1-3(-5) flowers in leaf scar axils of previous year’s growth, or one to several flowers at the base of new growth in scar axils of caducous bracts; pedicels 1-3 mm long, densely straight hairy, the hairs reddish, black, and white; pedicel bracts 2, 2-2.5 mm long, + linear. Male flowers (4-)5(-6)-merous; calyx infundibuliform-campanu- late, deeply lobed, interior glabrate or scantily minute white hairy, exterior sparsely minute deltoid scaly; calyx tube |.5-2 mm long, 2.5-3 mm wide, sparsely subappressed puberulent; sepals lance-ovate to triangular, 2-7 mm long, l-4 PROVANCE AND SANDERS, NEW SPECIES OF DIOSPYROS FROM MEXICO AND CENTRAL AMERICA 285 mm wide, exterior glabrous to pubescent basally, subappressed pubescent apically, the hairs glossy, black to reddish, simple and glandular, epidermis vis- cous; corolla urceolate-campanulate, white in living material; corolla tube 3- 5.5 mm long, 3-6 mm wide, exterior densely pubescent, most hairs distally ori- ented, some hairs irregular, interior sparingly hirtellous, hairs sometimes erratically directed; corolla lobes (4-)5, spreading, oblate to squarish or widely obovate, often emarginate, 2-4 mm long, 2-3.5 mm wide, several short black hairs near the apex, interior sericeous. Stamens 20, equal to, or barely surpass- ing tube, most adnate to corolla, some inserted on the receptacle; filaments 0.3- 0.8 mm long, ascending puberulent; anthers basifixed, oblong-ovate to ovate, 2 mm long, opening by short slits near apex, ay by cone: lateral ne granuliferous, apex acute or rounded an wheel-shaped, short, wide, corrugate. Female irifloreseenee flowers eee cauline near the base of current season's growth, rarely from leaf scar axil of old growth (e.g. Ferguson 15), sometimes two opposite inflorescences, but two mature opposite fruit not seen. Female flowers not seen, but styles (4-)5 as rem- nants on mature fruit, appressed white puberulent. Fruiting pedicels (4-)5-8(- 11) mm long, densely hirtellous. Fruiting calyx 5-6-lobed. Fruiting calyx tube 3- 4 mm long; fruiting sepals accrescent, + spreading, 19-20 mm long, 8-11 mm wide, lanceolate to narrowly elliptic or obovate, glabrous to sparsely appressed hairy, apices rounded, sometimes ascending, surfaces often bluish-white glau- cous, non-glaucous areas tawny, puncticulate. Fruit berry, subglobose to slightly obovoid, 3cm long, 3-4.5 cm in diameter, green when immature, turning black (and then presumably ripe), atropurpureous in dried specimens, locules 10; mesocarp orange when dry; hypodermis thin, not very stony, often broadly rippled in specimens; epidermis orange-peel-textured, scintillant, glaucous- pruinose. Seeds 16-19 mm ee 9-10 mm wide, rugose, shiny, light brown. Paratypes: MEXICO. Hidalgo. Mpi B le Meztitlan, 9 km al NE de Mesquititlan a lo largo de una canada, 20.74°N, 98. 98° W, 2200 m, 3 May 1975, FG. Medrano et al. 7932 (SD, US, XAL). Queretaro. Mpio Arroyo Seco: 2 km al W de El Jardin, 21° 25' 24"N, 99° 41' 42"W, 1380 m, 4 Jul 1989, F. Carranza 1852 (1EB). Mpio. Jalpan de Serra: 2-3 km al Poniente de San Isidrio, La Parada, 21° 30.5'N, 99° 10'W, 1300 m, 14 May 1990, Benito Servin 206 (CIIDIR, IEB); 2-3 km al S de La Parada, 21° 32.3'N, 99° 10'W, 1200 m, 26 Mar 1990, Benito Servin 58 IEB); 6-7 km al Oriente de La Parada, 21° 30.5'N, 99° 6.8'W, 1400 m, 5 Apr 1990, Benito Servin 96 (EB); 2-3 km al N de La Parada, 21° 32.3'N, 99° 10'W, 1100 m, 13 Aug 1990, Benito Servin 412 (EB); + 2 km al W de La Parada, Cuesta de los Lirios, 21° 30.5'N, 99° 11.2'W, 1180-1250 m, 13 Mar 1990, E. Carranza 2392 (C oie IEB). Mpio. Colon: cerca vado rio Colon, 20° 48'N, 100° 03'W, oe m, 2 Apr 1982, Elizabeth Arquti 1761 ae ies Pinal de Amoles: | km S de Escanelilla, sobre la carretera a Pinal de ee 21° ON, e 33'W, 1250 m, 18 May 1987, J. Rzedowshi 43398 (HUAA, IEB); + 4 km al SE de Santa Agueda, 21° 14' 30" N. 99° 37'54"W, 1190 m, = Apr 1989, E. Carranza 1635 (IEB); 3 km al S de Escanelilla, 21° 10.2'N, a 34'W, 1100 m, 19 Mar 1985, Ferndndez N. 2829 (US); same location, 30 Jul 1984, R. Fernandez N. 2459 (LEB). Mpio. Landa Matamoros: |.5 km al SW de El Naranjo, 21° Ol 18"N, 99° 27' 42"W, 900 m, 10 Mar 1990, Hiram a 1542 (IEB); 1 km al Poniente de El Cerro de La Palma, 21° 12' 28"N, 99° 04' 32"W, 1500 m, 2 Mar 1990, Hiram Rubio 1520 (IEB); cerca de Tres Lagunas, 21° 19' 36"N, 99° 12' 12"W, 1700 m, 22 Jun 1988, J. 286 BRIT.ORG/SIDA 22(1 Rzedowski 46695 (IEB); 2 km al Norte de Neblinas, 21° 13' 58"N, 99° 06'18"W, 1060 m, 29 Sep 1988, Hiram Rubio 185 (EB); El Humo, 2 km a TOuest d’El Humo, 21° 18'N, 99° 05'W, 1320 m, | Oct 1994, J.N. Labat & FE. Carranza 2562 (EB); 2 km al SE de El Humo, 21° 13'55"N, 99° 06'15"W, 1100 m, 7 Jun 1989, Hiram Rubio 775 (HUAA, IEB); 2.5 km NE de El Humo, [21° 18'N, 99° 05'W], 1000 m, 11 Aug 1990, Hiram Rubio 1865 (EB); 1.5 km SE de El Naranjo, 21° Ol' 18"N, 99° 27' 42"W, 1000 m, 25 Jun 1990, Hiram Rubio 1746 (EB), 4-5 km al S de Tres Lagunas, 21° 19’ 33"N, 99° 12'12"W, 1820 m, 3 Mar 1990, E. Carranzd 2359 (HUAA, IEB), El Calvario, | km al Poniente de El Aguacate, 21° 16' 06", 99° 14' 42", 1620 m, 26 Jun 1989, Hiram ae 838 (EB); + 8 km de desviacion, camino Tres Lagunas, 21° 19' 35"N, 99° 06'15"W, 1900 m, 7 Nov 1988, E. Carranza 1144 (EB); 2 km al SE de Neblinas, 21° 15'N, 99° 03'12"W, 900 m, 23 Jun 1989, Hiram Rubio 826 (EB). San Luis Potosi. Mpio. Rayon: 3.5 mi by road Sof km 81.5 onhwy 70 Wol Valles at microwave tower,[+ 21° 53'N, 99° 30'W], 1340 m, 23 May 1981, George Fergu- son 15 (UTEP). eae Mpio. Gomez Farias: Rancho del Cielo, a 11 km de Gomez Farias, [23° O4'N, 99° 12'W], 1110 m, 22 Sep 1974, EG. Medrano 7416(ARIZ); Rancho del Cielo, La Sierra de Gomez Farias, 1100 m, 1 May 1982, H. Narave F. & T. Dent 134 (XAL); Rancho El Cielo, F CATR E ridge of the Sierra Cucharas, 1200 m, 23 Aug 1950, E. Hernandez X.& F. Harrison X-586] (CHAPA Distribution and Ecology.—This specie urs in the Sierra Madre Oriental, in- cluding the Sierra Cucharas, et north of Gomez Farias in southwest Tamaulipas, t thwest of Tamasopo in southeast San Luis Potosi, northeast Queretaro and Barranca de Meztitlan in eastern Hidalgo. Col- lections are from a variety of vegetation types at elevations between 900 and 2200 m. More material, by far, has been collected in the state of Queretaro than any other state. In Queretaro, it has been come as abundant in pine-oak forest, oak forest, oak forest with Liquid and Tilia forests. Other settings in which it has been reported but deserved. as scarce include gallery forests with Platanus, and canyons with bosque mesofilo de montana (+ cloud forest). An unusual association occurs in Hidalgo, where it was described as very abun- dant in a canyon with matorral espinoso and matorral crasicaule at 2200 m (EG. Medrano et al. 7932). The five male flowering collections we examined were all taken in March. Some of these specimens were just leafing out at the time. The fruits ripen between September and early November. Ethnobotany.—Queretaro: ‘zapote prieto’ (Rubio 1746), ‘zapote del monte’ (Servin 206), zapotillo (Carranza 2000). Etymology.—The epithet is intended to honor both Arturo Gomez-Pompa, who discovered and described Diospyros riojae, and has also been a great inspi- ration to our work on Latin American Diospyros, and also the late J. ‘Carmelo’ Gomez, who assisted the first author in the field on many occasions, and was very knowledgeable on local plant use in the Sierra Tlachichila, Zacatecas. Collections of D. gomeziorum have often been confused with D. riojae. A conspicuous feature that differentiates D.gomeziorum from D. riojae is the leaf margin, which is beveled outward from the upper to the lower surface at about 45° in D.gomeziorum. The margin formed tends to be sharp, shinier, and greener, while that in D. riojae is thick, not as shiny, and a paler green. Other differences include the upper leaf surface, which is often copiously glaucous-pruinose in D. gomeziorum, while typically only the abaxial leaf surface is slightly prui- PROVANCE AND SANDERS, NEW SPECIES OF DIOSPYROS FROM MEXICO AND CENTRAL AMERICA 287 nose-scintillant in D. riojae. The epidermal cells of the upper leaf surface in D. riojae are large, with thickened anticlinal walls, a feature absent in D. gomeziorum. Leaf shape in D. gomeziorum ranges from elliptical to lance-ovate or ovate, while the leaves of D. riojae often tend to be oblong or obovate. How- ever, this character overlaps, since both taxa may have elliptical leaves. In fruit- ing collections, the shorter, less stout, fruiting pedicels, the thinner, less scleren- chymatous fruit wall, and the glaucous-pruinose to scintillant fruit distinguish D. gomeziorum. The pedicel bracteoles of the male inflorescence in D. gomeziorum are twice as long as those in D. riojae, and are nearly folded length- wise (navicular). The anthers in D. gomeziorumare minutely granular, lacking the apical constriction and smooth texture seen in D. riojae. Carranza (2000) described male flowers of this species (as D. riojae) as cy- mose. We did not find cymes in material referable to either species. Develop- mentally, the inflorescences may be related to cymes, but, they appear to be fascicles. Some flowers may appear to be in dense terminal clusters (e.g., H. Rubio 1542). However, this appearance is superficial, since small terminal shoot api- ces can be found, although sometimes only with difficulty. 3. Diospyros conzattii Standl., J. Wash. Acad. Sci. 1217):399. 1922. (Fig. 3). Tyee: MEXICO. OAXACA: Distrito de Pochutla, Cerro Espino, Cafetal oa ead a a 1917 (fr), C. Conzatti 3167 (HOLOTYPE: US-1014759 not seen; ISOTYPES: MO-8 6! Gi 1100 m, with Reko & Makrinus), MO-1039787!, fragment US-892600! (in ene 900 m, Gch Reko & Makrinus). Trees or shrubs, 8-10 m tall, probably facultatively deciduous, trunk not re- ported; aged stems subterete to angular, bark verrucose, fissured, gray, stemwood off-white to yellowish or pale orange; 2nd—-3rd year stems angular to subterete, bark rimose, epidermis glabrous to sparsely hirtellous, sometimes pulverulent, sparingly clavate glandular hairy, densely lenticellate, puncticulate, chestnut to tawny, becoming silvery gray; current year’s stems quadrangular, finely sul- cate, glabrate to hirtellous, sometimes sparsely strigillose, the hairs off-white to brown, the epidermis shiny, viscous, olive to nearly black. Petioles usually thin, sometimes flexuous, 5-8 mm long, pale green to green-brown, rounded below, glabrous to appressed puberulent, conspicuously winged part way above, the wings becoming vertically oriented and narrow along the petiole, petiole slightly convex and minutely V-grooved above, to widely flat-channeled, to 3- channeled (main channel raised and skirted by side channels formed by verti- cal wings), glabrous to erect puberulent, sometimes clavate glandular hairy, especially in young leaves. Leaves alternate, simple, entire, pergamentaceous to chartaceous, lanceolate to elliptic to ovate, 5-14(-14.7) cm long, (2.5-)3-5(-6) cm wide, immature leaves membranaceous, apex acuminate to an acutely or obtusely rounded point, margins flat to subrevolute, curved downward near the base of the leaf, thickened intramarginal zone seen when viewed abaxially, hyaline and minutely ciliolate in immature leaves, base acutely or obtusely 288 BRIT.ORG/SIDA 22(1) di A Fruiti } hlet RS H £] . L h C. Adayial £ £ ct GPEA CA tr apu Fic.3.A nate petal lobe (eft) ant Hplal anther (ight). D.S 41.] + eee ten anaes aac edon JL. Matinee A, Hernandez 1302. B- E. Based on F. Ventura A. 7131. PROVANCE AND SANDERS, NEW SPECIES OF DIOSPYROS FROM MEXICO AND CENTRAL AMERICA 289 rounded, someti te, decurrent onto petiole, sometimes loosely recurved near the petiole; lower leaf surface usually glabrous, very rarely minutely white glandular hairy, puncticulate, dull, olive; upper leaf surface glabrous, glaucous, densely puncticulate, dull, olive. Venation arcolanguid; midrib very prominent below, slightly undercut, terete, glabrous, straw to light-green, weakly raised above, and then caniculate within the raised midrib, very sparsely minute erect hairy; 2°-3° venation reticulate, veins narrow, raised below and above, lateral veins 9-14 pairs, 2° arches formed with superadjacent lateral veins typically obscure. Laminar extrafloral ies abaxial, conspicuous, round to oval, dark- red to black, often evenly spaced and near the midrib. Male inflorescences cymes, I(-2) near the base of current year’s stems, or (1-)2 leaf scar axils of the previ- ous year’s growth, 1-3-flowered, black to umber velutinous, sparsely covered with minute clavate glandular hairs; peduncles 4-6 mm long, velutinous, um- ber to dark brown; pedicels slender, 5-7 mm long, vestiture as in peduncles; pedicel bracts 1-2, alternate or opposite, narrowly oblong, 1 mm long, 0.8 mm wide, umber. Male flowers 5(-6)-merous; male flowering calyx funnelform, dry- ing very dark-brown to black, glabrate to appressed-pubescent, coterminous with pedicel and uninterrupted; male flowering calyx tube 2-3 mm long, 1.5-2 mm wide; sepals 5, acute-triangular, 2-4 mm long, ciliolate, apically vermiform glandular hairy, sinuses rounded; corolla 5-6-lobed, long-urceolate, drying very dark-brown to black, reportedly whitish in life; corolla tube widest above middle, 7-8.5 mm long, 4-5 mm wide, abruptly constricted distally, exterior densely minute white puberulent, hair density + increasing distally, interior sparsely white puberulent, the hairs concentrated in regions of filament attachment; corolla lobes quadrate to oval, 2-2.5 mm long, 1.5-2 mm wide, asymmetrical, apex obtuse-rounded, truncate, emarginate, or trifid, often with several short dark hairs, exterior densely minute white puberulent, interior inflexed at the distal right margin, left margin slightly involute, pubescent. Stamens 30, some- times attached in pairs, adnate to corolla from midpoint to bottom of tube, some- times inserted on the receptacle; filaments 3 mm long, minutely hairy; anthers basifixed, lanceo-apiculate, 3-3.5 mm long, opening by terminal pores or short lateral slits, connective minutely deltoid-scaly, pistillode minute, obturbinate, smooth, nearly unlobed, having a few smooth very small basal lobes, glabrous. Female inflorescence {lowers solitary, cauline, at the junction of current and previous year’s growth. Fruiting pedicels stout, (7-)12-15 mm long, glabrous to sparingly minute hirtellous. Fruiting calyx 5-lobed; fruiting calyx tube 3 mm long, explanate to ee usually ae a conspicuous enlarged basal re- gion encircling the pedicel joint; fruiti t spreading to reflexed slightly, apices straight, coriaceous, narrowly to linearly triangular, 24-3 Lin long, 5-8 mm wide, margins sometimes sharp edged, sometimes glaucous, gla- brous, puncticulate. Fruit berry, depressed-globose slightly obovoid, up to 4cm long, 4.5 cm in diameter, reportedly reddish (Chazaro 3969) or green (Standley 290 BRIT.ORG/SIDA 22(1) 1922) when ripe, mostly atropurpureous in herbarium material. Locules 10-12; mesocarp reportedly black when ripe, usually orange in dried specimens, hy- podermis 0.2-0.5 mm thick, stony; epidermis bullate, glaucous-scintillant, usu- ally wrinkled when dry. Seeds 15-16 mm long, 11-12 mm wide, 5.5-9.5 mm thick, sculpture cerebriform, chestnut. Additional material examined: MEXICO. Veracruz. Mpio. Calcahualco: 4.2 km W of Escola, 19° LO'N, 97° 10'W, 2200 m, 12 Jan 1981 (fr), M. Nee & G. Schatz 19777 (XAL). Mpio. Chiconquiaco: abajo de el reuvane Arroyo Colorado, 19° 46'N, 96° 45'W, 1650 m, 25 Oct 1988 (fr), C. Gutierrez B. 3292 XAL). Mpio Coscomatepec: 3 km al SE de la antigua Xicola, Cima del Cerro La ee i; O6'N, 97° 04'W, 1900 m, 29 Apr 1987 (fr), J.L. Martinez & A. Hernandez 1302 (XAL). Mpio. Jalacingo: El Cuizalin, ca. 19° 50'N, 97° 16'W!, 1500? m, 22 May 1982 (pist. fl), F Ventura A.7131 (IEB, MO, XAL). Mpio. Tatatila: camino de herradura de Tatatila a Escalone (o Puente Caballos), [+ 19° 42'N, 97° 6'W, 1500 mJ, 14 Jan 1986 (fr), M. Chazaro & Roberto Acosta 3969 (WIS, XAL). Mpio. Tlacolulan: Abajo del Saucal, direccion Agustin ie 19° 45'N, 96° 57'W, 1180 m, 14 Aug 1990 (fr), C. Gutierrez B. 4056 (XAL). Distribution and Ecology—The type locality is in the Pacific coastal ranges of the Sierra Madre del Sur, Oaxaca. It occurs there in dry tropical forest on coastal slopes (Anonymous 1927). According to Conzatti (in Standley 1922), the fruit is ripe in April. In Veracruz, occurrences are near Pico de Orizaba, the Sierra de Tezuitalan, and the Sierra de Chiconquiaco, between 920 and 2200 m altitude. It occurs on slopes with pine-oak woodland and Alnus, oak forests, and decidu- ous forests. Ethnobotany.— Veracruz: ‘zapotillo’ (C. Gutierrez B. 3292). Oaxaca: ‘zapote negro montés’ (Standley 1922 [quoting Conzattil, and on US-892600), ‘zapote negro silvestre’ (Conzatti 3167 [in his own handwriting). The wood is consid- ered valuable (Anonymous 1927 [paraphrasing Conzatti)). Only one of the isotypes we examined, MO-879066, included a fruit. Un- fortunately, the ges atropurpureous fruit isin rather poor condition. However, on close examination it was clear that the epidermis is bullate, glaucous, al- though ae towards brown in some areas. The fruit wall is thick and quite stony. A fragment packet attached to another type, MO-1039787, contains a spreading fruiting calyx tube bearing the basal 5 mm of one sepal, and much less of the base of another. The sepals appear to have been quite narrow, ca. 4-5 mm wide at the base and further narrowed distally. Characteristics seen in the fruit of D. conzattii collections from Veracruz are consistent with the charac- teristics found in the isotype. An enlarged region below the fruiting calyx en- circling the pedicel joint is only seen in the Veracruz material, but not the isotype. At this point, we are unsure of its significance. 'Coordinates based on F. Ventura A. collection of Rhamnus capreaefolia var. capreaefolia from Cuizalin, 22 June 1970 (LL) accessed through REMIB. www.conabio.gob.mx/remib_ingles/doctos/ remib_ing.html. The label indicates the collection was made at 150 m. This seems to be a typ he . The coordinates f isali thi th Hi + for Cuizalin p | PROVANCE AND SANDERS, NEW SPECIES OF DIOSPYROS FROM MEXICO AND CENTRAL AMERICA 291 4. Diospyros costaricensis M.C. Provance & A.C. Sanders, sp. nov. (Fig. 4b, e). Tyre: ICA. GUANACASTE. Canton de Liberia: Parque Nacional de Guanacaste, Cordillera de Guanacaste, Cerro Cacao, Estacion Cacao, 10° 55' 45"N, 85° 28' 15"W, 1100 m, 14 Jul 1991 (fr), Carlos Chavez 569 (HOLOTYPE: MO-5316680!; IsoTyPEs: “4 duplicates” indicated on label, but not seen by us). poet D ttii Stand. similis sed ides-faciente, usque ad 35 m alta et 1.5m diametro; petiolis lil neque elevata in Baend abana 1 ] ( ] \ costa cao ad erroniel adaiale, | longictroreim cricnantiin ti pk Trees, probably pelea dy deciduous, reportedly colonial, reportedly up to 35 m tall, and 15 m in diameter (Espinoza 54); trunk buttress-forming, chan- neled, smooth, greenish-black to greenish-brown, slash yellow and aromatic; aged stems somewhat angular, half-netted, black to gray above, lower layer beige to golden-brown, sometimes + mottled, sometimes shallow-fissured, lenticellate; 2nd-3rd year stems irregularly half-netted, beige beneath, black and grey above, hirtellous, pubescence ee persisting on three year old stems, lenticellate; current year’s st sulcate to minutely ridged, sparsely subappressed Bolden hairy, the hairs fine and = straight, sparingly to very densely erect puberulent, dark-green to black, somewhat glandular, be- coming lenticellate. Petioles usually thin, somewhat flexuous, 3-6.5 mm long, wings tapering gradually from the decurrent lamina, and twisting abruptly into a vertical orientation along the margin, glabrous to hirtellous below, rug- ose to invaginate, light olive to very dark brown, concave above, often minutely V-grooved, usually golden hirtellous, often glandular, epidermis often nearly black. Leaves alternate, simple, entire, chartaceous to subcoriaceous, cartilagi- nous, ovate to elliptic, rarely widely-obovate, 6.5-9.5(-10.2) cm long, 2.8-4(-5) cm wide, very often tattered around the margins, wrinkled, apex usually asym- metric, tapering to an obtusely rounded tip, margin curved under near the base of the leaf, base mostly obtuse and tardily abrupt-acuminate, sometimes acute, long decurrent onto the petiole; lower leaf surface sparsely appressed puberu- lent, puncticulate, sometimes clavate glandular hairy near base, dull, brown to brown-green; upper leaf surface glabrous, very rarely glaucous-scintillant, pap- illose, puncticulate, brown-green. Venation brochidodromous; midrib promi- nent below, glabrous to sparsely appressed puberulent or hirtellous, epidermis green to brown-green, shallowly concave above, glabrous to golden hirtellous, sometimes partly shrouded by the lamina, then canaliculate and hairs criss- crossing, epidermis of basal half often black, otherwise greenish; 2° venation fine below, usually prominent, lateral veins 9-12 pairs, forming definite 2° arches with superadjacent lateral veins, arch apices 3 or more mm from the margin, venation tending to be obscure above, but variable; 3°-4° venation re- ticulated below, fine, usually apparent, but varying from obscure to prominent. Laminar extrafloral nectaries often up to 30, sometimes more, scattered on the abaxial side of the lamina, minute, peripherally rimmed with a narrow band 292 BRIT.ORG/SIDA 22(1) Fic. 4, A-E. Di taricensis M.C. Provance & A.C. Sanders, sp. nov. A. Adaxial view of staminate male calyx with corolla removed. 8 ean pian G elypicl stamen. ie Staminiats flaws: Es aaa with leaves. A, C, and D. Based on Zobeida Fuentes PROVANCE AND SANDERS, NEW SPECIES OF DIOSPYROS FROM MEXICO AND CENTRAL AMERICA 293 of raised tissue. Male inflorescence cymes, 2-3-flowered, 1-2 cymes per axil, from growth of the previous year, or from the base of new growth, densely subulate puberulent; peduncles up to 4 mm long, sparsely to densely pubescent to stri- gose with light brown to black hairs, epidermis dark brown; pedicels 0.5-3.5 mim long, vestiture and epidermis as in peduncles, pedicel bracts rarely persist- ing in herbarium material, 0.3-1 mm long, deltoid, densely hairy, the hairs dark- brown and straight. Male flowers 5-merous, calyx infundibuliform, exterior short-appressed gold to white puberulent, sometimes with clavate glandular hairs, epidermis dark brown; calyx tube 0.8-1.3 mm long, interior with ring of straight appressed white hairs; sepals usually erect, rarely slightly recurved, 2- 3.5mm long, 0.8-1.5 mm wide, interior glabrous to glandular setulose, exterior pubescent, often densely so at the apex, the hairs wavy and amber, appearing glandular; corolla long-urceolate; corolla tube 3.5-7.5 mm long, 3 mm wide, in- terior lower-third of tube short golden pubescent, hairs + erratically directed, exterior evenly and densely short appressed to ascending golden hairy, also clav- ate glandular hairy; corolla lobes 5, quadrate to oval, 2-2.8 mm long, 1.2-2.5mm wide; apex sometimes bearing a long flagellate trichome, interior right mar- gins involute, interior glabrous, exterior appressed to ascending white puberu- lent, the left side of lobe densely appressed fine puberulent, sometimes with several minute black hairs near the apex. Stamens 17-19, adnate at various lev- els at and below the basal third of the corolla; filaments 1-1.5 mm long, nar- rowed distally, glabrous to densely erect hispidulous; anthers basifixed, 2.3-2.5 mi long, several minute hairs at the apex, surface light-yellow apically, open- ing by short lateral slits confined to the distal half to two-thirds of the anther, sometimes opening by a complete lateral slit, pistillode + sub-conical, slightly wavy peripherally, + 8 long straight hairs originating from the central apex, otherwise glabrous. Female inflorescence flowers solitary in leaf scar axils of 2nd year stems, or cauline at junction of previous and current growth. Fruiting pedicels stout, 3-18 mm long, sparsely hirtellous, reddish-gold, and gray, lenticellate. Fruiting calyx 5-lobed, sinuses acute; fruiting calyx tube explanate, sparsely appressed puberulent, scintillant, + 3.5 mm long; fruiting sepals accrescent, coriaceous, lorate to lanceolate, (16-)22 mm long, (5-)6-8(-9) mm wide, spreading to moderately reflexed, apex straight to quite incurved, puncticulate, shiny, golden brown, sparingly scintillant basally. Fruit berry, sometimes pendulant, depressed-globose, 2.5 cm tall, 4 cm wide, reportedly green in living material, dull, atropurpureus to dark-brown in herbarium ma- terial, locules 10; mesocarp reportedly yellow in living immature fruits, dark orange in herbarium material; hypedermis + 0.5 mm thick, stony, not wrin- kling in herbarium material; epidermis bullate, glaucous to pruinose-scintillant, either locally or over most of the fruit. Seeds not seen. ParatypEs: Costa Rica. Guanacaste. Canton de Liberia: Parque Nacional de G caste, Estacion Ca- cao, 10° 55'45"N, 85° 28'15"W, 1100 m, 24 Nov 1990, R. Espinoza 54 (kK, MO); Cordillera de G t 294 BRIT.ORG/SIDA 22(1) Estacion Cacao, cna: a Estacion uMeniteas 10°55’ N, 85° 28'10" Wy, ie m, oe 996, José Gonzdlez et al. LIO8(K). Canton de La C 5 . V. Orosi, Sector Orosi oe Maritza), sendero ton Fram. 10° 57' 40"N, ne 29 45'W, 600 m, 13 Jul 1994, José Gonzdlez et al. 301 (MO [2 accessions]). Puntarenas. Canton de Puntarenas: Reserva Bioldgica, Monteverde, se de Tilaran, Altos de San Luis, Los Leitones, 10° 17'25"N, 84° 48' LO"W, 1200 m, 4 Sep 1991, E. Bello 4014 (MO); sendero a la Catarata, por el rio, sendero Miguel Leitén, 10° 16'20"N, 84° 49' 30"W, 1100 m, Ll Mar 1993, Zobeida ade 248(K. MO). Distribution and Ecology.—As far as known, this tree is endemic to northwest- ern Costa Rica, where it occurs in the Cordillera de Tilaran and the Cordillera de Guanacaste, between 600 and 1200 m in elevation. Details concerning asso- ciated vegetation are mostly lacking on the collections from the Cordillera de Guanacaste, a drier and more seasonal mountain range than the Cordillera de Tilaran (Hammel et al. 2004). This species could be associated with deciduous forests of the region. The collections from Puntarenas come from the Pacific side of the Cordillera de Tilaran. Label data from one of the collections, Bello 4014, indicates that the tree was growing in charral, or young secondary forest (Kleinn et al. 2002). The other collection does not provide details about the as- sociated vegetation. Ina broad sense, the region of the occurrences in Puntarenas has been mapped as Costa Rican seasonal moist forests. These are deciduous forests that obtain 90% of their annual precipitation, (+ 1500 mm total), dur- ing the months of April through October (World Wildlife Fund 200D. The for- mation of new leaves and male flowers apparently occur during March (only one flowering collection was examine Ethnobotany.—Known by the common name ‘guacalillo’ in Puntarenas, Costa Rica (Fuentes 248). Etymology.—The epithet refers to the only country in which the species is currently known to occur. This species is different from D. conzattii in several respects. First, it is a buttress-forming tree attaining 35 m in height; this is quite exceptional among Diospyros from Central America and Mexico. Dine. conzattii is not known to reach over 10 m in height. The abaxial leaf venation is conspicuously brochi- dodromous, the lateral veins forming well defined secondary arches with the superadjacent lateral veins. Another interesting difference involves the distance from the outer-perimeter of secondary arches to the leaf margin, which is usu- ally around 3-6 mm at about mid-leaf. This distance is 1-3 mm in D. conzattii. The marginal loops of D. conzattii often become difficult, but not impossible, to discern, hence the aforementioned distance is measurable. Some other veg- etative differences useful for separating these species include yellowish to or- ange hairs on the adaxial midrib, and the converging adaxial lamina along the depressed midrib in D. costaricensis. The inflorescences of D. conzattii and D. costaricensis are clearly cymes, with peduncles around 6 mm and 4 mm long respectively. PROVANCE AND SANDERS, NEW SPECIES OF DIOSPYROS FROM MEXICO AND CENTRAL AMERICA 295 5. Diospyros tuxtlensis M.C. Provance & A.C. Sanders, sp. nov. (Fig. 5). Tyre: MEXICO. VERACRUZ: Mpio. San Andrés Tuxtla, borde de la cima del Cerro Mastagaga, al N del Ejido Ruiz Cortinez, Sierra de los Tuxlas, 30 May 1985, J. Calzada 11855 (HOLOTYPE: IEB-48870; ISOTYPE: XAL). Arbor usque ad 10 m alta, D. rigjae Gomez- pempe emis sed dintert tole eliptcls et non -glaucis marginibus non crassis et inter fructiferis 26-40 mm longis oO faa Q Glia ercau ans : a : 1 ea eae ay i 14 mm ioaeis Trees 8-10 m tall, probably facultatively A caenois aged terete to subterete, + glabrous, half-netted to squamose, dark-brown to grey-brown with beige, short-fissured stem wood yellowish to orange-brown; 2nd-3rd year stems an- gular to subterete, glabrous, sulcate to shallow-fissured or half-netted, in com- binations of dark-brown, gray, and orange-brown; current year’s stems angular, smooth to finely sulcate, very cream to grey, shiny, lenticellate, glabrous to densely minute hirtellous, appressed off-white puberulent near apex, puncticulate. Petioles 4-7 mm long, glabrous, flat to slightly concave above, golden-brown, below widely rounded, glaucous-scintillant, dark-purple. Leaves alternate, simple, entire, pergamentaceous to chartaceous, + elliptic, 8-15 cm long, 3-5(-5.8) cm wide, widest at or just above the middle, both sides very sparsely scintillant, but leaf surface dull golden-brown to bronze between in- dividual scintillae, apex usually acuminate, the acumen long, narrow, obtusely rounded at the tip, margin flat with slight intramarginal thickening, base acumi- nate to attenuate, decurrent onto petiole; lower leaf surface sometimes spar- ingly appressed puberulent near base and midrib, conspicuously puncticulate; upper leaf surface glabrous, papillose. Venation arcolanguid to brochidodromous; midrib conspicuously flat and wide below, sulcate, purplish basally, becoming orange-red to golden-brown and somewhat keeled apically, sometimes sparsely appressed puberulent, glaucous, shallowly concave above, glabrous, not darkened; 2° venation raised above, obscure, lateral veins 8-9 pairs, shiny, chartreuse to golden-brown and conspicuous; 3° venation obscure be- low, granular-papillose, apparent above. Laminar extrafloral nectaries abaxial, common, sometimes along 2° veins, rimmed with the same bright color seen in 2° veins Male inflorescence unknown. Female inflorescence flowers solitary, emerging at the junction of old and new growth. Fruiting pedicels stout, 14 mm long, 3mm wide, wavy-rugose, minutely fissured, lenticellate, black and tan mottled. Fruiting calyx with lobes and distal portion of tube strongly reflexed, sinuses rounded: fruiting calyx tube exterior sparsely appressed puberulent; fruiting sepals 5, very accrescent, 26-40 mm long, 7.5-11 mm wide, ovate to widely lanceolate, narrowed basally, apex acutely pointed, sometimes rounded sepals glabrous, golden-brown, sepal nerves distinct. Fruit berry, depressed-glo- bose, up to 4 cm tall, 4.5 cm in wide, atropurpureous, number of locules indis- cernible; mesocarp dark-brown in live material and in immature fruits of her- barium material; hypodermis + 0.5 mm thick, sclereidic; epidermis 296 BRIT.ORG/SIDA 22 1) Fic. 5. Diospy lensis M.C, Provance & A.C. Sanders sp. nov. Fruiting branch. Based on J./. Calzada 11855, glaucous-scintillant, wrinkled. Seeds immature, sizes indiscernible (broken seeds only), dark reddish brown, granulate to minutely ruminate (sensu Stearn 2000, fig. 38). PARATYPES: Mexico, Veracruz. Mpio. San Andrés Tuxtla: senda para el Cerro Baxin, al N de San Andrés Tuxtla, Sierra de Los Tuxtlas, 6 Mar 1985, J. Calzada 11929 (IEB, XAL). Mpio. Soteapan: Ejido Santa Marta camino a la Ventana, 18° 22'N, 94° 54'W, 920 m, 19 Sep 1986, R. Acosta P & C. Gonzalez R. 1312 (LSU, XAL). Distribution and Ecology.—Apparently endemic to the Sierra de Los Tuxtlas, PROVANCE AND SANDERS, NEW OFOLILI VE VIVO AMERICA 297 Veracruz, Mexico. Only one elevation recorded, 920 m (R. Acosta P. & C. Gonzalez R. 1312). The associated vegetation types for the known collections have included primary selva mediana perennifolia, selva baja perennifolia, and deciduous for- est. It was considered common in selva baja perennifolia with elements of de- ciduous forest (Calzada 11855). Ethnobotany.—A common name has not been reported, but since the fruits are likely to be edible, the species may be well-known to local residents. The species may eventually be found as a conserved tree near homes or pastures. Etymology.—The epithet refers to Sierra de Los Tuxlas, Veracruz, Mexico, from where the only collections of this species have been made. SPECIES OF UNCERTAIN STATUS si pergamentacea Lundell, Contr. Univ. Michigan Herb. 7:44. 1942. Type. EXICO. CHIAPAS: Pico de Loro, near Escuintla, in advanced forest, 2200 m, 25 Jun 1941, Eizi ny 4278 (HOLOTYPE: MICH not seen; ISOTYPES: photo A!, photo CAS! (sterile), photo FI, IT not seen, photo LL-372460!, MO- 1213673) ae i 3). A redescription of this taxon did not seem appropriate, given that there are no new specimens. We do not think we can add much to the work of Lundell. Over- all, this species looks quite similar to D. conzattii, with which some authors have considered it conspecific’. This species is known only from the type speci- men, collected on Pico de Loro, near Escuintla, Chiapas. The petioles are very long and flexuous, reported to be up to 15 mm long by Lundell (1942). There also seem to be differences in the sepals, they being strongly reflexed, and hav- ing a shape that is suggestive of some populations of D. rosei sensu lato. In fact, these differences are significant enough that we are not entirely convinced that itis synonymous with D. conzattii. The leaves of the isotypes we examined had a thin coating of clear glue on much of their surface, which may obscure some characters. We recommend re-evaluating the taxonomic status of this taxon when additional material from the Chiapas-Guatemala borderland becomes available. For the time being, we do not recommend reduction to synonymy with D. conzattii. DISCUSSION Neither the holotype nor the two isotypes of D. riojae were available for a first- hand examination. Unfortunately, there are no paratypes for D. riojae, D. conzattii, or D. pergamentacea. Microfiche of an isotype and the detailed de- scription of D. riojae by Gomez-Pompa (1964) were valuable in completing this paper. The original description made use of tables and text in contrasting dif- ferences between D. riojae and D. conzattii. Additionally, it was supplemented >For example: A. Gomez-Pompa (1964); F. White in 1968 and C.Whitefoord & S. Knapp in 1996, in both cases by annotation of the isotypes. 298 BRIT.ORG/SIDA 22(1) with an illustration prepared from the original collection (Gomez-Pompa, pers. comm.). The illustration is consistent with the type description, and the isotype at US, and in accordance with the Code, Article 9.2: Note 2 (Greuter et al. 2000), the illustration in the protologue represents original material. Because fruiting D. riojae material keys to D. conzattii in Standley’s treat- ment of Diospyros of Mexico (1924), Gomez-Pompa (1964) provided a table of leaf and fruit characters that could be used to separate these taxa. Admittedly, some of these characters overlap to some degree. When it is considered that Gomez-Pompa had one fruiting collection of D. riojae, one fruiting collection of D. pergamentacea, and probably only one collection of D. conzattii toexam- ine, it is to his credit that the characters he emphasized are often still useful for separating these taxa. We have provided an updated character table (Table 1) that should be helpful in the differentiation of the three new species described here. Redetermination of the available material resulted in roughly a 70% re- duction in the number of documented occurrences of D. riojae from what might have been reported based on an uncritical review of locations based on her- barium material. At the least, this constitutes a reaffirmation of the rarity of this species. The realization that D. conzattii isan element of the Veracruz flora is very interesting. As far as we can tell, these are the first reports of the species for Veracruz. Material from the Cordillera de Guanacaste, and the Cordillera de Tilaran, Costa Rica, represents a distinct new species, D. costaricensis. The dis- covery of a new species, Diospyros tuxtlensis, from the Sierra de Los Tuxtlas does not come as a shock, since this region is renowned for having numerous endemic species of plants and animals. The distributions of the closely allied taxa, D. tuxtlensis and D. costaricensis, are notable in light of some recent inter- est in Los Tuxtlas-Costa Rica disjunctions (Hammel 1997). In addition to occur- ring in native stands of vegetation, individuals of this taxon should be sought as conserved trees in local gardens and pastures. This taxon seems to be a rare endemic of the Sierra de Los Tuxtlas, and could be in need of formal protection. We have not seen female flowers of any of these species. The only descrip- tion of a female flower that we have seen in the literature is by Carranza (2000) and refers to material from Queretaro (D. gomeziorum). The lack of flowering Ebenaceae material in herbaria, as pointed out by Gomez-Pompa (1964) and Wallnofer (2001), and clearly demonstrated by our sample, makes searching for taxonomically informative vegetative characters a particularly attractive proposal (Gomez-Pompa 1964). Below we provide a key to the species described in this paper that emphasizes vegetative characters. Male flower and inflores- cence characters (Table 2) can be used to supplement the following key if de- sired. It should be noted that D. tuxtlensis is not treated in Table 2, since flower- ing material is not currently known. Taste 1.Vegetative and fruiting morphology across the described taxa. gomeziorum costaricensis conzattii riojae tuxtlensis Life form and size Leaf shape Leaf margin Leaf luster (both sides) Leaf vestiture below Leaf bloom above Epidermal cells of lamina above Epidermal cells of lamina below Midrib vestiture above Midrib topography above Number of major lateral veins Emergences on venation above Petiole color above Trees or shrubs to 20 m tall Feige to ovate to elliptic beveled, sometimes ciliolate, not thickened shiny glabrous to sparsely subappressed hairy glaucous to pruinose- scintillant not large and not thickene puncticulate sparsely to densely hirtellous to deltoid caly narrowly caniculate (8- Ww 11-13(-14) 2°-3° granular papillose yellow-green to olive- green Buttressed trees to 35m _ Trees or shrubs, 8-10 m tall ovate to elliptic, rarely widely-obovate curved under near base, not thickene dull sparsely appressed hairy, sometimes glandular very rarely glaucous- scintillant not large and not thickened puncticulate glabrous to golden hirtellous shallowly concave 9-12 none nearly black tall lanceolate to elliptic to ovate urved under near base, intramarginal thickening dull usually glabrous, very rarely glandular glaucous not large and not ickened puncticulate very sparsely hirtellous caniculate, sometimes raised 9-14 none pale green to green- brown Trees, rarely shrubs, to 25m tall elliptic, oblong, obovate, or nearly oval cs urved, thickened, so ie shiny metimes glabrous sometimes slightly pruinose or scintillant large and thickened puncticulate glabrous, sometimes sparingly glandular hall | DHIAHOVWIY COMCAVE 8-12 2°-3° granular papillose light green to chartreuse to amber Trees, 8-10 m tall elliptic flat, slight intramarginal thickening dull glabrous to sparingly appressed puberulent slightly scintillant somewhat large and thickened conspicuously puncticulate glabrous shallowly concave 8-9 3° granular papillose golden-brown ICNIN 4N SaNnadS MAN “SHIGNYS aNv JNVAOUd VOIUIWY 662 00€ Taste 1. continued gomeziorum costaricensis conzattii riojae tuxtlensis Petiole color below Fruiting calyx posture Fruiting sepal apices Fruiting sepal length Fruiting sepal shape Fruiting pedicels Fruit hypodermis olive-green to dark brown, gli tening spreading apices ascending 19-20 mm long lanceolate to narrowly elliptic or obovate 4)5-8(-11) mm long, not stout thin light olive-green to very dark brown spreading to moderately reflexed apices straight to quite incurve (16-)22 mm long lorate to lanceolate 3-18 mm long, stout thick pale green to green- brown spreading to slightly reflexed apices straight 24-31 mm long narrow to linear triangular (7-)12-15 mm long, stout intermediate light green to chartreuse to amber arcuate-reflexed to spreadin oblong, narrowly deltoid or ovate 7-12 mm long, stout thick dark purple strongly reflexed apices straight 26-40 mm long ovate to widely lanceolate 14mm long, stout thick (L)@Z VaIS/DYO"LINS Taste 2, Reproductive morphology of four similar taxa from Mexico and Costa Rica. Diospyros gomeziorum Diospyros costaricensis Diospyros conzattii Diospyros riojae Male inflorescence Male corolla tube shape Male corolla tube interior vestiture Stamens Anthers Filaments fascicles on previous year’s growth, solitary on new growth urceolate-campanulate sparingly hirtellous, hairs sometime erratic 20 2mm long, mer ovate to ovate, minutely granular, qe slits near apex 0.3-0.8 mm long, ascending-puberulent cymes from previous year’s growth, cymes from new growth long-urceolate lower-third of tube short golden pubescent, hair direction erratic 17-19 2.3-2.5 mm long, several minute hairs at apex, short lateral slits in distal 2/3 to 1/2 1-1.5 mm long, glabrous to densely erect hispidulous cymes from previous year’s growth, cymes from new growth long-urceolate sparsely puberulent, densest from mid-tube to tube bottom 30 3-3.5 mm long, lanceolate, apiculate, short lateral slits near apex 3 mm long, minutely hairy fascicles from previous year’s growth, fascicles from new growth urceolate-campanulate deltoid scaly at mid-tube 18 + 2.5 mm long, lance-ovate, constricted near the apex 1 mm long, hirtellous, especially along margin ICHIN 40 CAINIAC AAIN SHIGNYS any JINVAOYd VoIdIWV LOE 302 BRIT.ORG/SIDA 22(1) A KEY TO DIOSPYROS RIOJAE, DIOSPYROS CONZATTII AND SOME ALLIED BLACK ZAPOTES 1. Leaves dull, laxly revolute (loosely rolled under) near the base of the leaf; neither the 2° nor the 3° upper leaf surface venation granular papillose. 2. Buttressed trees up to 35 m tall and 1.5 min diameter; petiole color above et ae leaf margin or intramargi ina zone not thickened (viewing bottom sur- e); midrib ab shallowly, and often ples concave, or crimped eae by the adjacent sides of the lamina (boxed i gl to golden hirtel- lous; currently known only from Costa Rica 4. Diospyros costaricensis 2. Trees or shrubs, 8-10 m tall, lacking buttresses as far as known; petiole color above pale green to green-brown; leaf margin or intramarginal zone thickened (on bottom surface); midrib above ately caniculate, the canal(s) raised or + even with the lamina; midrib very sparsely hirtellous; currently known only from Veracruz, Oaxaca, and possibly from Chiapas 3. Diospyros conzattii . Leaves shiny or dull, margins flat or sometimes subrevolute, but never loosely rolled under near the base; 2° or 3° venation, or both, usually granular papillose on upper leaf surface 3. Leaf margin beveled outward from the upper leaf surface, down to the lower leaf surface;epidermal cells of the upper lamina surface not large and conspicu- ous with thickened anticlinal cell walls; leaves often copiously glaucous-prui- nose; fruiting calyx spreading, fruiting sepals with ascending sees fruiting edicels generally slender 2. Diospyros gomeziorum 3. Leaf margin never beveled; epidermal cells of the upper lamina yes large and conspicuous, having thickened anticlinal cell walls; leaves sometimes some- what pruinose or scintillant, but never with copious bloom; fruiting calyx spread- ing to strongly reflexed, apices various; fruiting pedicels very stout . Leaves shiny, often tending towards oblong, oval or obovate, but may also be elliptic; leaf margins thickened, rounded from the top leaf surface to the bot- m; petiole color below light green to chartreuse or amber; fruiting sepals 7-25 mm long, spreading to arcuate-reflexed, the apices incurved to ascend- ing; fruiting pedicels 7-12 mm long 1. Diospyros riojae . Leaves dull, elliptic; leaf margins flat or with a slight intramarginal thickening visible on the bottom surface; petiole dark purple below; fruiting sepals 26— 40 mm long, strongly reflexed, the apices straight; fruiting pale 14m long eae tuxtlensis tS ACKNOWLEDGMENTS The authors are grateful to Lia DeMarco for preparing the Latin diagnoses, Clau- dia Ross-Ibarra for translation of the abstract in to Spanish, Edward Plummer for technical support and editorial assistance, Victor Steinmann for reviewing an early version of the manuscript, and Araceli Aguilar for her assistance. Two anonymous reviewers made excellent suggestions that ultimately improved the final manuscript. Access to microscopes was kindly made possible by Martha L. Orozco-Cardenas at the UCR Plant Transformation Research Center. We ex- tend our gratitude to the herbaria that provided material for use in this study: ARIZ, CAS, CHAPA, CIIDIR, DES, HUAA, IEB, K, LSU, MO, SD, UCSB, US, UTEP, WIS, and XAL, and also Arturo Gomez-Pompa, the Field Museum, Chicago, and PROVANCE AND SANDERS, NEW SPECIES OF DIOSF AMERICA 303 the Plant Resource Center, Austin, for providing images useful in this study. The first author would like to extend a warm thanks to Juanita and Seychelle Provance for their financial support, logistic tolerance, and patience. We would like to thank Giles Waines and Arturo Gomez-Pompa for their insight and en- couragement. We are grateful to the Remib Database and Jose L. Panero (TEX, LL), the caretaker of the node that helped pinpoint El Cuizalin, Veracruz. Fi- nally, we thank UCMEXUS and the UCR Center for Conservation Biology for generously supporting a visit to XAL to photograph specimens. REFERENCES ANONYMOUS. 1927.Book review:Las regionas botanico-geogréficas del Estado de Oaxaca, Sep 1st:27-29. Carranza G,, E.2000. Ebenaceae. In: Fl. Bajio. Rzedowski, J.and G.Calderén R.(eds.). Instituto de Ecologia A.C., Centro Regional del Bajfo, Patzcuaro, Michoacan, México fasc. 83:1—9. Conoe, B.N. 2000. Germinacion de Diospyros riojae Gomez Pompa. Thesis. Universidad Ve- racruzana. México Gomez-Pomea, A. 1964. A new Diospyros from the Misantla region in México. J. Arnold Ar- bor.45:464-470. GreuTer, W., J. MCNeiLt, F.R. Barrie, H.M. Burbet, V. Demoutin, T.S. Fitcueiras, D.H. NicoLson, P.C. SiLvA, J.E. Skoc, P. TREHANE, NJ. TURLAND, and D.L. Hawkswortu (eds.). 2000. International code of botanical nomenclature. Regnum Veg. 138. Hammet, B.E. 1997. Three new species of Celastraceae from Costa Rica, one disjunct from Mexico. Novon 7:147-155. Hammel, B.E., M.H.Grayum, C. Herrera, N. Zamora (ed5.). 2004. Manual de plantas de Costa Rica: Volumen 1:Introduccién. Monogr. Syst. Bot. Missouri Bot. Gard. Vol. 97. Missouri Botani- cal Garden, St. Louis. Jackson, B.D. 1916. A glossary of botanic terms: with their derivation and accent. Second ed. London. Pp. 209. Keity, L.M.2001.Ebenaceae. In: Fl. Valle Tehuacan—Cuicatlan. Kelly, L.M., Ochoterena, H.,and R. Medina L. fasc. 34:1—5. Instituto de Biologia, Universidad Nacional Aut6noma de México. KeINN, C., L. Corraces, and D. Morates. 2002. Forest Area in Costa Rica: A comparative study of tropical forest cover estimates over time. Environ. Monit. Assessm. 73(1):1 7-40. Lunbett, C.L. 1942. Studies of American spermatophytes—ll. Plants of México, British Hon- duras, Guatemala, Costa Rica, Panama, and Puerto Rico. Contr. Univ. Michigan Herb. Pacueco, L. 1981. Ebenaceae. In: Fl. Veracruz. Gomez-Pompa, A. and V. Sosa, eds. fasc.16:8- 16. Instituto Nacional de Investigaciones sobre Recourses Bidticos, Xalapa. Provance, M.C. and A.C. Sanpers. 2005. Diospyros torresii (Ebenaceae): a new black zapote from tropical Mexico. Sida 21:2045-2050. SANCHEZ-COELLO, N.G. 2002. Induccién de respuestas morfogenéticas en Diospyros riojae Gomez-Pompa en la poblacién de Cruz Blanca, mpio. de Alto Lucero, Veracruz. Thesis. Insituto de Genetica Forestal, Universidad Veracruzana. Xalapa, Veracruz, México 304 BRIT.ORG/SIDA 22(1) Sosa, V., A.P. Vovioes, and G. Castitto-Campos. 1998, Monitoring endemic plant extinction in Veracruz, México. Biodiversity & Conserv. 7:1521-1527, STANDLEY, PC. 1922. Diospyros conzattii,a new species of persimmon from México.J.Wash. Acad. Sci. 12(17): 399-400. STANDLEY, PC. 1924. Trees and shrubs of México. Contr. U.S. Natl. Herb. 23(4):849-1312. STEARN, W.T. 2000. Botanical Latin, 4th ed. Portland, Oregon. Timber Press. P. 438. Toozia, C.A.and R.C. Keatinc. 1991. Leaf architecture of the Chloranthaceae. Ann. Missouri Bot. Gard. 78:476-496. TrasuT, D.L. 1926. Les Diospyros comestibles. Rev. Int. Bot. Appl. Agric. Trop. 6(63):675-678. Wattnorer, B. 2001. The biology and systematics of Ebenaceae: a review. Ann. Naturhist. Mus. Wien, Ser. B, Bot. Zool. 103:485-512. Wuite, F. 1983. Ebenaceae. In: Fl. Zambesiaca. E. Launert, ed. Vol. 7(1):248-300. Wauiteroorp, C. and S. Knape. 2001. Ebenaceae. In: Fl. Mesoamericana—lInternet Version, Davidse, G., Sousa S.,M., and S. Knapp, eds. 1998. Universidad Nacional Autonoma de México, México, Missouri Botanical Garden, St.Louis and The Natural History Museum, London. http://www. mobot.org/mobot/fm/welcome.html. Worto Witouire Funp. 2001. Powell, G., Palminteri, S.,and J. Schipper, preps. Costa Rican sea- sonal moist forests (NTO119), unrevised document. http://www.worldwildlife.org/ wildworld/profiles/terrestrial/nt/nt0119_full.Atml (29 June 2005 a TAXONOMY OF NORTH AMERICAN SPECIES OF OLDENLANDIA (RUBIACEAE) Edward E. Terrell! Harold Robinson Research Associate Curator Department of Systematic Biology-Botany Department of Systematic Biology-Botany Nati ee tae n of Natural ton oe Museum of Natural History titution mithsonian Institution ee DC. 20013-7012, USA. maa DC 20013-7012, U.S.A. ABSTRACT This paper presents a taxonomic treatment of nine peas of Oldenlandia occurring as native or adventive in United States or Mexico. The subg Oldenlandia is represented by four species: O. corymbosa, O. lancifolia, O. uniflora, and O eer Subgenera for the remaining species are undeter- mined. Four species are native to Mexico: O. pringlei, O. microtheca, O. ovata, and O. drymarioides. A ninth species, O. salzmannii, is native to South domed and aavenEcy in northwest Florida and adjacent Alabama. Keys, descriptions, and g g f the seeds are provided. RESUMEN Este articulo presenta un tratamiento taxonomico ode nueve especies, de oo wie que ocurren como nativas o adventicias en Estados por cuatro especies: O. corymbosa, O. lancifolia, O. eer eae Los subgéneros ae la especie restante son eieceuen Cuatro especies son nativas de Meco O. Parnes O. microtheca, O. ovata y O.drymarioides. ,O. salzmannii ti UVOCTILICIA el noroeste ond y pee ae. Se proporcionan claves, pea : eee a arrido de las semillas. Oldenlandia L. (Hedyotideae; Rubiaceae) is a genus of about 100 species (Verdcourt 1976) distributed worldwide in warm, subtropic, and tropic regions. It was named by Linnaeus for Henrik Bernard Oldenlan, a Danish physician and botanist of the 17th century. Bremekamp’s monograph (1952) of the African species of Oldenlandia rec- ognized 61 species in 16 subgenera. Verdcourt (1976) treated 37 species of Oldenlandia in the Flora of Tropical East Africa, and 25 species (1989) in the Flora Zambesiaca region. Other authors have recognized 12 species in West Tropical Africa (Hepper & Keay 1963) and 7 in Gabon (Hallé 1966). Lewis (1965) commented that Africa has more Oldenlandia species than exist in all other tropical regions combined and is the center of its morphological diversity. enlandia was considered as only a subgenus of Hedyotis by Fosberg (1943) in his study of the Polynesian Hedyotis species; however, the foreign flo- 'Address for correspondence: 14001 Wildwood Drive, Silver Spring, Maryland 20905, U.S.A SIDA 22(1): 305 — 329. 2006 306 BRIT.ORG/SIDA 22(1) ras cited here and recent American regional floras have all treated Oldenlandia as a distinct genus. Terrell (1996) discussed problems of generic limits among Hedyotis, Houstonia, and Oldenlandia while revising Houstonia, a genus of 20 North American species. Terrell and Robinson (2003) circumscribed Hedyotis subge- nus Hedyotis,an Asian subgenus, and recognized the genus Exallage as a new subgenus of Oldenlandia. A synopsis of the United States species of Oldenlandia (Terrell 1990) treated five species. Molecular studies found Oldenlandia to be paraphyletic (Bremer 1996) or polyphyletic (Andersson & Rova 1999). Its species were widely distributed in the rubiaceous lineage in what Bremer (1996) and Bremer and Manen (2000) treated as the tribe Spermacoceae, that name having priority over the Hedyotideae within which it was nested. Tested members of Oldenlandia are dispersed in the slightly paraphyletic part of that lineage that Terrell and Wunderlin (2002), Andersson and Rova (1999) and the present authors would continue to treat as a separate tribe Hedyotideae. Lewis (1965) described the pollen of Oldenlandia and other genera in de- tail. In Oldenlandia the pollen are small or medium-sized and (2-)3(4-5) aperturate. MATERIALS AND METHODS The taxonomic treatment is based on loans of many herbarium specimens, as cited in the representative collections. All species except O. pringlei and O. drymarioides were seen in living condition. Oldenlandia microtheca and O. ovata were studied and collected in Mexico. Oldenlandia salzmannii was ob- served in greenhouse plants. The North American species of subg. Oldenlandia were collected in various parts of their ranges. Previous work on Hedyotideae has shown that capsule and seed morphol- ogy are especially important in taxonomic studies. In the present work we ex- amined the seeds of each species by scanning electron microscopy. The results of this study are presented below in the summary of the more significant char- acters of each species. RESULTS AND DISCUSSION Oldenlandia subgenus Oldenlandia The type species of Oldenlandia, O. corymbosa, isa worldwide weed well estab- lished in eastern North America. Other members of subgenus Oldenlandia in- clude O. lancifolia, adventive in Mexico, and the native US. species, O. uniflora and O. boscii. The latter two species differ from the other two in having axillary and terminal glomerules and slightly smaller corolla and other flower parts, but otherwise have similar characteristics (Table 1). TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 307 Taste 1.Characters of species in subg. Oldenlandia. corymbosa lancifolia uniflora boscli Duration annual perennial/ann. annual perennial Habit erect/ prostr. erect/decumb. erect/prostr. sprdg./prostr. Leaves, Width mm ~9 - 2-11 1-5 Stipules, L, mm to2 to 1.5 to4 to2 Inflorescence pedicellate cymes pedicellate cymes glomerules glomerules Corollas,L, mm 1-2 15-2. 0.7-1.3 0.7-1.3 ape rotate/tubular rotate/tubular rotate rotate Tube L,mm 0.5-1.0 1-0.3 0.1-0.3 Lobes, L, mm 0.5-1.0 0.5-1.5 0.7-1.2 to 0.7 Anthers, L, mm 0.2-0.3 0.4--0.6 0.1-0.3 0.1-0.3 Filament, L, mm 0.1-0.3 0.2-0.5 0.2-0.4 0. Stig. Lobes, L, mm 0.3-1.0 0.5-1 .2 less than 0.6 less than 0.5 Capsules, L, mm 1-2.2 1.6-3.5 1-2.5 1.5-3.0 Seeds, L, mm 0.2-0.4 0.25-0.35 0.2-0.3 0.1-0.3 The following description of subgenus Oldenlandia is based mainly on the North American species. Small annual or perennial herbs usually less than 80 cm tall. Stems slen- der, erect, decumbent, or prostrate. Leaves 3-60 mm long, opposite, sessile or petiolate, usually linear to ovate. Stipules 1.5-4 mm long, interpetiolar, adnate to leaf bases, with 1-few marginal teeth, setae, or fimbriae. Inflorescences are of two types, either open, pedicellate, with few-flowered cymes from axillary and terminal nodes or else the flowers sessile or subsessile in axillary and terminal glomerules. Flowers tetramerous, isostylous (homostylous) in our species, but species in this subgenus elsewhere may be heterostylous. Hypanthium (calyx cup) cup-shaped or hemispheric. Corollas 0.7-2.5(-4) mm long, rotate or tubu- lar, white or tinted pink or purple, glabrous externally; tube 0.1-1.0 mm long, shorter or longer than the 4 lobes; lobes 0.5-1.5 mm long. Anthers 0.1-0.6 mm long, dorsifixed, sessile or on short filaments inserted on corolla tube at or be- low corolla sinuses. Stigmas 2-lobed, 0.2-L2 mm long, styles filiform or thick- ened, glabrous. Ovules on peltate placentae. Capsules 1-3.5 x 1-4 mm, bilocu- lar, usually subglobose, fused with hypanthium, 3/4 to fully inferior, dehiscing loculicidally or also septicidally. Seeds 50-100 or more per capsule, 0.1-0.4 mm long, black, brown or tan, trigonous or conoidal-trigonous, basal face oval or elliptical, often more or less flat, lateral faces slightly or somewhat concave, hilum punctiform at apex of the apical angle, testa reticulate, areoles (cells) usually polygonal, areole walls low, rounded, indistinct, testa surface often ver- rucose (densely covered with minute papillae). Placenta fused with septum about 1/3-1/4 of the distance above its base. Chromosome number x = 9. When compared with the other species (Table 2) the unique or unusual 308 Taste 2. Comparison of four Oldenlandia taxa. BRIT.ORG/SIDA 22(1) subg. Oldenlandia Pringlei Salzmannii Microtheca No.of species examined four one one three Habit erect/prostrate ascdq./prostr. creeping erect/decumbent rhizomatous Leaves Lxw,mm 3-60 * 1-12 3-15 XK 03-2 1.5-5.2 X 0.7-3 3-40 * 1-20 Stipules L 1.5-4mm 0.3-1.0 to 0.5 0.5-2 Inflorescence cymose/glomer. cymose solitary cymose Corollas L - 4.0-8.5 2-5.5 25-7 Cor. shape rotate/tubular subsalv./funnel. subsalverform — funnelform Cor. tube L 0.1-1.0 mm 1-2.2 1-3 Cor. lobes L 0.5-1.5 mm 1.5-3.5 1-3.2 1.5-4 Anthers L 0.1-0.6 mm 0.8-1.3 0.5-0.8 0.4-1 Capsules L x W 1—3.5 x 1-4 2-3 X 2-2.5 1.5 X 1.5 1-3 X 1.5-3.5 Seeds no./caps. 50-100 + 30-50 4-14 10-34 Seeds L/dia.mm 0.1-0.4 2-0. 0.3-0.5 0.4-0.7 Seeds shape trigonous subglob/ovoid trigonous angulate/irreg conoidal Seeds, areoles polygonal polygonal polygonal none Areole walls low, rounded thick, sinuous distinct none Testa reticulate reticulate reticulate not reticulate Testa surface verrucose/other — verrucose smooth entangled str Chrom.no. x= 9 ? 15 11,12 features of subg. Oldenlandia include the following: Inflorescence either pedicel- late in small cymes at the nodes or else flowers subsessile in glomerules; flowers usually isostylous (homostylous); corollas small, often 0.7-2.5 mm long, rotate or tubular; anthers 0.1-1.0 mm long; seeds 50-100 or more per capsule, 0.1-0.4 mm long, trigonous, areole walls low, rounded, indistinct; chromosome num- ber x =9 in many species worldwide. Seed data for this subgenus are described and illustrated in Figures 1-3. Figure | shows typical trigonous oldenlandioid seeds in O. corymbosa and O. lancifolia witha reticulate surface composed of polygonal areoles. A basal face is often flat (Fig. 1C) and the lateral faces are flat or slightly concave. The apical hilum is shown in Fig. 1D. An enlargement of part of an areole (Fig. IF) has a verrucose testa with low areole walls. Two Linnaean species, the Asian O. herbacea and the African O. umbellata (Fig. 2) show trigonous seeds with concave lateral faces lacking the verrucose surface; instead with a coarsely papillose (Fig. 2C) and an apparently smooth surface (Fig. 2D). Oldenlandia uniflora has trigonous seeds (Fig. 3) with a concave lateral face (Fig. 3B,C), a densely papillose surface and low indistinct areole walls. For td TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 309 Fic. 1. Seeds of Oldenlandia speci ined by SEM. A—D. O/deniandia corymbosa, A. Proctor 10493 (US), Jamaica; B, D. Lelong 6891.2 (NCU), Florida; C. Standley 52679 (US), Honduras. E-F. Olden/andia lancifolia, Vasquez et al. V-1324 (XAL), Mexico. A. Views of 5 seeds; B, E. End views; C. Tilted showing basal and lateral surfaces; D. Hilum at apex; F. Areoles enlarged. 310 BRIT.ORG/SIDA 22(1) Fic. 2. Seeds of O/denlandia species examined by SEM. A, C. Oldenlandia herbacea, Fosberg 40749 (US), Nigeria. B, D. Oldentandia umbellata, Fosberg 51929 (US), Ceylon. A—B. End views; C—D. Areoles enlarged. comparison with seeds of subgenus Oldenlandia two trigonous seeds are shown (Fig. 3D,E) from the Asian species, O. lapeyrousii, belonging to the subgenus Exallage lerrell & Robinson 2003). Oldenlandia species not in subgenus Oldenlandia Table 2 summarizes the more important characters of the remaining species as well as those of subgenus Oldenlandia. The remaining species are not readily assignable to new subgenera until more data are known about other genera. Oldenlandia pringlei, is restricted to San Luis Potosi, Mexico. Its more sig- nificant characters are as follows: Plants small, herbaceous, rhizomatous, as- cending or prostrate; leaves linear; flowers in open, few-flowered cymes, heterostylous; corollas 4+.0-8.5 mm long, subsalverform in contrast with the small, rotate or tubular corollas present in subg. Oldenlandia; anthers 0.8-1.3 mm long; seeds 30-50 per capsule, subglobose, ellipsoid, or ovoid instead of TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 311 Fic. 3. Seeds of Oldenlandi ined by SEM. A—C. Oldenlandia uniflora, Leonard 7450 (FSU), South Carolina. D- E. Ofdenlandia lapeyrousii, Smith 9079 (US), Fiji. A, D, E, end views; B-C. Two views of enlarged areoles. trigonous, and areole wall rather thick and sinuous, chromosome number not known. Ventral and dorsal views of the ellipsoid seeds with their sinuous thick- walled areoles are shown in Figure 4 A,B. The ventral view shows the centric punctate hilum. Oldenlandia salzmannii is a South American species established in north- western Florida and adjacent Alabama. The plants are herbaceous, creeping perennials, whereas other Oldenlandia species may be prostrate but rarely or not creeping; leaves small, 1.5-5.2 mm long; stipules small, to 0.5 mm long; flow- ers solitary, pedicellate; corollas subsalverform; capsules small, 1.5 < 1.5 mm; seeds only 4-14 e, 0.3-0.5 mm long, trigonous, areole walls distinct; peer — per capsu 312 BRIT.ORG/SIDA 22(1) Fic. 4. Seeds of Oldenlandia species examined by SEM. A-B. Oldenlandia pringlei, Pringle 3758 (US), Mexico. C-F. Oldenlandia salzmannii, Burkhalter & Hand 6537 (UWFB), Florida. A. ventral view; B. dorsal view; C. side view; D. hilar area; E. basal surface; F. areoles. TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 313 chromosome number n = 15; 2n = 30, a unique number in Hedyotideae (Lewis 1966b). Figure 4 C-E shows a trigonous seed with concave lateral faces, an api- cal hilum, and polygonal areoles with straight walls. The Oldenlandia microtheca group includes three Mexican species, O. microtheca, O. ovata, and O. drymarioides. Their relationships to other species or groups are somewhat in question, but they have been named Oldenlandia and appear closer to that genus than to any other. Oldenlandia drymarioides is a rarely collected species with a close resemblance to O. ovata. It is tentatively considered a distinct species. The characteristics (Table 2) of the three species may be summarized as follows: Small herbs, annual or perennial; flowers in cymes, heterostylous; co- rollas 2.5-7.0 mm long, funnelform; capsules 1-3 mm long; seeds 10-34 per cap- sule, 0.4-0.7 mm long, angulate or irregularly conoidal, surface irregularly hon- eycombed or alveolate, areoles lacking, coalesced, replaced by entangled vermiform strands; chromosome number x = 11, 12. These species are notable for their unusual seeds (Fig. 5), which differ conspicuously from those of all other Oldenlandia species. The seeds lack the usual reticulate testa and instead have a jumbled mass of entangled vermiform strands, a condition here referred to as coalescent areoles in reference to their running together (Fig. 5 C.F). These seeds are larger and fewer per capsule as opposed to subgenus Oldenlandia and show a range of shapes from irregularly and obtusely angulate to irregularly conoidal. Coalescent areoles and a chromosome number of x= 11 are known in Houstonia subgenus Chamisme section Ericotis of southwestern U. S. and Mexico (Terrell 1996), but those species are otherwise quite distinct from the O. microtheca group. _ TAXONOMIC TREATMENT OF NORTH AMERICAN SPECIES Oldenlandia L., Sp. Pl. 119.1753. Gerontogea Cham. & Schlecht. Linnaea 4:154.1829. TyPE: O. corymbosa L., designated by Hitchcock and Green (1929). Hedyotis ae bosa (L.) Lam., Tabl. Encycl. 1:272. 1792. Lectotype: Plumier, Nov. Pl. Amer. t.36. 1703, cited by Jarvis et al. (1993) and Verdcourt (1976). Four other generic synonyms were listed by eae (1918) KEY TO UNITED STATES SPECIES OF OLDENLANDIA, NATIVE OR ADVENTIVE 1. Creeping, mat-forming perennials; leaves 1.5-5.2 mm long; adventive in northwest Florida and adjacent Alabama 6. O. salzmannii Erect, spreading, decumbent, or prostrate annuals or perennials; leaves (3—)5— mm long; native or widely established species. 2. Flowers and capsules on pedicels more than 3 mm long, not in glomerules 1.0.corymbosa 2. Flowers and capsules in axillary or terminal glomerules, sessile or pedicels less = 3. Annual:leaves 2-11 mm wide, ovate to elliptic 3.0. uniflora 3. Perennial with woody tap root; leaves1—3(-5) mm wide, linear to narrowly elliptic 4.0. boscii 314 BRIT.ORG/SIDA 22(1) Fic. 5. Seeds of Oldenlandia ae) Semines by oe A- 2 eae peli Mg B. Schiede 390 (K), Mexico; C. Pennell 17920 (US), Mexico. D—F | Mexico. A—B. side views; D-E. ventral views including hilar areas; C—F. pen of coalesced areoles. TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 315 KEY TO MEXICAN SPECIES OF OLDENLANDIA, NATIVE OR ADVENTIVE 1. Corollas 0.7-2.5 mm long, rotate or tubular; anthers 0.2-0.6 mm long;seeds 0.1—0.4 mm long, trigonous. 2. Plants often 10-24 cm tall; inflorescence with 1-6 pedicels per node, pedicels usually 2-13 mm long; anthers 0.2-0.3 mm long; capsules 1-2.2 X mm locules not saccate, apices truncate or retuse 1.0. corymbosa 2. Plants often 15-60 cm tall; inflorescence with 1(-3) pedicels per node, pedicels (-30) mm long ; anthers 0.4-0.6 mm long; capsules 2-3.5 X 2.5-3.5(-4 mm, locules appearing inflated or saccate, apices rounded or beaked 2.0. lancifolia . Corollas 2.5-8.5 mm long, subsalverform or funnelform; anthers 0.4-1.3 mm long seeds 0.2-0.7 mm long, subglobose, ovoid, or irregularly angulate. 3. Plants often with rhizomes; leaves 0.3-2 mm wide; seeds subglobose, ellipsoid, or ovoid, testa reticulate, areoles polygonal; endemic to San Luis Potosi, Mexico 5.0. pringlei 3, Plants without rhizomes; leaves 1-20 mm wide; seeds angulate or irregularly conoidal, testa not reticulate, areoles lacking, coalescent, replaced by entangled — trands., 4, Perennials; plants 9-42 cm tall; leaves 2-20 mm wide; flowers heterostylous 7,0. microtheca 4. Annuals, plants less than 20 cm tall; leaves 1-9.5 mm wide; flowers isostylous. 5. Basal leaves absent or smaller than cauline 8.0. ovata 5. Basal leaves larger than cauline 9.0. drymarioides 1. Oldenlandia corymbosa |. Sp.P1.119.1753. Hedyotis corymbosa (L.) Lam, Tab. Eneycl. 72.1792. TyPE: Plumier, Nov. PI. Amer. Gen. 42, t.36.1703. (Verdcourt 1976; Jarvis et al. 1993). Small annual herb. Stems 10-24(-40) cm tall, slender, tetragonal, erect, spread- ing, decumbent, or prostrate, glabrous or puberulent toward base or at nodes, with l-many branches. Leaves (5-)10-40 x 1-7(-9) mm, sessile or short-peti- olate, l-nerved, narrowly elliptic, elliptic, narrowly oblong, or linear, glabrous, margins glabrous or ciliolate near leaf bases, apices acute or apiculate. Stipules to 2 mm long, to 3 mm wide, whitish, truncate or rounded, with 0-few mar- ginal teeth or setae to 3mm long. Inflorescence cymose, flowers isostylous, pe- duncles absent or present, similar to pedicels, usually 5-15 mm long, filiform, pedicels from most nodes, 1-6 per node, usually 2-13 mm long, filiform. Hy- panthium glabrous or scabrous; calyx lobes 0.5-1.3 mm long, slightly exceed- ing capsule, lanceolate, sometimes ciliolate on margins and sinuses, acute. Co- rollas 1-2 mm long, rotate or tubular, white or occasionally faint lavender or pink, partly obscured by calyx lobes; tubes 0.5-1 mm long, throat with ring of white, sometimes clavate, hairs; lobes 0.5-1 mm long, ovate, spreading. Anthers 0.2-0.3 mm long, ovate, whitish or purplish, on filaments 0.1-0.3 mm long, at- tached to sinuses of corolla tube. Stigma lobes 0.3-1.0 mm long, style thickened. Capsules 1-2.2 x 1.3-2.8 mm, subglobose or slightly wider than long, not or slightly compressed, glabrous, 4/5 to fully inferior, thin-walled, fragile, apices truncate or retuse. Seeds 50-100 or more per capsule, 0.2-0.4 mm long, usually brown, trigonous, hilum apical, punctiform, testa reticulate, areoles polygonal, 316 BRIT.ORG/SIDA 22(1) their walls low, rounded, indistinct, walls and testa surface densely covered with minute papillae. Flowering all year in tropical climates. Chromosome number: n= 9,18, 27: 2n = 18, 36, 54 (Lewis 1964, 1965, 1966c). Distribution and habitats—Pantropic weed. United States: adventive and becoming more common in Atlantic and Gulf Coastal Plains and Mississippi Embayment. North Carolina, South Carolina, Georgia, throughout Florida, southern parts of Alabama, Mississippi, and Louisiana, to eastern Texas. Mexico: Nayarit, Tabasco, Chiapas (probably more frequent in Mexico than indicated by available records). Standley (1918) did not give any records for Mexico and United States, listing only West Indies, Central America, and South America in the Western Hemisphere. Hawaii: Oahu, Hawaii, Maui. Eastern Hemisphere. Disturbed places, lawns, roadsides. Discussion.The description refers to the diploid race of var. corymbosa. Further comments on varieties in Lewis 1965, 1966c, Verdcourt 1976; Sivarajan and Biju 1990. Specimens cited below are considered to be var. corymbosa. . collections. U.S.A. FLORIDA. Alachua:, lawn, McCarty Hall, ie! of Florida, Gainesville, D’Arcy 2160 (FLAS). Collier: | mi S of Naples, ne et al. 9071 (USF). Dade: Homestead, Radford & Leonar fei ). Escambia: West Brainerd St, near Pensacola, Burk aa “LAS). Glades: 3 mi SW of Palmdale, Ward 5185 (FLAS). Hillsborough: oF of International Airport, Tampa, Lakela 24642 ee USF). Manatee: Bradenton, Cuthbert 1442 (FLAS). Orange: Orlando, Schallert 1590 (FLAS). Palm Beach: 836 Biscayne Drive, West Palm Beach, Cassen 559 (FLAS). Pinellas: Gulf- port, Thorne 13874 (US). Sarasota: opposite entrance to Pine Park west of US 441, Ward & Burch 3110 (FLAS). LOUISIANA. Orleans: Tulane University campus, Sundell 2028 (NO). SOUTH CAROLINA. Florence: Wynn Owens a Evergreen, Swails 83/971 (USCH). Richland: 6511 Helena Road, Colum- bia, Nelson 402 (FLAS) 2. Oldenlandia lancifolia (Schumach.) DC., Prodr. 4:425.1830. Hedyotis lancifolia Schumach. [in Schumach. & Thonn], Beskr. Guin. Pl. 72.1827. Type: GHANA: Valley of Aquapim, Thonning 210 (LECTOTYPE: C; ISOLECTOTYPE: S, n.v.). Hedyotis commutata Schult. & Schult. f., Mant.3:134.1827. TyPE: PUERTO RICO: Bertero s.n., nv. Perennial herb (rarely annual). Stems 15-60 cm tall, slender, sometimes flat- tened after drying, erect, decumbent or sprawling on other vegetation, often rooting at base, glabrous to densely puberulent. Leaves 10-60 x 2-12 mm, sessile or subsessile, l-nerved, linear, lanceolate, or elliptic, glabrous, margin revolute, apices acute. Stipules to 1.5 mm long, to 2.5 mm wide, whitish, truncate or rounded, with 1-few marginal teeth or setae to 3-4 mm long. Inflorescence cy- mose, flowers isostylous, pedicels from most nodes, 1-2(-5) per node, 5-15(-30) mm long. Hypanthium glabrous or pubescent; calyx lobes 0.7-L6 mm long, exceeding capsules, lanceolate or deltate, scaberulous to puberulent, acute. Co- rollas 1.0-2.5 mm long, rotate or tubular, white; tube 0.5-1 mm long; lobes 0.5- 1.5 mm long, usually slightly longer than tube, ovate, spreading. Anthers 0.4- 0.6 mm long, oblong, on filaments 0.2-0.5 mm long exserted ca. 0.5mm beyond throat, attached at sinuses of corolla. Stigmas lobes 0.5-1.2 mm long, linear, TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 317 exserted ca 1 mm beyond throat, styles 0.5-1 mm long, filiform. Capsules 1.6-3.5 x 2.5-4 mm, subglobose or slightly wider than long, slightly compressed, gla- brous, 3/4 to 9/10 inferior, thin-walled, fragile, apices rounded or somewhat beaked especially after dehiscence, locules appearing saccate. Seeds numerous per capsule, 0.25-0.35 mm long, trigonous, hilum apical, punctiform, testa re- ticulate, areoles polygonal, their walls low, rounded, indistinct, walls and testa surface densely covered with minute papillae. Chromosome number: n= 18, 2n = 36 (Lewis 1965). Distribution and habitat.—Tropical Africa; introduced or adventive in South America, Central America, West Indies, and Mexico. Recorded from Mexico in Tamaulipas, Veracruz, Oaxaca, Chiapas, and Tabasco. Low altitudes in moist places along streams, swamps, forest openings. Discussion.—Verdcourt (1976) provided data on African varieties of O. lancifolia. Many American collections of this species have been misidentified as O. herbacea (L.) Roxb, an Asian species; see key below. The occurrence of any O. herbacea collections in Mexico has not been substantiated by us. KEY TO OLDENLANDIA LANCIFOLIA AND O. HERBACEA 1. Stems weak, often reclining on other eed sometimes flattened in drying; leaves 2-12 mm wide; corollas 1.0-2.5 mm long; capsules 2.5-4 mm wide, subglobose or wider than long, locules a capsules often appearing inflated or saccate O. lancifolia ee erect, solid, tetragonal; leaves 1-4 mm wide; corollas 2.5-4.7 mm long; cap- sules 1.5-2.5 mm wide, subglobose or longer than wide, locules not appearing ee or saccate O. herbacea Both Hedyotis commutata and H. lancifolia were created in 1827, and apparently have equal priority. The epithet lancifolia has been the choice of Verdcourt (1976), Howard (1989), and others in floras. Representative specimens. COSTA RICA. Cartago: Rio Reventazon, Catie Agric. Station, Turrialba, Hillet al. 17735 (HILL). San Jose: Vic. ne General, Skutch 3932 (GH, MO, US). Limon: Vic. of Guapiles, iid 37296 (US), GUATEMALA. Izab c. of Quirigua, penta 24267 (GH, US). HONDURAS. mayagua: Vic. of tae ai sane a ons US). MARTINIQUE. 1 1/2 km S of Ajoupa-Bouil- ns Kimber 687 (WIS). MEXICO s: Mpio. of see i Agua, W of ee on road to Villahermosa, Breedlove e ute 5228 (CAS, MICH, TEX). Tamaulipas: 8 mi from Tampico on Mante highway, Jol tham 4077 (MEXU, MICH, TEX). Veracruz: Rio Sea aorillas del Campamento nos Cedillo, Haalgeetlen Brigada Vasquez 172 (XAL). NICUARAGUA. Zelaya: Pu- erto Isabel, Narvaez S. 2890 (GH). PANAMA. Bocas del Toro, Changuinola to 5 mi S$ at jet. of Rios Changuinola and Terebe, Lewis et al. 796 (GH). 3. Oldenlandia uniflora es SP. Bie 91753: aaa unilona (L.) Lam.,Tab.Encycl. 272. Ed i ) Raf., Actes Soc. Lin 9.1834. TYPE: VIRGINIA: Clayton 587 (LINN -155. 3) ISOTYPE: BM!). Hedyotis auricularia Walter, Fl. Carol.85.1788. (non H. auricularia L., Sp. PL. 101.1753). TYPE: n.v. Oldenlandia glomerata Michx., Fl. Bor.-Amer. 1:83. 1803. Hedyotis glomerata (Michx.) Elliott, 318 BRIT.ORG/SIDA 22(1) Sketch Bot. S. Carolina 1:188. 1816. Stelmotis glomerata (Michx.) Raf., New FI. 4:10L. 1830. Stelmanisglomerata (Michx.) Raf., Autik. Bot. 13.1840. Type: “in humidis Carolinae inferioris,” nv, Hedyotis virginica Spreng., Pl. Min. Cogn. Pug. 2:34. 1815. TYPE: n Hedyotis fasciculata Bertol., Mem.Reale Accad. Sci. Ist. Bologna 2: oi 1850.0. fasciculata (Bertol.) small, FIS.E.U.S. 1106. 1903. H. uniflora var fasciculata (Bertol.) WH. Lewis, Amer. J. Bot. +9:865. 1962. TYPE: nv. Oldenlandia aa C. Mohr, Bull. Torrey Bot. Club 24:27. 1897. Type: ALABAMA: Mobile, low wet places, borders of ponds, Grand Bay [near Mobile], Sep 6, 1888, C. Mohr s.n. (LECTOTYPE, here designated: UN Small annual herb. Stems 2-70 cm tall, slender, erect, spreading, decumbent, or prostrate, eee white-hirsute or villous to glabrous, much branched. Leaves 3-20(-28) x 2-11 mm, sessile or short-petiolate (to 2 mm), ovate or elliptic, hir- sutulous or pubescent to glabrous above or pubescent only on midribs and nerves, glabrous or pubescent on nerves beneath, margins ciliolate to glabrous, apices acute or obtuse. Stipules to + mm long, to 3mm wide, whitish, divided near base into l-few (sometimes branched) ciliate, linear or lanceolate, curved or straight teeth or setae toca 5mm long. Inflorescence with |-many isostylous flowers in axillary and terminal glomerules, sessile or on pedicels to ca 3mm long, sometimes flowering when plants are very small. Hypanthium hirsute to glabrous; calyx lobes 0.8-2.5(-4) x 0.5-2 mm, strongly one-nerved near base, ovate or ovate-lanceolate, pubescent and ciliate to glabrous. Corollas 0.8-1.3 mm long, rotate, white or pale blue; tube 0.1-0.3 mm long; lobes 0.7-L.2 x 05-11 mm, usually shorter than the calyx lobes, ovate. Anthers 0.1-0.3 mm long, ovate, purple, on filaments 0.2-0.4 mm long attached to corolla tube at sinuses. Style and stigma 0.2-0.6 mim long, stigma lobes included in tube, style thickened. Capsules 1-2.5 x 1-3 mm, subglobose or slightly wider than long, densely hir- sute with whitish hairs or pubescent to glabrous, 7/8 or fully inferior, walls medium thick, apices truncate, dehiscence loculicidal. Seeds numerous per cap- sule, 0.2-0.3 mm long, black, brown, or tan, trigonous, hilum apical, punctiform, testa reticulate, areoles polygonal, their walls low, rounded, indistinct, walls and testa surface densely covered with papillae. Flowering in United States in spring (Florida), summer, fall. Chromosome number: n = 18, 36; 2n = 36 (Lewis 1962). Distribution and habitats—United States: Mainly in Atlantic and Gulf Coastal Plains and Mississippi Embayment. New York (Long Island) south to southern Florida, west to eastern Oklahoma, eastern Texas, southern Arkansas, southeastern Missouri, western Kentucky, and western Tennessee. West Indies in Cuba, Puerto Rico, and Jamaica. Not recorded from Mexico. Wet or moist places, lake shores, swamps, stream banks and sand bars, pine and deciduous woods (often in openings), savannahs, fields, roadsides, gravel pits. Discussion.—Variation in leaf shape and plant vesture does not appear to be taxonomically significant. This species produces flowers at an early age: one collection from Virginia was flowering when plants were 4 cm tall. TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 319 apenas collections. CUBA. Isle of Pines: San Pedro and vicinity, Britton & Wilson 14703 (CM). DA. Glades: 8.6 mi SE of Palmdale, Lewis 5682 (TEX). Highlands: 6.7 mi S of US 27 and Florida ae intersection, Ray et al. 10420 (USF). Manatee: 2 1/2 mi N of Duette and road 62, Shuey 2113 (DS). Sarasota: 6 mi SE of Sarasota, Ward & Burch 3196 (MO). Walton: 4 mi S of Freeport, Davis & Davis eee (CM). GEORGIA. McIntosh: NE part of Sapelo Island, Duncan & Adams 17858 (MICH). LO . Beauregard: 1 1/2 mi Sof Bayou Amacoco, Ewan 20089 (NO). East Baton Rouge: Magno- lia, rate ene U). St. Tammany: near jct of hwys. 1-10 and I-59, Darwin & Sundell 918 (NO). MARY- D. Prince Georges: Magruder Landing, Smith 5131 (MT). MISSISSIPPI. Hancock: 0.5 mi W of St. Louis Bay and hwy. 90, Lewis 5688 (TEX). NEW YORK. Long Island: near Long Pond, Wading River, Miller s.n., 8/23/1878 (CM). NORTH CAROLINA. Hoke: 4 mi SSW of Ashley Hts., Ahles 36373 (ASU, CM, MICH). OKLAHOMA. Atoka: 0.5 mi NW of Boehler, Taylor 20365 (KANU). SOUTH CAROLINA. Aiken: Montmorenci, Ahles & hae 55058 (ARIZ, ASU, BALT, CLEM, CM, DS, KANU, KY, LL, MICH, TEX, US, VPI). TEXAS. Cherokee: Larissa, Palmer 8613 (CAS). Nacogdoches: 3.5 mi S of Nacogdoches, Lewis & Oliver 5510 (Ds, TEX). VIRGINIA. Accomac: Parksley, Norton s.n. 9/11/02 (MARY) oO. Bs Z 4. Oldenlandia boscii (DC.) Chapm., Fl. South.U.S.181.1860. Hedyotis boscii DC., Prodr. 4:420. 1830. Tyre: U.S.A: Carolina, 1798-1800, Bosc s.n. (G-DC (n.v.): microfiche US). Small perennial herb with woody taproot to ca 6 mm thick. Stems often nu- merous, 5-30 cm tall, slender, spreading, decumbent or prostrate, forming mats or mounds toca 4+ dm wide, glabrous to minutely papillose or puberulent, much branched. Leaves 10-25(-30) x 1-3(-5) mm, sessile, linear to narrowly elliptic, glabrous to minutely papillose or puberulent above, glabrous or scabrous on midrib beneath, margins glabrous or scabrous, sometimes revolute, apices ob- tuse or acute. Stipules to 2 mm long and wide, whitish, truncate or deltate, with l-several marginal teeth or setae to 2 mm long. Inflorescence with l-several (- many) isostylous flowers in axillary and terminal glomerules, sessile or subsessile. Hypanthium glabrous or minutely papillose; calyx lobes 0.8-2 x 0.3-0.8 mm, strongly l-nerved, deltate to lanceolate. Corollas 0.8-1.3 mm long, rotate, white, pink, or lavender; tube 0.1-0.3 mm long; lobes 0.7-1L.2 mm long, ca. 1-2 mm wide, shorter than the calyx lobes, ovate. Anthers 0.1-0.3 mm long, broadly elliptic, on filaments ca. 0.2 mm long attached to corolla at sinuses. Stigma lobes included in tube, style and stigma less than 0.5 mm long, style thickened. Capsules 1.5-3 x 1.5-2.5mm, subglobose or often slightly longer than wide, verrucose or papillose varying to glabrate, truncate or retuse, 7/8 to fully inferior, walls medium thick, dehiscence loculicidal. Seeds numerous per cap- sule (to at least 120), 0.1-0.3 mm long, black or brown, trigonous, hilum apical, punctiform, testa reticulate, areoles polygonal, their walls low, rounded, indis- tinct, walls and testa surface densely papillose. Flowering April to November. Chromosome number: 2n = 36, based on three collections (Lewis 1962). Distribution and habitats —United States: Mainly in the Atlantic and Gulf Coastal Plains, Mississippi Embayment, and Ozarks in the southeastern states: Southeastern Virginia, North Carolina, South Carolina, Georgia, northern Florida, and west to central and western Tennessee, southeastern Missouri, east- 320 BRIT.ORG/SIDA 22(1) ern Oklahoma, and eastern Texas. Wet or moist places, stream banks, lake shores, roadsides, ditches, fields, woods, savannahs, disturbed open places. Representative collections. U.S.A. ALABAMA. Escambia: Conecuh River bottoms 6 mi E of Boykin on US 29, Kral 44857 (K ANU). ARKANSAS. Faulkner: Conway, Haas 678 (US). FLORIDA. Gadsden: Flood- plain Appalachicola River at Chattahoochee, Godfrey 67526 (FLAS, TEX, USF). Leon: 10 mi S of Talla- gee on Spring Hill Road, Godfrey 62904 (MT, TEX, USF). Suwannee: Live Oak, Curtiss 6649 (ISC, US). GEORGIA. Miller: 7 mi ESE of Colquitt, Thorne 5199 (MT, US). LOUISIANA. Allen: 7.2 mi W of re Shinners 21522 (MICH). Grant: Louisiana road 8 ca 41/2 mi NE of Colfax, Allen et al. 7802 (NO). Sabine: Zwalle, Toledo Bend Reservoir, Sabine River, Demaree 48352 (NO). Washington: Along Pearl River E of Angie, Brown 17866 (LSU). MISSISSIPPI. Jackson: Biloxi, Tracy 6422 (US). Oktibbeha: 10 mi Sof Starkville, McDaniel 2661 (NO). NORTH CAROLINA. Scotland: Sink Hole Bay, SF side of SR 1622, Berg 1156 (NCU). OKLAHOMA. McCurtain: | mi N of Bethel, Waterfall 14776 (DUKE). SOUTH CAROLINA. Aiken: Montmorenci, Ahles & Crutchfield 55054 (ASU, CM, FLAS, KANU, KY, MICH, TEX, USCH, VPI. TEXAS. Bastrop: Bastrop, Tharp s.n., 8/7/38 (CAS, DS, MICH, US). Jefferson: Beau- mont, Palmer 12720 (US). Nucces: Padre Island, Jones 2244 (WWE). VIRGINIA. Southampton: Predler’s Pond, Nottoway Swamp, SW of Sedley, Fernald & Long 7625 (US). Sussex: Airfield Millpond, SW of Wakefield, Fernald & Long i (US). 5. oneal prnsiet BL. HOR Proc. se aca. Arts 27169. eee Hedyotis line eins Halicnde de nes a, 7 Jun 189], CG. eile! 3758 ( (LECTOTYPE, fae ee nated: GH: SOL ECTOTYPES: BMI, Fl, K!, MICH!, MO!, MSC, PHI, US-3), VT! Small perennial herb from slender scaly rhizomes and slender tap root. Stems 5-20 cm tall, slender, subterete, scaly toward base, ascending or prostrate, densely puberulent to glabrate, branched from base or upper nodes. Leaves 3- 15 x 0.3-2 mm, thickened, slightly fleshy, sessile, linear, glabrous or granular- puberulent. Stipules usually 0.3-1.0 mm long, somewhat sheathing, deltate or ovate, with marginal gland-tipped teeth or lobes.Inflorescence in terminal or axillary few-flowered cymes, pedicels to ca. 10 mm long, flowers heterostylous. Hypanthium glabrous; calyx lobes to 2 mm long, 0.2-1.0 mm wide, lanceolate or ovate-lanceolate, obtuse. Corollas 4.0-8.5 mm long, subsalverform or nar- rowly funnelform, white, with dark nerves; tube 2-5 mm long, 0.7-2 mm wide at throat, slightly widened distally, glabrous within proximally, puberulent distally; lobes 1.5-3.5 0.7-1.3 mm, lanceolate-ovate, puberulent within. An- thers 0.8-1.4 mm long, narrowly oblong. Stigma lobes 1-1.3 mm long, subglobose. Pin flowers with stigmas exserted to 2.5 mm beyond throat; anthers included, filaments 0.51.0, attached near midpoint of tube to just below throat. Thrum flowers with anthers sessile or on short (0.5 mm) filaments, exserted at throat, stigmas located in distal part of tube or becoming slightly exserted. Capsules 2-3 x 2-25 mm, usually slightly longer than wide, subglobose or broadly ob- long, slightly to somewhat compressed, thin- or rather thick-walled, fully infe- rior, apex rounded, retuse or truncate, tardily dehiscing loculicidally by a nar- row opening. Seeds ca 30-50 per capsule, (0.2-)0.3-0.4 mm diam., subglobose, ellipsoid, or ovoid, hilum punctiform, centric, testa reticulate, areoles polygonal, TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 321 their walls thick, sinuous, testa surface verrucose. Flowering May to September. Chromosome number not known. Distribution and habitats—Mexico: endemic in Rio Verde area, southern part of San Luis Potosi. Fourteen collections have been seen. Rio Verde is some- times spelled Rioverde. Saline or alkaline flats or plains, sometimes in gypseous soil, associated with Prosopis and Acacia. Discussion.—B.L. Robinson cited Pringle 3758in the protologue, but did not designate the GH specimen as holotype. Robinson worked at the GH and we are designating the GH specimen as lectotype, from among at least nine ieaiccl eee D MEXICO. San Luis Potosi: 5.6 mi E of jct. to El Centro in Rio Verde on hwy 70/86, mpio.Rio Verde, 21°54 N, 99° 53' W, Nesom 6635 (TEX); 0.7 mi N of Las Tablas (RR prrces mpio.Ciudad del Maiz, Nesom 6674 (TEX); Media Luna, near Rio Verde, Palmer 68 (CM, F GH, 2 US); Minas de San Rafael, Guascama, Purpus 5014 (BM, F GH, MO, NY, US); San Bartolo, mpio. Rio erde, Soe ki 5821 (ENCB, MICH); 5 km S of Rio Verde, wae el camino a El Zapote, ae sa2e “B); 9 km E of Rio Verde, carretera a Rayon, Rzed 1 2478, cee RIZ, ENCB, MICH sa ee Las Tablas, Takaki 44 (ENCB); 2 km W of Estacion pene a, 30 km N of Rio Verde, akaki a (ENCB); 9 km SE of Villa Juarez, mpio. Villa Juarez, Takaki oe (ENCB). 6. Oldenlandia salzmannii (DC.) Benth. & Hook. f. ex B.D. Jacks., Index Kew. 1:142. 1893; 2:336. 1893. Anotis salzmannii DC., Prodr. 4:433. 1830. Hedyotis salzmannii (DC.) Steud., Nom. Bot. ed. 2.1:728.1840. TYPE: “circa Bahiam requens,” Salzmann s.n., nv Hedyotis thesiifolia A. St. Hil., Voy. Distr. Diam. 1:397. 1833. Oldenlandia thesiifolia (A. St. Hil.) K. Schum. in Mart., FI. Bras. 6, 6:270. P1127, £.1.1889.TYPE: n.v. Additional synonyms cited by Lewis (1966a:44). Small perennial herb, rooting at nodes and forming mats. Stems creeping, very slender, tetragonal-rounded, glabrous, often much branched. Leaves 1.5-5.2 x 0.7-3 mm, rather thick, sessile or subsessile, broadly elliptic, elliptic, or ovate, glabrous to sparsely hirsute with pointed white hairs to 0.8 mm long, margins glabrous to ciliate especially at or near apex, base rounded, apex rounded. Stipules to ca 0.5 mm long, truncate, with O-few marginal fimbriae to ca 0.7 mm long and O-few minute dark sessile glands. Inflorescence with flowers heterostylous, +-5-merous, solitary on slender pedicels 2-12 mm long. Hy- panthium densely to sparsely hirsute with widely spreading, white, pointed hairs ca’ 0.5-1 mm long; calyx lobes numbering 4-5(-6), 1-22 x 0.7-1.0 mm, lanceolate, ovate, or oblong, glabrous or glabrate, apices obtuse or rounded. Co- rollas 2.5-5 mm long, subsalverform, pink, lavender, light purple, or white, usu- ally with white center; tubes 1-2.2 mm long, ca 2 mm wide at base, broader than long, 2-3 mm wide at throat, tube and throat within densely hirsutulous or pubescent with spreading white hairs about 0.5 mm long; lobes numbering 4(-5), 1-3.2 x 0.9-2 mm, as long as or longer than corolla tubes, ovate, spread- ing 90 degrees and apices recurved, striate-puberulent within with minute white hairs. Anthers numbering 4(-5), 0.5-0.8 x 0.2-0.4 mm, dark blue, oblong 322 BRIT.ORG/SIDA 22(1) or elliptic. Stigma lobes 0.5-L.5 mm long, elliptic or linear, whitish, diverging at maturity. Pin flowers with anthers on filaments 0.3-0.7 mm long, attached in distal 1/3 of corolla tube; stigma lobes exserted 1-2 mm beyond throat, styles ca 2-3mm long; slender, greenish or whitish. Thrum flowers with anthers exserted beyond throat on filaments 0.8-1.1 mm long; stigma lobes included in tube, style ca 1mm long. Capsules ca 1.5 mm long and wide, subglobose, hirsute with stiff white hairs, 7/8 to fully inferior, walls medium thick, dehiscence loculicidal. Seeds 4-14 per capsule, 0.3-0.5 mm long, black, trigonous, hilum apical, punc- tiform, testa reticulate, areoles polygonal or rectangular, areole walls distinct, sometimes double, testa surface apparently featureless. Flowering June to Au- gust in western Florida and adjacent Alabama. Chromosome number.—n = 15, 2n = 30. Lewis (1966b) as Hedyotis salzmannii, based on Lewis 6448 (MO), also in FTG, US! Pond edge in botanical garden, Rio de Janeiro, Guanabara State, Brazil. Lewis stated that meiosis was irregular and about 2/3 of pollen grains were shriveled. Terrell examined the US specimen and concluded that the plant does not closely resemble the plants from Florida. Thus, this chromosome number must be considered as tentative. Distribution and habitats.—South America: Brazil, Argentina, Chile, Uru- guay, Paraguay. United States: Locally adventive in Florida near Pensacola in Escambia and Santa Rosa counties and in adjacent Baldwin Co., Alabama, in roadside ditch, by ponds, and at edge of a marsh. First discovered by J. R. Burkhalter in Escambia Co. (Fosberg & Terrell 1985). Burkhalter sent Florida plants to Terrell, who grew them in a greenhouse for several years. Discussion.—The name, O. salzmannii, is here accepted as correct pending study of the type specimens. In Index Kewensis, vol. 1, B.D. Jackson, editor, listed “Anotis Salzmanni DC,, Le. =Oldenlandia Salzmanni.” In volume 2, p. 336, Jack- son listed “Oldenlandia Salzmanni Benth. & Hook. f. Gen. ii. 58.—Bras.” It was pointed out by Fosberg and Terrell (1985) that Bentham and Hooker did not validly publish the combination Oldenlandia salzmannii because they merely listed it under Oldenlandia, and did not state anything further about their in- tentions or viewpoints. Jackson validated the name Oldenlandia salzmannii by publishing it in volume | of Index Kewensis. We may alternatively cite only B.D. Jackson. The description of the later name Hedyotis thesiifolia A. St. Hilaire 1833) states that the leaves are linear, which casts some doubt on its identity, as the subject plants have ovate leaves. Later, Schumann (1889; see above) published a very good description of H. thesiifolia along with an excellent drawing, citing several synonyms, including Hedyotis salzmanni. Unfortunately, we have to stick with the earliest name, O. salzmannii, the one chosen by Fosberg & Terrell (1985). The taxonomy and nomenclature of this species need further study in- volving the variation in the native South American plants. —_— TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 323 Collections from U.S.A. ALABAMA. Baldwin: Near Orange Beach, just N of W end of Cotton Bayou, E of Ala. 161, Burkhalter 8753 (UWFP). FLORIDA. Escambia: near Pensacola, S of US 98, 0.2 mi W of Fairfield Drive, Burkhalter 5408 (FLAS); Gonzalez, N of Chemstrand Road (FL C-297), E of Judy St. R30W, TIN, Sec. 14, Burkhalter & Hand 6537 (FLAS, FSU, UWFP); SW of Pensacola, E side of Herrion Bayou, N of US 98, Burkhalter 8838 (FLAS, UWFP). 7. Oldenlandia eee ee & Cham.) DC,, Prodr. 4:428. 1830. re microtheca Schltdl. & , Linnaea 5:169. 1830. Hedyots fe ee (Schltdl. @ Cham Steud., Nomencl. Bot. ees TYPE: MEXICO. V Uz: “Barranca de erie in ee ae Sep 1828, Schiede & Deppe 390 one HAL); see further notes on isotypes in text below. Gerontogea deppeana Schltdl. & Cham., Linnaea 5:169. 1830. Oldenlandia deppeana (Schltdl. & Gon ) DC, Prodr. 4:428. 1830. Hedyotis oo (Schltdl. & Cham.) Steud., Nomencl. Bot. ed. 2. 1:727.1840.TYPE: MEXICO. VERACRUZ: San Andres, Aug 1828, Schiede & Deppe 391 (HO- LOTYPE: HAL} IsOTYPES: HAL! lca es Perennial herbs with roots sometimes thick and woody. Stem base of ten woody, to 5mm thick. Stems 9-42 cm tall, slender, erect or decumbent, glabrous, pu- berulent, or densely scabrid, branches to 30 cm long. Leaves subsessile or with partly winged petioles to 5 mm long, blades (6-)10-30(-40) x 2-13(-20) mm, l- nerved, quite variable in size and shape, ovate, lanceolate, or elliptic, varying to linear in Chiapas populations, glabrous to scaberulous or puberulent above, or densely scabrid in Chiapas populations, glabrous beneath, bases cuneate, rounded, or tapering, apices obtuse, acute, or subacuminate. Stipules toca 2 mm long, to3mm wide, scarious, whitish, deltate to lanceolate, glabrous or puberu- lent, with marginal gland-tipped teeth to ca 1 mm long, apices truncate to acuminate or lobed. Inflorescence terminal, in few-flowered cymes, flowers heterostylous, pedicels to 15 mm long, slender or filiform, glabrous or scaberulous. Hypanthium glabrous or with minute hairs on margins of sinuses; calyx lobes 0.5-13 x 0.3-0.5 mm, usually 1/4 as long to subequal to corolla tube, deltate to lanceolate, apices obtuse or acute. Corollas 2.5-7 mm long, fun- nelform, white or lined with purple, glabrous externally; tube 1-3 mm long, somewhat widened distally, 0.5-1.3 mm wide at base, 1-2 mm wide at throat, puberulent within with dark gland-tipped hairs 0.1-0.2 mm long: lobes 1.5-3(- 4) x 0.8-18 mm, ovate or lanceolate, glabrous or puberulent within. Anthers 0.4-1 mm long, 0.2-0.3 mm wide, elliptic or oblong. Stigma lobes to 1.4 mm long , linear, often somewhat coiled at maturity. Pin flowers with corolla lobes 1-2(-3) times longer than corolla tube, anthers located just below corolla throat, sessile, stigma lobes exserted 1-2 mm beyond corolla throat, styles ca 2-3.2 mm long. Thrum flowers with corolla lobes 2/3-1 times as long as corolla tube, an- thers exserted beyond throat on filaments ca 0.3-1 mm long, stigma lobes lo- cated near midpoint of corolla tube. Capsules 1-3 x 1.5-3.5 mm, usually wider than long or subequal, subglobose to obovate, slightly to rather strongly com- pressed, glabrous or glabrate, 3/4-9/10 inferior, thin- or rather thick-walled, 324 BRIT.ORG/SIDA 22(1) apices retuse, truncate, or broadly rounded, dehiscing loculicidally and then septicidally. Seeds 10-32 per capsule, 0.4-0.7 mm long, black or dark brown, irregularly and obtusely angulate or conoidal-angulate, hilum appearing ei- ther apical or centric, punctiform, testa not reticulate, areoles lacking, strongly coalesced, replaced by entangled vermiform strands. Flowering throughout the year. Chromosome number: n = 11 (Lewis in Terrell et al. 1986). Distribution and habitats —Mexico: Chiapas, Hidalgo, Oaxaca, Puebla, San Luis Potosi, Veracruz. Standley (1918) included Yucatan in the range, but I have not seen any specimens from there. Gravelly, rocky or sandy soil, slopes, banks, crevices of cliffs, in open or in pine or deciduous forests, and evergreen cloud forest in Chiapas, alt. 1000-2700 m. Discussion.—This species is closely related to O. ovata and O. drymarioides as shown by its morphology. Chiapas collections by Breedlove and Raven are linear-leaved and scabrid and occur in cloud forests. They were judged to be a possible new variety, but they appear to be otherwise similar to O. microtheca and, lacking more collections, are tentatively included in O. microtheca s.l. The type specimens for O. microtheca have been seen in the Halle (HAL) and other herbaria, but have presented problems in interpretation of label data. In addition to the holotype cited above at HAL, other collections included pos- sible isotypes, as follows: (1) a specimen at K labelled as 390, but lacking a local- ity; (2) a specimen at HAL labelled 390 and “Barranca de Tioselos,” but dated August 1829, not September 1828; (3) a type at B presumably destroyed, shown in photos of “Types of Berlin Herbarium” deposited at F MO, NY, US; this la- belled as “O. micrantha n. sp.” and as Hedyotis micrantha Schlecht. a name un- published in these combinations, (4) a specimen at HAL and MO, supposedly not a type, labelled from “In sylvis Papantlae,” thus agreeing with the citation under O. microtheca in Linnaea 6:414. 1831. The protologues of Gerontogea microtheca and G. deppeana appeared under the numbers 390 and 391 on the same page. Any question of priority seems to have been settled by Standley (1918), who adopted O. microtheca in his monographic treatment of Oldenlandia for the North American Flora. Standley (1918) listed O. latifolia Martens & Galeottii as a synonym of O. microtheca, but these are two separate species that are superficially so much alike, even in floral details, that it is difficult to identify them without the seeds. Like O. ovata and O. drymarioides, the seeds of O. microtheca are irregularly angulate or irregularly conoidal and have coalescent areoles as described here. In Hedyotis latifolia (Martens & Galeotii) Walpers, however, the seeds are flat and reticulate. The latter has a chromosome number of n = 17, compared to n =l1 in O. microtheca. A detailed study of the morphology of all parts of these two species in separate genera indicated that the best distinguishing charac- ters are those in the following key. TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 325 KEY TO OLDENLANDIA MICROTHECA AND HEDYOTIS LATIFOLIA . Seeds 0.4-0.7 mm long, ices arly angulate or conoidal-angulate, testa not reticu- late, areoles appeari sced, replaced by entangled vermiform strands; hilum not or scarcely Poeible corolla tube 1-3 mm long, 1-2 mm wide at throat; corolla lobes in pin flowers 1-2(-3) times longer than tube, lobes in thrum flowers 2/3-1 times as long as tube; anthers in pin flowers attached just below corolla throat Oldenlandia microtheca 1. Seeds 0.5-1 mmlona flat, testa finel | ith | | : tiform, near center of ventral Gee of Cae corolla tube 1 5-5 mm long, 1.8-3 mm wide at throat; corolla lobes in both flower types 1/2-1 times as long as tube; anthers in pin flowers attached near midpoint of corolla tube Hedyotis latifolia Representative collections. MEXICO. Chiapas: Sumidero de Tuxtla, 22 km N of Tuxtla Gutierrez, 00 ft., Raven & Breedlove 20123 (DS, NY); same loc., Breedlove 10650 (DS, NY); Summit of Chuchil Ton, NE of Bochil, mpio. of San Andres Larrainzar, 2700 m, Breedlove 29269 (DS, MICH, MO); 7 km NE of Bochil along road to Simojovel, mpio. of Bochil, Breedlove & Smith 32308 (DS); 11-15 km NW of Soyalo on road to La Bombana and Chicoasen, mpio. of Osumacinta, 1000 m, Breedlove 33756 (DS, MICH, MO); 6-8 km WNW of Soyalo, mpio. of Soyalo, pei 37168 (DS, ENCB, MICH, MO). Hidalgo: Vic. of Molango, road to Lolotla, Distr. Molango, Moore 2399 (GH, MICH). Oaxaca: Mpio. San Pedro Ixcatlan, Cerro Quemado, al W del Poblado de Cerro Quemado, Calzada 10351 (XAL). Puebla: 15 k adelante de Xicotepec de Juarez rumbo a Poza Rica, Chazaro B. 569 (XAL); Near Metlaltoyuca, ean. man 26 (GH, US); near Ocostoc below eee eas ised va TENN, US); Mpio. Cuetzalan, San Miguel Tzinacapan, Turra 1524 (ENCB); El Reparo, Mpio Ico, E ue 414 (CAS, CHAPA, ENCB, US); Carretera aoe San Andres, een Cuetzalan, Zola B. 253 (XAL). San Luis Potosi: Las Canoas, Pennell 17920 (F GH, MEXU, MICH, NY, PH, US); Tamasopo Canyon, Pringle 3510 (FE GH, VT). Veracruz: Rancho La es a5 km al N de Banderilla, mpio. Banderilla, Calzada 4327 (XAL): 1 km al E de San Pablo, mpio. de Papantla, Gutierrez R. 26 (ENCB); El Esquilon, mpio. Jilotepec, Ortega O. 75 (XAL); Monte Rey, Ejido Coetzalan, Mpio. Axocuapan, Robles H. 171 (ENCB, XAL); Below Txolo Falls near Jalapa, Sharp 45802 (GH, MEXU, TENN, US); 6 km SW of See mpio. Acatlan, road to Misantla, Terrell & Koch 5379 (CAS); Encinos, mpio. de Totutla, F Ventur 7627 (CAS, ENCB); Coacoazintla, mpio. Coacoazintla, F Ventura A. 8292 (CAS, CHAPA, co Filipinas, mpio. de Tlapacoyan, E Ventura A. 12316 (CAS, CHAPA, ENCB). 8. Oldenlandia ovata S. Watson, Proc. Amer. Acad. Arts 18:97. 1883. Hedyotis watsonii WH. Lewis, Rhodora 63:222. 1961 (non H. ovata Thunb. ex Maxim., Bull. Acad. Imp. Sci. Saint- Petersbourg 29: 161. 1883). TYPE: MEXICO. NUEVO LEON: Guajuco, Mar 1880, E. Palmer 399 (LEC- TOTYPE, here designated: GH; ISOLECTOTYPES: FI, K!, NA!, US!, VT!); LECTOTYPE mislabeled later as O. microtheca. PARATYPES: Ervend berg 200 (GH! atypical or not O. ovata); Parry and Palmer 676 1/2 (GH); these in GH are on the same sheet as the lectotype. Small annual herb. Stems to ca. 20 cm tall, slender, weak, ascending, decum- bent, or diffusely spreading, glabrous or puberulent. Leaves with petioles to ca 5mm long, blades 3-14 x 1-9.5 mm, thin, l-nerved, ovate, broadly ovate, or up- per leaves elliptic, glabrous to sparsely pubescent above, glabrous beneath, bases rounded, subcordate, or subtruncate or upper leaves tapering to base, apices obtuse or acute. Stipules ca. 0.5 mm long, truncate, with gland-tipped teeth. Inflorescence terminal and axillary in groups of 1-3 flowers, cymose, pedicels filiform, 2-16 mm long, flowers isostylous. Hypanthium glabrous; calyx lobes 0.4-1.2 mm long, deltate to lanceolate, obtuse or acutish. Corollas 2-4 mm long, 326 BRIT.ORG/SIDA 22(1) short-funnelform, white, glabrous or puberulent within; tubes 1-2.2 mm long, scarcely wider distally; lobes 1-1.5 mm long, ovate. Anthers 0.4-0.7 mm long, sessile, elliptic or narrowly elliptic, attached at corolla sinuses, slightly exserted. Stigma lobes 1-1.2 mm long, linear, slightly exserted at throat, not exceeding corolla lobes, style 1.3-1.7 mm long. Capsules 1.3-2.2 x 2-3 mm, subglobose or broadly oblong, slightly wider than long, usually widest at apex, glabrous, thin- walled, fragile, 3/4-4/5 inferior, subtruncate at apex, dehiscing loculicidally and then septicidally. Seeds ca. 17-34 per capsule, 0.4-0.7 mm long, black or brown, irregularly and obtusely angulate or conoidal-angulate, hilum scarcely visible, apical, punctiform, surface often warty and bumpy, testa not reticulate, areoles lacking, strongly coalesced, replaced by entangled vermiform strands. Flowering March to December. Chromosome number: 2n = 12 (Lewis 1962 as Hedyotis watsonii). Distribution and habitats.-MEXICO: Central and southern Nuevo Leon and Tamaulipas south to eastern San Luis Potosi, northern Hidalgo, and Ver- acruz (one collection). Moist shaded rocks and outcrops along streams, rock crev- ices, and on canyon walls, alt. 100-1500 m, 300-5000 feet. Discussion.—Additional nomenclatural data are provided as follows. The lectotype is labelled “March 1880,” but isolectotype F is labelled “March 1-8, 1880” and states that Guajuco is 27 miles southeast of Monterrey; K and US are labelled “February to October 1880.” A paratype listed above is L.C. Ervend berg 200 (GH), Wartenberg, near Tantoyuca, prov. Huasteca, 1858, en route from San Luis Potosi to Tampico, December 1878 to February 1879. Ervend berg 200 is ei- ther atypical O. ovata or not O. ovata; it is difficult to know which because of the condition of the specimen. eae: collections. MEXICO. Hidalgo: Near Chapulhuacan, Sharp 441822 (GH, MEXU, TENN, US). Nuevo Leon: Canyon Denuncio, Rancho La Bolla, Mpio. de ie a Santi: Haas Muel ler 2020 (F GH, MICH); Sierra Madre Orientale 21 mi W of Linares, Ripley & Barneby 13559 (NY); 0.5 mi SE of Canoas, 5.5 mi SE of Allende, E. & B. Terrell 4426 (CAS): Trail below base of ey ig ee Monterrey, E. & B. Terrell 4430 (CAS); Hacienda Vista Hermosa, 35 mi S of Monterrey, White 1555 (MICH): Queretaro: 4 km SW of Ahuacatlan, mpio. de Pinal de Amoles, Rzedowski 27744 (ENCB); San Luis Potosi: Roeaune it Fisher 45180 (BRIT, US). Tamaulipas: 22 mi SW of Victoria, Crutchfield & Johnston B( TEX); 3 mi us of Villagran on Victoria-Linares hwy., yeh nston & Graham 4672B (TEX); Mpio. Sener 43m of Adelaida-Guayabas jet., Nesom et al. 6351 (TEX); Cerro del Picacho, mpio. de Villa de Casas, ae 4098 (ENCB); ee ies Sabinas, opposite i 619 on Pan American hwy. N of El Limon, Sharp & Hernandez X. 50393 9. Oldenlandia drymarioides (Standl.) Terrell, Phytologia 59:80.1985. Houstonia drymarioides Standl., J. Wash. Acad. Sci. 18:162.1928. Hedyotis drymarioides (Standl.) WH Lewis, Rhodora 63:221.1961.T ype: MEXICO. TAMAULIPAS: mountains S of Victoria, alt. LOOO m, 9 Apr 1926, Robert Runyon 870 (HOLOTYPE, designated by Standley, US!: ISOTYPE: Fl). PARATYPE: same loc. and date, Robert Runyon & B.C. Tharp 4039 (TEX!, US). Annual herb (protologue states perennial, but plants appear annual). Stems to cal5cm tall, slender, erect or decumbent, glabrous, simple or sparsely branched, TERRELL AND ROBINSON, TAXONOMY OF NORTH AMERICAN OLDENLANDIA 327 with the 3-4 internodes much longer than the leaves. Basal leaves to 12 mm long, to5 mm wide, much larger than cauline leaves, forming a rosette, ovate or oval, rounded at base, with short petioles, glabrous or minutely puberulent: cauline leaves to 6 mm long, to 3 mm wide, few and much reduced, elliptic to narrowly elliptic. Stipules 0.5 mm long, with short marginal teeth. Inflorescence in few-flowered, terminal cymes, flowers apparently isostylous, pedicels to ca 18 mm long, filiform. Hypanthium glabrous; calyx lobes 0.5-1.2 mm long, equal- ling or slightly exceeding mature capsules, deltate. Corollas 2-4 mm long: short- funnelform, apparently white; tubes 1.3-2.3 mm long, scarcely wider distally: lobes ca 1-1.8 mm long, ovate. Anthers ca 0.8 mm long, sessile, oblong, attached at sinuses of corolla lobes, slightly exserted. Stigma lobes not seen Capsules 13-2 x L7-28mm, slightly wider than long, slightly I d walled, fragile, 1/2-3/4 inferior, broadly rounded at apex, dehiscing loculicidally and then septicidally. Seeds 0.3-0.45 mm long, black, obtusely and irregularly angulate or conoidal-angulate, hilum scarcely visible, apical, punc- tiform, testa not reticulate, areoles lacking, strongly coalesced, replaced by en- tangled vermiform strands. Chromosome number not known. Discussion.—This description is shortened due to the few collections. Fur- ther collections are needed to determine whether this species is distinct from O. ovata. glabrous, thin- Recent collection. MEXICO: Tamaulipas: Mpio. San Carlos, Sierra de San Carlos, ca. 5 mi S of San Carlos, N side of Bufa El Diente, igneous bedrock, N-facing steep slope; scattered oaks and shrubby vegetation of Rhus, Croton, Chiococca, Helietta, Vauquilinia, 770 to 1100 m, 24°31.5' N, 98° 57.6’ W. Flowers white, very common on moist shady bank, 17 Jun 1987, Nesom 6049, with Norris, Martinez, Woodruff (MEXU, TEX). ACKNOWLEDGMENTS We thank Scott Whittaker and Susann Braden, SEM Laboratory, Smithsonian Institution, for preparation of illustrations, and Marjorie Knowles for format- ting the illustrations. We express appreciation to the curators of the herbaria cited for many loans of specimens or access to collections. Richard Wunderlin and Bruce Hansen provided facilities at University of South Florida where some work on this manuscript was carried out. Joseph Kirkbride and John Wiersema provided helpful discussions concerning difficult nomenclatural problems. Steven Ginzbarg, Collections Manager, UNA, assisted in selection of a lectotype for Oldenlandia littoralis. REFERENCES Anpersson, L.and J.H.E. Rova. 1999. The rps16 intron and the phylogeny of the Rubioideae (Rubiaceae). Pl. Syst. Evol. 214:161-186. ANDERSSON, L., J.H.E. Rova, and F.G. Atzate.2002.Relationships, circumscription, and biogeog- raphy of Arcytophylium (Rubiaceae) based on evidence from cpDNA. Brittonia 54: 328 BRIT.ORG/SIDA 22(1 = 40-49. Bremekamp, C.E.B. 1952. The African species of Oldenlandia L. sensu Hiern et K. Schumann. Verh. Kon. Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect. 48:1-297. Bremer, B. 1996. Phylogenetic studies within Rubiaceae and relationships to other families based on molecular data. Opera Bot. Belg. 7:33-50. Bremer, B.and J.-F. Manen. 2000. Phylogeny and classification of the subfamily Rubioideae (Rubiaceae). PI. Syst. Evol. 225:43-72. FosperG, F.R.1943. The Polynesian species of Hedyotis 1-102. Fospera, F.R.and E.E. Terrett. 1985. A recently established exotic in west Florida and Alabama (Hedyotis salzmannii or Oldenlandia salzmannii; Rubiaceae). Castanea 50:49-51. Hate,N.1966. No. 12, Rubiacées (1° part), Hedyotidées. In: Aubréville, A., editor. Flore du Gabon. Mus. Hist. Nat., Paris. Pp. 75-124. Hepper, FN. and R.W.J. Keay. 1963. Rubiaceae (part), vol. 2, Oldenlandia. In: FN. Hepper, ed. Flora of West Tropical Africa. 2nd ed. Crown Agents, London. Pp. 210-212. HitcHcock, A.S.and M.L.Gareen. 1929. Standard-species of Linnaean genera of Phanerogamae. Intl. Bot. Congress, Cambridge, 1930. Nomencl. Prop. Brit. Bot. 110-199. London. Howarb, R.A. 1989. Flora of the Lesser Antilles. Vol. 6, Part 3. Arnold Arboretum. Jarvis, C.E., FR. Barrie, D.M. ALLAN, and J.L. Reveat. 1993. A list of Linnaean generic names and their types. Koeltz, KOnigstein Lewis, W.H. 1962. Phylogenetic study of Hedyotis (Rubiaceae) in North America. Amer. J. Bot. 49:855-865. Lewis, W.H. 1964. Oldenlandia corymbosa (Rubiaceae). Grana Palynolog. 5:330-341. Lewis, W.H. 1965. Cytopalynological studies of African Hedyotideae (Rubiaceae). Ann. Mis- souri Bot. Gard. 52:182-211. Lewis, WH. 1966a. The Asian genus Neanotis nomen novum (Anotis) and allied taxa in the Americas (Rubiaceae). Ann. Missouri Bot. Gard. 53:32-46. Lewis, WH. 1966b Chromosome numbers of Phanerogams. 1. Ann. Missouri Bot. Gard. 53:100-103. Lewis, WH. 1966c. Chromosome numbers of Oldenlandia corymbosa (Rubiaceae) from southeastern Asia. Ann. Missouri Bot. Gard. 53:25 7-264. Sivarasan, VV. and S.D. Buu. 1990. Taxonomic and nomenclatural notes on the Hedyotis corymbosa-diffusa complex (Rubiaceae) in India. Taxon 39:665-674. STANDLEY, P. 1918. Rubiaceae, Oldenlandieae, in, North American flora 32, part 1:17-39. Terrett, E.E.1990. Synopsis of Oldenlandia (Rubiaceae) in the United States. Phytologia 68:125-133. TerreLt, E.E. 1996. Revision of Houstonia (Rubiaceae-Hedyotideae). Syst. Bot. Monogr. 48. |- : Terrett, E.E., W.H.Lewis, H-Rosinson, and J.W.Nowicke.1986.Phylogenetic implications of di- verse seed types, chromosome numbers, and pollen morphology in Houstonia (Rubiaceae). Amer. J. Bot. 73:103-115. — Rubiaceae). Bishop Mus. Bull. 174: TERRELL AND ROBINSON, TARAVINUIVIE UE 329 Terrett, E.E. and H. Rosinson. 2003. Survey of Asian and Pacific species of Hedyotis and Oldenlandia (Rubiaceae) with nomenclatural notes on Hedyotis types. Taxon 52: Terrett, E.E.and R.P. Wunber.in. 2002. Seed and fruit characters in selected Spermacoceae and comparison with Hedyotideae (Rubiaceae). Sida 20:549-557. Verdcourt, B.1976. Rubiaceae (Part 1), Oldenlandia.|n:R.M. Polhill, ed. Flora of Tropical East Africa. Crown Agents, London. Pp. 268-315. Verdcourt, B.1989. Rubiaceae, Oldenlandia. In: Flora Zambesiaca, vol. 5, part 1. Flora Zambesiaca Managing Committee, London. Pp. 120-142. BRIT.ORG/SIDA 22(1) Book REVIEW Ricarpo AyerzA Jk.and Wayne Coates. 2005. Chia: Rediscovering a Forgotten Crop of the Aztecs. (ISBN 0-8165-2488-2, pbk.). University of Arizona Press, Tuscon, AZ 85719, US.A. (Orders: http://ww ae ona.edu/). $14.95, 215 pp., b&w enna tables, index, 6" x The plant | Salvi hispanica and Salvidcol } i \} part of the human diet, medicine chest, and culture for 4000 years. Although its’ importance to humans has waned over time, there is currently a renewed interest in chia both as a food and a good source of omega-3 fatty acids. Authors Ayerza Jr and Coates have created an interesting primer on the history and uses of chia, as well as the pabake values of the species. luce the reader to th lithi learl so-American diets and compare oe toa ace western diet. Although these early diets were ‘high in animal products like ours today, theirs relied heavily on lean game animals from the wild; ours consists mostly of grain fed domesticated animals. This comparison of diets and the introduction of omega-3 and omega- 6 fatty acid ratio prepare the reader for a detailed discussion of fats in the human diet. uthors have written the text on fats and oils in the human diet ina very readable way. The information is well referenced with research and related literature. Each type of lipids, cholesterol and fatty acids is discussed. Information is presented on coronary heart disease as well as the detri- mental increase of omega-6 fatty acids and decrease of more healthful omega-3 fatty acids in the western cliet. After this in-depth discussion of how the omega-3/6 fatty acids fit into our diet, the authors delve into chia even further. The text includes a chapter on the history and uses of chia through information collected from codices, written histories and related cultural uses of today. Chia was important in the Aztec empire as a food, beverage, medicine, soup, and religious ceremonies. The S] anish ¢ nquest caused a decline in the use of chia because the invaders banned chia to help deter non-Christian religious ceremonies. After so many « uries, Cl s still gr inM 1 Soutl America, but i wnotas impor rfantac rop acir once was. 12 : is ] ee The remaining chapters of the | vays to determine the different specs botanical description, ecology, and cuevation information on the chia pene Rel ated tO the earliet discussion of human die SL as a good source of omega-3 fatty acids. They also compare it to other omega- ge sources and discuss the potential use of chia in enriching other foods sources such as eggs with omega-3 fatty acids. One very interetne section n included an introduction to the Nahua system of botanical classification. CI a Forgotten Crop of the ii tecsisa poo filled “ ae anaes about chia, its’ s history. uses, and nutritive value. ] ifot mative as a functional food with ee nutrient ae substantial amounts of vita- mins and miner a ow sodium content, a substantial source of omega-3 fatty acids, and are a non- allergenic food with antioxidants and fiber. This book can be recommended to anyone interested in learning more about omega-3 fatty acids, and chia’s nutritive value or history.—Lee Luckeydoo, Her- barium, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. SIDA 22(1): 330. 2006 NEW TAXA AND COMBINATIONS IN CULTIVATED BAMBOOS (POACEAE: BAMBUSOIDEAE) C.M.A. Stapleton Royal Botanic Gardens, Kew Richmond, Surrey UK chris_stapleton@onetel.com ABSTRACT A new species in Fargesia Franch. from China, and a new variety in Drepanostachyum falcatum (Nees) Keng f. from the eastern Himalayas, are described. Foun: new Son ee are Eade in Borinda Stapleton for species from western China, previously irge new com bination is made in Indocalamus Nakai for a Japanese species deeaed: in Bune Nakai RESUMEN te J st * ] om ‘ | 4 h. de China, y una nueva variedad de Drepanostachyum faxed (Nees) Keng f. ‘del Este del Himalaya. Se hacen crane combinaciones nuevas en Borinda Stapleton para especies del Oeste de China, p en el género Fargesia, y se hace una combinacion nueva en Indocalamus Nakai para una especie japonesa descrita en el género Pseudosasa Nakai. Fargesia apicirubens Stapleton, sp. nov. Type: UK: Ness B ic Garden, C.M.A. Stapleton 1060, 11 May 1996 (HOLOTYPE: K!). Fargesia robusta foliis crassis lanceolatis vel vaginorum culmorum glabris differt. Culms 2-4 m, 0.3-1 cm in diam., pendulous; internodes 5-18 cm, smooth, lightly white-powdery initially, becoming glossy dark green with purple spots, often apically purple-red, later orange-yellow, wall to 5 mm thick; nodes with weakly prominent supra-nodal ridge, sheath scar level and tall, mid-culm branches 6-8. Culm sheaths slowly deciduous, basally leathery and smooth, light brown with prominent red-brown ribs, apically asymmetrically rounded with a persistent red band below the blade, much shorter than internodes, glabrous, margins densely yellow-brown-setose; auricles absent; oral setae absent; ligule ca. 1 mm, truncate, initially ciliolate; blade lanceolate, erect or reflexed, glabrous, margins setose. Leaves 3-7 per ultimate branch; sheath thick, glabrous with outer margin densely white-ciliate at first, usually apically red with yellow veins; auricles and oral setae absent ligule ca. 1 mm, truncate, purple, shortly tomentose, external ligule not evident, tomentose; blade lanceolate to fusiform, 7-10 = 0.6-1.5 cm, slightly thickened, glabrous, basally cuneate, apically long-acuminate, secondary veins 4-5-paired, one margin shortly spinescent, the other obscurely serrulate, transverse veins distinct, petiole yellow. Inflorescence racemose, unilateral; spike- lets shortly pedicellate, usually subtended by densely pubescent reduced sheaths. Introduced from Daba Shan, Shaanxi Province, China, and grown under the misapplied name Fargesia dracocephala TP. Yi. SIDA 22(1): 331 — 332. 2006 332 BRIT.ORG/SIDA 22(1) Etymology.—The epithet “apicirubens” refers to the distally red-colored sheaths and culm internodes. sd aaa Oia falcatum var. sengteeanum Stapleton, var. nov. Type: U.S.A. OR- N: Gold Beach, Aug 1996, S.G. Cooper s.n. (HOLOTYPE: K!). =— Drepa ie ee THK Keng f. var. falcato spiculis bifloris vel trifloris, culmis porcatis, foliis angustis similis, sed vaginis culmorum praeter annulum pilosum circumcirca fundum omnino slabris, lemmatibus scabris, paleis vix bifidis, rhachillis minus puberulis, pilis costae foliarum longioribus unifariis dispersis, vaginis foliorum haud cilatis differt. Culms 2-4 m, 4-10 mm in diam, internodes 10-21 cm, uniformly finely porcate, lightly white-waxy at first; nodes substantially raised, with ring of persistent, initially orange-brown, later colorless, 1-3 mm hairs. Culm sheaths abaxially gla- brous except for a dense ring of 1-3 mm red-brown hairs around basal 1-2 mm; margins distally with ca. 1mm white cilia; auricles absent; oral setae absent; ligule long-fimbriate, abaxially basally puberulous and apically glabrous. Leaf sheaths glabrous; external ligule pronounced, thin, with even ca. 0.1 mm cilia; auricles glab ,elevated; oral setae absent; ligule ca. 1mm, rounded to acute, shortly laciniate, shortly ciliate, basally puberulous. Leaf blade to 12 x | cm, basally narrowly cuneate; abaxially glabrous except for sparse ca. | mm hairs on one side of the proximal midrib; adaxially glabrous. Spikelets with (1-)2-3(- 4) florets. Lemma 7-10 mm, deeply furrowed, abaxially distally scabrous, distal margins shortly ciliate. Palea 8-10 mm, abaxially distally shortly scabrous, apex acute or very shortly bifid, shortly ciliate; keels distally scabrous. Rhachilla sec- tions to5 mm, mainly glabrous, apically bearded with ring of ca. 1 mm hairs. Cultivated in U.S.A. and Europe under misapplied names that will be dis- cussed in a further article. Presumably originating from the Himalayas. Etymology.—This taxon is named in honor of Dr. Lee Seng Tee. absent, Selected specimens. UK (cult.): Kew, Temperate House, 1894, Anonymous (K); Kew, Sep 1895, Anony- ous (K); Kew, Temperate House, 5 Jan 1897, Gamble s.n. (x); Kew, oe louse, 14 Sep 1901, oe (K). U.S.A. California: Bonny Doon, Stapleton 1136, 6 Oct 1999 (kK — angustissima (T.P. Yi) aes comb. nov. BasionyM: Fargesia angustissima Yi, J. Bamboo Res. 4(2):21. 198 Borinda contracta (TP. bee saplton comb. nov. BasIoNyM: Fargesia contracta TP. Yi, J. mboo Res., 7(2):60 Borinda nujiangensis ae & C.M. Hui) Stapleton, comb. nov. Basiony™: Fargesia nujiangensis Hsueh @ C.M. Hui, Bull. Bot. Res., Harbin 18:261. 1998. Borinda utilis or Stapleton, comb. nov. BASIONYM: Fargesia utilis T.P. Yi, J. Bamboo Res. 7(2):28 a hamadae ou ) Stapleton, comb. nov. Basionym: Pseudosasa hamadae atus., J. Geobot. 15:86. 196 ACKNOWLEDGMENTS [thank Mary Barkworth (UTC) and Kanchi Gandhi (GH) for promptly review- ing this note. ESTUDIOS EN LAS APOCYNACEAE NEOTROPICALES XXIV: BAHIELLA (APOCYNOIDEAE: ECHITEAE), UN DESAPERCIBIDO NUEVO GENERO ENDEMICO DE BAHIA, BRASIL J.Francisco Morales Instituto Nacional de Biodiversidad (INBio) pto 22-3100 Santo Domingo de Heredia, COSTA RICA RESUMEN BahiellaJ.F Morales (Apocynoideae Echiteae) 6 dé ] lo de Bahi Brasil es descrito, proponiéndose una nueva combinacion, B. Blanchet (A. DC.) J.F Morales y una nueva especie, B. infundibuliflora J.F Morales. Se incluyen descripciones completas, ilustraciones y Specunenes eapouinages Seen ee se realiz6 un analisis cladistico para determinar las la, el cual incluy6 un total de 44 especies (19 géneros) de i tribu Echiteae como Bae interno y 10 especies (7 géneros) de las tribus Apocyneae, Malouetieae y Mess como pen unOss externos. Hees resultados sugieren que Bahiella pertenece a las Echiteae, Ee p g ser determinadas por ahora. ABSTRACT ] - oy R a ee pa a P| 4 Bahiella J.F Morales (Apocynoideae, Echiteae), a new g illustrated. A new Reconstr B. hana: (A. Peel Morals: sane a new species, B. infundibulif d ] Aditionally. a iL 13-14 : ] : ft if J . 1} cg 2 : ] : Ks ft es 44 species a9 genera) as ingroup of the tribe Echiteae, aod 10 species (7 ae of the ribes Apocyneae, Malouetieae y Mesechiteae as outgroup. The results suggests that Bahiella belong to the Echiteae, but their intergeneric relationships can’t be established by now. Desde su gecaion os Pale de eu (1810), la familia Asclepidaceae fue tradici if tedelas A Apocynaceae, pero estudios moleculares en la tltima década (e.g., Judd et al. 1994; Potgieter @ Albert 2001; Sennblad & Bremen 1996) sugirieron que ambas debian tratarse como una sola unidad. En forma reciente, Endress & Bruyns (2000), propusieron un nuevo sistema de clasificacion tratando en forma conjunta los géneros anteriormente divididos en Apocynaceae y Asclepiadaceae, reconociendo un total de cinco subfamilias. De estas, la subfamilia Apocynoideae esta dividida en cinco tribus, Apocyneae, Echiteae, Malouetieae, Mesechiteae y Wrightieae, separadas principalmente por las caracteristicas de la cabeza estigmatica. Sin embargo, la conformacion definitiva de estas tribus atin se encuentra en disputa, dado que analisis moleculares recientes (e.g., Simoes et al. 2004) sugieren que al menos una de ellas no es monofilética (Apocyneae) y mas estudios son necesarios con SIDA 22(1): 333 - 353. 2006 334 BRIT.ORG/SIDA 22(1) el objeto de definir apropiadamente sus limites. Ahora bien, la mayoria de géneros neotropicales de las tribus Echiteae y Mesechiteae fueron monografiados por Robert Woodson (1933, 1935, 1936), siendo amos grupos relativamente faciles de distinguir por la ausencia 0 presencia de coléteres en la superficie adaxial del nervio central, asi como por las caracteristicas de la cabeza estigmatica. En los miembros de Mesechiteae, las laminas foliares tienen coléteres en la superficie adaxial del nervio central, ya sea agrupados en la base o distribuidos en forma irregular a lo largo de su entera longitud, aunque también pueden estar } en las venas secundarias (i.e., Mandevilla) y las cabezas estigmaticas tienen cinco costillas o proyecciones longitudinales, las cuales pueden estar restringidas a la base o distribuidas a lo largo de la cabeza. En contraposicion, los miembros de la Echiteae, tienen laminas foliares sin coléteres en la superficie adaxial de los nervios, asi como cabezas estigmaticas sin costillas longitudinales y con forma de carrete de hilo o mas o menos fusiformes y con un anillo membranoso basal. Dado que las monografias publicadas por Woodson se encuentran en su mayoria desactualizadas, se ha iniciado un proceso paulatino para hacer nuevas revisiones de los géneros de las tribus Echiteae y Mesechiteae que no hayan sido monografiados en los Ultimos anos. Por el momento, esto ha producido la revision total o parcial de varios grupos (e.g., Hansen 1985; Morales 1997 c, 1999 b, 2002 a, 2002 b, 2003; Williams 1998), la descripcion de nuevos géneros (Mo- rales 1999 a; Morales & Williams 2004; Williams 2002) y la publicacion de varias nuevas especies. En el proceso de revision del género Prestonia, se procedié a examinar la mayoria de colecciones tipos disponibles, tanto de especies descritas en ese género, como de aquellas ubicadas en otros diferentes (e.g., Echites), pero que han sido sinonimizadas a través del tiempo en Prestonia. Echites blanchetii fue descrito por Alphonse de Candolle (1844), basadoen una coleccion hecha por Blanchet (3223 a) en 1840 en el estado de Bahia, Brasil. Este nombre fue reducido a la sinonimia de Prestonia coalita (Vell.) Woodson por Woodson (1936), quien probablemente nunca vio el tipo, ya que no cité dicha coleccion entre la lista de especimenes examinados. Luego de concluir la monogratia de Prestonia, era claro que Echites blanchetii fue incorrectamente sinonimizado bajo P. coalita y que ademas no podia ser ubicado incluso en ese género, pues sus corolas carecian de estructuras coronales accesorias (i.e., Co- rona anular alrededor de la fauce 0 corona epiestaminal, compuesta por cinco I6bulos coronales libres en forma opuesta a cada antera), uno de los principales caracteres diagnosticos de Prestonia. Ahora bien, mientras trabajaba en la elaboracion de una nueva monografia para los géneros Mandevilla y Mesechites, noté que varias colecciones provenientes del estado de Bahia en Brasil, identificadas a través de los anos como M. mansoanus (A. DC.) Woodson y morfoldgicamente algo similares a esa especie, tanto en la forma de las hojas, comoen la estructura de la inflorescencia, tenian hojas eglandulares, careciendo MORALES, BAHIELLA F f ECTAN oe de coléteres en la superficie adaxial del nervio central. Aunque la presencia de coléteres en el nervio central es algo comtin para varios géneros de la tribu Mesechiteae, muy raramente algunas especies pueden carecer de coléteres (e.g, Mandevilla angustifolia (Malme) Woodson). Sin embargo, al proceder con el analisis de las flores, pude descubrir que la cabeza estigmatica de este grupo de especimenes tiene forma de carrete de hilo, mas angosta medialmente y con un anillo membranoso basal, sin costillas o crestas basales longitudinales, esto ultimo un caracter diagnéstico que poseen el resto de miembros de la tribu Mesechiteae, a la cual pertenece Mesechites (Simoes et al. 2004). Este material coincide en todos sus caracteres morfol6gicos con el tipo de E. blanchetii, asi como a otras escasas colecciones depositadas en diferentes herbarios europeos. De esta manera, siguiendo a Endress & Bruyns (2000) y tomando el cuenta las caracteristicas de la cabeza estigmatica, est pertenecen a latribu Echiteae, pero no estan relacionados con ninguno de los géneros hasta ahora conocidos dentro de esa tribu. Por lo tanto, un nuevo género se propone a continuacion, incluida una descripcion de sus caracteres morfologicos, con discusidn de sus relaciones intergenéricas, una nueva combinacion y la descripcion de una nueva especie. MORFOLOGIA Habito y tallos Las dos especies de Bahiella son lianas, con los tallos viejos conspicuamente lefosos. En forma general, las ramitas son algo aplanadas bajos los nudos cuando jovenes, volviéndose teretes a subteretes con la edad. En algunos especimenes y en forma mas rara, tallos jovenes levemente angulados son encontrados. La secrecion de las ramitas y otras partes de la planta al cortarse es blanca. En contraposicion con especies de géneros de la tribu Mesechiteae (e.g., Mandevilla, Mesechites), donde los coléteres de los nudos pueden ayudarnos a reconocer ciertos grupos de especies, en Bahiella (asi como en el resto de los géneros de la tribu Echiteae) los coléteres intrapeciolares son diminutos, fusiformes e inconspicuos y no tienen utilidad para distincion especifica a nivel genérico. Hojas Las hojas son siempre pecioladas, no peltadas y eglandulares, sin coléteres en el nervio central adaxialmente, con la lamina conspicuamente coridcea, basicamente glabra y los margenes conspicuamente revolutos marginalmente. En los demas miembros de la tribu Echiteae, es muy raro encontrar especies con los margenes revolutos, estando limitados principalmente a Hylaea y ocasionalmente y en forma muy rara a unas pocas especies del género Prestonia y Echites. En forma general, solo la vena principal esta impresa en ambas su- perficies, siendo las venas secundarias y terciarias usualmente no evidentes y no impresas. 336 BRIT.ORG/SIDA 22(1) Inflorescencias Las inflorescencias son cimosas, axilares, ramificadas, con numerosas flores densamente aglomeradas en las partes distales de la inflorescencia, diminutamente puberulentas a glabrescentes, pedunculadas, con los pedicelos siempre conspicuos y evidentes. Las bracteas florales son siempre escariosas e inconspicuas y las bracteolas son ausentes. La presencia de bracteolas no es algo comun en las Echiteae y esta limitadoa unos pocos géneros (e.g., Macropharynx, Peltastes). Flores El caliz esta compuesto por cinco sépalos escariosos e inconspicuos, con el apice erecto o suberecto (no reflexo), los cuales albergan en la base de la cara adaxial un coléter solitario, irregular y profundamente lacerado apicalmente. En Bahiella blanchetii, los coléteres estan algunas veces profundamente coco por lo que erroneamente pueden ser tomados como una linea de colé como los presentes en el género Peltastes. Sin embargo, el grado de laceracion de los coléteres no parece tener ninguna utilidad taxonomica, dado el alto nivel de variacion intraespecifica presente en varios géneros de las Echiteae. La forma de la corola varia de hipocrateriforma a infundibuliforma, glabra a inconspicuamente puberulenta exteriormente, sin estructuras coronales interiores (en forma de corona anular alrededor de la fauce o de una corona epiestaminal conformada por cinco lobulos coronales libres en forma opuesta a cada antera). El limbo esta conformado por cinco lobulos, usualmente extendidos en la antesis, pero algo reflexos marginalmente, con estivacion dextrosa. Los estambres estan ubicados en la mitad distal del tubo y son totalmente incluidos, con las anteras glabras dorsalmente, fuertemente aglutinadas a la cabeza estigmatica en dos puntos, con sus bases estériles y sagitadas, con cada auricula conspicuamente acuminada El ovario es apocarpico y esta conformado por dos carpelos, glabros y unidos apicalmente al estilo, cada carpelo con numerosos vulos. La cabeza estigmatica tiene forma de carrete de hilo, relativamente fusiforme, con un conspicuo anillo membranoso en la base, siendo mucho mas ancha en su parte basal. El nectario es anular, pero con el borde superior irregularmente pentalobulado. La lobulacion del nectario puede ser muy leve y casi no evidente o estar dividido en lobulos relativamente bien marcados, incluso en el mismo especimen. Este tipo de variacion intraespecifica no es algo raro en las Echiteae, ya que tal y como se ha comentado en forma previa (Morales 2004), el grado de lobulacion del nectario puede ser muy alto es algunos géneros y por lo tanto, es un caracter que debe utilizarse con reserva a nivel de distinci6n interespecifico. Frutos y semillas Los frutos estan compuestos por dos foliculos, relativamente divaricados, continuos y glabros, libres, continuos, secos y levemente lenosos al secar, con MORALES, BAHIELLA 3 337 diminutas costillas 0 protuberancias longitudinales. Foliculos divaricados tampoco son comunes en las Echiteae, estando presentes en unos pocos miembros de unos cuantos géneros (e.g., Echites, Prestonia, Stipecoma). Las semillas son numerosas, secas, glabras, comosas apicalmente y como muchos otros miembros de las Echiteae, son conspicuamente rostradas, con el rostro delgado. MATERIALES Y METODOS Con el fin de determinar las posibles relaciones intergénericas de Bahiella, un total de 44 especies pertenecientes a 19 géneros de la tribu Echiteae (sensu En- dress & Bruyns 2000) con distribucion restringida al Nuevo Mundo, fueron incluidos en un analisis cladistico. Como grupos externos fueron escogidos dos representantes de las tribu Apocyneae (Beaumontia, Odontadenia) y seis de la Mesechiteae (Allomarkgrafia, Tintinnabularia, Mandevilla), basado en estudios moleculares previos (e.g, Sennblad et al. 1998; Sennblad & Bremen 2002) que han sugerido las relaciones cercanas de Echiteae con esas tribus. Finalmente se utilizaron dos representantes de las Malouetieae (Pachypodium) con el objeto de enraizar el arbol. Cincuenta y seis caracteres representado ciento treint ay unestados fueron evaluados para cada uno de las especies Gaile i cae caracteres fueron seleccionados de estudios previos (Endress et al. 1996; Simoes et al. 2004; Will- iams 2002, 2004), mientras que otros son incluidos por primera vez. Los estados de caracteres fueron evaluados en especimenes de herbario, flores preservadas y material fresco cuando estuviera disponible, los que a su vez fueron corroborados en forma paralela con descripciones de literatura (Morales 1997 a, 1997 c, 1997 d, 1999 a, 2002 a, 2002 b, 2005 a, 2005 b; Morales & Williams 2004, Williams 1998, 2002; Woodson 1933, 1936). La explicacion de los caracteres que no hayan sido utilizados anteriormente en analisis cladisticos previos, asi como algunas discrepancias encontradas entre lo citado en la literatura y lo presente en los especimenes son brindados en el apéndice 1. Los caracteres morfologicos evaluados fueron organizados en una matriz (Tabla 1) utilizando Winclada, version 1.00.08 (Nixon 2002). Todos los caracteres fueron tratados con un mismo peso y en forma desordenada. Especies con caracteres multiestados fueron reconocidos como polimorficos. El andlisis heuristico incluy6 una ronda de 500 replicaciones de secuencia y adicion aleatoria, con la opcién “Multi TBR + TBR” seleccionada y con un maximo de 10000 arboles a mantener, salvando un arbol por réplica. El soporte relativo de los nudos del arbol de mayor consenso estricto fue obtenido mediante el procedimiento de “bootstrap” implementado en Winclada, con un numero to- tal de 1000 réplicas, 10 réplicas de busqueda aleatoria para secuencias adicionales, un arbol conservado en cada réplica y con la opcion “TBR” activada, salvando un maximo de 100 arboles en cada r€plica. ] Taa.a 1.Matriz de datos morfoldgicos de 56 caracteres usados en el andlisis cladistico. Especie Pachypodium lamerei Pachypodium geayi Beaumontia grandiflora Odontadenia matogrossana Tintinnabularia gratissima Tintinnabularia mortonii Alllomarkgrafia brenesiana Allomarkgrafia plumeriifolia Mandevilla hirsuta Mandevilla sagittarii Allotoonia turbinata Asketanthera dolichopetala Asketanthera ekmanniana Bahiella blanchetii Bahiella infundibuliflora Laubertia contorta Estados de caracter BEE wn wn Sa cep TS LS ee es ee es eS SS > > se (L)@2 WaIs/9¥O' LINE TaBLa 1.continued Especie Macropharynx renteriae Macropharynx spectabilis Neobracea bahamensis uteum Prestonia falcatosepala Prestonia quinquangularis Prestonia ipomaeifolia Prestonia longifolia Prestonia vana Rhabdadenia biflora Rhabdadenia macrostoma Thoreauea paneroi Estados de caracter 01010 01010 25 30 00001 42110 001 42110 00001 42110 00001 12110 00001 32110 00001 42110 00001 32110 00001 02110 00001 02110 01111 42110 01101 42110 0111] 42110 01111 42110 00001 b2110 00001 12110 00001 12110 Caracteres polimorficos representados por letras de acuerdo a la siguiente disposicién: /3 a=0/1,b=1 ? = datos desconocidos. Vilainvad Sd TVaOW 6£€ 340 BRIT.ORG/SIDA 22 =a 1) RESULTADOS Un total de 45 arboles parsimoniosos de 184 pasos fueron obtenidos por el analisis heuristico, con un indice de consistencia de 0.4 y un indice de retencion de 0.75. El arbol de mayor consenso estricto, asi como los valores de “bootstrap” mayores 0 iguales a 50 son presentados en la Figura 1. En general, al igual que otros analisis cladisticos basados unicamente en datos morfoldgicos que han involucrado miembros de las Echiteae y Mesechiteae (e.g., Simoes et al. 2004; Williams 2004) el soporte de los clados basales es bajo y una gran politomia prevalece; solamente algunos clados y subclados terminales estan soportados con valores de “bootstrap” mayores a 50%. Por lo tanto, la discusi6n subsecuente se orientara a comentar la conformacion de los clados y subclados del arbol de mayor consenso estricto. En primera instancia es interesante la posicién de Neobracea sugerida en este cladograma. Este género ha sido tradicionalmente ubicado dentro de las Echiteae, pero aca aparece ubicado en un subclado relacionado con el grupo externo y no con el interno (las Echiteae). Aunque el propésito de este analisis no es probar la monofilia de las Echiteae, la ubicacion de Neobracea sugiere la necesidad de evaluar la monofilia de la tribu tal y como se encuentra conformada en la actualidad (segtn Endress & Bruyns 2000). Dentro del grupo interno, el primer clado esta conformado por los géneros Stipecoma, Cycladenia, Rhabdadenia, Angadenia y Pentalinon, dentro del cual Cycladenia y Rhabdadenia conforman un subclado y Angadenia y Pentalinon otro subclado. Esta misma relacion entre Angadenia y Pentalinon, habia sido sugerida anteriormente por Williams (2004), quién obtuvo resultados similares en un analisis orientado a determinar la monofilia del género Echites. El siguiente clado esta conformado por 4 subclados que conforman una gran politomia sin resolver. Un total de 4 subclados son reconocidos: el subclado de Fernaldia, el subclado de Asketanthera, el subclado de Echites y el ultimo de ellos conformado por 10 géneros, que a su vez se encuentran agrupados en algunos subclados: Iemnadenia, Allotoonia, el subclado de Bahiella y Hylaea, el subclado de Macropharynx y Peltastes, el subclado de Thoreauea y Thenardia y el subclado de Prestonia y Laubertia. Sin embargo, a pesar de que los resultados sugieren que Bahiella esta relacionada con el resto de las Echiteae, basado en el bajo soporte de los clados basales y la gran politomia sin resolver prevaleciente, no es posible determinar por el momento y de manera concisa los géneros mas cercanamente relacionados con Bahiella. Partiendo de los resultados sugeridos por este analisis, es claro que son necesarios estudios moleculares complementarios para definir apropiadamente las relaciones intergénericas dentro de las Echiteae, asi como para probar la monofilia de la tribu. Analisis filogenéticos basados en caracteres morfoldgicos en otras tribus de las Apocynoideae (e.g., Mesechiteae), han resultado en cladogramas pobremente resueltos, en contraposicion con analisis basados en estudios moleculares (o MORALES, BAHIELLA E stipecoma peltigera mbellatus Echites yucatanensi Temnadenia stenantha a by oe ) i A, if, 77 Allotoonia turbina Allotoonia noodsonan 65 Bahiella blanchetii Bahiella ifnuitor 83 lylaea a Hylaea le, ie Macropha ete 38 Macropharynx Esuebie 85 Peltastes p Itatu. a gf i S Thenardia galeottiana Pestone falcalosepala 90 La nee boissien e ta ee ie so ifol ia Hise sk ue a ae Allomarkarai Bienesiane Tintinnabularia aan 50 Tin tinnabulan ja mo. hireuta Mandevilla eagle SE geayi Pachypodium lamerei Fic. 7. Arbol d ‘oe er eae eee ad | AC deh nl y 4s parsimoni Valores de “bootstrap” mayores 0 igual a 50 son indicados sobre las ramas. combinados), que han propuesto ee con mucho mas soporte (Simoes etal. 2004). En todo caso, los mismos estudios han conllevado a la necesidad de variar la configuracion original propuesta por Endress & Bruyns (2000) para conformar tribus naturales y monofiléticas. Sin embargo aun falta trabajo para determinar la conformacion definitiva de otras (e.g., 342 BRIT.ORG/SIDA 22(1) Apocyneae), dado que se ha determinado que su constituci6n actual no es monofilética. TRATAMIENTO TAXONOMICO Bahiella J.F. Morales, gen. nov. Tipo: Bahiella blanchetii (A. DC.) JF Morales Fruticosa volubilis; folia opposita petiolata, coriacea, eglanduligera, revoluta; inflorescentia corymbosa, lateralis, pedunculata, multiflora; corolla regularis 5-partita, infundibuliformis vel hypocraterimorpha, corona et coronae lobis absens, antheris sagitattis, basi acuminata. Fructus folliculis divaricatus. Lianas con tallos viejos lenosos, con secrecion lechosa al cortarse, las ramitas ee porns teretes a subteretes, algo aplanadas en los nudos, mas rara- 3 lé i iolares inc onspl¢ uos Hojas opuestas, peciola- das, sin soiree enel nervio central fae aieneate sin domacios abaxialmente. Inflorescencias cimosas, axilares, pedunculadas, con muchas flores, densamente aglomeradas distalmente, el raquis usualmente noduloso, bracteas florales escariosas, bracteolas ausentes; sépalos escariosos, con un coléter solitario, episépalo, dispuesto en la base de la cara adaxial, usualmente profunda e irregu- larmente lacerado, a veces casi totalmente dividido en varios segmentos; corola infundibuliforme o hipocrateriforma, glabra, glabrescente 0 inconspicuamente puberulenta exteriormente, sin estructuras coronales interiormente, el limbo actinomorfico, dext luto, estambres incluidos, las anteras con- niventes y aglutinadas en dos puntos a la cabeza estigmatica, las bases estériles conspic das y sagitadas, ovario bicarpelar, los carpelos unidos en forma apical al estilo, los 6vulos numerosos, la cabeza estigmatica fusiforme, con un anillo membranoso basalmente, nectario anular, irregularmente pentalobulado. Frutos apocarpicos, compuestos por dos foliculos, mas o menos divaricados, continuos, glabros, con diminutas costillas longitudinales; semillas numerosas, secas, rostradas, eee comosas apicalmente. Bahiella es un génerod especies, endémico a las formaciones costenas del estado del Bahia, Brasil. Su nombre genérico hace referencia al estado de Bahia, Brasil, la (nica localidad conocida por el momento. CLAVE PARA LAS ESPECIES DE BAHIELLA 1. Corola hipocrateriforma, con la fauce de 2,5-3,5 mm de didmetro; ldbulos de la corola de 7-11 X 4-6 mm; anteras de 5,5-6 mm de largo; sépalos de 1-1,3 X J1-1,2 mm B. blanchetii a DC.) J.F. Morales . Corola infundibuliforme, con la fauce de 9-10 mm de didmetro;lébulos de de 14-16 X 10-13 mm, anteras de 77,5 mm de largo; sépalos de 2,2—3 mm et ae J.F. Morales 1. Bahiella blanchetii (A. DC.) J.F Morales, comb. nov. (Fig. 2). BAsIONIMo: Echites blanchetii A. DC,, Prodr. 8:448. 1844. Mitozus blanchetii (A. DC.) Miers, Apocyn. S. Amer. 219. 1878. Tipo: BRASIL. BAHIA: Bahia, 1840 ({D, Blanchet 3223a (HOLOTIPO: G-DC, foto F neg. 7567, INB). as MORALES, BAHIELLA 343 E > mm lcm ; f \ ene RY \\ Ih 5mm : iis Wi , = YY a Y VY Fic. 2 Bahiella blanchetii (A-G de Santos 2219, CEPEC; H—J de Amorin et al. 1386, CEPEC). A. Ramita con infl Natalla do lal pee dee pee ean [eee pranted} tnt 1.4 Ce | Lig fetal 4 Snalo.mostrando I colét lat D.Tubo del lal g linal biert t Jo | estilo y nectario. E Antera ista dorsal y lateral. F. Cab tica. G. Nectario. H. Foliculos. 1. Detalle de | Je los foli J. Semilla J 344 BRIT.ORG/SIDA 22(1) Liana, los tallos viejos lenosos, ramitas usualmente teretes a subteretes, glabras a glabrescentes, coléteres interpeciolares ca. | mm de largo. Hojas: peciolos de 6-13 mm de largo; laminas de 8-16,5 x 4,5-7,5 cm, coriaceas a subcoridceas, elipticas a obovadas u obovado-elipticas, redondeadas, obtusas a inconspicuamente retusas 0 apiculadas apicalmente, redondeadas, obtusas a raramente cuneadas basalmente, Seen conspicuamente revolutas marginalmente cuando viejas, esencial ,sin coléteres en el nervio central ad sialmente la venacion secundaria apenas evidente o inconspicua en ambas superficies, la venacion terciaria no evidente. Inflorescencias inconspicua y esparcidamente puberulentas a glabrescentes, pedinculo 4-65(-100) mm de largo, el raquis conspicuamente noduloso, pedicelos de 12-25 mm de largo, bracteas de 0,5-1 x 0,5 mm, escariosas, sépalos de 1-1,3 x 1-1,2 mm, levemente imbricados basalmente, angostamente ovados, agudos a obtusos apicalmente, muy esparcida e inconspicuamente puberulentos lé 0,5mm de largo, irregularmente lacerados corola hipocrateriforma, el tubo ptirpura intenso, rosado a rojizo, los lobulos lila a rojizos, pero blanquecinos marginalmente, glabra, glabrescente a inconspicua y esparcidamente puberulenta exteriormente, tubo de 15-17 mm de largo, de 2,5-3,5(-4) mm de diametro en el orificio, inflado sobre la posicion de los estambres, oe de 7-11 x 4 ° mm, angostamente obovados, extendidos ero vari t dorsalmente, cabeza estigmatica dé 15 L75 mm de largo; ovario de 1-2 mm de largo, glabro, nectario ca 1,6-2 mm de largo, anular, leve e irregularmente q glabrescen tesexteriormente -anteras de 5,5-6 mm de largo glabras pentalobula o0.F t por dos folic ulos apoca arpicos, de 14- 19x 0,5- 0,7 cm, mas 0 menos divaricados, BIAPOS; continuos, con numerosas costillas longitudinales e i IncOnNnsf 1 uas; semi! llasdel5- 20mm de largo, coma apical de 2 8- 4 cm de largo, color canela Distribucién, habitat y ecologia.—Endémica del estado de Bahia, Brasil, donde se encuentra en bosques humedos, vegetacion costera, vegetacion de restinga, y areas de vegetacion alterada asociada, en elevaciones de 0-100 m. Especimenes con flores han sido recolectados entre enero y febrero y de septiembre a noviembre. Material con frutos se reporta en enero, febrero, septiembre y noviembre. Bahiella blanchetii es una especie ayant comun, que se reconoce al instante por sus flores con corolas hipocrateriformas. Es interesante notar como este género y en particular esta especie, la cual fue descrita desde hace mas de 150 anos, ha permanecido desapercibida desde entonces. En forma gen- eral, pocos taxOnomos vieron el tipo, pues ni Miers (1878) ni Woodson (1936) lo examinaron y aparentemente solo Miller Argoviensis (1860) lo hizo, pero en todo caso lo traté en el entonces inclusivo género Echites. En todo caso, las colecciones disponibles antes de 1970 eran muy escasas y ayaa esta especie se conocia solo por el tipo y as por Regel MORALES, BAHIELLA E : 345 y Riedel, ambas sin localidad ni fecha exacta y depositada en el herbario de la Universidad de Cambridge, Inglaterra (CGE) y en el Museo de Historia Natural de Viena (W). La semejanza de esta taxon con un Mesechites (i.e, Mesechites mansoanun (A. DC.) Woodson) es muy notable, ya que ambas especies son morfoldgicamente muy similares; inclusive, el especimen depositado en Viena fue examinado por Woodson y catalogado como M. mansoanus; sin embargo, como se explicé anteriormente, las caracteristicas de la cabeza estigmatica de ambas géneros son bien distintas y excluyentes, asi como otros caracteres morfologicos adicionales. En los ultimos 30 anos el incremento de la actividad botanica en Bahia, ha aumentado considerablemente el numero de especimenes disponibles hoy en dia, lo que ha permitido determinar en forma definitiva, el estatus ce de esta especie. BRASIL. Bahia: [|héus, Fazenda Jaguariba, al O de carretera Ilhéus-Olivenga, 31 ie 1997 (fl), Arbo et al. 7835 (CEPEC, CTES), ie reserva biolégica do Mico-leao (IBAMA), entre Ilhéus y Una, 14 Sep 1993 (£1, fr), Amorin et al. 1386 (C C), Marat, carretera BR 030, entre Ubaitaba y Marat, 25 Feb 1980 (fl, fr), Carvalho et al. 166 ae. Cabralia, reserva biologica DO Pau-Brasil, 5 Ene 1972 (£1, fr), ee 126 (CEPEC): S de Cumuruxatiba, ca Prado, 18 Ene 1977 Harl ey 18089 (CEPE ce Z),e Una y Ilhéus, 23 Ene 1977 (£1), ee et a 18178 (CEPEC, INB, K, NY); Santa Cruz de ae carretera antigua, 21 Oct 1978 (f)), Mori et al. 10919 (CEPEC, a Nov 1971 (£1, Santos a2) (CEPEC); - Valenca, entre Valenca y Guaibim, 21 Feb 1975 (£1, fr), ae 2888 (CEPEC): Itacaré, Nd le, caminoa Itacaré, 15 Nov 1992 (fl), Thomas et al. 9507 (CEPEC NY, USF). Datos perdidos: (f1), Regel s.n. (CGE); (fD, Riedel s.n. (U, W). 72 pene pen dibabtloea Morales, sp. nov. (Fig. 3). Tipo: BRASIL. BAHIA: Ilhéus, nco da Vitoria (5.7 km al O del puente sobre el rio Fundao, camino a Ita en camino ee sector O de Mata da Esperanga, 15 Ene 1995 (fl), Thomas et al. 10765 ees CEPEC). A B. blanchetii (A. DC.) J.E Morales, cui affinis llae infundibulif is, lobis 14-16 mm longis, et antheris 7-7,5 mm longis differt. Liana, los tallos viejos lenosos, ramitas usualmente teretes a subteretes, glabras a glabrescentes, coléteres interpeciolares ca. 1 mm de largo. Hojas: peciolos de 5-8 mm de largo; laminas de 8-16,5 x 3,8-5,5 cm, subcoriaceas, elipticas a obovado-elipticas, obtuso-apiculadas apicalmente, obtusas y cortamente atenuadas basalmente, revolutas marginalmente cuando viejas, glabras, eglandulares, sin coléteres en el nervio central adaxialmente, la venacion secundaria apenas evidente en ambas superficies, la venacion terciaria no evidente. Inflorescencias diminuta y esparcida a moderadamente papilado- puberulentas, pedtnculo de 95-96 mm de largo, el raquis inconspicuamente noduloso, pedicelos de 10-13 mm de largo, bracteas de 1-1,5 « 0,5-1 mm, escariosas, ae de 22. e x 1,9- e 6 mm, aha dunloyaler eles basalmente, angostamente ovad esparcida a moderadamente puberulentos exteriormente, mas raramente glabrescentes, coléteres ca. 1 mm de largo, irregular y profundamente lacerados; corola infundibuliforme, magenta basalmente, con los lébulos verde palido, glabra, 346 BRIT.ORG/SIDA 22(1) Wi 2mm 1G a a mm 1.5 cm 5cm Fic. 2. Rahiella infiundibylifl (Tt tal. 10765, CEPEC).A.R +. gd ia R Detalle delah davial dala lamina fali + dnl d lot C. Vista adaxial ded inal + dal lot | base. D. Detalle del tubo de | la. E. Antera, vista d ly lateral. F. Cab ti Atica. G. Ovario y nectario é MORALES, BAHIELLA, FE E 347 glabrescente a inconspicua y esparcidamente puberulenta exteriormente, la parte inferior del tubo de 10-12 x 2,7-3,2 mm, recto, la parte superior de 10-12 mm de largo, de 9-10 mm de diametro en la fauce, l6bulos de 14-16 x 10-13 mm, obovados, extendidos pero variadamente reflexos marginalmente; anteras de 7-7,5 mm de largo, glabras dorsalmente, cabeza estigmatica de 2-2,2 mm de largo; ovario de 1,8-2,1 mm de largo, glabro, nectario de 1-1,3 mm de largo, anular, leve a moderadamente pentalobulado. Foliculos desconocidos. Distribucién, habitat y ecologia.—Bahiella infundibuliflora es conocida unicamente de la localidad tipo, donde crece en bosques hmedos y margenes de bosques, en elevaciones bajo 50 m. El unico especimen con flores conocido fue recolectado en enero. Bahiella infundibuliflora es morfologicamente muy similar a B. blanchetii y aunque de primera instancia, ambas especies pueden confundirse, se pueden diferenciar con facilidad. De esta manera, B. infundibuliflora se puede distinguir facilmente por su corola infundibuliforme (vs. hipocrateriforma), con la fauce del tubo de 9-10 mm de ancho (vs. 2,5-3,5(-4) mm), sépalos mas largos y anchos, anteras de 7-7,5 mm de largo (vs. 5,5-6 mm de largo) y lobulos de la corola mucho mas grandes y desarrollados, de 14-16 x 10-13 mm (vs. 7-11 x 4-6 mm). APENDICE 1 CARACTERES Y ESTADOS DE CARACTER UTILIZADOS EN EL ANALISIS CLADISTICO, CON ALGUNAS NOTAS EXPLICATORIAS A continuacion se citan los caracteres y estados de caracter utilizados en el analisis cladistico. Notas explicatorias son incluidas para aquellos que sea necesario, asi como cuando se hayan encontrado discrepancias entre lo citado por la literatura y lo encontrado entre los especimenes. Los caracteres 1, 3, 7, 8, 20, 23, 26, 38, 44, 45, 46, 47, 48, 49, fueron tratados por Simoes et al. (2004), mientras que el 12, 13, 14, 16, 28, 29, 36, 40, 50, 52, 53, 55 por Williams (2004). Por lo tanto, no seran explicados aqui. 1. Habito 4. Tallos inferiores 0 Liana o hierbas escandentes 0 Herbaceos, no lignificados 1 Hierba perennes 1 Variadamente lignificados 2 Arboles 0 arbustos Casi sin excepcidn, la mayoria de los 2. Tallos con espinas miembros neotropicales de las Echiteae o 0 Si Mesechiteae son lianas, con los tallos viejos variadamente lenosos, algunos veces con la En general los miembros de las Echiteae y corteza conspicuamente suberosa. Sin em- Mesechiteae no tienen espinas 0 aguijones bargo, Cycladenia es una hierba erecta o en el tallo, pero este en un cardcter que se suberecta, que se caracteriza por sus tallos presenta en forma ocasional en géneros de herbaceos, no lenosos (Woodson 1936). otrasa tribus (e.g., Malouetieae) 5. Yemas axilares cubiertas por grupos de 3. Latex coléteres filiformes 0 Lechoso 0No 1 Acuoso 15S! En general, la pubescencia de los tallos y brotes jovenes de las Apocynaceae es muy variable, pero en las Echiteae Unicamente en Stipecoma se da la presencia de coléteres tiliformes envolviendo las yemas axilares 1 Opuestas 7. Hojas con coléteres en nervio central 0No 1Si 8. Hojas con domacios No 1Si 9. Hojas peltadas 0 Ausentes 1 Presentes 10. Consistencia de las hojas 0 Coriaceas 1 Membrandceas 11.Margenes revolutos 0 Ausentes 1 Presentes 12. Venas secundarias impresas 0Si 1No 13. Venacion terciaria en la cara abaxial 0 Conspicua, bien impresa 1 Inconspicua, apenas impresa O no impresa 14. Venas terciarias finamente reticuladas 0s f 1No 15.Venas terciarias perpendiculares al | nervio centra Si 1No Las venas terciarias Cuando presentes, tienen patrones irregulares en cuanto goon ag. en ‘aia ee (e.g., BRIT.ORG/SIDA 22(1) En forma general, de igual manera que se presenta en otras tribus relacionadas (e.g,, Mesechiteae), los miembros de las Echiteae tienen una sola inflorescencia por nudo, pero en unos pocos géneros (e.g., Macro- pharynx, Peltastes), dos inflorescencias por nudo usualmente estan presentes 18. Tipo de inflorescencia acimo 1 Cima sin ramificar o con una sola division basal 2 Cima ramificada Williams (2004) menciond que las inflorescencias en Laubertia contorta (M. Martens & oe ede no son ramificadas, per fl encias con al menos una ramificacion también son encontradas en esta especie, por lo que este estado de caracter debe tratarse como Eon eNice. 19. Fl g I pect beli forme 0Si 1No 20. Bracteas 0 Escariosas, inconspicuas 1 Foliaceas, conspicuas 21. Bracteolas 1 Presentes La presencia de bracteolas es raro en las Echiteae y Mesechiteae, presentandose Shes en Macropharynx y Peltastes, nque si bien, bracteolas inconspicu ie presentarse en unas pocas especies de Mandevilla o Neobracea 22. Consistencia de los sépalos Hlalleida 1 mas © menos perpendicular en relacion al nervio centr 16. Posicion de : inflorescencia 0 Axilar 1 Terminal 17. Numero de inflorescencias por nudo na sola inflorescencia en alguna de las dos axilas 1 Dos inflorescencias, una en cada axila 0 Escariosos 1 Foliaceos o subfoliaceos 23. Disposicioén de los coléteres 0 Solitarios y epicentricos 1 Filas continuas e ea tee en la base 2 Alternos con los sépalos Woodson (1936), en su monografia de Pentalinon (tratado entonces como p= ae Ure hi tes 5) me nci ion ie, que e los léteres en P. andrieuxti (Mall. Arg.) B. Hansen & Wunderlin eran alternos con los sépalos, pero la evaluacion de dicho caracter en diferentes MORALES, BAHIELLA especimenes han revelado que estos se encuentran distribuidos de manera con- tinua en una hilera de forma irregular a lo largo de la base del sépalo. 24. Numero de coléteres por caliz 0 Cero 1 Cinco 2 Seis O mas 25. Forma de la corola 0 Hipocrateriforme 1 Infundibuliforme, parte superior conica o tubular re q | 3 Urceolada 4 Rotada Tanto Simdes et al. (2004) como Williams (2004) han propuesto estados de caracter que difieren en la clasificacién de corolas infundibuliformes, siendo homédlogos en lo referente a corolas hipocrateriformes o rotadas. En este andlisis, las oe infundibuliformes fueron divididas S estados,el primero para clasificar las ee con la parte superior del tubo cédnica y el segundo para aquellas con la supe- rior campanulada. Dentro de las Echiteae, 7 t ~ ' con facilidad por la parte superior de la corola (e.g.,Angadenia, Pentalinon),la cual es siempre campanulada 26. Parte inferior del tubo 0 Recto 1 Curvado 27.Boton floral con union Id6bulo-tubo 1 Presente La presencia de coronas anulares, limitadas a unos pocos géneros en las Echiteae (Mo- rales 1997 a, b, 2002 a; Williams 2002; woodsell2s ) provecan ginacion | | if UNO con el tubo en gorones iietales que se presenta de Laubertia y Prestonia, pero que no es perceptible en Thoreauea (Morales 2005 b; Williams 2002). 28. Corona anular 0 Ausente 1 Presente 29. Corona supraestaminal 0 Ausente 1 Presente En ie Echiteae. a colon epiestaminal J C k F por cinco Sn dj 4 folia ubicados en forma opuesta a cada antera 30.Margen de los Iébulos en botones florales 0 Enroscados en espiral sobre su eje varias ces 1 Enroscados en espiral una sola vez Los Idbulos de la corola en las Echiteae y Mesechiteae, se encuentran enroscados en espiral, pero de manera regular estan invaginacos hacia el interior,de manera que el margen externo de los ldbulos se ven sll cba en ik ee externa Uni- traposicion en Asetonthera, los guile estan enroscados sobre su eje hacia el apice del botédn varias ee manera que el margen externo de lo ldbulos se ven enroscados varias veces sobre su eje 31. Color predominante de los Idbulos de la orola 0 Amarillo 1 Rosado 2 Rojo o morado 3 Verde 4 Blanco o crema 32. Forma de los I6bulos 2 Obovados En forma general, la mayoria de las géneros y especies de las Mesechiteae y Echiteae tienen Idbulos oblicuamente obovados, rf + | + 7 lAkK | A ofiliformes es caracteristico para unos pocos Woodson 1936; Williams 2002; Morales & Williams 2004 — 33.L6bulos erectos o suberectos 0 Si 1 No Los Idbulos de la corola se encuentran usualmente extendidos de manera irregular en la mayoria de los géneros de las Echiteae y Mesechiteae. Sin embargo, lobulos estricta- 34. 36. 37. 38. 39. mente erectos o suberectos, estan limitados a unas pocas especies de Mandevilla y al género Thoreauea (Williams 2002; Morales 2005, b). Lobulos torcidos en espiral distalmente Si 1No Los oe en ie eee eae son antésis, pero Ponelnnneets reflexos de manera irregular en sus bordes; los Idbulos erectos o suberectos son menos comunes. Por otro lado, la presencia de lObulos enroscados en espiral distalmente solo esta presente en Allotoonia (Morales & Williams 2004 ee fe city gaa 0 Ausente o inconspicua, los pelos hasta 1 mm de largo 1 Presente y conspicua, los pelos mas de 2 e largo Aunque la pubescencia de la corola como tal puede ser variable en las Echiteae, unicamente en Fernaldia se da la presencia de pelos largas y blancos en la cara adaxial de los ldbulos (Morales 2002 a). Tubo de la corola 0 Diminuto, menos de 6 mm de largo 1 Pequeno, 8-19 mm de largo 2 Mediano, 20-33 mm de largo 3 Grande, mas de 35 mm de largo Insercion de los estambres 0 Mitad distal, cerca de la boca 1 Mitad basal, cerca de la base Filamentos nteras + sésiles 1 Menos de 1 cm de largo 2 Mas de 3 cm de largo Filamentos envolviendo totalmente al fo) 0 Si 1No Los filamentos de la mayoria de las Echiteae y otras tribus de las Apocynoideae son muy cortos y casi ausentes, por lo que apenas tienen contacto con el estilo en la regién distal de este, ue antes de la posicioén de la cabeza estig generos (e.g., Forsteronia, Thenurdia: Thoreauea), dado el pequeno tamano del Cin embarg enotro BRIT.ORG/SIDA 22(1) tubo y estilo envuelven a este ultimo casi en la totalidad de su longitud. 40. Anteras 41. 42. 44. Canales 0 Incluidas 1 Apicalmente exsertas asi totalmente exsertas Pubescencia del apice de las anteras 0 Ausente 1 Presente La pubescencia dorsal de las anteras ha sido utilizado en otros andlisis (e.g., Simdes et al. 2004), pero este cardcter puede variar hotablementes en as eae Sin em- bargo, | restringida a la region estéril y apical de las anteras, que se puede incluso prolongar en un apéndice. Apice de las anteras 0 Triangular,no prolongado en un apéndice filiforme 1 Prolongado en un apéndice filiforme En general, la mayoria de las Apocynoideae neotropicales tienen los apices de las anteras variadamente triangulares, ya sea agudos 0 acuminados. Sin embargo, la prolongacion de estos apéndices por varios milimetros es raro y esta limitado a unos pocos geéneros 43.Apéndices filiformes de las anteras ier en espiral 0 Aus 1 ee Los ee filiformes de las anteras ando | pueden ser erectos (€.g., = er e) de, entrelazados en espiral (Pentalinon). guia de las anteras 0 Compuesto principalmente de engrosa- mientos endoteciales 1 Compuesto principalmente de esclerén- present quima 45. Base de las anteras 0 Acuminadas 1 Agudas 2 Obtusas a redonde adas 46. Union antera/cabeza estigmatica 0 Anteras unidas por un parche circular de células con apariencia de tricomas 1 Anteras unidas por un aro de pelos en forma de herradura MORALES, BAHIELLA f 7 TeTARANED es 2 Anteras unidas por un aro de pelos en 53. Grosor de los foliculos forma de herradura y una tira longitudi- 0 Gruesos, mas de 10 mm de ancho nal delgada 1 Delgados, hasta 9 mm de ancho 3 Anteras unidas por fusién celular 54. Folicul I tillas longitudinal 47. Cabeza estigmatica en corte transversal 0 Si 0 Circular o subcircular 1N 1 Pentagonal 55. Semillas 2 Con cinco costillas prominentes 0 Rostradas 48. Costillas de la cabeza estigmatica 1 Truncadas 0 Restringidas a la base 56. Extremo micropilar de la semilla 1 Distribuidas a lo largo de la cabeza estig- 0 Extendida mds alla del punto de naci- matica miento de la coma 49. Anillo en la base de cabeza estigmatica 1 No extendida mas alla del punto de 0 Ausente nacimiento de la coma 1 Presente La presencia de una coma micropilar en la 50. Nectario semilla es algo comun en muchas Apocy- 0 Anular noideae.Generalmente, el extremo micropilar 1 Dividido en cinco glandulas termina en el punto en el cual la coma nace. 51.Gineceo En muy pocas especies, el extremo 0 Apocarpico micropilar se prolonga por varios mm por 1 Sincarpico encima del punto de insercién de la coma, 52. Folicul convirtiéndose en un caracter potencial 0 Moniliformes o torulosos para la delimitacién genérica e incluso tribal. 1 Cilindricos AGRADECIMIENTOS Un reconocimiento y agradecimiento profundo al Herbario del Centro de Pesquisas do Cacau (CEPEC) en Bahia, Brasil, por el numeroso envio de mate- rial en préstamo, que permitio definir y clarificar el status definitivo de este nuevo género, lo que de otra manera no hubiera sido posible. Por otro lado los siguientes herbarios colaboraron con el préstamo y uso de colecciones: CGE, G- DC, K, NY, P, W. Un agradecimiento sincero a Fabian Michelangeli (NY) por su paciencia y disposicion a clarificar varias dudas del andlisis cladistico aca presentado. También reconozco la revision critica y numerosas sugerencias de Mary Endress (Z) en muchos aspectos de este trabajo, lo que permitio su correccion y mejoria notablemente. También se agradece la ayuda de Lucille Allorge por facilitar el acceso y estudio del material en los herbarios del Museo de Historia Natural de Paris, Francia, asi como de Bruno Wallnéfer por las mismas razones en el Museo de Historia Natural de Viena (W), Austria. Finalmente, quiero agradecer a Cecilia Ezcurra (BCRU) ya un revisor anonimo por la revision critica del manuscrito. REFERENCIAS Brown, R. 1810. On the Asclepiadeae, a natural order of plants separated from the Apocineae of Jussieu. Mem. Wern. Nat. Hist. Soc. 1:12-78. 352 BRIT.ORG/SIDA 22(1) Canoolte, A d&. 1844. Apocynaceae. In: A de Candolle, ed. Prodromus systematis naturalis regni vegetabilis, Vol. 8. Treuttel & Wurtz, Paris. Paris. Pp. 317-489. Enoress, M.E., B. Sennetao, S. Nicsson, L. Civeret, M. CHASE, S. Huysmans, E. Grarstrom & B. BREMER. 1996. A phylogenetic analysis of Apocynaceae s. str. And some related taxa in Gentianales:a multidisciplinary approach. Opera Bot. Belg. 7:59-102. Enoress, M.E. & P. Bruyns. 2000. A revised classification of the Apocynaceae s.l. Bot. Rev. 66:1—56. Hansen, B. 1985.A monographic revision of Forsteronia (Apocynaceae). Ph.D. dissertation, Univ. of South Florida, Tampa. [University Microfilms International, Ann. Arbor] Jupp, W., W. Sanoers & M. Donochue. 1994. Angiosperm family pairs: preliminary phyloge- netic analyses. Harvard. Pap. Bot. 5:1-51 Miers, J. 1878. On the Apocynaceae of South America. Williams & Norgate, London, Edinburgh. Mora es, J.F.1997a.A synopsis of the genus Prestonia (Apocynaceae) seccion Jomentosae in Mesoamerica. Novon 7:59-66. Morates, J.F.1997b.A reevaluation of Echites and Prestonia section Coalitae (Apocynaceae). Brittonia 49:328-336. Morates, J.F. 1997c. A synopsis of the genus Allomarkgrafia (Apocynaceae). Brittonia 49: 337-345 Morates, J.F.1997d [1998].A synopsis of the genus Macropharynx (Apocynaceae).Rhodora 99:58-68. Morates, J.F. 1999a. Hylaea (Apocynaceae-Apocynoideae), a new genus from South America. Novon 9:83-85. Morates, J.F. 1999b. A synopsis of the genus Odontadenia (Apocynaceae), In AJ.M. Leeuwenberg, ed. Series of revisions of Apocynaceae XLV. Bull. Jard. Bot. Nat. Belg. 67: 381-477. Morales, J.F. 2002a. Studies in Neotropical Apocynaceae |:A revision of the genus Laubertia Rhodora 104:170-186. Monrates, J.F.2002b. Studies in Neotropical Apocynaceae ll:A revision of the genus Fernaldia. Rhodora 104:186-200. Morates, J.F. 2003. Studies in Neotropical Apocynaceae Ill: A revision of the genus Secondatia, with discussion of generic classification. Candollea 58:305-319. Morates, J.F.2004. Estudios en las Apocynaceae Neotropicales VIl: Novedades taxonomicas en Prestonia (Apocynaceae, Apocynoideae) para Colombia y Ecuador,con comentarios sobre el grado de lobulacién del nectario. Candollea 59:159-165. Monrates, J.F. 2005a. Estudios en las Apocynaceae Neotropicales XIll: revisién del género Temnadenia (Apocynoideae, Echiteae). Candollea 60:207-231. Morates, J.F. 2005b. Estudios en las Apocynaceae Neotropicales XV: Sinopsis del género Thoreauea (Apocynoideae, Echiteae), con una nueva especie de Veracruz, México. Brittonia 57:258-263. Morates, J.F. & J. Wittiams. 2004. Allotoonia,a new neotropical genus of Apocynaceae based on a subgeneric segregate of Echites. Sida 21:133-158. MORALES, BAHIELLA E E Al 353 Mutter Arcoviensis, J. 1860. Apocynaceae. En: C.F.P. von Martius, ed. Flora Brasiliensis 6(1). Munchen, Wien, Leipzig. Pp. 1-180. Nixon, K. 2002. Winclada, version 1.00.08. Programa de computacidn distribuido por el autor en, http://www.cladistics.com. Cornell University, Ithaca, NY. Poraieter, K & V. Avsert. 2001.Phylogenetic relationships within Apocynaceae s.|. based on trmL intron and trnL-F spacer sequences and propagule characters. Ann. Missouri Bot. Gard. 88:523-549, SENNBLAD, B. & B. Bremer. 1996. The familial and subfamilial relationships of Apocynacecae and Asclepiadaceae evaluated with rocL data. Pl. Syst. Evol. 202:153-175. SENNBLAD, B. & B. Bremer. 2002. Classification of Apocynaceae s.|. according to a new ap- proach combining Linnaean and phylogenetic taxonomy. Syst. Biol. 51:389-409. SENNBLAD, B., M.E. Enpress & B. Bremer. 1998.Morphology and molecular data in phylogenetic fraternity: the tribe Wrightieae (Apocynaceae) revisited. Amer. J. Bot.85:1143-1158. Simoes, A., M.E. Enpress, T. vAN DeR Niet, L. KinosHita & E. Conti. 2004. Tribal and intergeneric relationships of Mesechiteae (Apocynoideae, Apocynaceae): evidence from three noncoding plastid DNA regions and morphology. Amer. J. Bot. 91:1409-1418. Wiuams, J.K. 1998. A revision of Thenardia (Apocynaceae, Apocynoideae). Lundellia 1:7 8 Wiuams, J.K. 1999. A phylogenetic and taxonomic study of the Apocynaceae subfamily Apocynoideae of Mexico with a synopsis of subfamily Plumerioideae. Ph.D. disserta- tion. Austin: The University of Texas. Wituams, J.K. 2002. Thoreauea (Apocynaceae: Apocynoideae), a new genus from Oaxaca, Mexico. Lundellia 5:47-48. WILLIAMS, J.K. 2004. Polyphyly of the genus Echites (Apocynaceae: Apocynoideae:Echiteae): evidence based on a morphological cladistic analysis. Sida 21:117-132. Woopson, R.E. 1933. Studies in the Apocynaceae IV. The Américan genera of Echitoideae. Ann. Missouri Bot. Gard. 20:605-790. Woopson, R.E. 1935. Studies in the Apocynaceae. IlV.The American genera of Echitoideae. Ann. Missouri Bot. Gard.22:153-306. Woopson, R.E. 1936. Studies in the Apocynaceae. IV. The American genera of Echitoideae. Ann. Missouri Bot. Gard. 23:169-438. 354 BRIT.ORG/SIDA 22(1) Book REVIEW SusAN McCiure.1996. The Herb Gardener: A Guide for All Seasons. (Paperback edition). (ISBN 0-88266-873-0, pbk.). Storey Publishing, 210 MASS MoCA Way, North Adams, MA, 01247. U.S.A. pOnlets: # ooer 1-800-441-5700; wwwstorey.com). $14.95, 236 pp., color photogray d drawings, appen- dices, index, 8 3/8" x 10". Gardeners and cooks alike will learn a a deal from The Herb gardener: A Guide for All Seasons. Author Susan McClure has created a comprehensive compendium for people interested in herb g dening; subjects include a eae on designing, garden duties throughout the year, crafts ee projects, as well asa growing gui The book is divided into five main parts concerning a such as herbs, herb gardening Basics, seasonal planning in the garden, herb projects, and a grower’s guide. gardening basics chapter shows the reader items to pete in planning a garden and it also ie) in oe a garden space. General preplanning considerations are discussed including a plant life cycles, and plant tolerances. Also included are environmental considerations before you plant (sun, soil, fertilizer, pH, water, etc.). The portion of this chapter on designing your garden will be incredibly helpful information for most readers. The author has included many different garden types/shapes such as hardscape considerations, included labeling suggestions and maintenance thoughts. The section about yearlong gardening divides the year up into the four seasons and provides gardeners with typical, well-organized inf h snot occu: quae rach period of the year. One of the very helpful f in this bool he side | PI through- out the text; examples are the seed sprouting facts chart in the Spring period, harvesting techniques and troubling pests and diseases charts in the summer period, first frost estimator in the fall period, and plants for an indoor garden 1 in ah winter pene od. The section on recip d d | cool benefits of the pla ning and herbal harvest. Tie portion iaeludes auultiple methods of preserving herbs for longer-term hj use, good h tions, recipes, herbal vinegars, cheese and butters, a wreaths. It also say oie a helpful mre of nutritional value for the common her nal section of the book includes a very useful grower's guide. Ne section vee read- ] ers with oe a along nh nace of 52+ eke al f J varieties as files for herbs are organized by Iso incl ] opagation, cultivars, potential problems, harvesting, use, and more. Each profile ae summaries of oo in the forms of colorful icons for quick reference. If you have a hankering to make your own pesto or create your own herbal seasoning blend, check out The Herb Gardener: A Guide for Ail Seasons and plan your own herbal garden. McClure has written a text that is easy to read and easy to use. It’s full of helpful side boxes, charts and herbal profiles. This book would be a ae aeditiont to the pEary of gardeners, cooks and herbal connois- seurs.—Lee Luckeydoo, Herbariu h Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. SIDA 22(1): 354. 2006 ESTUDIOS EN LAS APOCYNACEAE NEOTROPICALES XxV: NOVEDADES Y NUEVOS REPORTES EN LAS APOCYNACEAE (APOCYNOIDEAE: RAUVOLFIOIDEAE) DE VENEZUELA J.Francisco Morales Instituto Nacional de Biodiversidad Apartado 22-3100 Santo Domingo, Heredia, COSTA RICA RESUMEN Se proponen varias Seine poniendiaturales en mere género Mandevilla, incluyendo una nueva especie (M. tristis J.F Morale anu dn (M. angustata (Steyermark) J.F Morales), y un nuevo sindnimo (M. bella a Woodson). Nuevas adiciones a la Flora de Venezuela en los géneros Aspidosperma (A. rigidum Rugby, A. schultesii Woodson), Mandevilla (M. vanheurckii (Mull. Arg.) Markgr.) y Odontadenia (O. killipii Woodson) son reportadas. ABSTRACT Several nomenclatural novellies are (piepered in the eeenus Mangers: la, including a new species (M. tristis J. Morales), a new k) J.-F. Morales) and a new synonym (M. bella (Pittier) Woodson). New additions to ache Venezuelan Flora in the genera Aspidosperma (A. rigidum Rugby, A. Sauultesty Wesdsen)) Mandeville AM NGDNEM TCE an Arg.) Markgr.), and OnE. clip lin the genus Mandevilla, incl species (M. tristis J.E Morales), a new combination (M. angustata Geom J.E Mo ouik a new report ((M. vanheurckii (Mull. Arg.) Markgr) and a new synonym (M. bella (Pittier) Woodson Venezuela es uno de los paises con una alta diversidad de Apocynaceae s. s.,con un estimado de 30 géneros y ca. 200 especies (Morillo 1989a). El numero de géneros puede variar dependiendo de la aceptacion de Iabernaemontana como una sola unidad genérica segtin lo propuesto por Leeuwenberg (1994) 0 dividido en una serie de grupos como lo propuesto por Allorge (1985). Asimismo, la relativa confusa taxonomia prevaleciente en algunos géneros como Aspi- dosperma y la divergencia de taxones aceptados de ese género en diferentes tratamientos floristicos puede incrementar o decrecer el numero estimado de taxones. En la ultima Buonep ae del género Mandevilla, Woodson (1933) report6 un total de 15 esy [M. congesta (Kunth) Woodson, M. bella (Pittier) Woodson, M. caurensis Marker, M. fendl eri (Mull. Arg.) Woodson, M. hirsuta (Rich) K. Schum., M. javitensis (Kunth) K. Schum., M. lancifolia Woodson, M. moritziana (Mull. Arg.) Donn. Sm., M. scabra (Hoffmanns. ex Roem. & Schult.) K. Schum., M. subcarnosa (Benth.) Woodson, M. subsagittata (Ruiz & Pav.) Woodson, M. subspicata (Vahl) Markgr, M. surinamensis (Pulle) Woodson, M. veraguasensis SIDA 22(1): 355 — 365. 2006 356 BRIT.ORG/SIDA 22(1) (Seem.) Hemsl., M. villosa (Miers) Woodson)| para Venezuela, de las cuales cuatro (M. bella, M. caurensis, M. lancifolia, M. moritziana) fueron consideradas endémicas. En forma posterior, mas especies f descritas (e.g., Markgraf 1971, 1978; Monachino 1958, 1961; Woodson 1948) aumentando a 23 el namero aproximado de taxones conocidos para el pais. En su tratamiento de los géneros de Apocynaceae de Venezuela, Clausnitzer (1968) reporto un total de 22 especies, mientras que Morillo (1989 b), en conjuncion con la descripcion de dos nuevos taxones, report6 la presencia de M. annularifolia Woodson (anteriormente conocida solo de Colombia) y propuso dos nuevos sindénimos. En el tratamiento de las Apocynaceae para la Flora en la Guyana Venezolana, Morillo (1995) trato un total de 24 taxones, estimando un total de 40 especies para Venezuela. Aunque si bien este ultimo trabajo se trata de un tratamiento de flora para un area geografica especifica (y no para toda Venezuela), tiene una relativa importancia, por el hecho de incluir algunas especies previamente no conocidas en el pais. El tratamiento preliminar de una nueva monografia de Mandevilla (Morales, datos sin publ.) contabiliza un total de al menos 37 taxones en Ven- ezuela, algunos recientemente descritos (Morales 2005). De ellos, tan solo cinco pertenecen al subgénero Mandevilla, mientras que el resto pertenecen al subgénero Exothostemon (Woodson 1933). Como resultado, varias novedades son propuestas, incluyendo nueva sinonimia, un nuevo reporte, una nueva combinacion, y una nueva especie. Por otro lado, hasta el momento, no existe un tratamiento completo de la familia (s.s.) para el pais y Unicamente tratamientos para los géneros (Clausnitzer 1968) 0 regiones geograficas determinadas (e.g., Morillo 1989a; Zarucchi et al. 1995) han sido publicados. Mientras revisaba material sin identificar de Venezuela, varios taxones ente no reportados para ese pais de los géneros Aspidosperma (papi ies yO ren (Apocynoideae) fueron identificados, asi como una serie de novedades taxonomicas en Mandevilla (Apocynoideae) estas altimas derivadas principalmente de la elaboracién de una nueva monografia para el género. Estas novedades son presentados a continuacion, agrupados en subfamilias y tribus de acuerdoa la clasificacion propuesta por Endress y Bruyns Mandevilla angustata (Steyerm.) J.-F Morales, comb. & stat. nov. (Fig. 1). BAsionimo: Mandevilla subcarnosa var. angustata Steyerm., Acta Bot. Venez. 2:318. 1967. Tipo: VENEZU- ELA. BOLIVAR: Auyan-tepui, cumbre de la parte Norte, entre campamento Llovizna y pie del segundo Muro, 1] May 1965 ({]), Stevermark 93737 (HOLOTIPO: VEN; ISOTIPOs: MO (fotocopia en J, US, VEN). Lianas o arbustos erectos; ramitas jovenes levemente anguladas o aladas, subteretes con la edad, tallos sdlidos a ligeramente huecos, glabros, coléteres interpeciolares inconspicuos, hasta 0,8 mm de largo. Hojas opuestas; peciolos MORALES : 357 Zi: De Fic. 1. Mandevilla angustata (Liesner & Holst 20762, VEN). A. Ramita fértil con flores y frutos. B. Detalle de la superficie Jo vtal da la hn} Pisani ea lot In] dol : tral. C. C4liz Jicelo v bra D. Vista adaxial de un sépalo, t Jo el col ici y teg t F. Antera, vista dorsal. G. Cat tig Stica.H Nectario y ovario. | Semilla J. Detalle de la pubescencia de las semilla. 358 BRIT.ORG/SIDA 22 1) 3-8 mm de largo; laminas foliares 2,6-10 x (1-)1,4-3,9(-5) cm, obovadas a obovado-elipticas, el Apice obtuso, obtuso-apiculado a agudo, la base cuneada, los coléteres distribuidos en forma irregular a lo largo del nervio central, firmemente membranaceas, glabras o glabrescentes, usualmente no revolutas, venas secundarias y terciarias apenas impresas en ambas superticies. Inflorescencia ligeramente mas larga que las hojas subyacentes, subterminal, pero rapidamente volviéndose axilar, diminuta e inconspicuamente puberulenta cuando joven, pero volviéndose rapidamente glabra o glabrescente, con 5a 15 flores, sésiles o subsésiles, el pedtnculo (cuando raramente presente) de 1-5 mm de largo, pedicelos 3-5 mm de largo (a veces hasta 8 mm en fructificacion), bracteas 1-2 x 0,5-1 mm, angostamente ovadas, escariosas; sépalos 2-3 x 1,4-1,7 mm, ovados, obtusos a agudos apicalmente, el apice no reflexo, escariosos, glabrescentes externamente, diminuta e inconspicuamente ciliolados marginalmente, el coleter solitario, irregularmente lacerado apicalmente; corola infundibuliforme, amarilla, inconspicuamente papilada a glabrescente externamente, el tubo giboso basalmente, la parte inferior 27-32 x 2-2,5 mm, la parte superior 13-19 mm de largo, conica, 14-22(-25) mm de diametro en la fauce, el apice del boton floral agudo o cortamente apiculado; lobulos 16-18(-21) x 9-13(-17) mm, obovados, relativamente extendidos, ligera e irregularmente reflexos marginalmente; estambres insertos en la base de la parte superior del tubo, anteras 6-7,4 mm de largo, glabras dorsalmente, la base auriculada, con las auriculas subtruncadas, cabeza estigmatica 2,7-3 mm de largo; ovario 1,3-1,7 mm de largo, diminuta y moderadamente puberulento; nectario anular, leve a moderadamente pentalobulado, ca. la mitad (o ligeramente menos) de la longitud total del ovario. Foliculos 15,5-18 cm x 1,5- 3,5mm, glabros, levemente moniliformes; semillas 7-7,5 mm de largo, diminuta y densamente hirsutulas, coma 1,4-2,1 cm de largo, canela-amarillo. Distribucion, habitat y ecologia.—Restringida al E de Venezuela (estado de Bolivar) y Guyana (region Potaro-Siparuni), donde crece en bosques enanos y ventosos, en elevaciones de (450-)950-2000 m. Especimenes con flores han sido recolectados en Mayo y Octubre. Especimenes con frutos fueron recolectados en Mayo. Ensudescripcion de M. subcarnosa Benth. var angustata, Steyermark (1967) menciono que esta diferia de la variedad tipica (la que ocurre en el S y SE de Venezuela, Guyana y el N de Brasil en el Estado de Roraima) por sus hojas con las bases “angostadas y cuneado-agudas” (vs. redondeadas u obtusas a subcordadas), asi como laminas foliares oblanceoladas u oblanceolado-oblongas (vs. oblongas). En forma general estos caracteres son persistentes, pero existen suficientes caracteres adicionales que justifican el elevar su estatus a nivel especifico. De esta manera, M. a neta se puede diferenciar de M. subcarnosa por sus hojas con las t uadas (vs dispuestas L MORALES 3 359 perpendicularmente), separadas por 4-10 mm entre si (vs. 2-5 mm), asi como inflorescencias sésiles o subsésiles, con el pedtnculo de 1-3 mm de largo (vs. pedunculadas y con el pedtinculo de 12-45 mm), foliculos de 16-18 cm de largo (vs. 7-13,5cm) y semillas diminuta y densamente hirsutulas, conel dumente erecto a suberecto (vs. densamente tomentulosas y cone Especimenes examinados. VENEZUELA. Bolivar: Piar, cima del Amaruay-tepui, 10 May 1986 (£1), Liesner & Holst 20762 (MO, VEN); Ptari-tepui, cercanias del campamenteo Misia Kathy, 28 Oct 1944 (fl), Steyermark 59473 (MO). GUYANA: Montanias Pakaraima, Aymatoi, 17 Oct 1981 (f{D, Maas et al. 5801 (K, MO, U), Maas et al. 5824 (U, WAG); region Potaro-Siparuni, rio Blackwater, 22 May 1991 (f1), McDowell et al. 4895 (US, WAG). Mandevilla boliviensis (Hook. f.) Woodson, Ann. Missouri Bot. Gard. 20:716. 1933. Dipladenia boliviensis Hook. f., Bot. Mag. 95:t. 5783. 1869. T1po: BOLIVIA: Datos perdidos (£1), Pearce 708 (HOLOTIPO: K, foto, MO). Mandevilla bella (Pittier) Woodson, Ann. Missouri Bot. Gard. 20:717. 1933, syn. nov. Dipladenia bella Pittier, J. Wash. Acad. Sci. 21:141. 1931. Tipo: VENEZUELA. DISTRITO FEDERAL: Hacienda rto La Cruz, region costera, 2300 m, 28 Ago-4 Sep 1918 (f), Pittier 8108 (HOLOTIPO: US). La unica diferencia mencionada por Woodson (1933) para separar Mandevilla bella de M. boliviensis (Hook. FE.) Woodson fue la longitud y forma del apice de los lobulos de la corola, asi como distribucion geografica disyunta. Sin embargo, no existen diferencias adicionales en otros caracteres morfoldgicos en los especimenes tipo que permitan reconocer ambos taxones. En todo caso, la longitud de los lobulos es muy variable en el material de M. boliviensis disponible hoy en dia, lo que en conjuncion con la alta variacion en la longitud y tamanio de la corola presente en otras especies del subgénero Mandevilla (eg., M. pohliana (Stadelm.) A. Gentry), impide reconocer ambos taxones. Otros sinonimos de M. boliviensis fueron anteriormente propuestos por Morales (1995, 98). Mandevilla tristis J.F. Morales, sp. nov. (Fig. 2). Tipo: VENEZUELA. TACHIRA: Uribante, S ase de Cerro El Morro, 21 Jun 1990 (f1, fr), Dorr et al. 7115 (HOLOTIPO: NY; foto, INB) A Mandevilla bracteata (Kunth) Kuntze, M. hirsuta (Rich) K., Schum. M. sagittarii Woodson et M. steyermarkii Woodson, cui similis, foliis sessilis vel subsessilis, petiolis 1-2 mm longis, et bracteis 2-2 mm latis differt Liana; ramitas algo aplanadas cuando jovenes, teretes a subteretes con la edad, ligeramente huecas, densamente tomentulosas cuando jovenes, esparcida a moderadamente puberulentas con la edad, mas raramente glabrescentes; coléteres interpeciolares inconspicuos, hasta 0,9 mm de largo. Hojas opuestas; peciolos 1-2 mm de largo; laminas foliares 2-7,3 x 1,5-3,2(-3,7) cm, ovadas a ovado-elipticas, el Apice cortamente cuspidado, la base cordada, los coléteres irregularmente distribuidos a lo largo del nervio central, membranaceas, diminuta y esparcidamente papilado-puberulentas en la superficie adaxial, la pubescencia mas densa sobre el nervio central, diminuta y densamente 360 BRIT.ORG/SIDA 22(1) F 4mm 2mm 4mm 5mm 2cm 2mm 2mm 2cm Fic. 2. Mandevilla tristis (A-H de Werff & Gonzdlez 5313, |NB;\—J de Dorr et al. 7715, NY).A. Ramita con inflorescencia. B. Detalle de la sapere adaxial de la a mostrando los coléteres a lo largo del nervio central. C. Caliz, pedicelo y bractea. D. Vista de la adaxial de un sépalo, mostrando # coléter. E. Tubo de un botén parcialmente abierto, mostrando la posicién i: ie anteras y el tubo basalmente g F, Antera, vista dorsal. G. Cabeza estigmatica. H. Nectario y ovario. 1. Foliculos.J. Semilla MORALES 3 361 tomentosas en la superficie abaxial, el indumento secando glauco, no revolutas, venas secundarias y terciarias conspicuamente impresas en la superficie abaxial, usualmente solo las venas secundarias apenas visibles en la superficie adaxial. Inflorescencia usualmente mas larga que las hojas subyacentes, axilar, densa a moderadamente puberulentas, con muchas flores, pedunculo 4-16(-20) mm de largo, pedicelos 1-1,5 mm de largo, raramente algunos pocos alcanzando los 3 mm de largo en fructificaci6n, bracteas 9-15 x 1,2-2 mm, linear-ovadas, subfoliaceas; sépalos 8,5-11 x 0,5-1 mm, linear-ovados, largamente acuminados apicalmente, el apice no reflexo, escariosos, densa a moderadamente puberulentos externamente, el coleter solitario, irregularmente laciniado apicalmente; corola infundibuliforme, amarilla, algo giboso, densamente a moderadamente hispidulosa externamente, sobretodo en botones florales, el tubo giboso basalmente, la parte inferior 13-18 x 2-3,6 mm, la parte superior 14-16 mm de largo, conica, 10-13 mm de diametro en la fauce, el apice del bot6n floral cortamente apiculado; lobulos 8-12 x 7-11 mm, obovados, relativamente extendidos; estambres insertos en la base de la parte superior del tubo, anteras 3 2. 4,1 mm de largo, glabras dorsalmente, la base auriculada, con las auriculas d ,cabeza estigmatica 1,6-1,9 mm de largo; ovario 1,3-1,5 mm de largo, inconspicua y diminutamente papilado-puberulento a glabrescente; nectario anular, moderada a profundamente pentalobulado, ca. la mitad de la longitud total del ovario. Foliculos 10,5-13,5 cm x 1-3,6 mm, densa a esparcidarnente hispidulos, conspicuamente moniliformes; semillas 7,5-8 mm de largo, glabras a glabrescentes, coma 1,7-2,4 cm de largo, canela. Distribucion, habitat y ecologia.—Endémica al estado del Tachira en el E de Venezuela, donde crece en formaciones de sabanas y vegetacion arbustiva asociada a afloramientos rocosos o arenosos, en elevaciones de 1100-1300 m. Especimenes con flores y frutos han sido recolectados en Junio, pero material con flores se conoce también de Julio. Especimenes examinados. VENEZUELA. Tachira: Uribante, entre Siberia y Pregonero, 1] Jul 1983 ({D), Werff & Gonzalez 5313 (INB, MO). Mandevilla tristis se puede confundir con M. bracteata (Kunth) Kuntze, M. hirsuta (Rich) K. Schum., M. sagittarii Woodson y M. steyermarkii Woodson, un grupo de taxones que se caracterizan por sus inflorescencias con bracteas foliaceas o subfolidceas y corolas infundibuliformes. Sin embargo, M. tristis puede distinguirse facilmente por sus hojas subsésiles (pero aparentando ser sésiles), con los peciolos inferiores a 2 mm de largo (vs. 4-60 mm de largo), inflorescencias con las bracteas florales mucho mas angostas (1,2-2 mm vs. 3- 9 mm), flores subsésiles, con el pedicelo hasta 1,5 mm de longitud (vs. 2-8 mm de largo) y corolas con el tubo mas pequeno y con un diametro mas angosto en la fauce. 362 BRIT.ORG/SIDA 22(1) SUBFAMILIA APOCYNOIDEAE Tribu Apocyneae Odontadenia killipii Woodson, Ann. Missouri Bot. Gard. 18:546-547. 1931. Tipo: PERU. LORETO: Iquitos, 26 Sep 1929 (f1), Killip & Smith 29847 (HOLOTIPO: MO; IsoTIPO: US). Odontadenia killipii es un taxon relativamente facil de reconocer por el tamano relativamente pequeno de sus sépalos, corolas y foliculos. Aunque ha sido confundida con la comun O. verrucosa (Willd. ex Roem. & Schult.) K. Schum. ex Markgr, O. killipii puede reconocerse por sus sépalos relativamente mas pequenos (2-4 mm vs. 3-9 mm), corolas glabras externamente (vs. esparcidamente puberulentas a glabrescentes), con la parte superior del tubo de la corola anchamente conica (vs. angostamente conica) y foliculos mucho mas pequenos (7,5-10,5 cm vs. (13-)19-30 cm) (Morales 1999). Odontadenia killipii era conocida del S de Colombia a través de Ecuador hasta Pert, laGuyana Francesa y Brasil, pero se reporta por primera vez para Venezuela, es el estado de Amazonas. Especimenes examinados. VENEZUELA. Territorio Federal Amazonas: Rio Negro, Serrania de Tapirapeco, May 1992 (fl), Martin 1909 (MO, PORT); Atabapo, Oct 1989 (f1), Velasco 730 (MO). Tribu Mesechiteae Mandevilla Mage (Mull. Arg.) Markgr., Notizbl. Bot. Gart. Berlin-Dahlem 9:87, fig. 2L. 1924. Heterothrix ea vase Mull. Arg., Observ. Bot. 164. 1871. Eriadenia obovata Miers, Apocyn. S. Amer. 117-118, t. 14 B. 1878, nom. superfl. Tipo: PERU. LORETO: cerca de Tarapoto, fecha perdida (£D, Spruce ae (HOLOTIPO: W, foto F neg. 31792; IsOTIPOs: BM, CGE, F (fragmento), G, foto F neg. 26855, K [2 laminas], P) Este taxon fue originalmente reportado por Clausnitzer (1968) para Venezuela (pero sin testigo citado). Posteriormente, Morillo (1995) trato este taxon como “Mandevilla sp. A.’ en el tratamiento de la Flora de la Guayana Venezolana, aunque hizo referencia que esta especie habia sido conocida como M. vanheurchii. El estudio del tipo, asi como de diversos especimenes no deja duda de que este taxon esta presente en Venezuela, donde se conoce de los Estados de Amazonas y Bolivar. En forma general, esta especie se puede confundir con M. subcarnosa (Benth.) Woodson, ya que ambas comparten varios caracteres tales como tallos algo angulados cuando jovenes, hojas con el indumento ausente o glabrescentes y no evidente, inflorescencias con bracteas escariosas, con pedicelos cortos y corolas infundibuliformes, pero M. vanheurckii se distingue por facilidad por sus laminas foliares con las venas secundarias y terciarias usualmente no evidentes. Especimenes examinados. VENEZUELA. Amazonas: Casiquiare, alrededores de Yavita, rio Temi, 6-19 Jul 1969 (£1, fr), Bunting et al. 3736 (U); Rio Negro, SE de San Carlos de rio Negro, 12 Nov 1987 (£1), Liesner & Carnevali 23012 (MO, WIS), cerro de la Neblina, rio Yatua, 14 Nov 1957 (fl, fr), Maguire et al. 42069 (F, NY, US); cerro de la Neblina, rio Yatua, 1 Ene 1958 (fl), Maguire et al. 42570 (K, NY, U, VEN), carretera San Carlos-Solano, 3 Feb 1977 (£1), Morillo 5153 (VEN): Atures, cano Piedra, SE de MORALES 3 363 Puerto Ayacucho, Sep 1989 (f1, fr), Sanoja et al. 2965 (MO, NY, FOR Bolen {alpen tepui, Feb 1948 (f1), Phel ps etal. 384 (NY); S Sororopan tepul, 14 Nov 1944 (£1) SUBFAMILIA RAUVOLFIOIDEAE Tribu Alstonieae El género Aspidosperma (Rauvolfioideae, Alstonieae) se encuentra ampliamente distribuido desde el S de México hasta Bolivia, Paraguay y el N de Argentina, asi comoen las Antillas, donde esta representado por pocas especies. En forma gen- eral, es uno de los grupos de las Apocynaceae neotropicales con una de las mas altas variaciones intraespecificas, en cuantoa la forma de las hojas e indumento de partes vegetativas e inflorescencias. Debido a esto, un sin numero de formas, variedades y taxones han sido propuestos para albergar diferentes estados dentro del rango de variacion de una misma especie, provocando una divergencia no- table en cuanto al numero de taxones aceptados en el género. De esta manera, Woodson (1951), reconocio un total de 52 especies, mientras que la revision de Marcondes-Ferreira (datos sin publ.) acept6 un total de 34 taxones. Clausnitzer (1968) report6 un total de 16 taxones para Venezuela, mientras que Morillo (1995), estimo que ca. 30 especies estaban presentes en el pais. Ahora bien, en el transcurso de identificaci6n de material sin identificar depositado en el Mis- souri Botanical Garden (MO), dos taxones anteriormente no conocidos en Ven- ezuela, pertenecientes al subgénero Aspidosperma (Marcondes-Ferreira & Kinoshita 1996) fueron encontrados, ambos restringidos a la cuenca baja amazonica. Aspidosperma rigidum Rusby, Mem. New York Bot. Sak 7:323. 1927. Tipo: BO- _LA PAZ: rio Bopi, 12 Sep 1921 (£1), Rusby 593 Aspidosperma jaunechense A. Gentry, Phytologia 47:98-99. 1980, syn. nov. Tipo: ECUADOR. Los Rios: Vinces, bosque de Jauneche, 26 Mar 1980 (f1), Dodson & Gentry 9920 (HOLOTIPO: MO; ISOTIPO: SEL) Aspidosperma rigidum pertenece a la seccion Rigida (Marcondes-Ferreira & Kinoshita 1996) y ahora se encuentra distribuida en Costa Rica, Colombia, Ven- ezuela, Ecuador, Pert, Bolivia y el NO de Brasil. En general, esta especie se puede reconocer con facilidad por la siguiente combinacion de caracteres: fuste prin- cipal conspicua e irregularmente acanalado, tallos jovenes con las yemas apicales sin catafilos, hojas de 4-11 cm de largo, elipticas y membranaceas, inflorescencias laterales, con los lobulos de la corola angostamente elipticos y foliculos hasta 7 cm de largo, dolabriformes, lisos externamente y con las semillas secando negras. De forma vegetativa, se puede confundir con facilidad con las especies de la seccion Excelsa (Marcondes-Ferreira & Kinoshita 1996), pero A. rigidum se puede reconocer por sus laminas ovado-acuminadas y venas secundarias arquedas y unidas a la siguiente en forma terminal, formando una vena terminal. En la descripcion de Aspidosperma jaunachense, Gentry (1980) tnicamente 364 BRIT.ORG/SIDA 22(1) clasific6é ese taxon dentro de la clasificacion intragenérica propuesta por Woodson (1951), pero sin discutir en forma directa sus relaciones con alguna especie en particular. Sin embargo, aunque él cité que este taxon era muy simi- lara A. rigidum, no profundiz6 en discutir las diferencias que podrian separar ambos taxones. De hecho, el estudio de los tipos ha revelado que no existen diferencias significativas que justifiquen la validez de A. jaunachense por lo que es reducido a la sinonimia. Especimenes examinados. VENEZUELA. Territorio Federal Amazonas: mision rio Mavaca, || Feb 1991 (El, fr), Stergios et al. 15335 (INB, MO, PORT) Aspidosperma schultesii Woodson, Ann. Missouri Bot. Gard. 38: 168, tl, f.6. 1951. } Tipo: BRA SIL: A | Ira i-Igarapeé ) V bit atluente del rio Taraira, 4-6 Jul 1948 ({1), Schultes & Lopez 10178 (HOLOTIPO: MO: ISOTIPO: F). Aspidosperma schultesii, pertenece a la seccion Schultesia (Marcondes-Ferreira & Kinoshita 1996), y dentro del género es un taxon bastante distintivo que se puede reconocer con facilidad por sus ramitas jovenes sin catafilos, tallos y peciolos secando negros, glabrescentes, laminas foliares con la venacion secundaria apenas evidente 0 no evidente, asi como corolas con los lobulos ca. 3 veces mas largos que el tubo, y foliculos negros al secar, con la superficie externa conspicuamente granulada. Esta especie se encuentra ahora distribuida en Co- lombia, Venezuela, N de Brasil y Peru. Es pees examinados. VENEZUELA. Territorio Federal Amazonas: San Carlos de Rio Negr al de la conf] lel Rio Negro y el brazo Casiquiare, 29 Feb 1980 (fr), Clark & Maquirino 7367 (MO). AGRADECIMIENTOS Los siguientes herbarios facilitaron el préstamo o uso de sus colecciones: BM, CGE, FE K, MO, NY, P, PORT, U, US, W, WIS, VEN. Quiero reconocer la ayuda prestada por Gerardo Aymard (PORT) por su disposiciOn a suministrar una referencia bibliografica especifica. Asimismo, agradezco al Missouri Botanical Garden (MO) por el apoyo brindado que ha hecho posible la visita de su herbario y por la cantidad de especimenes enviados como regalo por identificacién y a Alina Freire-Fierro y Rosa Ortiz-Gentry, por su colaboracién en mis visitas a dicho herbario. Finalmente, se agradece la revision y comentarios por parte de David Goyder y un revisor anonimo REFERENCIAS A.vorce, L. 1985. Monographie des Apocynacées—Tabernaemontanoidées Américaines. Mém. Mus. Natl. Hist. Nat., B, Bot.30:1-216 CLAUSNITZER, |. 1968. Los géneros de Apocynaceae de Venezuela. Acta Biol. Venez. 3: 4 Enoress, M. & PV. Bruyns. 2000 A revised classification of the Apocynaceae s.|. Bot. Rev. 66: MORALES 3 365 LeeuwenserG, A.J.M.1994.A revision of Tabernaemontana two.The New World species and Stemmadenia. Royal Botanic Gardens, Kew, U.K. Pp. 213-450. Marconbes-F erreira, W.& L. KINOSHITA. 1996.Uma nova divisdo infragenérica para Aspidosperma Mart. (Apocynaceae). Revista Brasil. Bot. 19:203-214. Markarar. F. 1971 [1972]. New Apocynaceae and Asclepiadaceae from Venezuela. Acta Biol. Venez. 6(1-4):65—74 Markarar. F. 1978. Novedades de Apocynaceae. Acta Biol. Venez. 13:353-355. MonacHino, J.1958.Apocynaceae. In:B. Maguire y J. Wurdack, eds. The botany of the Guyana Highland Ill. Mem. New York Bot. Gard. 10:117-138. MonacHino, J. 1961.Apocynaceae.In:B. Maguire y J.Wurdack, eds. The botany of the Guyana Highland lV.Mem. New York Bot. Gard. 10:58-65. Morates, J.F. 1995. An evaluation of the Mandevilla boliviensis complex. Phytologia 78: 197-198. Mora es, J.F.1998.A synopsis of the genus Mandevilla (Apocynaceae) in Mexico and Cen- tral America. Brittonia 50:214-233. Morates, J.F. 1999. A synopsis of the genus Odontadenia (Apocynaceae). In: AJ.M. Leeuwenberg, ed. Series of revisions of Apocynaceae XLV. Bull. Jard. Bot. Nat. Belg. 67: 381-477. Monrates, J.F.2005.Estudios en las Apocynaceae Neotropicales X:definicién de la verdadera identidad de Mandevilla leptophylla, con la descripcién de dos nuevas especies y una nueva combinacion de Mandevilla (Apocynoideae, Mesechiteae) para Sur América. Sida 21:1537-1548. Moritto, G. 1989a. Las Apocynaceae del estado Lara. Biollania 6:193-237. Moritto,G.1989 b. Notas preliminares sobre el género Mandevilla Lindley (Asclepiadaceae) en Venezuela. Ernstia 53:15—20. Monritto, G. 1995. Aspidose In:J.A.Steyermark, P Berry y B. Holst, eds. Fl. Venez. Guayana 2:481-490. Timber Press, Portland, OR and Missouri Botanical Garden, St. Louis. Moritto, G. 1995 Mandevilla. In: J.A. Steyermark, P. Berry y B. Holst, eds. Fl. Venez. Guayana 2:518-529. Timber Press, Portland, OR and Missouri Botanical Garden, St. Louis. STEYERMARK, J. 1967. Flora del Auyan-Tepui. Acta Biol. Venez. 2(5-8):5-370. Woopson, R.E. 1933. Studies en the Apocynaceae IV. The American genera of Echitoideae XXVI. Ann. Missouri Bot. Gard. 20:605-790. Woooson, R.E. 1948. Miscellaneous new Apocynaceae and Asclepiadaceae. Ann. Missouri Bot. Gard. 35:233-237. Woopson, R.E. 1951. An interim revision of the genus Aspidosperma Mart. & Zucc. Ann. Missouri Bot. Gard. 38:1 19-206. ZARUCCHI, J.,G.N. Moritto, M.E. Enpress, B.F. HANSON & A.J.M. LEeUWwENBERG. 1995. Apocynaceae. In: JA. Steyermark, P Berry & B. Holst, eds. Fl. Venez. Guayana 2:471—571.Timber Press, Port- land, OR and Missouri Botanical Garden, St. Louis. 366 BRIT.ORG/SIDA 22(1) Book REVIEW Davip STUART. 2004. Dangerous Gardens: The quest for plants to change our lives. (ISBN 0-674-01104-X, hbk.). Harvard Press, Cambridge, MA (Orders: TriLiteral LLc, 100 Maple Ridge Dr, Cumberland, RI 02864-1769, U.S.A., 800-405-1619 http://www.hup.harvard.edu/). $35.00, 208 pp., color drawings, color and b&w photo, bibliography, and index, 7 1/2" x 10 3/8" Do poisonous plants and aphrodisiacs interest you? What about medicinal plants, life elixirs, and “mind expanding” plants? Then you should venture into the enjoyable new book called Dangerous Gardens: The i st for plants to change our lives. David Stuart has created this intriguing text about mankind's use of plants to pursue h, haps and long life. The book is divided into chapters that focus on different subject ar- eas to which plants have been see by humankind: “Great Afflictions,” “ The Vital Organs,” “The Flight a Pain,” “Chasing Venus,” “The Killing Plants,” “The Seven Ages of Man,” “The Mind” and “Mysteries of the Gods.” The author ee both historical background and historical reasoning regarding why plants were ana for different ance or desires. For example, the book begins witha history of the pl Other oan ve through- ~_ healt out the book ne now plants were aed oe used for one reason and accidentally or eventu- ally learned that the same plant actually cured other illnesses. Foxglove (Digitalis dela for example, in the 1600/1700s was used for wounds and lung ailments; not until the late 1700s was it learned that this same plant had dramatic effects on the heart. The text also describes many ae that were liberally used in history until their true and often-dangerous behavior was unearthed. One example is ee w mach up until he late ee was used liber oF as a cure for au kinds of ailments. The I th { famous poisoners, “cure-all” quack pills, ee use in rituals ad the use of various - plants as aphrodisiacs. The author also included information in s chapter what oS pas labelled "shadow plants,” plants that are often used for treatment, sucl ] f| pp.), but that science and/or the medical community a not ceed the true benefit and risks, if any. I found Dangerous Gardens hard to put down, it reads like a very good cable TV beumenay on plant use by man, complete with photographs and/or drawings of the plants being discussed, practitioners of herbal medicine, perpetrators of poisoning, quackery, and many other historical images related to plant use. This book was insightful regarding the rationale that was used to decide which Cea to use for different illnesses. It was fascinating to learn about how various plants were used diff y throughout time. This book is recommended to anyone with an interest in the topic of plant use by mankind; it is easy to read and engaging.—Lee Luckeydoo, Herbarium, Botanical Re- search Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A SIDA 22(1): 366. 2006 TAXONOMIC OVERVIEW OF THE HETEROTHECA VILLOSA COMPLEX (ASTERACEAE: ASTEREAE) Guy L.Nesom Botanical Research Institute of Texas t Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT ieterthecs villosa (as treated by senile 1996, 2006) is a comipiex species with nine varieties, most of Heterotheca villosa var. nana and H. villosa var. scabra are ena silonarrc and intergrade Tittle, but each is widely sympatric with H. villosa and distinct from it. Recognition at specific rank accurately reflects the status of var. nana and var. scabra, and they are treated here, respectively, as Heterotheca horrida (Rydb.) Harms and Heterotheca polothrix Nesahy nom. et stat. nov. pieleoiiees senoptiila sensu stricto is distinct from H. stenophylla var. g with it,and the latter is appropriate mere villosa Pele oe Fe (Rx741 var. angustifolia ydb.) Harms. The New Mexico endemic Heterotheca vil osa var. sierrablancensis Semple is raised to specific rank as Heterotheca sierrablancensis Sane Ny ,comb et stat. nov. Identifications of vars. villosa, foliosa, ballardii,and minor (all judge- ents because of their broad sympatry and extensive intergradation. The disimeten between var, el nculata and H. zionensis is not clear, and both taxa apparently intergrade broadly with more typical H. villosa. Variety depressa is maintained at specific rank as H. depressa (Rydb.) Dorn. Maps show the generalized distributions of the taxa of the H. villosa complex sensu Semple, and a nomen- clatural summary outlines an alternative taxonomy. RESUMEN Heterotheca villosa (as treated by sae e 1996, 2006) pl ho variedades la mayoria de las cuales son sim varios eee Hehe rothec villosa var. nana y ie L villosa P g ] ae A oO ye i simpatrica con H. villosa y dik le ella. El imi ivel ifi flej yi el estatus de var. nana y var. scabra, y son tratadas aqui Gonieleiete ol ecelion r ida (Rydb.) Harms y Heterotheca po othrix tstat. nov. Heterot! es diferente de H. stenophylla var. ene ean Semple) y simpatrica con ai, y esta ultima es euiada Apropcamien( como H. Valo var. sangusyol ia aay Harms. ea pcicemies de Neevo villosa var. sierrablancensis (Semple) Neco comb. et stat. nov. Las identificaciones ae las vars. villosa, foliosa, pl picen at bido a su amplia simpatria e ballardii, y minor (todas sensu intergradacion extensa. La diferenci tre var. pedunculata y H. zionensis no es clara, y ambos axa aparentemente se intergradan con la mas tipica H. villosa. La variedad depressa se mantiene en el rango especifico como H. depressa (Rydb.) Dorn. En los mapas se muestran las distribuciones gen- erales de los taxa del cas H. villosa sensu Semple, y un resumen nomenclatural delinea una taxonomia alternativa Intermediates between taxa of Heterotl ften 1 and unequivo- cal identifications are correspondingly difficult. Phenotypic plasticity, small SIDA 22(1): 367 — 380. 2006 368 BRIT.ORG/SIDA 22(1) degrees of differentiation, and hybridization apparently underlie the taxonomic difficulties. Tetraploids are common and gene exchange among them appar- ently occurs frequently. A monograph of Heterotheca sect. Phyllotheca (Semple 1996) provides a detailed view of the taxonomic structure of this group and addresses difficulties in identification Heterotheca villosa, as treated by Semple (1996, 2006), is a widespread and complex species with nine varietal taxa, most of which are geographically over- lapping or almost completely superimposed. Typical H. villosa (var. villosa) oc- cupies essentially the northern half of the species range, but it is broadly sym- patric with var. minor, var. foliosa, var. ballardii, and var. nana. In part of Nebraska and South Dakota, five varieties (sensu Semple) of H. villosa occur sympatrically; four varieties occur sympatrically in Saskatchewan; three oc- cur sympatrically in much of Wyoming, Colorado, and Utah. Two are relatively narrow endemics (within a single state); all the others occur in at least four states. All taxa were mapped in detail by Semple (1996, Figs. 39 and 40); gener- alized outline maps shown here (Figs. 1, 2, 3) are derived in most part from Semple’s dot maps. Regarding the Heterotheca villosa complex, Semple (1996, p. 114) noted that it might be “logical to merge all ... varieties together with no infraspecilic taxa being recognized ... This would result in the loss from the formal nomenclature of a great deal of information on variation and distribution in what is admit- tedly a difficult species complex.” His approach (as noted by Nesom 1997) has been to recognize sympatric conspecific varieties, using formal varietal names to identify what many (or most) systematists might regard as geographical trends or populational variants. “A variety is characterized by all members of a population exhibiting a particular morphology distinct from other individu- als in the species. The distribution of these populations is sympatric with popu- lations whose members are not within the same variety, and also many popu- lations of morphological intermediates exist [citing various references]. .. Van Steenis described varietal level variation as being continuous with other vari- eties, although the continuum would have pronounced modes” (Semple 1974, p. 8-9). Similarly, and with regard to Heterotheca villosa, “the races fit well with the concept of variety in that each occurs in pure form in some populations, and the overall ranges are sympatric to a considerable degree with at least one other variety” (Semple 1996, p. 108). Semple has used a similar taxonomic approach for Heterotheca sessiliflora (Nutt.) Shinners sensu lato, a primarily Californian complex, and formal changes in taxonomic rank were proposed by Nesom (1997) to clarify its sys- tematics. The taxonomic situation with H. villosa (Pursh) Shinners is consid- ered here. Harms (1968, 1970) also has addressed aspects of the taxonomy of the H. villosa complex. NESOM, TAAUNUMIC OVERVIEW MI 369 | ae ® + Fic. 1. Generalized distribution of Heterotheca villosa (A) var. villosa and (B) var. minor, both Semple, and (C) H villosa var.an gustifolia. Illustrated is the broad sympatry of I {th | sympatry/alloy try of var. angustifolia. The range of H. villosa d | minor, includ I N da (Clark Co., Excelsior Canyon, 7 Sep 1941, Clokey 8759-MO).Th ge of ifolia includ I lowa (Lyon Co., prairie, among Sioux Quartzite exposures, nw corner of county, 25 Aug 1924, Shimek s.n.-MOQ). 370 BRIT.ORG/SIDA 22(1) Fic. 2. Generalized distribution of Heterotheca villosa (A) var. foliosa, (B) var. ballardii, (C) H. zionensis, and (D) var. pedunculata, all sensu Semple. Compare with Fig. 1 to see broad sympatry of var. foliosa and var. ballardii with var. villosa and var. minor. Status of Heterotheca villosa var. pedunculata Heterotheca villosa var. pedunculata occurs in complete sympatry with H. villosa var. minor in the Four Corners region, as indicated by Semple (1996) (var. pedunculata with upper cauline leaves densely to extremely densely strigose, thus pale green to white, eglandular to very sparsely glandular; 2n = 18, 36; var. minor with upper cauline leaves sparsely to moderately densely strigose, sparsely to densely glandular; 2n = 18, 36). Variety minor “is the most variable linfraspecific taxon] in the species and includes some local distinctive morphotypes that grade into other forms. Plants intermediate between this and all other varieties occur in areas where the ranges are sympatric, and they make infraspecific taxonomy of the species difficult” (Semple 2006). In a study of NESOM, TAXONOMIC VVYECRVICVY VI 371 ~ \ A ian bet Fic. 3.G lized distributi f (A) Heterott | lothrix, (B) H. horrida, (C) H. stenophylla (D) H. depressa and (E) H. £4 2 Pes + bE L sierrablancensis. Ill ha dictrihut; 3 Four Corners Heterotheca, | have been able to make only arbitrary morphologi- cal distinctions between var. pedunculata and var. minor. Variety pedunculata is very similar to Heterotheca zionensis Semple and partly sympatric with it. Plants of H. zionensis are said to be identified by stems and leaves appearing silvery to whitish from densely strigose vestiture and pe- duncles and phyllaries usually densely glandular but otherwise glabrate. Such plants are distinguished by Semple from more densely pubescent individuals of H. villosa var. minor or from plants identified as var. pedunculata (with gray to silvery leaves) by glandular peduncles and phyllaries, even though plants of var. minor may have glandular phyllaries and populations of H. zionensis may 372 BRIT.ORG/SIDA 22(1) include individuals with eglandular peduncles and phyllaries—“non-glandu- lar forms of H. zionensis occur in north-central Utah; glandular and non-glan- dular forms occur in the Utah-Arizona border region” (Semple 1987, p. 385). The geographic range of H. zionensis overlays the range of the western portion of var. minor as well as of var. pedunculata, and distinctions among these are un- clear. It does seem clear that silvery-leaved plants are concentrated in the south- west part of the range of H. villosa, but if the silvery-leaved plants are represen- tative of a geographic trend in vestiture, widely intergrading with more typical plants, then it is likely that the names H. zionensis and H. villosa var. pedunculata represent plants shaped by the same trend and the same underly- ing genetics. Status of Heterotheca villosa var. nana and var. scabra Heterotheca villosa var. nana (diploid) is almost completely overlaid in its geo- graphic range by var. foliosa (diploid and tetraploid but tetraploid in its area of overlap with var. nana, many chromosome counts, fide Semple 1996) and by var. minor (diploid and tetraploid, many counts, fide Semple 1996). Variety nana varies considerably and probably hybridizes with H. villosa and H. fulcrata (Greene) Shinners (Semple 2006), but even so, var. nana is morphologically con- sistent and recognizable throughout its range and has previously been treated at specific rank by various botanists as H. horrida (Rydb.) Harms (e.g., Harms 1968, 1970; Great Plains Flora Association 1986; Dorn 1988). Heterotheca villosa var. scabra is the westernmost taxon treated by Semple within H. villosa, except for the south-reaching extension of the species in Cali- fornia (var. minor fide Semple). Semple (1992) noted that var. scabra is closely related to the Californian endemic H. shevochii (Semple) Semple (originally described as H. villosa var. shevochii). It seems equally plausible that var. nana is closely related to var. scabra. All three taxa tend to have spreading leaves and strongly developed glandularity. The geographic ranges of var. nana and var. scabra are slightly overlap- ping along the Utah/Colorado and northern Arizona/New Mexico borders (Fig. 3). Semple (2006) observed that “In the Four Corners area, var. nana grades into var. scabra, and collections often are difficult to place into one of the two taxa”— but I find the distinction clearer and intermediates not numerous, especially in view of the broad range and relative morphological constancy of var. scabra. Plants of var. scabra and “aff. var. scabra” cited by Semple at a southeastern ex- tension of its range in trans-Pecos Texas are tentatively identified here as forms of H. viscida (A. Gray) Harms. New Mexico collections of var. scabra well east of the range illustrated in Fig. 3 (San Miguel Co., Rio Arriba Co. cited but not mapped by Semple 1992) are better identified as var. nana. | have not seen records of var. scabra from Idaho or Colorado (as indicated on Fig. 3), but it is a distine- tive entity and Dr. Semple’s citations document its presence there. — NESOM, TAXONOMIC VYLAVICW UE 373 Variety nana and var. scabra are distinguished by Semple by various dif- ferences (mostly foliar), and my study corroborates and emphasizes their dis- tinction, as in the following couplet. 1. Cauline leaves usually not contorted upon drying, upper mostly lanceolate to tri- angular-lanceolate, acute at apex; leaf surfaces moderately to densely glandular, the glands ue elie ~~ the sparsely strigose (nonglandular) vestiture, nonglandular hai t ricted to midvein area (middle 1/3) of adaxial surfaces or sometimes nearly absent on both surfaces, especially on more distal leaves; phyl- laries glabrous to very sparsely strigose, glandular; 2n = 18, 36 (many counts, fide Semple 1996) Heterotheca villosa var. scabra Cauline leaves often contorted (margins undulate) upon drying, upper mostly ob- long or oblong-lanceolate, acute to obtuse at apex; leaf surfaces moderately to gosely Senauiay me glands conspievous a the ee hispid to lyt telyd ie Goce. glandular; ae = 18 (many counts; 2n = 27, one count, ide Semple 1996) eter Shee villosa var. nana Including some features more difficult to precisely contrast, plants of var. scabra (in contrast to var. nana) have stems and leaves with much reduced non-glan- dular vestiture but with denser and more conspicuous glandularity, leaves dif- ferently shaped, smaller, and more widely spaced, heads relatively fewer, and phyllaries glabrous (lacking non-glandular hairs) but glandular. The two taxa are different in geography. In their area of sympatry, var. scabra is distinguished from a regional, more typical form of Heterotheca villosa (var. minor fide Semple) by the following contrasts. 1. Upper cauline leaves lanceolate to triangular-lanceolate, acute at apex; leaf sur- faces moderately to densely glandular, the glands little obscured by the sparsely Sstrigose (nonglandular) vestiture, nonglandular hairs often restricted to midvein area (middle 1/3) of adaxial surfaces or sometimes nearly absent on both surfaces; phyllaries glabrous to very sparsely strigose Heterotheca villosa var. scabra Upper cauline leaves linear to oblanceolate, oblong-lanceolate, or narro vate, acute to obtuse at apex; leaf surfaces eglandular or sparsely to cones glan- dular, moderately to densely strigose evenly over both surfaces; phyllaries sparsely strigose Heterotheca villosa var. minor — In view of their morphological integrity and apparent reproductive isolation from elements of more nearly typical Heterotheca villosa, both H. villosa var. nana and H. villosa var. scabra are treated here at specific rank. Heterotheca polothrix Nesom, nom. et stat. nov. Chrysopsis villosa var. scabra Eastwood, roc. Calif. Acad. Sci. 2.6:294. 1896. Heterotheca villosa var. scabra (Eastwood) Semple Phytologia 73:453. 1992. Type: U.S.A. UTAH. San Juan Co. Willow Creek, 14 Jul 1895, A eee 38 (HOLOTYPE: CAS, digital image!). Non Heterotheca scabra DC. 1836. The species is named for a resemblance of the medial strip of adaxial leaf vestiture to the mane of a pony (Greek, polos, pony, and thrix, hair). Blake's varietal epithet (‘cinerascens,’ at varietal rank, as below) is poorly descriptive of the taxon (as also noted by Semple 1987). 374 BRIT.ORG/SIDA 22(1) Chrysopsis viscida A. Gray subsp. cinerascens Blake, Proc. Biol. Soc. Washington 35:173. 1922. Heterotheca horrida subsp. cinerascens (Blake) Semple, Brittonia 39:381. 1987. TYPE: U.S.A. UTAH. [Beaver Co. fide Welsh 1982] Beaver Creek, among rocks in the oak region, 2 Sep 1901, I. Tidestrom 2873 (HOLOTYPE: US, internet image). Heterotheca horrida (Rydb.) Harms, Wrightia 4:17. 1968. Chrysopsis horrida Rydb, Bull. Torrey Bot. Club 31:648. 1905. Type: U.S.A. COLORADO. |Weld Co. New Windsor, 8 Aug 1900, G.E. Osterhout 2326 (HOLOTYPE: NY internet image! ISOTYPES: RM, WIS). Chrysopsis canescens var. nana A. Gray, Mem. Amer. Acad. Sci. 4[PI. Fend1.:78. 1849. Heterotheca villosa var. nana (A. Gray) Semple, Novon 4:54. 1994. TyPE: U.S.A. NEW MEXICO. [Mora Co.?!: elevated rocky region 2 mi E of the Mora River, Aug 1847, A. Fendler 391¢ (HOLOTYPE: GH). Status of Heterotheca villosa var. sierrablancensis Heterotheca villosa var. sierrablancensis was described by Semple (1996) from the White Mountains and Sacramento Mountains of Lincoln and Otero cos., New Mexico. It is similar to H. horrida in its conspicuously glandular vestiture and congested and consistently corymbiform capitulescence but distinct in its leaves more densely strigose, the upper cauline ascending, larger, and differ- ently shaped (lanceolate or oblong-lanceolate to ovate), ray corollas longer, and habitats at higher elevation. It occurs at the southeastern corner of the range of H. horrida (H. villosa var. nana) and Semple indicated (1996, Fig. 3) that the two taxa are most closely related to each other. Both are primarily diploids. The ex- tended population system of var. sierrablancensis is morphologically variable but it lies inside the geographic range of H. horrida and apparently is isolated from it as well as the regional expression of H. villosa, both of which occur at lower elevations. Recognition at specific rank is appropriate for this taxon, es- pecially in view of the parallel rank of H. horrida. Heterotheca sierrablancensis (Semple) Nesom, comb. & stat. nov. Heterotheca vil var. sierrablancensis Semple, Univ. Waterloo Biol. Ser. 37:146. 1996. TYPE: U.S.A. NEW MEXICO. Lincoln Sierra Blanca, NM 532 8.5 km W of NM 48, rocky outcrop and face of roadcut at shback directly below scenic overlook, 2 Oct 1995, J.C. Semple and Semple 10513-A (Ho- LOTYPE: WAT, ISOTYPES: MO!, NMC, RM). OSd Status of Heterotheca villosa var. depressa. Semple treated this narrowly endemic taxon at varietal rank because it hybrid- izes with var. minor. Both taxa are tetraploid in their area of sympatry. As noted by Semple (2006), “Var. depressa grows in geyser basins in Yellowstone National Park and is locally common in rocky soils in Teton National Park; it is very rare farther south in Wyoming. It is distinguished by its small stature, small heads, and linear-oblanceolate leaves with many hairs and usually some glands. It is similar to some forms of var. minor, which occur throughout the range of var. depressa and with which it hybridizes. ... Typical var. depressa occupies the vi- cinity of hot springs and geyser basins and nearby river flood banks, while var. minor occupies other drier or non-geyserite nearby habitats.” Markow (2001, NESOM, TAXONOMIC VVYCAVICYY UT 375 2004) found that the occurrence of H. depressa is strongly correlated with that of another narrow endemic, Stephanomeria fluminea Gottlieb. The geographic range of Heterotheca villosa var. depressa is relatively dis- crete, like that of H. sierrablancensis, and is essentially imbedded within the much larger distribution of H. villosa sensu lato. Although var. minor (sensu Semple) hybridizes with var. depressa, the latter appears to have its unique eco- logical niche, maintains its morphological identity, and is justifiably treated at specific rank. Heterotheca depressa (Rydb.) Dorn, Vasc. Pl. Wyoming (ed. L) 295. 1988. Chrysopsis depressa Rydb., Mem. New York Bot. Gard. 1:381. 1900. Heterotheca villosa var. depressa (Rydb.) Semple, Novon 4:53. 1994. TyPE: U.S.A. WYOMING. Teton Co.: Yellowstone National Park, Lower Geyser Basin, 7000 ft, 4 Aug 1897, PA. Rydberg and E.A. Bessey 5067 (HOLOTYPE: NY internet imagel; ISOTYPES: CAN, NY, US internet image!). — Status of Heterotheca stenophylla var. angustifolia. Semple (1996, 2006) has treated Heterotheca stenophylla (A. Gray) Shinners as comprising two varieties, var. stenophylla and var. angustifolia (Rydb.) Semple, “based on field studies and the results of multivariate analyses .... The type of var. angustifolia is morphologically closer to many indi Is of [H. stenophylla] var. stenophylla than it is to either H. canescens or typical H. villosa. ... At the very least, the type of var. angustifolia belongs in H. stenophylla, regardless of where the more canescens-like and the more villosa-like plants are placed” (Semple 1996, p. 94). Previous botanists have treated var. angustifolia within Heterotheca villosa (e.g., Harms 1968, 1970; Great Plains Flora Association 1986). Harms observed (1968, p. 16-17) that “over most its range, including Texas, Heterotheca stenophylla appears to represent a quite distinct diploid (n = 9) species, but some tetraploid (n = 18) populations are known from a narrow band along the east- ern edge of the species’ overall distribution reaching from south-central Kan- sas tothe Wichita M tains of south-central Oklahoma (to ?Archer Co., Texas) which hybridize and intergrade somewhat with [tetraploid] H. villosa. Asa re- sult, the tetraploid H. stenophylla populations as a whole are usually distinguish- able from the diploid populations by the presence of such introgressed charac- ters from H. villosa as more pubescent involucres, stems, or leaves, but it seem impossible to assign individual plants to a particular ploidy level on this basis. Thus it appears impractical to attempt to taxonomically distinguish the tetra- ploid from the diploid H. stenophylla populations despite the the presence of an obvious reproductive barrier based upon different ploidy levels. Neither does it seem logical to taxonomically merge H. stenophylla with H. villosa [var. angustifolia] merely because certain tetraploid populations of the former hy- bridize more or less extensively with the latter, when most of the H. stenophylla taxon is diploid and clearly distinct morphologically and biologically.” 376 BRIT.ORG/SIDA 22(1) Heterotheca stenophylla (sensu stricto) is broadly sympatric with var. angustifolia and I (with Harms) also find that the two are distinct in the field and herbarium. Triploid putative hybrids have been reported (Semple 1996, 2006). In contrast, var. angustifolia intergrades with broader-leaved forms of H. villosa, especially in Nebraska and the Dakotas, and is reasonably treated within H. villosa. “Inclusion of var. angustifolia ... only slightly increases the morpho- logical complexity of H. villosa, as defined by Semple, and [it] occupies a part of the overall geographic range es its sympatry with conspecific varieties is relatively less” (Nesom 1997, Harms (1970, p. 1567) digea “a ‘most of the Texas material [of Heterotheca villosa] belongs to var. foliosa (characterized by dense canescent appressed pu- bescence of involucres and herbage, and larger heads) or to var. angustifolia (characterized by narrower leaves, coarser and sparser herbage pubescence and smaller heads); these varieties intergrade on a broad scale, and much Texas material appears to be more or less intermediate between them.” Still, a trend within H. villosa toward narrow leaves is pronounced in Texas, Oklahoma, and Kansas, and identification of these plants as var. angustifolia is justifiable. i . Stems eglandular; leaves oblong-oblanceolate to elliptic-oblanceolate, 3-6 mm wide, eglandular or sessile-glandular on abaxial surface, moderately to densely stri- gose, hairs usually without inflated basal celss; phyllaries eglandular or inconspicu- ously and minutely glandular; chromosome counts of 2n = 36 (fide Semple 1996) Heterotheca villosa var. angustifolia . Stems sessile- oe leaves linear to narrowly ol oblanceolate, 2-3(—4.5) mm wide, sessile-glandular, sparsely hispid-strigose, hairs with conspicuously inflated and vis cid basal =alle aS eile glandular to eglandular, sparsely strigose or essentially without nonglandular hairs; many chromosome counts of 2n = 18, few of 2n= (fide Semple 1996) Heterotheca stenophylla aaa Consistent distinction between Heterotheca villosa var. angustifolia and H. canescens (DC.) Shinners is sometimes more difficult than between var. angustifolia and H. stenophylla. Heterotheca canescens is mostly diploid, but some tetraploids have been reported, and as noted by Semple (1996, p. 97), “tetraploids [of H. canescens and H. villosa var. angustifolia] apparently hybrid- ize forming local swarms of parent-like and hybrid individuals.” And “in Texas, it sometimes is difficult to clearly distinguish herbarium specimens of [H. canescens] from certain canescent, strigose-sericeous, smaller-headed, more narrow-leaved forms of H. villosa (Harms 1970, p. Heterotheca villosa (Pursh) Shinners var. angustifolia (Rydb.) Harms, Wrightia 16. 1968. Chrysopsis a Rydb., Bull. Torrey Bot. Club 37:128. 1910. Chrysopsis villosa var. angustifolia (Rydb.) Crongq., Bull. Torrey Bot. Club 74:150. 1947. Heterotheca senophla vat. iis See (Ryd) Semple, Novon 4:53. 1994. Type: (Semple 1990): U.S.A. NEBRASKA. Hooker Co.: Middle Loup River, near Mullen, on sandhills, 14 Sep 1893, PA. Rydberg 1766 (LECTOTYPE: NY internet image! ISOLECTOTYPES: GH, NY, US). NESOM, TAXONOMIC OVERVIEW OF 377 ee stenophylla (A. Gray) Shinners, ao & Lab. 19:68. 1951. Chrysopsis var. stenophylla A. Gray, Boston J. Nat. Hist. 6 (PL. Lindh.):223. 1850. Chrysopsis villosa var. epee (A. Gray) A. Gray, Synopt. FI. : Amer. (ed. 2) 1(2):123. 1884. ees stenophylla (A. Gray) Greene, Erythea 2:96. 1894. Type: U.S.A. TEXAS. [Llano Co, Llano, from strong ligneous roots in crevices of smooth granite rocks, Nov 1847, EJ. Lindheimer 631 LOTYPE: GH; ISOTYPES: CAN, DS, GH, K, MO!, ND-G, NYI4 sheets, internet image], US). Taxonomic summary of the Heterotheca villosa complex Four taxa treated by Semple as varieties within Hetorotheca villosa are recog- nized here at specific rank: H. horrida, H. polothrix, H. sierrablancensis, and H. depressa. Intergradation and sympatry among var. villosa, var. foliosa, var. ballardii,and var. minor make it impractical and biologically unrealistic to rec- ognize them as more thana single entity. The identity of var. pedunculata vs. H. zionensis and vs. other densely hairy plants in the region of var. minor is not clear. Variety angustifolia appears to be justifiably recognized within H. villosa (vs. within H. stenophylla) Distinct morpl hic entities are recognized here with formal names. Relatively stable morphology and sympatry with close relatives indicate that such taxa are reproductively isolated, and they are treated at specific rank. In addition to typical Heterotheca villosa, var. pedunculata (sensu lato) and var. angustifolia appear to be geographically distinct but each intergrades with more typical plants where ranges meet—these latter two are tentatively regarded here at varietal rank. It might be useful to recognize widespread morphological “nodes” within var. villosa sensu lato at the rank of forma or with an informal name; the contrasting taxonomic approach (formal recognition of sympatric conspecific varieties) requires a majority of typological and arbitrary judgments. Local and regional floristic studies may encounter and identity stable sym- patric races that call for a more complex nomenclature than proposed here. But because of the geographic mix of hybrids and ploidy levels, biological and morphological patterns in the Heterotheca villosa complex may remain more complex than can be adequately circumscribed by traditional nomenclature _ eK 1. Heterotheca villosa (Pursh) Shinners [1951 Amellus villosus Pursh (1814, NEOTYPE: Missouri River, almost certainly North Dakota] la. Heterotheca villosa var. villosa Including laa, lab,and lac. The sympatry and intergradation of var. villosa sensu Semple with var. foliosa, var. ballardii, and var. minor suggest that these taxa are arbitrarily identified. [laa]. Heterotheca villosa var. iosa (Nutt.) Harms [1968] Chrysopsis foliosa Nutt. (1841, TYPE: sw WYOMING] Chrysopsis' villosa var. see iosd ut ) ssa [1947] Chrysop: ata Is. [190 PE: COLORADO. El Paso/ Teller Co.] eee ae var. ae 7 oe Nels. [1909 378 BRIT.ORG/SIDA 22(1) If Chrysopsis foliosa var. imbricata is treated as synonymous with Heterotheca villosa var. foliosa (as done by Semple 1996, 2006), the earliest name at varietal rank is “var. imbricata.” In this context, H. villosa var. foliosa is an invalid name. [lab]. Heterotheca villosa var. ballardii (Rydb.) Semple [1994] Chrysopsis ballardii Rydb.[1931, TYPE: MINNESOTA. Carver Co] “This is the generally robust, larger-headed, many-rayed, eglandular, oblong- leaved race of the species occurring on the northeastern prairies of Canada and the United States” (Semple 1994, p. 53). “Forms of var. ballardii are sufficiently similar to forms of var. foliosa that the two taxa might be treated as convarietal races under the name var. foliosa” (Semple 1996, p. 114). The geographic range of var. ballardii lies within that of var. villosa sensu Semple. [lac]. Heterotheca villosa var. minor (Hook.) Semple [1994] Chrysopsis villosa var. minor t look. [1847, TYPE: WYOMING. Sweetwater Co] Diplopappus hispidus Hook eee TYPE: SASKATCHEWAN. Carlton House] Ch rysopsis villosa var. hispida (Hook.) A. Gray [1884] Heterotheca villosa var. hispida (Hook.) Harms [1974] Canadian botanists have commonly recognized two varieties of H. villosa, var. villosa (leaves and involucres with appressed, nonglandular vestiture) and var. hispida (leaves and involucres with spreading hairs intermixed with glands), but none has indicated that the taxa are geographically or ecologically distinct or that intergradation and typological identifications are not prevalent in this ~~ region. 1b. Heterotheca villosa var. pedunculata (Greene) Harms ex Semple [1987] Chrysopsis pedunculata G [1900, TYPE: COLORADO: Archeluta Co.] ?Heterotheca zionensis Sem ple [1987. TYPE: UTAH. Washington Co]] Representing the densely strigose, silvery-sericeous expression of the H. villosa complex; common in Utah, northern Arizona, and the Four Corners region. As noted above, I have not been able to distinguish H. zionensis from var. pedunculata, based on criteria established by Semple. 1c. Heterotheca villosa var. angustifolia (Rydb.) Harms [1968] Chrysopsis angustifolia Rydb. cea TYPE: NEBRASKA. Hooker Co] Chrysopsis villosa var. angustifolia (Rydb.) Crongq. [194 Heterotheca stenophylla var. quantal (Rydb.) Semple [1994] Allopatric and distinct from other expressions of H. villosa, at least in Texas, Oklahoma, and most of Kansas, intergrading in Nebraska, South Dakota, and North Dakota. 2. Heterotheca horrida (Rydb.) Harms [1968] Chrysopsis horrida Rydb. [1904, TyPE: COLORADO. Weld Co.] Heterotheca villosa var. nana (A. Gray) Semple [1994] Chrysopsis canescens var. nana A. Gray [1849, TyPE: NEW MEXICO, ?Mora Co. NESOM, TAXONOMIC OVERVIEW OF MPLEX 379 Ww . Heterotheca sierrablancensis (Semple) Nesom [2006] Heterotheca villosa var. sierrablancensis Semple [1996, TYPE: NEW MEXICO. Lincoln Co. > . Heterotheca polothrix Nesom [2006] Chrysopsis villosa var. scabra Eastwood [1896, TyPE: UTAH. San Juan Co] Heterotheca villosa var. scabra (Eastwood) Semple [1992 . Heterotheca depressa (Rydb.) Dorn [1988] Chrysopsis depressa Rydb.[1900, TYPE: WYOMING. Teton Co|] Heterotheca villosa var. depressa (Rydb.) Semple [1994] WN ACKNOWLEDGMENTS All pertinent specimens at BRIT/SMU, MO, and TEX/LL were studied, and a loan from SJNM was helpful in providing a base of study for plants of the Four Corners region. In all, more than 1350 specimens of the Heterotheca villosa com- plex were examined. Additionally, I have examined all collections of all taxa of Heterotheca at these herbaria. Staff at CAS provided an image of the holotype of Chrysopsis viscida subsp. cinerascens and made observations of details of its vestiture. Robert George lent his digital proficiency toward production of the maps. Constructive review comments of Vernon Harms and an anonymous reviewer are greatly appreciated. Postcript.—Dr. John Semple has read some or all of the manuscript, and in any case, he is intensely opposed to the approach and its conclusions. But it has not seemed necessary to re-assemble and re-cite the massive number of specimens (10,300) Dr. Semple had on hand for his monographic study of sect. Phyllotheca. He has noted that his treatment is based on multivariate analyses (yet unpub- lished), and no statistics are found here (nor do I believe that taxonomic con- clusions based on such an approach must necessarily be correct). I do not have the long and focused field experience he has had with this group. Still, because of our strong and basic differences in taxonomic philosophy (regarding con- cepts of species and varieties) and apparently because of differences in the way in which we perceive and interpret patterns of variation, it obviously is pos- sible to differ in taxonomic conclusions regarding the same group of plants. We agree on the delimitation of some of the taxa but not others. For those we agree should be formally recognized, we disagree at what rank some should be treated. If this taxonomic overview is seen as no more than a “superficial opinion-piece,” at least I have tried to provide a clear rationale for the basis of my opinions and why they differ from those of Dr. Semple. The alternative taxonomy proposed here allows me to identify plants of the Heterotheca villosa complex in a con- sistent and biologically meaningful way, from my point of view. Others also have a choice. 380 BRIT.ORG/SIDA 22(1) REFERENCES Dorn, R.D. 1988.Vascular plants of Wyoming (ed. 1). Mountain West Publishing, Cheyenne, Wyoming. GREAT PLAINS FLorA Association. 1986. Flora of the Great Plains. Univ. Press of Kansas, Lawrence. Harms, V.L. 1968. Nomenclatural changes and taxonomic notes on Heterotheca, including Chrysopsis,in Texas and adjacent states. Wrightia 4:8-20. Harms, V.L. 1970. Heterotheca. In: D.S. Correll and M.C. Johnston. Manual of the vascular plants of Texas. Texas Research Foundation, Renner, Texas. Pp. 1563-1569. Markow, S. 2001.Report on a survey for Stepbhanomeria fluminea in Grand Teton National Park. Prepared for the Greater Yellowstone Network Inventory and Grand Teton Na- tional Park. Wyoming Natural Diversity Database. Accessed February 2006. Markow, S. 2004. Survey for Stephanomeria fluminea on the Bridger-Teton National Forest. Prepared for the Bridger-Teton National Forest ae the BealOe 4 is Service Office. Wyoming Natural Diversity Database. Accessed Fabris 2006. Nesom, G.L. 1997. Review: “A revision of Heterotheca sect. Phyllotheca (Nutt.) Harms (Compositae: Astereae)" by J.C. Semple. Phytologia 83:7-21. SemPte, J.C. 1974. The phytogeography and systematics of Xanthisma texanum DC.: proper usage of infraspecific categories. Rhodora 76:1-19. Sempte, J.C. 1987. New names, combinations, and lectotypifications in Heterotheca (Compositae: Astereae). Brittonia 39:379-386 Sempte, J.C. 1990. Neotypification of Amellus villosus, the identity of a Bradbury collection, and typification of some other goldenasters (Compositae: Astereae). Brittonia 42: 22) 5228. Sempte, J.C. 1992. The goldenasters of California, Heterotheca (Compositae: Astereae names and combinations. Phytologia 73:449-455, SemPte, J.C. 1994. New combinations in the Heterotheca villosa (Pursh) Shinners complex (Compositae: Astereae). Novon 4:53-54. Sempte, J.C. 1996. A revision of Heterotheca sect. Phyllotheca (Nutt.) Harms (Compositae: Astereae). Univ. Waterloo Biol. Ser. 37:i-iv, 1-164 Sempe, J.C. 2006. Heterotheca (Asteraceae: Astereae). In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. Oxford Uni- versity Press, New York and Oxford. Vol. 20. WetsH, S.L. 1982. Utah plant types——historical perspective 1840 to 1981-—annotated list, and bibliography. Great Basin Naturalist 42:129-195, WS new TAXONOMY OF LANTANA SECT. LANTANA (VERBENACEAE): 1 CORRECT APPLICATION OF LANIANA CAMARA AND ASSOCIATED NAMES Roger W. Sanders Botanical Research Institute of Texas 09 Pecan Street Fort Worth, Texas 76102-4060, U.S.A. rsanders@brit.org ABSTRACT The previous lectotypification of Lantana camara L. is evaluated by examination of characters of the lectotype, review of oo original material, and documentation of current usage of the name. The current usage is analyzed by surveying Peranieat acu and by eeu specimens for an- notations between 1753 and the q t f those specimens. Current usage of L. camara includes a eadely cultivated and naturalized cultigen species of hybrid origin that is taxonomically distinct from L. camara. To determine the correct name of the cultigen, al names in Lantana’ sect. Lant d, typil Hes if TISCESSAE YE and Bessils and son amucaMy disposed. h il i I | is K W. Sanders. ace of the oaea De brids in the L.camara ee is hypothesized. oa new com- p. aculeata (L S R. me paneets and Lantana nivea Vent. Subse. mutabilis (WJ. Hook ) R.W. Sanders. Nineteen | RESUMEN La lectotipificacion previa de Lantana camara L. se evaltia mediante examen de los caracteres del lectotipo, revision de otro material original, y documentacion del uso actual del nombre. El uso ac- tual se analizé revisando la bibliografia adecuada y por muestreo de especimenes entre 1753 y el nte por comentarios en relacion a caracteres criticos de esos especimenes. El uso actual de L. y naturalizada de origen hibrido que es pres camara incluye una especie ampliamente cultivada smi dif L. camara. Para determit el nominee correcto ne la planta cultivada los | bre de Lantanasect. Lantana osible, y taxonomicamente decidido. No hay nombres disponibies as . mana eu a nas se describe como Lantana strigocamara R.W. Sanders. Se hace una en el complejo L. camara. Se hacen dos an ees ok Daven camara subsp. aculeata (L.) R.W. Sanders y Lantana nivea Vent. subsp. mutabilis (WJ. Hook.) R.W. Sanders. Se realizan diez y nueve lectotipificaciones y siete epitipificaciones. Plants that have gone under the Linnaean name Lantana camara L. are well known, not only as hardy summer ornamentals worldwide but also as natural- ized weeds having devastating economic impact in humid tropical areas of the Old World (Wolfson & Solomons 1964; Howard 1970; Stirton 1977; Swarbrick et al. 1995; Day et al. 2003). The systematics of these cultivated and naturalized plants is not well understood, even though considerable effort has been made to elucidate their biological control (Day et al. 2003). This confusion arises in SIDA 22(1): 381 — 421. 2006 382 BRIT.ORG/SIDA 22(1) part from the convoluted history of exploration, cultivation, hybridization, and artificial selection that began at least 60 years before the publication of Linnaeus’ Species Plantarum (1753). Records that have been compiled (Howard 1969; Stirton 1977) suggest that, during the eighteenth century, fanciers hybridized different wild species and infraspecific taxa of Lantana L. sect. Lantana from Mexico, the West Indies, and Brazil. As shown in this paper, the early validly published names were based on garden-grown material (almost entirely so until 1817 and com- monly so into the 1850s), much of which consisted of hybrid combinations. As herbarium specimens of wild-collected species of sect. Lantana became available after the early 1800s, many of the available, poorly distinguished names were frequently misapplied to them. Adding to this confusion was the introduction of cultivated hybrids into neotropical regions where indigenous taxa occur. Due to the propensity of lantanas to undergo polyploidy and the partial fertility of odd polyploid levels (Natarajan & Ahuja 1957; Khoshoo & Mahal 1967; Spies 1983, 1984; Spies & Stirton 1982a, b, c; Sanders 1987a, b), even more complex hybrids formed between indigenous taxa and the escaped hy- brid cultigens (Sanders 1987a, b, c, 1989a). Thus, the limits of natural variation have been obscured, a the ability of taxonomists to develop effective classifications for the grou Schauer (1847), Br ae (1895), and Troncoso (1974) developed current sec- tional concepts. Species of Lantana sect. Lantana (=sect. Camara Cham.) are characterized by predominantly narrow floral bracts that are somewhat in- conspicuous among the tubular corolla bases, by usually yellow or orange pig- mented corollas (white-flowered populations known in several species [unpubl. observ. see also discussion below concerning purplish pigments in the section), and by blackish drupes. Each drupe contains a characteristically inflated com- pound endocarp that resembles a horse’s skull in which the seed chambers are in the position of the eye sockets. Some of the species of the other major section, Lantana sect. CallioreasCham., might be confused with those of sect. Lantana. Generally, species of sect. Callioreas are described as involucrate, usually hav- ing ovate to reniform conspicuously imbricate floral bracts and purplish co- rollas. However, fruits are needed to insure correct sectional placement—drupes are usually white or purplish and endocarps are subglobose, bilobed (seed chambers fill each hemisphere), noninflated, and reticulately ornamented. Schauer (1847) published the only worldwide revision of Lantana in De Candolle’s Prodromus. All more recent work has been limited to regional and garden floras. The foremost student of Verbenaceae in the twentieth century, Harold N. Moldenke, never produced a revision of Lantana, but he did describe a number of new species and infraspecific taxa. Horticulturalists, ecologists, and some floristic taxonomists have effectively submerged much of Lantana sect. Lantana into L. camara, treating this wide spectrum of variation asa single SANDERS, TAXONOMY OF LANTANA 383 species, (e.g., Kuntze 1891; Troncoso 1965, 1974; Bailey Hortorium 1976; Schemske 1976, Huxley et al. 1992; Cullen et al. 2000). Sanders (1987a, b, c, 1989a, b) undertook a study of the variation of wild and naturalized populations of Lantana sect. Lantana in Florida and parts of the West Indies. By correlating chromosome numbers and meiotic behavior with morphology, he was able to distinguish the natural (often diploid) wild taxa from the naturalized and spontaneous hybrids and to develop morphological taxonomic criteria. Sanders found that plant architecture, leaf and bract size and shape, indument features, inflorescence/infructescence development and size, fruiting bract persistence, and floral pigment classes and dosage effects correlate well with cytology, geographic distributions, and sf delimitations. On the other hand, he argued that development of prickles and specific corolla colors, characters relied upon by many other authors, vary too widely within and among taxa to be effective criteria, a conclusion recently supported by molecular studies (Scott et al. 1997; Day et al. 2003). Further support comes from the work of Isidro Méndez S. of Cuba who took up the study of Cuban and West Indian Verbenaceae (1992, 1993, 2002). Méndez explicitly accepted the taxo- nomic criteria proposed by Sanders for Lantana sect. Lantana. The natural taxa of sect. Lantana fall into three separate phenetic groups (to be validated in a subsequent paper; cladistic status not yet determined) on the basis of trichome structure on the abaxial surface of the leaf blades, as well as shape, adaxial luster, and venation of the leaf blades. The “pilose-morph” species (Pilose Group, Fig. 1) are cl terized by soft, filiform, spreading (erect or curly) hairs usually densely disposed on all veins and intervening tissue. Most of these have leaf blades that are broadly ovate to narrowly deltate, rounded to cordate at the base and acuminate to obtuse at the apex, dull above (epidermis poorly reflective), and more or less pinninerved. The “strigose-morph” species (Strigose Group, Fig. 2) have the abaxial hairs developed as conspicuous strigae, i.e. the hairs are stout, conical, antrorsely geniculate, and are scattered and re- stricted to the midrib, and secondary, tertiary, and sometimes higher order veins, but not to the intervening tissue. Leaf blades are usually ovate-elliptic, often tapering at base and apex, lustrous above, and more or less triplinerved. The “setose-morph” species (Setose Group, Fig. 3) abaxially bear well-separated, long, flexible to subrigid, spreading, setiform trichomes that are more or less restricted to the midrib and secondary and tertiary veins. Otherwise they are like the pi- lose-morph species. Thus, even though all three groups have adaxial strigae, the Stigose Group is the only one with strigae on both surfaces. Despite Méndez’s (2002) acceptance of Sanders’ criteria, he disagreed with Sanders over the application of the name L. camara. Because L. camara is the proposed type of the genus (Jarvis 1992; Jarvis et al. 1993) and one of the two oldest names in the secton, the correct application of most other names in the BRIT.ORG/SIDA 22 = a LM ey oy | =! Fic. t nen, paises INE ie A), ee Group. a. inflorescence. b. rey tative leaf blad section hinge upon the correct application of L. camara. Asserting that Sanders had committed two errors, Méndez stated: Sanders (1989a]) attributed this binomial [L. camara] to a phenotype very different from prevailing opinion among the authors later to Linnaeus and not in agreement with the lectotype selected by Moldenke & Moldenke (1983) However, until the present study, analysis of the lectotype and subsequent ap- plication of the name have not been straightforward. The critical characters were not visible on the lectotype since all of its leaves were mounted adaxial side up. Sanders (unpubl.) was not able to determine whether the lectotype matched the pilose-morph or the strigose-morph plants, especially given the quality of the microfiche and xerographic images available at the time. How- ever, because H. N. Moldenke had selected the type, Sanders (unpubl.) relied primarily on the numerous annotations of Dr. Moldenke (Table 1, App. B, see also Moldenke 1980a) to develop a concept of the species and apply the name. Sanders’ concept was further reinforced by annotations of C.D. Adams, and the 1) SANDERS, TAXONOMY OF LANTANA 385 Fic. 2.Lantana scabrida,| (Soland ?,,BM) ing Stri G a.infl { ae £. L on | £ £ c Scale bars:a = 1cm;b&c=1mm. leaves. b. treatments of Lantana in The Flowering Plants of Jamaica (Adams 1972) and A Flora of Tropical Florida (Long & Lakela 1971), as well as on identified culti- vated material received under that name on the U.S. National Arboretum, Longwood Gardens, and tabl | nurseries. In all cases, the plants were characterized by leaves dominated by strigose-morph hairs, cordate-ovate blades, and dull upper surfaces! (here subsequently called the “Strigose-Cor- date-Dull-” or “SCD cultigen”, Fig. 4). In the West Indies, some wild-collected specimens bear codominant mixtures of both hair morphs. However, cytologi- cal study confirmed these to be hybrids between the SCD cultigen and native taxa of the Pilose Group (Sanders 1987b). Thus, Sanders concluded that the lec- totype had the strigose-morph hairs, and he applied the name L. camara spe- cifically tothe SCD cultigen. Many of the described species in the Pilose Group (including L. horrida, L. tiliifolia,and L. arida) he lumped into a single species and applied the name L. urticifolia Mill. Méndez appears to have studied more recent, higher quality images in con- nection with a project to lectotypify Linnaean species of Verbenaceae of the West Indies in collaboration with Steve Cafferty of the Linnaean Typification Project (Méndez & Cafferty 2001). Although he, too, was unable to see the abaxial surfaces, he concluded on other grounds that the lectotype matched the pilose- 'Adams included in his concept of L.camara the SCD cultigen and L. scabrida. The few specimens of L. scabrida that | studied and bear his annotation as L. camara actually have lustrous upper leaf surfaces. 386 BRIT.ORG/SIDA 22(1) al al Fic. 3.1 hii (Hi) 20499, TEX) ing $ a.inflorescence.b J t leaf surface. Scale bars:a=5 mm;b&c=1mm. morph, not the strigose-morph plants. He applied the name L. camara to the taxon to which Sanders applied the name L. urticifolia. If Méndez’s two assertions merit redress, then another, later-published name should be applied to the SCD cultigen that Moldenke, Adams, and Sanders called Lantana camara. Of course, current usage, regardless of usage by authors of the early post-Linnaean period, must be established. If current usage of L. camara has clearly replaced the Linnaean concept with the concept of the SCD culti- gen, then conservation of the nomenclatural type could serve as an alternative solution. Therefore the purpose here is to |) evaluate the lectotypification by review- ing its publication, by clarifying the characters of the lectotype, by understand- ing current usage of L. camara, and by determining any variance between the type and current usage; 2) affirm the correct application of that name, 3) if nec- essary, determine which other name is to be applied to the SCD cultigen, and 4) dispose of all other names, known to me, that are applicable to the complex. Typification of Lantana cama Moldenke and Moldenke (1983) effectively lectotypified Lantana camara by citing LINN 783.4 as “type” (ICBN, Art. 7.11, Greuter et al. 2000). This lectotypification was accepted by Sanders 1989b) and Méndez (2002). In March, 2004, | visited BM, and Dr. Charles Jarvis, long-time participant in the Linnaean Typification Project, consented to break the lower stem to permit study of the abaxial surface of the lowest two leaves (Fig. 1). The type, indeed, is of the pi- lose-morph and is verified by leaf shape and inflorescence structure as an ele- ment of the naturally occurring species distributed from the Bahamas and Greater Antilles, through Mexico south to northwestern South America. Thus of Méndez’s two assertions, the one that “Sanders (1989) attributed this bino- mial toa phenotype...not in agreement with the lectotype selected by Moldenke and Moldenke (1983)” is correct. Taste 1.Data documenting history of annotation by H.N.Moldenke and other taxonomists familiar with the classification of Lantana."v.moritz” indicates plants annotated as L. camara var. moritziana. See text for further discussion. Years Authorities N Pilose % Mixed % Strigose % Setose % 1981-2005 Moldenke (incl. v. moritz) 62 15 24% 20 32% 21 44% 0) 0% Moldenke (excl. v. moritz) 47 0 0% 20 43% 27 57% 0 0% tner 45 ¢) 0% 25 56% 20 44% @) 0% All (incl. v. moritz) 107 15 14% 45 42% 47 44% 0 0% All (excl. v. moritz) 92 0 0% 45 A9% 47 51% 0) 0% 1956-1980 Moldenke (incl. v. moritz) 153 ifs) 8% 56 37% 83 54% i 1% Moldenke (excl. v. moritz) 14] ] 1% 56 40% 83 59% ] 1% Other 60 0 0% 4] 68% 19 32% 0 0% All (incl. v. moritz) 213 13 6% 97 46% 102 48% ] 0% All (excl. v. moritz) 201 ] 0% 97 48% 102 51% ] 0% 1931-1955 Moldenke 16 2 13% 6 38% 7 44% ] 6% Other ] 0 0% 0 0% ] 100% @ 0% All 17 2 12% 6 35% 8 47% ] 6% 1906-1930 All 16 0 0% 10 62% 6 38% 0 0% 1881-1905 All 4 0) 0% 3 75% ] 25% Q 0% 1856-1880 All 2 0 0% 0 0% 2 100% 0 0% 1831-1855 All 3 1 33% ] 33% ] 33% 0 0% 1753-1830 All 15 10 67% 2 13% 1 7% 2 13% VNVINV1°L35 VNVLNVT JO AWONOXVL‘SUIONYS L8€ 388 BRIT.ORG/SIDA 22(1) b. inflorescence Development of current usage Linnaeus’ concept of L.camara was developed from an array of cultivated and horticulturally selected plants. The synonyms and illustrations that he cited in the protologue are based on vouchers that are primarily elements of the Pi- lose Group. All Hortus Cliffortianus (1737) specimens are either the same spe- cies as LINN 783.4 (319 Lantana 1-B!, Linn. Herb. specimen at S [a cut-down Herb. Cliff. specimen, C. Jarvis, pers. comm., dig. photo!]), are hybrids between that taxon and the Strigose Group (319 Lantana I), or are elements of L. horrida SANDERS LANTANA SECT. LANTANA 389 Kunth (319 Lantana 1-C), 1-D!, and 320 Lantana lol[the latter possibly hybrid- ized with the Strigose Group]). Moreover, an unnumbered Herb. Cliff. sheet (!) is an element of the Strigose Group (L. splendens Medik.). Linnaeus did not see the vouchers for the syntype illustrations. One of these (icon in Plukenet, Phytographia 385. t. 114, f. 4. 1691. [Voucher: 98:143 top-left specimen, BM-SL!]), belongs to the Setose Group (L. hirsuta M. Martens & Galeotti). The Commelin syntype has no known voucher (C. Jarvis, pers.comm.), illustrates only an inflorescence, and cannot be placed to trichome-morph. Even so, the trichome characters may not have been of much concern to Linnaeus. For example, in 1767, he did segregate L. mista, which has distinctly hispid twigs, as depicted in a Dillenius plate (see App. B), which he cited. However, Linnaeus did not use this character to differentiate the new species, but rather used the protracted leaf-blades, longer bracts, and capitula with mixed corolla colors. According to C. Jarvis (pers. comm.), no other original material is known. To determine how this somewhat broad Linnaean concept was modified by later botanists, I present two lines of evidence. The first isa survey of sampled specimens to determine annotation patterns by taxonomists (Table 1, Fig. 5). The specimens are all those annotated by H. N. Moldenke as L. camara in LL/ TEX (including the Moldenke Herb.) and BRIT/SMU, a selection of those at K, and a few for which | obtained photographs at BM, LINN, and OXE Besides Moldenke, annotating authorities include: Linnaeus, Medikus (implied by cita- tion, 1775), Schauer, Urban, Merrill, Hutchinson, R. Meikle, J. K. Morton, R. Fernandes, B. Verdcourt, G. Bromley, and S. Atkins. Dr Méndez and I are excluded. Plants annotated as L. camara (or in a few cases as L. aculeata, see App. B) were scored for whether trichomes on the abaxial surface were pilose-morph only, strigose-morph only, setose-morph only, or codominant mixtures of stri- gose- and pilose-morphs (including rare mixtures of strigose- and setose-mor- phs). Between the Linnaean period, when L. camara was applied primarily to pilose-morph plants, and the late nineteenth century, specialists began to ap- ply the name primarily to the strigose-morph plants or those with mixed tri- chome morphs. For those annotated specimens from the neotropics, the mixed trichome plants are presumed hybrids primarily between the naturalized SCD cultigen and native species of the Pilose Group. In the paleotropics, the mixed- trichome plants appeared to be either 1) hybrids between escaped pilose-morph plants and the SCD cultigen, 2) escaped cultivars selected from hybrids between L. camara and various species of the Strigose Group, or 3) naturalized plants consisting of complex spontaneous hybrids between plants of the preceding two categories. As this annotation trend developed, the plants of the Pilose Group began to be annotated with other later names. In particular, note the quotation of H. Moldenke [apparently in personal correspondence] by Howard (1969): 390 BRIT.ORG/SIDA 22(1) BO% 70% ‘ | wn 60% a ! | a} . ra x | 3 \ ig . | hen * ~ 50% s < | seed ? ee -_=- * * = 2 : oe. Ren ee =A £ nad 4 fa o- T 1763-1830 1831-1905 1906-1930 1931-1955 1956-1980 1981-2005 Year —e—Pilose - @ - Mixed —a — Strigose — - Setose —e—Pilose +mrz - & - Mixed+mrz —tz, — Strigose + mrz Fic. 5. Plot of portion of data from Table 1, showing historical pattern of specialists’ application of the name Lantana camara to pilose-morph taxa (mostly L. camara, some L. horrida and their hybrids) vs. strigose-morph taxa (mostly L. strigocamara, L. nivea, and L. sanslouel vs, oe eibiibab the two groups une morph plants) vs. setose-morph plants (mostly L. hirsuta). “+ mrz f L. camara var. moritziana. You accuracy of identification on the labels of plant a marked ‘Lan- tana camara’ in ne | have found that a large percentage of such material is actually L. moritziana, L.glandulosissima, L. scorta, L. horrida, L. arida, L. glutinosa, etc. etc In other words, authorities (e.g... Moldenke and others) attributed specimens of L. camara to other species names referable to the Pilose Group. Surprisingly, in the mid 1970s, Dr. Moldenke accepted Lopez-Palacios’ (1974) reduction in rank of L. moritziana asa variety of L. camara. If that anomaly is excluded from the analysis, then the application of L. camara to strictly pilose-morph plants by taxonomic specialists is negligible over the last 50 years. The second line of evidence is a survey of the literature (App. A). Adequate descriptions of leaf indument are scarce prior to the 1850s, but those by Medikus (1775), Sprengel (1825), and Schauer (1847, 1851) all indicate a prevailing accep- tance of plants with the abaxial surfaces soft-hairy. The first publication di- verging and treating strigose-morph plants as L. camara is by Otto and Dietrich (1841). This reflected a growing confusion as to what L. camara really is. By the SANDERS, TAXONOMY OF LANTANA 391 beginning of the twentieth century, most horticultural taxonomists took a broad concept of L. camara to include both pilose- and strigose-morph (or even se- tose-morph) plants in the species. This sentiment is reflected in Bailey’s Cyclo- pedia (1900) Lantanas have been long in cultivation, and it is difficult to refer the garden f botanical spe- cies. The species themselves are confusing. Most of the garden kinds are of the L. Car mara type. There are several Camara-like species which probably have pbudized to produce these forms; but Voss ] (1896], the latest garden monographer, regards these s] forms of L. Camara (preferring, however, to use the name L. aculeata). Accepting L. Camara in Voss’s sense, the garden lantanas may be said to be derived from that species; and this view is adopted below. Over the last 100 years, most floristic taxonomists (and hence, ecologists) have developed a similar attitude and broad concept or have followed Moldenke (1970) and Adams (1972), applying L. camara to the SCD cultigen and its hy- brids, as has Sanders. Therefore, Mendez’s other assertion that “Sanders (1989) attributed this binomial toa phenotype very different from prevailing opinion among the authors later to Linnaeus” is inaccurate. Disparity between the lectotype and current use The disparity between the lectotype and current use can be viewed in two ways. Either the current usage 1) includes the type and encompasses both pilose- and strigose-morph plants, or 2) excludes the type and is applied only to the SCD cultigen (and its hybrids with other species). The first view is problematic be- cause the SCD cultigen behaves as a species distinct from the species named L. camara by Linnaeus. The second view raises the issue of relectotypifying L. camara to fit current usage. This might be preferable if LINN 783.4 were a hor- ticultural hybrid no longer extant. However, the type is an element of a wide- spread wild species to which the name L. camara has been properly applied, at least in part, by taxonomists Standley (1924), Leon & Alain (1957), Macbride (1960), Gibson (1970), Nash & Nee (1984), Lopez-Palacios (1977), Méndez (2002), and many ecologists. Futhermore, neither original material nor vouchers for cited illustrations consists of the SCD cultigen, and most of these specimens are of the pilose-morph (see preceding section). Therefore, overturning the choice of LINN 783.4 is not supported. Correct name of the widespread SCD cultigen To rectify the common practice of misapplying the name Lantana camara to the widespread SCD cultigen and to implement Méndez’s recommendation that “the system established by Sanders around L. camara should be reinterpreted,” an attempt has been made to locate, to examine and, when needed, to choose nomenclatural types of all other published names in sect. Lantana (App. B). The results demonstrate that other names, also, have been misapplied by a num- ber of specialists, including Schauer, Moldenke, Sanders, and Méndez. With regard to the abaxial leaf indument, the SCD cultigen is essentially 392 BRIT.ORG/SIDA 22(1) outside the range of variation of Lantana camara but within the range of the Strigose Group (Fig. 4, 12). Thus, it would be imprudent to submerge this widely naturalized, aggressive species of hybrid origin as an infraspecific taxon of L. camara. Rather, the SCD cultigen more likely has a complex parentage involving L. nivea Vent. L. scabrida Sol. in Aiton, and L. splendens of the Strigose Group, as well as L. camara, and possibly other species (noteably, L. hirsuta of the Setose Group). Morphologically, it retains only the leaf shape and adaxial surface dull- ness of L. camara. Given the protracted history of collection and publication of new taxa, it is surprising that none of the available names can be verified as ap- plying to this widespread cultigen. Apparently, this is due to there being so many published names and to the confused state of the taxonomy of sect. Lantana. Through uncritical analysis of vague descriptions, one could argue that a few of the names that lack known types could be applied to the cultigen. How- ever, taxonomic stability is more important than unsubstantiated priority. As- signing a neotype toa name whose concept can never be known with certainty in order to honor early taxonomists, who never understood the complexity of these plants, would be imprudent. Such names were never taken up and have no precedence of usage in the horticultural and ecological communities that taxonomists attempt to serve. As a species of hybrid origin, of which that origin remains obscure, the SCD cultigen must be treated as a species nova. To provide taxonomic and bib- liographic continuity, thus maximizing stability, the new epithet proposed here is based on the root “-camara.” To indicate its hybrid nature and complex hy- pothesized parentage involving several species of the Strigose Group, it is given the prefix “strigo-"to produce Lantana strigocamara. At least two infraspecific epithets, and perhaps others, could serve as basionyms. However, epithet prior- ity is mandatory only within rank, and I chose a new species name. Lantana camara L. var. rubella Moldenke (1949) is not widely known, and applying it to the whole of the species would broaden Moldenke’s concept considerably. Lan- tana flava Medik. f. sandersii Méndez (2002) is only four years old and has not been taken up outside Méndez’s own work. Raising it to species level would be tantamount to honoring myself. — Lantana strigocamara R.W. Sanders, sp. nov. (Fig. 4). Type U.S.A. Floripa: Dade Co. near intersection of Montgomery St. and Old Cutler Rd, Montgomery Botanical Center, dis- turbed edge of rock pineland, 23 Sep 1981, Sanders 1450 (HOLOTYPE.: FTG! [Dig. photo! see Fairchild Tropical Botanic Garden 1999}, IsoTYPE: NY!; Drawing, Sanders 1987, fig. 9). Credita hybrid le Lant d L.etL.nivea Vent. et L.scabrida Sol. in Aiton et L. splendenti Medik. et L. hirsuta M.} ns& ti mixta simulat speciem. Differta 1 ‘a laminis paces euis str dees s, ichomatibus remotis non nisi nervis mediis secondariis tertiariisque geniculatis; a L. nivea et L. scabrida et L. splendenti laminis magis rugosis, ee abruptius contracti ypInnInery ibus, oe adn hebetibt bus ma nitidis; a L. at Ns nivea trichomatibus | vel aureis; a o x t SANDERS, TAAUNVIVIT UF LANTANA SECT. LANTANA 393 amnelieaue | Strict to lax, ae erect, iid or ene shrub to 3 m tall (climbing much higher in some areas of the paleotropics); twigs hirtellous, angled, with or without prickles. Petioles 5-10 mm long; blades cordate to ovate, 3-10 x 2-6 cm, mostly 1-L7 times longer than wide, the base abruptly narrowed onto peti- ole, the apex acuminate, moderately rugose between higher orders of veins, dull above; hairs of adaxial laminar surface 0.3-0.7(-1.2) mm, scattered, in the form of antrorse strigae, often from pustulate bases; hairs of abaxial laminar surface usually 0.5 mm or less, stout, in the form of attenuately conical, geniculately antrorse strigae, but not closely appressed to surface, not deciduous, usually restricted to and scattered on midrib, secondary and tertiary veins (in some populations small erect hairs 0.1-0.2 mm long weakly developed on smallest veins and along crevice under larger veins); abaxial sessile glands green or not visible with 10 lens; teeth 15-25 per side, obtuse to short-acuminate, not revo- lute. Inflorescences 2-3 cm in diameter; peduncles 2-10 cm long; receptacle fistulose; bracts mostly 3-8 mm long, subulate, long-triangular or narrowly lanceolate (single outer series sometimes longer and narrowly spathulate or rarely subfoliaceous), apically attenuate, puberulent or hirtellous abaxially only, deciduous in fruit. Calyx 3 mm long, membranous; corolla yellow, orange-red, white, or pink to deep reddish purple and often mixed with orange or crimson, the throat often yellow, the tube 7-12 mm, slightly curved, the limb 6-10 mm across. Drupe ca. 5 mm across, blue-black; endocarp obovoid, inflated. Paratypes: BURMA: Chantaburi, Makam forest, 26 Aug 1966, Larsen et al. 1841 (AAU, LL!). CHINA. Kwangsi: Kweilin, cult., 1979, Wan & Chow ates L!). INDIA. Lucknow, cult., 28 Oct 1974, Shivarajan 85750 (LL!). PAPUA NEW GUINEA. East Sepik . Wewak, 31 Aug 1968, Krauss 1287 (LL!). SRI LANKA. Sabaragamuwa Prov.: Kegalle: Klnahatene 12 Feb 1974, Moldenke et al., 28331 (LLLUSI). UGANDA. Paraa, 29 Jul 1972, Moldenke & D ke 26073(LL!). USA. Florida. Dade Co.: Homestead, 18 Jul 1966, Smith FI. 6 (BRI, LL!). New York. nate Co.: Yonkers, cult., 23 Sep 1950, Moldenke 21133 (LL!) Hypothesis of the history of the cultivated Lantana camara complex Prior to the Linnaean period, the following species (citations in App. B) were already in cultivation in Europe, as deduced from the Sherard specimens at OXF, the Sloane and Clifford Herbaria at BM, and the Linnaean Herb. at LINN and S: Pane a (Fig. 1, West Indies and Mexi h outh America) Pilose group Lan Lantana. horrida (Fig 8, aes aa id, Mexico to eons Strigose group Lantana nived (Fig. 7, southern Brazil to ae Lantana scabrida (Fig. 2, West Indies, Mex Lantana splendens (Fig. 6, Bahamas) Setose group — Lantana hirsuta (Fig. 3, Mexico) All these species, except L. nivea, are characterized by capitula single in the leaf axils with stout peduncles and with corollas opening yellow and changing 394 BRIT.ORG/SIDA 22(1) Fic. 6. Lantana splendens, epitype (Herb, Sherard 1269, OXF), a. infl f i b. adaxial leaf surface.c. abaxial leaf surface. Scale bars:a = 1 cm;b &c= 1 mm. Photogray pplied by yf to darker yellow, orange, or red-orange, and are aculeate or not. To my knowl- edge, Lantana nived is exceptional among the natural species in having capitula commonly paired in the leaf axils with filiform peduncles and with corollas opening white and aging bluish, rose or purple, or opening pink and aging pur- plish. The plants are usually aculeate with recurved prickles, sometimes fiercely so. Therefore the only source in the cultivated complex for capitula opening yellow or cream and changing to purplish reds or orange plus purple would be hybridization between L. nivea and one or more of the other species. Parentage involving L. nived is also consistent with stoutly recurved prickles in some of the cultivated hybrids. Apparently by backcrossing L. camara into such hybrids, plants with the mixed-colored capitula were developed within the range of variation of L. camara with respect to other characters (Fig. 9). Such plants | consider to be the subspecies combination L. camara subsp. aculeata. lam rec- ognizing subspecies here and elsewhere in the complex for the following rea- sons: |) the rank varietas in Lantana sect. Lantana has been overused for culti- vars, and the varietal names are largely misapplied; 2) there may be yet undiscovered variety epithets that would take priority and further destabilize the nomenclature; and 3) | recognize the subspecies rank for geographically widespread infraspecies that comprise varieties, and further research will likely reveal localized geographic races that may warrant recognition at the rank of varietas. According to the ICBN (Greuter et al. 2000), this hybrid taxon cannot be treated as a nothosubspecies, but rather Art H3.3 would require treating it at the nothospecies level. However, it does not behave as a species distinct from L. camara, and it is the major representative of the species where it is naturalized, thus, simulating a widespread natural subspecies. Lantana camara L. subsp. aculeata (L.) R.W. Sanders, comb. & stat. nov. (Fig. 9). BASIONYM: Lantana aculeata L., Sp. Pl. 627. 1753. Lectotype designated by Méndez & Cafferty (2001, see App. B). SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 395 “s ry »* “* a Fic. 7. Lantana nivea, ef tative | infl itt jJuncle, and inset of ve {Cioht t t eres fy £ , C),a. stem I inflorescence. b f surf c. abaxial leaf Scale bars:a =1cm;b&c=1mm. Likewise, backcrosses of L. nivea into the original mixed-color hybrids produced multicolored plants within the range or extending the range of variation of L. nivea with respect to other characters. Such plants I consider to be the subspe- cies combination L. nivea subsp. mutabilis. It is not treated as a nothotaxon by similar reasoning. Lantana nivea Vent. subsp. mutabilis (WJ. Hook.) R.W. Sanders, comb. & stat. NOV. BASIONYM: Lantana nived Vent. var. mutabilis WJ. Hook., Bot. Mag. 5: pl. 3110. 1831. LEc- TOTYPE (here designated): icon in WJ. Hook., Bot. Mag. 5: pl. 3110. 1831. Lantana nivea subsp. mutabilis may have been an important genetic vector for the origin of L. strigocamara. Even so, this taxon is not to be identified with L. strigocamara regardless of Bailey’s 1900) and Moldenke’s (1980b) having com- bined this taxon under L. camara and applying it to L. strigocamara (Moldenke in sched.) (see L.amethystina and L. mutabilis Lippold ex Otto & A. Dietr, App. B.). The array of species names, dating from the Linnaean and early post-Lin- naean period, that apply to hybrids (App. B, Fig. 12) shows the extent to which horticultural hybridization was complicating the systematics of this group. Lantana camara subsp. aculeata was hybridized at least with L. horrida (pro- ducing L. x mutabilis C. E. Weigel), with L. hirsuta (L. X mista L., Fig. 10), and with L. strigocamara or some undetermined species or species hybrids from the Strigose Group (L. x purpurea Hornem.). Lantana horrida was crossed with 396 BRIT.ORG/SIDA 22(1) K Se . Fic. 8. Lantana horrida.a. portion of lectotype (Humboldt & Bonpland 4149, P-HBK, photograph provided by the Herbier National Paris [P], used with permission). b—c. representative specimen (Laughlin 1535, LL). b. adaxial leaf surface. c. abaxial leaf surface. Scale bars: a = 1cm;b&c=1mm some of the species or hybrids of the Strigose Group to produce L. xflava L. (Fig. Ll). This latter hybrid appears to have mixed with backcrosses of L. nivea into L.camara subsp. aculeata (L. x antidotalis Schumach. & Thonn.). Further- more, within the Strigose Group, L. nivea was hybridized probably with L. scabrida or L. splendens to yield L. x multiflora Otto & A. Dietr. At about this time, these various hybrid plants were being introduced and naturalized throughout the tropics, especially as colonists settled in the West Indies, India, South Africa, Indonesia, Australia, and Pacific Islands. In some cases, the hybrids underwent polyploidy resulting in cultivated and escaped plants that grew aggressively and genetically swamped the characters of other cultivated and escaped plants with which they crossed. Thus, over the next two centuries, the diversity of the wild escaped plants was reduced to a few aggres- SANDERS LANTANA SECT, LANTANA 397 a Fic. 9. Lantana camara subsp. aculeata, epitype (BM-SL 98: 143 bottom center ).a. whole specimen. b. adaxial leaf sur- face. c. abaxial leaf surface. Scale bars = 1 mm sive cultigens and the recombinant variation expressed in hybrids between cultigens and in those between cultigens and indigenous species. Apparently having originated along with the other early cultigens, Lan- tana strigocamara became one of the most widespread and aggressive. Clearly it was already escaped and crossing with native L. urticoides Hayek of Texas and L. depressa of Florida by the 1820s (see L.x rubra Berland. and L.xfloridana Raf., App. B). Lantana strigocamara probably originated as a diploid, because diploid cultivars, usually given names such as ‘Dwarf Yellow, ‘Dwarf Pink, etc., are still available (Sanders 2001). However, post-origin allopolyploidy certainly has contributed to its aggressive growth and success in the wild. In introduced areas, it appears to have steadily increased in numbers, and by the early 1900s it came to dominate the occurrences among pantropical naturalized plants. Howard (1969) pointed out that during the later half of the nineteenth cen- tury, lantanas lost popularity among horticulturalists. Most of the early hybrid cultivars apparently went extinct, except where escaped and naturalized in the absence of competition with Lantana strigocamara. However in the 1950s, lan- tana cultivars, developed primarily from a base of L. strigocamara, began to re- 398 BRIT.ORG/SIDA 22(1) a Pe pe rT +A € ct +t J I last £ GHOMAIOUS I€dl Fic. 10. Lantana x mista, epitype (Herb. Sherard 1272, OXF). a. whol ‘ hap nas Javinl laaf curt Scale bars:a = 1¢m;b,c 2. h b. inflorescence. c gain popularity, especially in seasonally dry subtropical areas where they can be used as drought tolerant borders. In particular, Monrovia Nursery in Califor- nia crossed L. strigocamara with the very drought tolerant, persistently bloom- ing L. depressa var. depressa of Florida to produce the now widely planted Callowiana Hybrids (Sanders 2001). The Callowiana Hybrids apparently originated after autotetraploidy was induced in Lantana depressa var. depressa. The resulting tetraploid L. depressa would hybridize with tetraploid L. strigocamara to produce hybrids with an even balance of characters from both parents. This hybrid was then hybrid- ized with various cultivars of L. strigocamara to generate an array of cultivars with a wide variety of floral color combinations, growth habits, and ploidal levels from triploid to hexaploid (Sanders 2001). The Callowiana Hybrids, thus, comprise a spectrum of variation completely connecting L. strigocamara and L. depressa var. depressa. This confusing variation led Moldenke (1975b) to name — SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 399 va } a : \ : 4 : Ney aa {\S . Ve ee oy 4 * Fic. 11. Lantana x flava, epitype (Herb. Sherard 1275, OXF). a. infl 1 rey ive | b. adaxial leaf surface. ¢. abaxial leaf surface. Scale bars:a = 1¢cm;b &c=1mm. one of these hybrids, probably Lantana cv. ‘Cream Carpet, as L. bahamensis f. albiflora. CONCLUSION AND SUMMARY The convoluted horticultural and natural history of Lantana sect. Lantana has resulted in highly complex systematic relationships of both cultivated plants and plants growing in the wild. Initial focus on cultivated forms during the Lin- naean period resulted in an early proliferation of names for closely related cul- tivated plants and hybrids. These names clouded the taxonomy of naturally occurring species as plant exploration penetrated remote areas away from hu- man population centers in colonial tropical America. Horticultural selection developed aggressively growing allopolyploid cultigen species and subspecies that became naturalized, often as pernicious weeds. Even though Lantana strigocamara (i.e., the SCD cultigen), one of the most widespread and aggressive of these cultigens, is characterized predominantly by technical features of the Strigose Group, it became confused with L. camara 400 BRIT.ORG/SIDA 22(1) S & i) @ S — Q wn = a Oo wn @ Or @ ’ HH » =e | PE RRO eas 2. sone ata a _ ‘ Mutabilis ; 2 . uv a cc Sean auc - : =s ‘antidotalis | + Fic. 12.Diag howing relationships I speci ' { hybrids. Dashed ovals indicate hybrids tt are extinct or rarely seen, either Itivated plant turalized pes. Solid indicate | likely + kK a aA Cah 4 1; £ |} , | nh L J J 1 bs J ede tec A dod d £ } } H | . £. 4s 4 Tl i re +h £+h + +h setose-morph plants on the left. The ‘‘is omitted from hybrid names. For full names see Appendix B. of the Pilose Group. Otto & Dietrich (1842) applied to one color variant of Lan- tana strigocamara the name L. mutabilis Lippold ex Otto & Dietr, but this isan illegitimate later homonym of L. mutabilis C. E. Weigel, a name published for the distinct hybrid combination L. camara subsp. aculeata X L. horrida (both of the Pilose Group). Sanders (1987a, b, c, 1989a, b, 2001) and Méndez (2002) both saw the need to treat the widespread SCD cultigen as a distinct species. Unfortunately, Sanders misapplied the name L. camara to the SCD cultigen and the name L. urticifolia to L. camara. Méndez correctly applied the name L. camara but misapplied L.xflava to the cultigen. Thus, the SCD cultigen is newly named and described here as L. strigocamara. SANDERS LANTANA SECT. LANTANA 401 As popularity of the original cultivars waned, later horticultural selection used a restricted set of aggressive polyploid hybrids, especially Lantana strigo- camara, as breeding stock; thus, this species of hybrid origin and its hybrids now strongly dominate the horticultural trade. Furthermore, where it has been introduced into tropical America, it competes or also hybridizes with indig- enous taxa to produce a spectrum of variation challenging taxonomic research, which may be resolvable only with cytological and molecular techniques. APPENDIX A Use of name Lantana camara L. in taxonomic and horticultural literature in which trichomes are sufficiently described, in historical sequence. Pilose-morph only Meoikus (1775) “folia...minus rugosa, mollioraque foliis Lantanae mistae” SpRENGLE (1825) “subtus albido-villosis” SCHAUER, (1847) “subtus pallidis pagina vel reti saltem brevi-villoso subcanescentibus” ScHAUER, (1851) diagnosis = “subtus pallidis pagina v. reti saltem brevivilloso subcanescentibus" and description ="Foliorum indumentum variabile, magis pilosum et (subtus imprimis) magis tomentosum, magis minusve d tamen semper breve et in facie foliorum scaberrimum,e setulis antrorsis callo insidentibus confectum, neque hirsutum’ Troncoso (1965) “densamente villoso-pubescentes abajo” Note:Lantana camara does not naturally occur in Argentina. Thus, this description applies to escaped cultivars or to plants of Lantana horrida (incl. L. tiliifolia), L. micrantha, and L. riedeliana, which are na- tive but submerged by Troncoso into L. camara. Pilose- and strigose-morphs both included Voss (1896) “oberseits scharf, unterseits blasser odor auf der Unterflache oder ihrem Adernetz kurz-grauweif-zottig.” [inclusion of wide range of hair morphs clear from synonymy, included varieties, and use of “scharf” to modify “zottig.”] BaiLey (1900) “pubescent beneath” [inclusion of wide range of hair morphs clear from synonymy and included varieties.] STANDLEY (1924) “variously pubescent or occasionally glabrate beneath HUTCHINSON & Datziet (1931) “shortly pubescent below” Baitey (1951) “pubescent beneath” [inclusion of wide range of hair morphs clear from synonymy and included varieties.] LEON & Atain (1957) “el envés pubescente” [term equivocal, could be applied to both morphs] Maceribe (1960) “often canescently short-villous beneath” [presumes less pubescent mor- phs also included] GiBson (1970) “variously pubescent beneath (densely viscid-tomentose to strigose or rarely glabrate)" Lopez-Patacios (1977) varying from “glabresentes’ to “esparcido-pilosos o estrigulosos en el envés" to “corto-pubescentes por ambas caras” " 402 BRIT.ORG/SIDA 22(1) NasH & Nee (1984) “el envés densamente viscido-tomentoso a estrigoso o hirsuto, raremente glabro" Huxtey (1992) “pubesc. above and beneath” [inclusion of wide range of hair morphs clear from synonymy and included varieties.] CULLEN ET AL. (2000) “downy or almost hairless underneath” Strigose-morph only (or also mixed) Orto & Dietrich (1841) diagnosis = “subtus hirtulis” and description = “oberhalb durch kleine Héckerchen scharf, etwas kurzhaarig, unterhalb ebenfalls [emphasis mine] zerstreut kurzhaarig, und etwas heller grun oder beinahe weiflich [glaucous?]” Note: They described Lantana moritziana, which is conspecific with true L. camara, as “Die Blatter... kurz-weichhaarig, nur wenig scharf und ziemlich weich anzufahlen.” Further- more, their description of L.camara is almost identical to theirs for L.crenulata, which Schauer (1847) described as “subtus pallidis, pubescenti-scabris.” Pearson (1912) “scabrid-pubescent above and on the veins beneath" Mo oenke (1970a) “short-pubescent, mostly on the venation beneath Lone & Laketa (1971) “strigose beneath" Apams (1972)"Lower surface of leaves thinly pubescent or almost glabrous" (as opposed to“with numerous short hairs’) Mo toenke (1973b) “usually only sparsely pilosulous or strigillose beneath (mostly only on the venation) with canescent or brownish trichomes, sometimes glabrescent” [as op- posed to “densely puberulent or short-pubescent’] Corrett & Corrett (1982) short pubescent, mostly on the venation beneath” MOLDeNke & MOLDENKE (1983) “Usually only sparsely pilosulous or strigillose beneath (mostly only on the venation) with canescent or brownish trichomes, sometimes glabrescent” Jansen-Jacoss (1988) “strongly or sparsely hirsute-strigose below, especially on the veins” Waaner €T AL. (1990) “lower surface sparsely short-pilose or strigillose, sometimes glabrate” Verocourt (1992) “+ pubescent or glabrescent beneath but scabrid on raised venation" Lioaier (1994) “algo pelositas en el envés, a veces glabrescentes" Poot (2001) “envés con tricomas estrigosos pequenos limitados a los nervios” [My work, as well as LOpez-Palacios (1977) cited.] 7] APPENDIX B Typification and history of application of names in Lantana sect. Lantana. Unless otherwise noted, herbaria given for authors and collectors in TL-2 (Stafleu @ Cowan 1976-1988) were consulted for original material. An exhaus- tive search for isotypes was not attempted. Unless cited otherwise, handwrit- ing comparisons based on Burdet (1979). A name is considered to be misap- plied if the cited author or annotator clearly misunderstood the concept of that species and applied it toa species in sucha way to exclude the type. A misappli- cation is inferred for publication of an infraspecific combination in which the infraspecific taxon actually belongs toa second species that excludes the type of the species under which the combination is made. The hybrid formulae for SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 403 nothospecies and species of hybrid origin are presumed parentages based on character combinations and oe 1753—Lantana camara L., Sp. P|. 6 Lectotype (Moldenke & Moldenke 1 i —Cult, probably Hort. Uppsala, Herb. Linnaeus 783.4 (LINN!) ig. ] Name misapplied to L. xantidotali (Verdcourt 1992) L. xflava (Moldenke 1980b) L. horrida ae 1962, 1982c; es 2002) L. X mista (Bailey 1900; Moldenke 1942) L. nivea (Bailey 1900; Moldenke eve 1980b) L. scabrida eae ide 1973b; Adams 1971) L. splendens (Molden L. strigocamara (Otto . te 1841; Moldenke 1940, 1949, 1970a, 1973b, in sched.; Long & Lakela 1971; Correll & Correll 1982; Sanders 1987a, b,c, 1989a, b, 2001, in sched; Verdcourt 1992; Liogier 1994) Taxonomic disposition.—L. camara (Pilose Group) Discussion.—Linnaeus' concept of L.camara was developed from an array of garden and horticul- turally selected plants. His reference in the protologue, in Sp. Pl. (ed. 2) 1763, and in Syst. Nat. (ed. 12 1767 to synonyms that include the phrases “floribus miniatus,"“flore variegata,” and “flore variabile’ indicates that garden hybridization had already introduced genes of at least L. nivea into the mate- rial he studied. However, the Moldenkes’ lectotypification associated the name with an element characteristic of native Jamaican populations that lack the wider Genene ae and have only flowers opening yellow and changing to darker yellow, or at most, to r Indeed, Proctor 18266 (NY!) and Yuncker 17043 (NY!), native material from Jamaica, are close orticg to LINN 783.4. Otto and Dietrich (1841) appear to have been the first to misapply L.camara to L. strigocamara They described L. camara as having leaf blades (my translation) “basally somewhat rounded or almost cordate and having surfaces above with small, rough, sharp, somewhat short hairs, and likewise below scattered short-haired.’They restricted the application to material of L. strigocamara with flowers opening yellow and aging orange (see also L. crocea [1804], L. moritziana [1841], L. mutabilis Lippold ex Otto & A. Dietr. [1842], and L. variegata [1842] below) rs (1987¢) explicitly excluded the type of L.camara by describing the species as having, “Hairs of abaxial laminar surfaces sparse, restricted mostly to midrib, secondary, and tertiary veins, stout, tapering-conical, geniculate toward base with distal 2/3 held parallel to lamina or vein surface.” 1753—Lantana aculeata L., Sp. P|.627. Lectotype (Méndez & Cafferty 2001).— icon in Plukenet, Phytographia t. 233, f.5. 1692. Epitype (here designated).—Herb. Sloan 98:143 bottom center specimen (voucher of Plukenet, ee 233, aa BM-SL! (Fig. 9) Name misapplied to a camara Greats 1775; Méndez 2002) a X L. strigocamara (Méndez 2002) hy horde Boch ees ann 1858; Méndez 2002) L. nivea (Méndez L. strigocamara ne a Taxonomic beatae on.—L. camara (as L.camara subsp. aculeata) Discussio omy knowledge ae (1847) was the first author to place L. aculeata in synonymy under ie camara, which has equal priority, and his choice is followed. sndez and Cafferty chose Plukenet's figure as the lectotype. The voucher of the lectotype (i.e, epitype) is more or less within the range of variation of L. camara with regard to indument. = —t — 404 BRIT.ORG/SIDA 22(1) Although the abaxial surface we are ot wee is typical praia material, this condition is found in occasional native collec nnaeus y developed the concept of L. aculeata to accommodate ae? that therwise like L.camara but differed only by the devel- opment of prickles. As with L. camara, a. proteleeds indicates both plants with yellow flowers changing only to dark yellow ate those with flowers opening yellowish changing to red or pur- plish.In particular, Plukenet’s ame includes “floribus miniatis.” Not only the floral color varia- tion (especially the late ee Salah pigments from L.nivea), but also the presence of prickles and sparse abaxial foliar hairs both point to hybridization influencing the wider variability in L. aculeata. Thus, my concept of the taxon is that of an infraspecific rank to account for all the culti- vated and escaped plants of hybrid as (regardless of floral color) falling within or slightly ex- sae the range of variation of L. camara. éndez described L. aculeata as mon surface with hairs relatively abundant, usually on all veins fa eres areoli), robust, conical, all, most, or at least a significant portion on the midrib, sec- ondary or tertiary veins geniculate toward base and with distal 2/3 held parellel to vein surface, all those that grow on lower veins shorter and erect.”To me, this describes hybrids between L. camara and L. strigocamara. Therefore, his application of L.aculeata to L.camara x | sn conform to the voucher of the lectotype, pointing out the need for epitypification. Aaa L. aculeata f. rubella (Moldenke) I.E. Méndez is to be identified with L. strigocamara, L. aculeata f.nivea (Vent.) LE. Méndez with L. nivea, and L. aculeata f. parvifolia (Moldenke) I.E. Méndez hae L.camara This taxon apparently was the main vector for infusing L. camara genes into the Strigose Group to produce L. strigocamara. It is not now widely available in the roticltutal trade but ap- pears to be rather widely naturalized in the paleotropics, especially Austra 1767—Lantana mista L., Syst. Nat.ed. 12, 2:417 Lectoryee (Méndez & Cafferty 2001).—icon in pienie Hort. Eltham. t. 56, f.64. 1732. Epitype (here designated).—Herb. Sherard 1272. (voucher for Dillenius, Hort. Eltham. t. 56, f.64) OXF! (Fig. 1 Name misapplied to L. hirsuta os hauer 1847, 1851 t. 42) L. horrida (Méndez 2002) eee disposition.—L. x mista (L. hirsuta x L.camara subsp. aculeata) Discussion.—Reliance on the Dillenius figure led Méndez (2002) to associate this name with the wild species, L. horrida, which occurs in the Greater Antilles. However, when the epitype (i.e, voucher of the figure) is examined with reference to critical trichome characters, it is shown to be a hybrid between . hirsuta and L. camara subsp. aculeata. It is not typical of L. horrida. 2s hx} ri oa et This hybrid combination is mostly of historical interest, as it is not now widely available in the horticultural trade and may have served as an agent for further hybridization and introduction of wider genetic variation into the cultivated complex 1768—Lantana pio Mill, Gard. Dict. ed. 8, Lantana 5. LecTorre (Sanders 1989a).—Mexico. Edo. Veracruz: Veracruz, Houstoun s.n., Herb. Sloane 6:84. (BM-SL photo!, mounted epHotes NY!, UCI). cme misapplied to L.can cara Gueldenke in sched.; Adams 1971; _ 1987b, c, 1989a, b; Liogier 1994) L. horrida (Adams 1971, Sanders 1987b, 1989 Taxonomic disposition.—L. camara (see 1753) Discussion.—Mark Spencer la Typification Project, pers. comm.) verified that Herb. Sloane 6:84, collected by Houstoun at Veracruz, Mexico, is the only specimen bearing Miller's polynomial and is the only known original oa of L. urticifolia. Nash & Nee (1984) cited the collection as “type” but questioned its existence at BM. Based on a mounted photograph at UC of the specimen at BM, Sanders (1989a) cited the specimen as “holotype” which is to be corrected to “lectotype.” oo = SANDERS LANTANA SECT. LANTANA 405 Because Sanders (papers cited pov) had lest L.camara so as to exclude its type, he applied the name L. urticifolia t ies name to all members of L.camara.|In his concept of L. urticifolia, he also included the species a horrida, ne regarded as distinct. 1775—Lantana flava Medik., Hist. & Commentat. Acad. Elect. Sci. ae a 3.Phys. 225, Lectotype (here designated).—icon in Dillenius, Hort. Eltham. t. 57, f.6 Epityee (here designated).—Herb. Sherard 1275 (voucher for aa on ENAaAy: t.57, f.66). OXF! (Fig. 11) Name misapplied to: L. camara an 1847; Koch & Fintelmann 1858) L. scabrida (Méndez 2002) L. strigocamara nee 2002) Taxonomic disposition.—L. flava (L.horrida X L. spp. Strigose Group) Discussion.—There is no indication on the Sherard specimen that Medikus saw it, and his descrip- tion closely parallels Dillenius: Because the protologue cites only Dillenius' name in synonymy, the Dillenius plate must be the type The protologue clearly ascedbes the lower leaf surface as “pallidiora, tomentoso-pilosa.” | bearing the mixture of long and short appressed hairs on the foliar upper surface and mixture of antrorse strigae with softer, filiform hairs on the lower surface, oe epitype (. e,, uch of the lec- totype) agrees with the protologue and confirms that the rrida and one or more species of the Strigose Group. Therefore, it is not clear why Méndez (2002) ae to apply this name he sate ta of L. ee He cited neither the plate nor its voucher. ti mostly of historical interest, as it is not now widely available in the ovticuturel trade and may ne served for further hybridization and introduction of wider ge- netic variation into the cultivated complex. 1775—Lantana splendens Medik., Hist.& Commentat. Acad. Elect. Sci. Theod.-Palat. 3. Phys. 226. Lectoryee (here designated).—icon in Dillenius, Hort. Eltham. t. 57, f.67. 1732 Epitype (here designated).—Herb. Sherard 1269 (voucher for Dillenius, Hort. Eltham. t. 57, f. 67), left hand stem. OXF! (Fig. 6) Name male p Rte to: ie pe foldenke & Moldenke 1983) Tepenonie dis pou on.—L. splendens (Strigose Group) Discussion.—The lectotypification by Moldenke & Moldenke ([Dillenius s.n., OXF] 1983) is rejected as both imprecise and incorrect. The protologue cites as original material only the Dillenius name and plate as a definite synonym and a Plukenet name and plate as a questioned synonym. Not only does Medikus’ description closely parallel Dillenius, but the voucher is not annotated by Medikus. Therefore, there is no evidence that Medikus studied the specimen, and the Dillenius plate is cho- sen as lectotype. Both the protologue and the epitype (i.e., the voucher of the aa rnatch native material from the Bahama Archipelago, the provenance given by Dilleni 1775—Lantana sanguinea Medik,, Hist. & Commentat. Acad. Elect. Sci. Theod.-Palat. 3. Phys. 229. Type.—Unknown. Name misapplied to: L. camara or L. X mista (Schauer 1847) Taxonomic disposition.—L. camara subsp. aculeata (see 1753) or L. X mista (see 1767) Discussion.—Medikus cited no synonyms, illustrations, or specimens that could serve as type. His own herbarium is unknown. His description of trichomes is scant, making clear application of the name difficult. He did relate it most closely to L. aculeata L.; thus, the assumption made here is that it is a color variant of that taxon or L. mista L., for which he gave a similar description, except for ultimate color of the corollas. 406 BRIT.ORG/SIDA 22(1) 1776—Lantana mutabilis C.£. Weigel, Physiogr. Sallsk. Hand. 1 Lectorype (here designated).—Cult., Greifsweld Bot. Gard. ("H. ae '), Pyl s.n., Aug. 1774 UE (dig. photo!)). Taxonomic disposition.—L. x mutabilis (L. camara aes aculeata ° x L.horrida) Discussion.—The protologue indicates that the n the Greifsweld Botanic Garden was the basis of the new species. The 1774 collection a Pyli is the a erieinal material found at JE and was annotated by Weigel U. Mueller, JE, per ly, Wei ited Camara melissae folio, on aaa Dill. (Hort. Eltham.65, t.56, f.65),in synonymy.The Het ee ee illustration (Herb. Sherard 1274, OXF!) also is a hybrid between L. camara subsp. aculeata and L. horrida. =f — 1789—Lantana scabrida Sol.in Aiton, Hort. Kew, ed. 1, 2:352. Lectoryee (here designated) —Cult. Royal Bot. Gard. Kew,"Hort. Kew 1777," (Solander s.n.?) (BM). (Fig. 2) Taxonomic disposition.—L. scabrida Clears Grou p) Discussion.—The specimen at BM apparently is the only original material. It is annotated “Hort Kew 1777" in an unknown hand, possibly Solander’s, and “Lantana scabrida Ait. Hort. Kew ii 352!" possibly by J. Britton (M. Spencer, BM, pers. comm.). The only material cited in the protologue is “Nat. of the West Indies, Mr. Gilbert Alexander, Introd. 1774." 1789—Lantana melissifolia Sol. in Aiton, Hort. Kew, ed. 1, 2:352. nom. illeg. (superfluous) Lectotype (here designated).—icon in Dillenius, Hort. Eltham. t.57,f.66.] Epitype (here designated).—Herb. Sherard 1275 (Voucher for Dillenius, re Eltham. t.57, f.66). OXF! Ieee rai —l. Xflava (see 1775) Discussion.—Solander cited Lantana flava Medik. in synonymy and cited the Dillenius plate. Evi- dence is oe that he studied the voucher in the Sherard Herbarium. 1796—Lantana mutabilis Salisb., Prodr. Stirp. Chap. Allerton. 107. nom. illeg. (later homonym of L. x mutabilis C.E. Weigel) Type.—None selected. Taxonomic ne tion.—L. camara subsp. aculeata (see 1753) Discussion.—Because this is a later homonym ake xmut gall is C.E. Weigel, it presents no problems. Based on the protologue, Salisbury app ial with heads that change colors from yellow to purple. He cited only L. camara Linnaeus in Sp. Pl. (ed. 2) 1763 in synonymy; so, it is not clear whether he was ae a substitute name or separating the material with multicol- ored heads as anew spec 1804—Lantana crocea Jacq,, P|. Hort. Schoenbr. 4:t.473. Lectotyee (here designated).—icon in Jacq., Hort. Schoenb. 4:t. 473. Name misapplied to: L. bahamensis (Schauer 1847; oo & Fintelmann 1858) L. camara (Otto & Dietrich 184 ibrida (Sck 1847 se . Fintelmann 1858) L. sple ndens cone 1847) Taxonomic disposition. —L. camara (see 1753 Discussion.—No original material other than the plate has been found. The plate is a good match for many wild collected specimens from Jamaica. These differ from the type of L. camara only by details of leaf shape. However, herbarium samples suggests (unpubl. observ.) that, among Jamai- can Penn: leaf-shape variation is not correlated with Baa ae or ecology. Bec e Otto and Dietrich (1841, 1842) applie d L.camara to L. strigocamara (see above), they applied i crocea to L. camara, restricting its use to ee with heads changing from yellow to orange and stems with prickles (see L. moritziana, 1841 w 1804—Lantana nivea Vent., Jard. Malmaison t.8. Lectoryee (Here designated).—icon in Vent., Jard. Malmaison t.8. — SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 407 Epirype (here designated).—Cult., Paris “ex h. Paris, mis. Siebke,” Siebke s.n, C! (Fig. 7) Taxonomic disposition.—L. nivea (Strigose Grou Discussion.—Ventenant provided a good description, excluding some details of leaf indument. Al- though the garden at Malmaison received material from gardens in the East Indies, it is clear that the description and plate apply to native material from southeastern Brazil, as noted by Schauer (1847). None of Ventenant’s ofginal matetlal sea in Paris was located in B-WILLD, C, G, or P (no response from WU), cultivated in Paris was annotated as L.nivea, but apparently not in Ventenant’s hand, sae it not to be original material. Because subsequent authors have misunderstood this species as evidenced by the several new names described for more recent wild collections of this species, the specimen in C is designated as to support the icon. It clearly matches the protologue and was at Paris about the time of Ventenant such that the annotator should have been familiar with Ventenant’s concept ae 1815—Lantana purpurea Hornem., Hort. Bot. Hafn. 2:583. Lectotype (here designated). —Cult. Hort. Reg. Bot. Hafn."1814" (Cl). Name ee ed to: ra (Schauer 1847; Koch & Fintelmann 1858; Bailey 1900) eee eee —L. X purpurea (L.camara subsp. aculeata x L. spp. Strigose Group) Discussion.—The lectotype is in a type folder marked as “specimina originalia” (also labeled as "IDC microfiche 111"'*). The verso is annotated “Lantana purpurea Horne. 1814,” of ean in Hornemann’s hand, and stamped “Hb. Schum..” A second specimen (IDC microfiche 1 a is anno- tated “Lantana purpurea h.h.” in a different hand and stamped “Hb. Liebm.” Both s mens are characterized by a mixture of erect and geniculate trichomes on the abaxial eae purplish corollas suggest that a purple-flowered cultivar of L. camara subsp. aculeata was crossed with a cultivar from The Strigose Group 1817—Lantana hispida Kunth in HBK, Nov. Gen. Sp. 2:260. Lectoryee (Nash & Nee 1984).—Mexico. Edo. Veracruz:"juxta Xalapam,” Humboldt & Bonpland s.n. (P- K (dig. photo!, Macbride Neg. oe Fl, BRIT!)). Name misapplied to L. hirsuta (Moldenke 1947, 1973b) L. hirta Grah., sp. sect. Callioreas (Schauer 1848; Koch & Fintelmann 1858; Standley 1924; rea 1963, roe 1970) L. velutind Mart. & Gal. sp. sect. Callioreas (Moldenke 1982c) Taxonomic ee —L. horrida (see next) Discussion.—Schauer’s misapplication of the name to L. hirta Graham in section Calleoreas has re- sulted in ee confusion. Furthermore, the epithet is easily confused with L. hirsuta M.Martens & Galeotti, a distinct species. Therefore, in combining L. hispida with L. horrida as conspecific,| choose the latter in accord with ICBN Article 11.5 (Greuter et al. 2000). 1817— pants ae Kunth in HBK., Nov. Gen. Sp. 2:2 Lectotype (Nash 84).—Mexico. Distrito Federal:“monte eChealpecees urbem Mexici,.” Humboldt &Bonpand4i 49 (P-HBK [dig. photo!]). (Fig. 8a) pals misapplied to rubra Cahaue 47) L pee ee 1942, nae . 1978) L. hirsuta XL. camara (Moldenke ) Taxonomic Deen as horrida ae Group) Discussion. —Berlandier 2310, annotated by Berlandier as Lantana rubra (L. urticoides X L. strigocamara), a syntype of L. horrida var. parviflora Schauer, and the epitype of L. x rubra (see be- low) was mistakenly included in L. horrida by Schauer (1847). As a result, Moldenke misapplied the name L. horrida to L. urticoides until correcting himself (Moldenke 1978). 408 BRIT.ORG/SIDA 22(1) 1827—Lantana antidotalis Schumach. S Thon in poeta eek ou PI.276. Lectotype Junghans 1962.p.94,as to collec h plicat Ghana, Thonning 25 (C [IDC microfiche ident. no. 64 | *"]!; soLectoryee: ClIDC. no.64'°"]!). ae disposition.—L. xantidotalis (L.nivea X L.camara * L. x flava?) Discussion.—The duplicate most in eae with the protologue (leaf blades 2-3 inches long) is chosen as the lectotype. Though given the same collection number, these two specimens may actually represent two different populations as judged from slight morphological differences. As- signing hybrid status to this naturalized plant is based on the variable leaf shape and size (narrowly ovate-oblong or narrowly triangular;length 1.7 to 2.7 X width), subsetiform trichomes on the up- per and lower leaf surfaces,and the mixture of erect and gemicuiste hairs on the lower leaf surfaces. The parentage probably includes L. nivea or L.splendens from the Strig nd L.camara and L. X flava from the Pilose Group. Y 829—Lantana albopurpurea Desf, abl. Ecole Bot., ed 3. 393. Lectotype (here designated).—Cult. Hort. Paris.,"H. p.” labeled “Herbarium Webbiana ex Herb. Desfontaines” (FI [dig. photo!]). Taxonomic disposition.—L. X mista (see 1767) Discussion.—The specimen at Fl is clearly original material, as the annotation is in Desfontaines’ hand (C.Nepi, pers.comm.). None other is known to me. The leaf indument suggests this specimen has the same parentage as does L. X mista. 1829—Lantana suaveolens Desf, Tabl. Ecole Bot., ed 3.393.nom. illeg. (non L. suaveolens Spreng. = L. angustifolia Mill, Type.—Not investigated. Taxonomic disposition.—probably L. camara or its hybrid (see 1753) Discussion.—The description of prickly stems, ovate leaves, lanuginose bracts, and yellow flowers suggests this is an element of L.camara te 32—Lantana rubra Berland. in Teran & Berland., Mem. Comis. Limites 15. LECTOTYPE . designated).—icon in Berlandier in Ohlendorf et al., transl. Journey Mex., t.5 (top, facing p.410).1 Epitype (here designated). —Berlandier 2310=880, Mexico. Tamaulipas: Matamoros, GH! (isoepitype: NY, none found at G under L. rubra). Taxonomic disposition. ie xrubra (L.urticoides X L. strigocamara) Discussion.—The only original material known to me includes the recently published plate, which Berlandier either executed or supervised, and three sheets (Berlandier 2114=697 [GH!], 2310=880 ([GH!, NY!]), which consist of hybrids. Although the protologue suggests L. urticoides (see 1906 be- low), the only native species in the region of the eastern Texas-Mexico boundary, all the original material lacks the rotund, large-toothed leaf blades and persistant fruiting bracts, characteristic of L. urticoides. Instead, they are intermediate between L. urticoides and L. strigocamara, which obvi- rial that actually belongs to L. urticoides should be found, such a pein could be easily overturned. However, stability of the application of L. rubra and rete Uke urticoides as the correct name of the wild species is best served by accepting the hie as i the two poten omen: aa A me, a 14= er" is more ane ee hybrids, espe- cially in the ho 1.5mm long. Berlandier 23 10=880 has pioeneeth ve supeersctent bracts more like the L. urticoides parent, but the indument shows it to be a hybrid. It is chosen as epitype because it is annotated “Lantana rubra B,” apparently in Berlandier's hand, is a syntype of L. urticoides var. parvifolia Schauer, and is represented by dupli- cates in major herbaria. SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 409 This hybrid combination is the typical cultivated plant in Texas and the Gulf Coast Plain of the Southeastern United States. It may have been propagated for the trade due to the cold hardiness ae from the Texas parent, L. urticoides x Kewensis Jackson 1895) cites’ agian rubra Perr.,”but this is a mistake as Perrottet (1825) ca referred to Latania rubra Jacq. (Arecaceae), the next entry in his list after Lantana. 1832—Lantana tiliifolia Cham., Linnaea 7:1 Lectoryee (Santos Silva 2001).—Brazil: Bahia. Si ae (B-WILLD 11502 [dig. photo! Berlin Dahlem. 005). Name misapplied to: L. strigocamara or its hybrids (Moldenke in sched) Taxonomic disposition.—L. horrida (see 1817) Discussion.—The protologue lists three syntypes—the Sieber collection from Bahia, Brazil, Sellow collections (“southern Brazil...copious numbers from many locations and times,” BR [dig. photo]; NY!, fragment NY!),and Lhotsky s.n (n.v,). | disagree with the lectotypification by Santos Silva (2001). In the digital photograph of B-WILLD 11502 provided by B, the specimen appears to have mostly glandular polis on sy stems ang petigl>: Ne Sellow specimen at BR bears mostly non-glandular hairs. Chamisso's diagnosis ly _ hirtis,"although his description iy ansietion) indicates that the age stems have Teac ie often but not always mixed in.” Later, Poeppig segregated the strongly glandular plants as L. glutinosa (see 1842 below). Modern usage follows Poeppig in restricting L. tiliifolia to the non- or weakly glandular specimens. However, because | treat both L. tiliifolia and L. glutinosa as synonyms of L. horrida, further research is needed to deter- mine whether a proposal to conserve a different type is neede 1832—Lantana floridana Raf., Atl. J. 148. Type.—Not found in DWC, FI, G, NY, P. P-DU, PH, Pl, WIS, or WS; no response from LE, NAP or W. Taxonomic disposition.—L. x floridana (L. depressa x L. strigocamara) Discussion.—The only native wild taxon of the upper Atlantic coast of Florida is L. depressa var. floridana Gone R.W. Sanders (see 1905, below). However, the protologue describes the flow- ers as “versicolor, yellow, orange, red, crimson on same shrub.” As discussed in Sanders (1987a), none of the varieties of L. depressa is characterized by the orange or red floral pigments. However, this does characterize hybrids between L.depressa and L. SE (the latter in Sanders 1987a as L. camara).In light of the hybridization sees introduce and native L. urticoides in Texas in the 1820s (see L. rubra 1832 above), L. strigocamara was likely also naturalized in Florida by that time, and Rafinesque's description refers to hybrids 1838—Lantana antillana Raf, Sylva Tellur. 82. Type.—Not found in DWC, FI, G, NY, P P-DU, PH, Pl, WIS, or WS; no response from LE, NAP, or W. Taxonomic disposition.—L. horrida (see 1817) or L. camara (see 1753) Discussion.—Rafinesque noted this to be Lantana camara of authors and an Antill b, which he saw living. He also noted met it cles from L. fon primarily by the stem and leaves hirsute and the bracts ovate-lan er of these characters appears to be consistent with this being an element of L. ees 1838—Lantana rosea Raf, Sylva Tellur. 83. Type.—Not found in DWC, FI, G, NY, P. P-DU, PH, PI, WIS, or WS; no response from LE, NAP. or W. Taxonomic disposition.—probably L. reticulata Pers. or L. involucrata L. (. Sect. Callioreas) Discussion.—Even though Rafinesque placed this species in his subgenus Camara Raf., his subgen- era are neither well demarcated nor consistent. He included L. camara L.in subgenus Camara Raf. and the conspecific L.aculeata L.in subgenus Periana Raf. (along with L. involucrata). Lantana rosea is described as a native Antillean plant with small roundish tout apical and basally acute) leaves, rounded cinerous stems, and short, ovate bracts, and rose-p all of which appear to place it in sect. Callioreas. 410 BRIT.ORG/SIDA 22(1) 1838—Lantana incarnata Raf, Sylva Tellur. 83. Type.—Not found in DWC, FI, G, NY, P. P-DU, PH, PI, WIS, or WS; no response from LE, NAP.o Taxonomic disposition.—probably L.nivea subsp. mutabilis (WJ. Hook.) R.W. Sanders 1804 and text 184 ee on. ae incarnate,’ Rafinesque presumably meant that the heads are flesh-colored in bud, p ream and turn ign or fleshy pink. This is consistent with some populations of L. strigocamara. However, the ovate-oblong leaf blades acuminate at both apex and base rule out L. eal The floral color, leaf shape, and long, linear-lanceolate bracts are all consistent with L. vea subsp. mutabilis. amethystina ) 1839—Lantana coccinea G. Don, Hort. Brit., ed. 3.245.nom. nud. Type.—Unknown. Taxonomic disposition.— camara subsp. aculeata? 1841—Lantana crenulata Otto & A. Dietr,, Allg. Gartenzeitung 9:363. Type.—Unknown (no original material known; if herbarium specimen from material cultivated in. Hort. Berlin [Otto s.n.,?] existed in B, now destroyed; Shauer [1847] indicated seeing a live specimen in Hort. Berlin). Name misapplied to: L. lar nei as r 1847) Taxonomic disposition.—L. Xantidotalis (see 1827) ar L. scabrida (see 1789 Discussion.—The See: cites no original material, but the description strongly suggests L. scabrida or related species or hybrids in the Strigose Group. Identity with L. strigocamara is ex- cluded because the leaves are described as pointed on both ends. 1841—Lantana pals Otto & A. Dietr.,, Allg. Gartenzeitung 9:369. Lectotype (Schauer 1857, as to gathering; here designated, as to specimen).—Venezuela. Caracas, Moritz 163 (G [dig. photol]; isotectorype: G [lacking inflor.,dig. photo!) Name misapplied to: L. camara (Otto & Dietrich, 1841, 1842) Taxonomic disposition.—L. camara (see 1753) Discussion. —The protologue seo nativity only by “Habitat in Caracas” without specifically cit- ng the Moritz specimen. In his treatment, Schauer (1847) cited only Moritz 163 from Caracas, Ven- ee but stated that he saw ce and wild-collected herbarium specimens at B. The only duplicates of Moritz 163 known to me are at G.The one chosen as lectotype bears two annotation labels besides the collection label, all apparently in Schauer’s hand. The duplicate at G lacks infloresences, is not annotated by Schauer, but is annotated by Moldenke as “Isotype.” Macbride photographed material at B (neg. no. 17478, destroyed [photo F!, BRIT!, GH!]), which he may have thought was type material. The specimen is not signed by either Otto & Dietrich or Schauer, and it bears the date’1844."Therefore it cannot be considered a type, even though Lopez-Palacios (1977 cited it as such. Because Otto and Dietrich (1841, 1842) misapplied L. camara to L. strigocamara, they applied L.moritziana to plants of L.camara with no prickles and flower heads changing from yellow to orange. 1841—Lantana multiflora Otto & A. Dietr, Allg. Gartenzeitung 9: oa Lectorype (here designated).—Cult. Hort. Berlin, Otto s.n.(B,d brid 17379 F!, BRIT!]). Taxonomic disposition. —L. X multiflora (probably L.nivea X L.scabrida or L. splenden Discussion.—Otto and Dietrich cited no specimen, but the destroyed specimen Seueehee by Macb oe appears to have been the only original material. In general, the protologue suggests L. nivea, although, the more distinct scabrosity of the leaves suggest L. scabrida or L. apIENGeN The floral color, described further in Otto and Dietrich (1842) as lilac (in bud?) changing to yellow, clearly implicates genes of both L. nivea and yellow-flowered species of the Strigose Group. SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 411 1841—Lantana amethystina Otto & A. Dietr,, Allg. Gartenzeitung 9:370 Type.—Unknown (no original material known; if herbarium specimen from material cultivated in. Hort. Berlin [Otto s.n.,?] existed in B, now destroyed). Taxonomic disposition.—L. nivea (as L. nivea subsp. mutabilis, see 1804 and text) Discussion.—Otto and Dietrich (1842) indicated that this is a replacement name for L. nivea var. mutabi lis. Hooker. Apparently, Otto and Dietrich were not aware of L. mutabilis C.E.Weigel and L. mutabilis Salisb. but did consider that Lippold had named L. mutabilis Lippold ex Otto & A. Dietr. for a distinct species such that the Hooker varietal epithet was unavailable at the rank of species (see next entry). 1842—Lantana mutabilis Lippold ex Otto & A. Dietr,, Allg. Gartenzeitung 10:314.nom. illeg. (later homonym of Lantana X mutabilis C. E. Weigel) Type.—Unknown (no original material known; if herbarium specimen from material cultivated in. Hort. Berlin [Otto s.n.,?] existed in B destroyed; no material of Lippold found at RB [R.Campostrini F,, pers eee series —L. strigocamara (see 2006 and text) or L. strigocamara x L. camara subsp. ulea ae eee noted under L. camara, Otto and Dietrich (1841) appear to have applied the name L.camara to the phenotypes of L. strigocamara having floral heads changing from yellow to orange. However, material of L. strigocamara with purplish and yellowish pigments occuring together they apparently described under L. mutabilis Lippold ex Otto & A. Dietr.As a later homonym, this name is not available for L. strigocamara. 1842—Lantana variegata Otto & A. Dietr., Allg. Gartenzeitung 10:314 Tyee.—Unknown (no original material known; if herbarium specimen from material cultivated in. Hort. Berlin [Otto s.n.,?] existed in B, now destroyed). Taxonomic disposition.—L. camara subsp. aculeata or L. camara subsp. aculeata x L. spp. Strigose ro Discussion.—No holotype or syntypes are cited; however, the protologue suggests cultivated ma- terial in the range of L. camara subsp. aculeata (see 1753). 1842—Lantana glutinosa Poepp. in Otto & A. Dietr,, Allg. Gartenzeitung 10:315. LECTOTYPE renee as 1977, as to gathering; here designated as to specimen).—Peru, Poeppig 1375 (G-DC [dig. photo!]; duplicate, B?, destroyed [Macbride Neg. 34340 F!, BRIT!]; soectorype: GH! Taxonomic disposition.—L. horrida (see 1817) Discussion. —Other than eee in eae no material is cited and lectotypification is required. The eee photograph (Neg. 34340) is presumed to be of a specimen at B, since destroyed, although no herbarium eae is visible in the image. If this be true, it would have been the preferred lectotype. oot 1842—Lantana asperata Hort. ex Vis., Orto Bot. Padova 142.nom. nud. Type.—Unknown. Taxonomic peel mes vee (see Discussion.—V i | | this name at Hort.Parisiense, without fAcading to ube it as his own.He also stated,"Non pare diversa della L. nivea Vent.” a 804) 1843—Lantana bartramii Baldwin, Reliq. Baldw. 247. Type.—Not designated. Taxonomic disposition.—L. X floridana? (see 1832) Discussion.—The protologue cites no material.In a search of DWC, LASCA, MANCH, NY, P-DU, PH, WECO and WLU, only two kee specimens were located. At NY originally in the Torrey Her- barium (T. Zanoni, pers.comm.) is a specimen (!) labeled “grown in Louis Lecontes’ Garden in Geor- gia” without collector or ie and annotated as the “Lantana camara [var] bartramii, cultivated in 412 BRIT.ORG/SIDA 22(1) Florida by Dr. Baldwin."This specimen may or may not have been seen by Baldwin. At PH, there is a sheet (dig. photo!) from the Muhlenberg Herbarium that has mounted on it several fragmentary specimens from different collectors.One such fragment is labeled,”1482 Lantana Hardy native’ and presumably was collected by Baldwin. Further research is needed to determine if this specimen is original material. Both specimens are elements of |. depressa Small var. floridana (Moldenke) R.W. Sanders (see 1905). Baldwin's protologue refers to plants along the Florida seacoast as being the same that Bartram described, without mentioning cultivated material. Bartram traveled only in the geographic range of var. floridana. Further, Baldwin describes the species as attaining the height of six feet. Material of L. depressa var. floridana that | have seen reaches three feet but not six. This, therefore, suggests Baldwin was describing the same hybrid that Rafinesque did under the name Lantana x floridana. 1844—Lantana hirsuta M. Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 11:326. Hovotyee.— Mexico. Edo. Veracruz: Jalapa, Mirador, 1840, Gallioti 749 ( BR [dig. photo!]; isoryee: G [dig. to!]). pho Taxonomic disposition.—L. hirsuta (Setose Group) 1846—Camara cau Benth., Bot. Voy. Sulphur 154 Lectorype (here designated).—Herb. Linnaeus 783.4 (LINN). Taxonomic disposition.—L. camara (see 1753) Discussion.—This was a replacement name for Lantana camara when placed in Camara Adans. However, the correct name in Camara. is C. aculeata (L.) Kuntze (Rev. Gen. Plant. 2:503. 189 184 ae anane amare Schauer, Linnaea 20:480. pel (her cited as “isotype” by Lopez-Palacios 1977).—Venezuela.’ad Caracas," Moritz 92 (BM, n.v.; HoLoTYPE NOW destroyed: B [Macbride Neg. 17463 FI, GHI]). . misePPe ed to: LA ] ia 97 2a) Tavonannic disposition —L. hirsuta (see 1844) 1847—Lantana polyacantha Schauer in DC., Prodr. 11:597. Hototype.—Mexico, Schiede s.n. (B, destroyed, ie c 17481 [F!]). Taxonomic disposition.—L. horrida (see 1817) 1847—Lantana aoe Schauer in DC,, Prodr. 11:599. Lectoryee (S —Brazil. Matto Grosso, Manso s.n., Mart.Fl. Bras. 1026 (M,n.v.ISOLECTOTYPES: BM, fe ) (dig. ee c DC [dig. photo!, Macbride Neg. 7873 F!, BRIT!], G [dig. photo!, Macbride Neg. 24632 FI, eis K,NY!). Name misapplied to L. nivea (Moldenk Ke ae L. viscosa (Moldenke 19 Taxonomic disposition. —L. en (Strigose Group) Discussion.—The are two paratypes known to me. One is Poeppig 1485 (Peru) at G. It apparently is annotated by Schauer (without his “!"). He cited Poeppig 1405 which is probably a typographical error for"1485."The other is Martius s.n. (Brazil. Rio ely at M (n.v.). 1847—Lantana robusta Schauer in DC., Prodr. Lectorype (Santos Silva 2001).—Brazil. Rio de cea i ae do Correia,” 1821, Poh! 40-5955 (W, nv). Taxonomic disposition.—L. robusta (Setose group) Discussion.—There is a Macbride photograph ae 17485 [F!, BRIT!, GH!, LL! NY!J) of Pohl 782 at B, now destroyed, that has been distributed as a photograph of a type. The specimen in the photo- graph should be disregarded as original material. This specimen neither appears to be annotated by Schauer nor bears sufficient inflorescence material to match the protologue. Furthermore, the paratype, Raben 509 (NY [fragment]! and BR [photo at LL!]), and Poh/ 182 are not the same species. SANDERS LANTANA SECT. LANTANA 413 Pohl 182 may belong to L. horrida or L. horrida X L. robusta. 1847—Lantana riedeliana Schauer in DC., Prodr. 11:601. Hototype.—Brazil, Riedel s.n. (LE, n.v.). Name misapplied to L. sp. sect. ee loreas (Moldenke 1970b) ce ep osion: —L.riedeliana or L.robus Discussion.—The only original material cited is a sigeate at LE, which did not respond to inquiries. There is a photograph in LL (perhaps by Moldenke) of Glazier 1657 that is annotated as L. riedeliana by Moldenke, as well as someone other than Schauer. This specimen appears to fit the description of L. riedeliana and suggests that L. riedeliana is conspecific with L. robusta, being a glabrescent, more narrow-leaved variant. 1847—Lantana pohliana Schauer in DC., Prodr. 11:601. Type (Lectotype not designated).—Brazil, Pohl s.n. (W?, n.v.; duplicate: B, destroyed [Macbride Neg. 17480 F!, BRIT!, GH!]). Taxonomic disposition.—L. pohliana or L. viscosa? (Setose Group?) Discussion.—Schauer cited no collection number or herbarium. Pohl material should be at W, but no response to inquires there was received. Further investigation is neede 1847—Lantana viscosa Pohl ex Schauer in DC., Prodr. 11:601. Lectotype (Santos Silva 2001).—Brazil. Goids:"Ad Concei¢do, prope Trahiras,” Poh! 1876 & 2680“D. n° 181." (left-hand specimen) (W, n.v.;isoLectoryee: B, destroyed [Macbride Neg. 17492 FI, BRIT!]) Taxonomic disposition.—L. viscosa (Setose Group) — 1847—Lantana multicolor Lem., Fl. Serres Jard. Eur. 3:239 O L. camara subsp. aculeata or its hybrid (Koch & Fintelmann 1858) Taxonomic disposition.—L. camara subsp. aculeata, L. x mista, or L. sp. sect. Callioreas? Discussion.—Seeds of this were sent to Brussels from Mexico, possibly of cultivated origin. Lemaire distinguished it from other cultivated lantanas by its unusually large leaves and by the rose and rimson flowering heads. Although Koch & Fintelmann (1858) recognized the species, they indi- cated that it may be a form of L. camara or L. Xmista. However, the protologue also describes the heads as involucrate with four large bracts and with the floral bracts cordate-rotund. These charac- ters would place the plants in section Callioreas 1857—Lantana hybrida Neubert, Deutsch. Mag. Garten- Blumenk. 10:98. nom. illeg. (nom. subnud.) Lectoryee (here designated). —icon in Neubert, Deutsch.Gart. Mag. 10:t. facing p. 1 Name diese ed to Le ara subsp. euleane or L. ee (Everett 1980) ee disposition.—L. X flava (see 1775) or L. x mista (see 1767) Discussion.—No specimens were found at ‘oe or STU to serve as lectotype or epitype. The de- Se pHenN: is very brief, indicating only the color of the flower heads. The plate shows nothing dis- tinctive except for spreading hairs on the twigs, petioles,and peduncles, suggesting that the plants may be elements of L. flava or L. X mista. 1858—Lantana es K. Koch & Fintelmann, Wochenschr. Gartnerei Pflanzenk. 1 :322.nomm.illeg., pro syn. sub L.cro Tyee—Unknown. ees ic disposition.—L. camara (see 1753) 1863—Lantana triplinervia Turcz,, Bull. Soc. Imp. Naturalistes Moscou 36:205. Hovotyee.—Java, Goering 225 (KW?, n.v.). 414 BRIT.ORG/SIDA 22(1) Name misapplied to L. nivea (Moldente 1974, 1977) L. nivea X L. spp. Pilose Group (Moldenke 1974, in sched; Sanders in sched.) Taxonomic disposition.—L. nivea (see 1804) Discussion.—Turczaninow eae material that was cultivated or escaped from cultivation. The protologue strongly suggests L. nivea by the leaves long petiolate, ovate-lanceolate, triplinerved, apically acuminate, and minutely scabrous on the veins below. 1863—Lantana bahiensis Turcz., Bull. Soc. Imp. Naturalistes Moscou 36:206. Hototype.—Brazil. Bahia, Sa/zmann s.n. (KW?, n.v.). Name misapplied to: L. nivea X L. spp. Setose meee mani in sched.) Taxonomic disposition. —L. sp. sect. Callior Discussion.—The protologue are ne species in section Callioreas, which is corroborated by the description of bracts ovate, briefly acuminate, and 5 ed. 1904—Lantana micrantha Briq., Annuaire Conserv. Jard. Bot. Geneve 7-8:299, Ho.oryre. ve taatley Asuncion, Balansa 1039 (G). Name misapplied to L. horrida (Moldenke 1948, 1975d) L. sp. sect. Call ie 1981c) Taxonomic disposition.—L. micrantha (Pilose Group) 1905—Lantana depressa mall, Bull. New York Bot. Gard. 3:436. HoLoryPe. ee. en a Co.:pinelands between Coconut Grove and Cutler, Small & Carter 747 (NY; isotype: F!). axonomic disposition.—L. depressa (Strigose Group) Discussion,—See Sanders (1987a) 1905—Lantana bahamensis Britton, Bull. New York Bot. Gard. 3:450. Hotoryee—Bahamas. New Providence: Ft. Montague, Britton & Brace 174 (NY!;\sotyees: F!, US! = Name misapplie die L. camara (Moldenke 1975a; Correll & Correll 1982 (pro parte)) L. depressa (Moldenke 1975c) L. depressa X L. strigocamara Callowiana Hybrids (Moldenke, 1975b) L. splendens (Moldenke in sched; Correll & Correll 1982 (pro parte); Sanders in sched.) Taxonomic disposition.—L. bahamensis (sp. hybrid origin between L. splendens and L.camar Discussion.—Apparently Lantana camara, occurring in the southern Bahamas, began to hybridize introgressively into L. splendens, centered in the central Bahamas, shortly after settlement by Euro- As a result, the native plants in the Bahamas are variable. The type of L. bahamensis shows the nitid surface and scattered geniculate hairs on the leaf undersurface venation typical of L. splendens, as well as the soft, erect hairs persisting along the undersurface veins, apparently ex- pressing genes of L. camara. The bracts vary from oblong-lanceolate as in L. splendens to obovate- elliptic as in L.camara 1905—Lantana ovatifolia Britton, Bull. New York Bot. Gard. 4:123. Hotorype.— Bahamas. Grand Bahama: Eight Mile Rocks, Britton & Millspaugh 2450 (NY1; soryee: FI). Name misapplied to: L. depressa (Long 1970; Long & Lakela 1971; oes 1982) L. strigocamara X L. depressa (Long & Lake 71) =L. xfloridana Raf. Taxonomic disposition. eo ovatifolia isos ni Discussion.—See Sanders 1987a. 1906—Lantana glandulosissima Hayek, Repert Spec. Nov. Regni Veg. 2:161 SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 415 Hotorvee.—Mexico. Jalisco: Tequila, Pringle 4431 (W, n.v.isotypes: BR, F!, MO!, NY!). Name Sa ere Wet 4 (KAS -] L. Taxonomic disposition.—L. camara (see 1753) 1906—Lantana cummingiana Hayek, Repert.Spec. Nov. Regni Veg. 2:161 Hototyee.—Chile. Cumming 1065 (W, n.v.) Taxonomic disposition.—L. horrida = 1817) or L. hirsuta (see 1844) 1906—Lantana urticoides Hayek, Repert.Spec. Nov. Regni Veg. 2:162. Tyee—Not selected, to be lectotypified in a subsequent paper. (Syntypes: USA: Texas. Lincheimer Exsic. No. 503, which includes two different Lindheimer numbered collections, F!, GH!, MO!, UCI, W, n.v.; Texas: Comal “ Matthes 19, W,n.v,). Name misapplied L. horrida ae 1981b) Taxonomic disposition. —L. urticoides (Setose Group) Discussion.—See L. rubra 1832 above 1907—Lantana foetida Rusby, Bull. NewYork Bot. Gard. 4:431. Hototyee.— Bolivia, Bang 2034=469 (NY [n.v., dig. photo not available]; isoryees: US!, GH!). Taxonomic disposition.—L. horrida (see 1817) 1909—Lantana weberbaueri Hayek in Urb., Bot. Jahrb. Syst. 42:166. Hotoryee.—Peru. Junin: Tarma: Palca, Weberbauer 2017 (GB?, n.v., W?, n.v., or B? [destroyed, Macbride Neg. 17493, LL). Taxonomic disposition.—L. weberbaueri or L. viscosa (see 1847) or its hybrid? (Setose group) Discussion.—This is a rare, poorly known species. Hayek's opinion was that it is near L. viscosa and L. robusta in the Setose Group of section Lantana 1910—Lantana arida Britton, Bull. Torrey Bot. Club 37:357. Hototyee.—Jamaica, Fort Henderson, Britton & Hollick 1824 (NY!). Name misapplied to: L. camara (Moldenke, 1982a) L. horrida (Moldenke, 1982a: Méndez, 2002) Taxonomic disposition.—L. horrida (see ) Discussion.—This is a small-leaved variant of Antillean L. horrida, perhaps as a growth response to drought conditions. 912—Lantana subcordata Urb. Symb. Antill. 7:351. Lectoryee (here designated)—Dominican Rep.near Santiago, Schomburgh 5 (K! [Hotoryee:B, destroyed]; CO ~~ ISOLECTOTYPE: P). Taxonomic disposition.—L. subcordata (Pilose Group) Discussion.—See Sanders 1989a 1922—Lantana pulchra Larranaga, Escritos D. A. Larranaga 1:406.nom. dub. Type.—Destr Taxonomic ae —Cannot be determined. Discussion.—Written in the style of the Linnaeans, the protologue contains only 14 words besides the ee and provides no features to distinguish it from other published species. As both the specimens and Larranaga’s drawings are destroyed (E. A. Paz, MVFQ, pers. comm.), the name must be treated as a nomen dubium 1934—Lantana tenuifolia Rusby, Phytologia 1:74. Hotoryee.—Bolivia. Bopi River, Rusby 653 (NY [n.v., dig. photo not available). Taxonomic disposition.—L. cujabensis? (see 1847) 416 BRIT.ORG/SIDA 22(1) 1940—Lantana insularis Moldenke, Caribbean Forester 2:16. Hovotyet.—Jamaica. trail from St. Helens Gap to Latimer River, Maxon & Killip 912 (NY!;\sorype: US! Taxonomic disposition.—L. insularis (Setose Group) 1940—Lantana eee lceeas Ssleae et 2:17. ee as L. “leonardorum” Hototype.—Haiti. near Jean Rabel, | 82 (NY!; isotypes: Al, GH! US!). eines ion.—L. heats (Pilose pee Discussion.—See Sanders 198 = 1940—Lantana scorta Moldenke, Publ. Carnegie Inst. Washington 522:161. Hototype.—Mexico. Queretaro: near San Juan de Rio, Rose et al. 9520 (NY!). Name misapplied to: L. horrida (Moldenke in sched.) L.camarax L. horridaX L. hirsuta (Moldenke 1940) Taxonomic disposition.—L. hirsuta (see 1844) 1940—Lantana notha Moldenke, Phytologia 1:422. Hotoryee.—Mexico. Sinaloa: Fuerte, Rose et al. 13573 (NY1). Taxonomic disposition.—L. notha? (section ? Discussion.—Although Moldenke (in sched.) applied this name to plants in section Lantana, the type appears to be in section Callioreas, but the critical character to distinguish sections, mature fruits, is lacking. 1941—Lantana scandens Moldenke, Phytologia 2:18. Hototype.—Mexico. Michoacan: Coalcoman: Villa Victoria, Pto. de Aire, Hinton 12315 (NY! isorypes: FI, LL!, MO! NY!, US). Taxonomic disposition.—L. scandens or L. scabrida (see 1789 Strigose Group) 1941—Lantana brittonii Moldenke, Phytologia 2:52. Hotoryee.— Jamaica. Tweedside, below Moody’s Gap, Harris & Britton 10541 (NY!:isoryee: US). Taxonomic disposition.—L. scabrida (see 1789) 1946—Lantana minasensis MICIEELING) Byoiegia 2:1 38. HoLoryee. pea Minas ‘| ¢ B Mexia 4448a (NY [dig. photo!]; TEX ISoTYPES: M Name ea to: L. nivea (Moldenke 1946) L. nivea X L. spp. Pilose Group (Moldenke 1966, 197 2a, 1973a, in sched.: Sanders in sched.) Taxonomic disposition.—L. nivea (see 1804) 1962—Lantana kingii Moldenke, Phytologia 8:161 orype.—Mexico, Oaxaca: 2 km south of Niltepec, King 1775 (TEX! isotype: US!) Taxonomic disposition. —L. kingii or L. scabrida (see 1789, Strigose Group) 1979—Lantana morii Moldenke, Phytologia 41.449. Hotoryee.—Brazil. Bahia: Mun. Camaca: Sao Joao do Panelinha, Santos & Mattos 3304 (LLI:isorype: NY! Taxonomic disposition.—L. nivea (see 1804) = 1987—Lantana hodgei R.W. Sanders, J. Arnold Arbor. 68:343. Hotoryer.—Dominica. Fresh Water Lake, elfin forest, Fosberg 48269 (USI; isoryees: F!, GHI, K, MO!, NY!). Taxonomic disposition.—L. hodgei (Strigose Group) 2006—Lantana strigocamara R.W. Sanders, in this publication. Hotorype.— USA. Florida: Dade Co., Sanders 1450 (FTGI; isotype: NY!). (Fig. 4) Taxonomic disposition.—L. strigocamara (L. spp. Strigose and Setose Groups L. camara subsp. aculeata) — SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 417 ACKNOWLEDGMENTS The Botanical Research Institute of Texas supported this research with institu- tional facilities and resources. The Queensland Department of Natural Re- sources and Mines provided a travel grant to support the author while in Eu- rope to visit BM, K, and OXF. Michael Day and Isidro Méndez encouraged the author to complete this research after a hiatus of several years. Logistic and library support provided by Charlie Jarvis is especially appreciated. Curators of the cited herbaria are thanked for loans, digital images, and historical infor- mation. Guy Nesom, John McNeill, Sandy Atkins, and Charlie Jarvis provided helpful discussion and comments. REFERENCES Aoams, C.D. 1972. Flowering plants of Jamaica. Univ. West Indies, Mona, Jamaica. 1972. BaiLey Hortorium. 1976. Hortus third:a concise dictionary of plants cultivated in the United States and Canada. MacMillan, New York. Baivey, LH. 1900. Cyclopedia of American horticulture. MacMillan, New York. Baivey, L.H. 1951.Manual of cultivated plants. MacMillan, New York. Berin-DAHLEM. 2005. Digital specimen images at the Herbarium Berolinense. April 15,2005. http://ww2.bgbm.fu-berlin.de/herbarium/ Briquet, J. Verbenaceae. 1895 ("1897"). In: A. Engler & K. Prantl, Nat. Pflanzenfam. IV. 3a: 132-182. Buroet, H.M. 1979. Auxilium ad botanicorum graphicem. Conserv. & Jardin Bot., Geneva. Corrett, D.S.and H.B. Corrett. 1982. Flora of the Bahama Archipelago. J. Cramer, Vaduz. CULLEN, J. eT AL., (eds.). 2000. The European garden flora. Cambridge Univ., Cambridge. 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Annecke, ed. Proceedings of the second national weeds confer- ence. Cape Town. Pp. 321-334 Swarerick, J.T., B.W.Wittson, and M.A. Hannan-Jones. 1995. The biology of Australian weeds. 25. Lantana camara L. PI. Protect. Quart. 10:81-95. Troncoso, N.S. 1965. Verbenaceae. In: A.L. Cabrera. Fl. Prov. Buenos Aires 5:121-140. Troncoso, N.S. 1974.Los géneros de Verbendceas de Sudamérica extratropical. Darwiniana 18:295- 412. Verpcourt, B. 1992. Verbenaceae. In:E. Milne-Redhead and R.M. Polhill, eds..Flora of tropical East Africa. 155 pp. Voss, A. 1896. Lantana. In: A. Voss, ed..Vilmorin’s Blumengartnerei. Ed. 3, 2 vol. Berlin. Vol. 1, pp. 822-824. Waaner, W.L., D.R. Hersst, and S.H. Soumer. 1990. Verbenaceae. In: Manual of the flowering plants of Hawai'i 2:1315-1327. Wo rson, S.L. and T.W. Sotomons. 1964. Poisoning by fruit of Lantana camara. Amer. J. Dis. Children 107:173-176. 422 BRIT.ORG/SIDA 22(1) Book REVIEW STEVEN Foster and REBECCA L. JOHNSON. 2006. Desk Reference to Nature’s Medi- cine. (ISBN 0-7922-3666-1, hbk.). National Geographic Books, 114517" Street, N.W,, Washington, D.C.,, 20036-4688, U.S.A. (Orders: 202-857-7335, http:// shop.nationalgeographic. eee $40.00, 416 pp., 200 color photos, 150 line drawings, 159 maps, 7 5/8" x 4" This hefty book is an excellent introduction to medicinal plants for people of all ages. Co-author Steven Foster is a noted medicinal plant expert and photographer. So, it is no wonder that beautiful photographs and clear factual writing grace this collaboration between Foster and National Geo- graphic writer and photographer Rebecca L. Johnson. Included are 150 medicinal plants arranged alphabetically by common name. Plants covered include those with common household uses like Aloe (Aloe vera), herbal medicines like Saint John’s Wore (Hypericum perforatum), and plants that are sources of drugs, such as sweet asia Artemisia annua), the source of the anti-malarial drug artemisinin. There is even a roted toa plant used as a medicine by non-humans. The section on bitterleaf (Vernonia oe describes it use by chimpanzees to rid themselves of parasites. For each plant, the authors include brief botanical descriptions, range maps, Latin names, traditional and current medicinal uses, and information on cultivation and preparation. A sidebar in each section includes interesting tidbits about bak aa intended to add to “..a better under- standing of human experience with the plant or its relatives.” The useful Latin name index includes illustrations of each plant for visual reference. There are also essays that highlight the medicinal plants and ethnobotany of regions from all over the world, including Africa, North America, and the Middle East. Although it is not a comprehensive text book, browsing through this book is an enjoy- able way to ground oneself in the basics of phytomedicine and ethnobotany.—Marissa Oppel, MS, Herbarium Technician, Botanical Research Institute of Texas, Fort Worth, Texas 76102-4060, U.S.A. SIDA 22(1): 422. 2006 CRATAEBGUS SERIES PARVIFOLIAE AND ITS PUTATIVE ELEY BRIDS IN THE SOUTHEASTERN UNITEDSEATES J.B.Phipps and K. Dvorsky Department of Biology University of Western Ontario 1154 Richmond St., North London Ontario N6A 5B7, CANADA ABSTRACT Dp D This paper deals with Crataegus series ifoli Rosaceae, subfamily Pyroideae) in the main part of its range, the southeastern United cues a species, C. uniflora and C. brittonii, are recognized together with one putative interserial hybrid, C. x vailiae. A number of other obscure entities are also discussed. The series and constituent species are provided with detailed descriptions, full syn- onymy, keys to taxa and are typified so far as possible. Line illustrations and county level distribu- tion maps, the latter derived from a database of over 1200 records, are also given for each species treated. Keyworbs: Crataegus, series Parvifoliae, Rosaceae ision, interserial hybrids, southeast- ern United States RESUMEN rE sar dae ie Parvifoliae(R subfamilia Pyroideae) en la parte princi- pal de su area, el sureste de I Os Bae: Unidos oS reconocen dos especies, C. kek ora re C. brittonii, junto con un hibrido putativo interserial, C._ vailiae. Se discuten Imbien a tidades poco claras. Tanto de | de | ipci letallad sinonimia completa, abe es de pes taxa, y se 3 tipfican en as posible para ose especie n tratada se dan ilustraciones y map mas de 1200 registros. INTRODUCTION The first author was asked to prepare the Crataegus treatment for the Vascular Flora of the Southeastern United States only vol. 1 (Cronquist 1980) of which, however, was completed. In spite of the soon defunct status of this flora JBP continued with this Crataegus project which is now being issued as a set of accounts usually dealing with individual ora few series, e.g. Phipps (1988, 1998). The third of these is presented here and deals with Crataegus series Parvifoliae. This series extends well beyond the borders of the flora area, to Texas and Mexico westwards and New York and New Jersey northeastwards; however, it is best represented in the southeastern United States. Here, there is one widespread and abundant species, C. uniflora, which is somewhat variable, plus the local C. brittonii (perhaps the same as C. rhodella of western North Carolina). Several of the names that have been synonymised with C. uniflora nevertheless repre- SIDA 22(1): 423-445. 2006 424 BRIT.ORG/SIDA 22(1) sent distinct nodes of variation and these will receive explicit comment here. Forms such as C. X vailiae and related types, however, appear to represent interserial hybrids with C. calpodendron (series Macracanthae) but are treated here for convenience. The primary function of this paper is to provide a taxonomic revision that illustrates the variation within series Parvifoliae and its putative interserial hybrids in a synoptic manner. Crataegus uniflora has often been illustrated but its more variant forms have not, as with C. brittonii, and both are illustrated here as well as the putative interserial hybrid C. x vailiae. None of the taxa rec- ognized in this paper, even C. uniflora, have ever received detailed range maps so far as can be determined. While the raw information to create dot maps did exist in our database, the scale of work necessary to provide each specimen with its mapping coordinates precluded such an ideal approach and instead, county maps, refined so as to indicate record frequency, were made. An appendix lists one representative specimen per county. A key to all taxa recognized here is also provided. The taxa recognized and their synonyms are typified so far as possible. However, lack of lectotypifiable material for the Ashe names involved has caused familiar difficulties for the typification of his names. In the cases encountered here, resolution through neotypification was not, however, adopted because of the arbitrary aspect of making type choices that would correctly match critically defective protologues. The situation with C. rhodella (discussed on p. 433) perfectly exmplifies this. Fortunately, although Ashe names consti- tute e the ae in series Parvifoliae, the variation encountered in the t d field material seen could all be accommodated within the names of other authors. The synonymy provided is based on names encountered in Palmer (1925). The paper will commence with a description of the series, fol- low with descriptions of the constituent taxa and close on the ¢ ive interserial hybrids. This work has been made possible by loans of over 1200 specimens from 43 herbaria and the first author has endeavored to see and collect representa- tive exemplars in the field of all the taxa treated in the course of over 20 field trips to the southeastern United States. However, the rarer forms, which appar- ently have always been rare, were never encountered in fieldwork, and possibly no longer exist. The database of specimens is held at UWO. The taxonomic de- cisions have been made by the first author J.B.P) and the second author (K.D.) has been responsible for creating the distribution maps. The ultimate resolution of variation in series Parvifoliae and its putative interserial hybrids invites a further generation of work in whic opu- lation studies are supported by chromosome counts, critical morphometric analysis as well as molecular studies. The revisionary treatment presented here thus erects the platform for such work. TAXONOMIC TREATMENT series eee (Loudon) Rehder, Man. Cult. Trees Shrubs, 2nd ed.1940. sect. ifoliae Loudon, Arbor. et Frutic. Brit. 2.1838. Group without rank Uniflorae Beadle, nom. a ple Bot. Stud. 1:117. 1902. TyPE SPECIES: C. uniflora Muenchh Small, + xeromorphic shrubs with fine straight thorns. Leaves deciduous; usu- ally nearly epetiolate, petioles glandular or eglandular; blades small, longer than broad, entire, and often with conspicuous black glands on the tips of the teeth; unlobed to shallowly lobed; margins crenate or crenate-serrate; + pubescent, coriaceous. Inflorescences 1-few flowered, bracteolate, the bracteoles small, ca- ducous, linear, gland-margined, membranous to thinly herbaceous. Flowers 10- 18mm diam.; hypanthium externally dense-tomentose; calyx-lobes foliaceous, us. glandular-serrate, often deeply so, as long as to somewhat longer than the petals; petals + circular, white; stamens 20, anthers us. ivory to cream; styles us. 5. Fruit up to 12 mm diam., + orbicular, us. quite hairy, yellowish to ruddy; ca- lyx-lobes conspicuous, patent, glandular-serrate, often deeply so; nutlets 4-5, dorsally grooved, sides plane. This is a small series confined to North America. It consists mostly of one somewhat variable species, C. uniflora, widespread and often common in the coastal plain from Texas to Long Island (N.Y.) with outliers further inland and in Tamaulipas, Mexico. There is also a second, much rarer species, C. brittonii, from North Carolina and adjacent states. In addition there are three dubiously placed entities, C. croomiana, C. grossiserrata and C. choriophylla that will re- ceive brief discussion. Of the 15 names assigned by Palmer (1925) to series Parvifoliae (as natural group Uniflorae), one, C. choriophylla, may not belong and is therefore only treated briefly here. This is a very rare and probably ex- tinct Florida endemic tree with distinctly petiolate, thin, strongly toothed, of- ten slightly lobed leaves, 3-5 flowered inflorescences, cream anthers and or- ange-red fruit. Sargent (1922) doubted that C. choriophylla really belongs in series Parvifoliae,a doubt which Iam endorsing, in which case it is a rare local species or hybrid. Crataegus X vailiae, a putative interserial hybrid with series Macracanthae, is treated here for convenience, on account of its foliaceous ca- lyx-lobes. Small stature, rather glandular plants, small + unlobed leaves, often uniflorous inflorescences, rather small flowers, + foliaceous calyx-lobes as long as or longer than the petals and normally non-anthocyanic anthers, are fea- tures that characterize this series. Anther color may in practice be described as white, ivory or cream, representing both degree of precision in observation and some natural variation. Series ee is pomeeS me closely related to series Triflorae, and at least in istic that flowering shoots are leafy shoots of the season and not permanent woody short shoots. However, the plant 426 BRIT.ORG/SIDA 22(1) parts are so much smaller than those of C. triflora, that even casual confusion is not possible. It is actually with smaller members of series Lacrimatae that confusion is most frequent. Members of both series are small to medium-sized xeromorphs that are rather glandular, often hairy and often have yellowish fruit. Series Lacrimatae, however, may be immediately distinguished by their zigzag twigs, non-foliaceous calyx-lobes much shorter than petals and fewer styles and nutlets. KEY TO SPECIES 1. Plants often only 1-2 m tall; leaf-blades + elliptic, 1.6:1 or narrower, unlobed; an- thers white to crea 1.C. uniflora Plants 3—4 m tall at ears leaf-blades broad-ovate or broad-obovate “broader larger leaves usually distinctly, though sometimes only are aoe anthers anthocyanic 2. Lobes of leaves obtuse; margins crenate; sides of nutlets smooth 2.C. brittonii 2. Lobes of leaves sharper; margins with sharp, distinct teeth; sides of nutlets + roded 3.C. Xvailiae 1. Crataegus uniflora Muenchh., Der Hausvater 5:147.1770; non C. ae ord Du Roi, Der Harbkesche Baumzucht, ed. 1184. 1771. (Figs. 1, 2, 3, 4, 7). Tyre U.S.A. GEORGIA. Crawford Co.: ca. 2 mi SW of Reber: 13 Apr 2006, J.B. Phipps ee YPE designated here: UWO; ISONEOTYPES: A, BRIT, GA, MO, NCU, US). Muenchhausen’s original material would be at LINN according to TL-2 but none could be located of this species. Crataegus parvifolia Aiton, Hort. Kew. 2:169. 1789. Type: Specimen labeled ‘C. tomentosa Linn. with ‘tomentosa’ crossed out; no further information except a ‘type’ label (LECTOTYPE desig- nated here: Banksian Herb. BM a parvifolia var. flor oe Loudon, Arbor. Frut. Brit. 2:42 (no Loudon herbarium exists and this variety has not been neotypified; see discussion). cee parvifolia var. grossulariaefolia Toudon, Arbor. Frut. Brit. 2:842 (this Loudon variety has also not been pear see discussion). ? Crataegus bisulcata Ashe, Bull. No iene Exp. Stn. ae 1900. Crataegus earlei Ashe, Bull. nee Carolina Exp. Stn. 175:112. 1900. Crataegus grossiserrata Ashe, Bull. North Carolina Exp. Stn. ne 112. 1900. Crataegus raleighensis Ashe, Bull. North Carolina Exp. Sm. 175.112. 1900. Crataegus armentalis Beadle, Biltmore Bot. Stud. 1:117. 1902. TYPE: U.S.A. ALABAMA Marshall Co.: near Albertville, no date, T.G. Harbison 3281, fruiting specimen (LECTOTYPE designated here: US). Crataegusgregalis Beadle, Biltmore Bot. Stud. L:118. 1902. Type: U.S.A. NORTH CAROLINA. Buncombe Co. flats of ae as River, 18 Sep 1900, Biltmore Herb. 1270b, in part (LECTOTYPE desig- nated here: US crates penance ee Elisha ane Sei. Soc. 19:12. 1903. Crataegus smithii Sarg, Trees & Shrubs 2 903. TYPE: U.S.A. PENNSYLVANIA. Delaware Co.: Lownes Run, Spr ingfield, 21 May 1902, B.H. Smith 201 (LECTOTYPE designated h Crataegus trianthophora Sarg,, Trees & Shrubs 2:11. 1907. TyPE: U.S.A. MISSOURL Ae Co. near G randin, (lectotype to be designated by Phipps et al., in press). ? Crataegus croomiana Sarg,, J. Arnold Arbor. 3:202. 1922. TyPE: U.S.A. FLORIDA. Leon Co.: near Tallahassee, 15 Apr 1920, T.G. Harbison 5710 (HOLOTYPE: A). Fic. 1.T eee bi | £¢ oar gore 4) k ° fap Pies | 1 i] £ bf amar £ € ith 1350 (MO, UWO); flowering st 1 flower details from St 214 (TENN). S. Laurie-Bourque del.; scale bars = 1 cm. 428 BRIT.ORG/SIDA 22(1) — kK + rae ad ai 7 J Ms P J 7 a from Chapman s.n. (US). S. Laurie-Bourque del.; scale bars = 1 cm. 429 Fic. 3. Parvifolia- or trianthophora-like form of C. uniflora showing several-flowered inflorescence form with pointed leaves; from Murrilf 7 (MQ). S. Laurie-Bourque del.; scale bars = 1 cm. Shrubs, us. under 2 m tall at maturity, less commonly to 3 m and occasionally reaching 5 m; extending shoots densely appressed-pubescent; one-year old twigs brown, older grayish; thorns 3-5(-8) cm long, usually plentiful, + straight, needle-like, occasionally absent, black at | yr old; young thorns commencing as soft, rapidly growing, needle-pointed-shoots with small linear bracts, often curved at first and then straightening out (Fig. 2). Leaves deciduous; petioles very short to lacking, glandular or not; blades (1-)1.5-3(-6) cm long, narrow- elliptic, elliptic, obtrullate, spatulate, oblanceolate or narrowly rhomb-elliptic, occasionally suborbiculate; cuneate at the base, obtuse to broadly acute at the tip; entire; margins crenate to dentate, with variably caducous black gland- tipped teeth though sometimes eglandular even young; venation semi- 430 BRIT.ORG/SIDA 22(1) Ss SS \\ JB VES \\ Witen ei Ave Fic. 4 VU 4 | L tyy POF a +t 9g be 9] lat L Ef g ‘3 hd = L J] f L p ll lnb A leaves with veins to si tension shoots. From left hat broad-leaved form—Phipps 7677 (UWO); pos- sible arenicola form—Shinners 12651 (GA); C. croomiana form—Murrill in FLAS 34624; C. grossiserrata form with two tension st | prigt Bozeman 7188 (NCU) §. Laurie-Bourque de I.; scale bar 431 camptodromous, lateral veins 3-4(-5) per side, + impressed; densely scabrous- pubescent above especially when young, below + densely pubescent on the veins, thinly to moderately pubescent on the surface; often somewhat glossy above at maturity, + coriaceous. Inflorescences 1-3(-5) flowered; pedicels pu- bescent, bearing us. one or two small, caducous, linear, membranous to herba- ceous, gland-margined bracteoles. Flowers 10-15 mm diam:; penta ] ns a strongl externally, sometimes bearing a linear, caducous, gland-margined bracteole; calyx- -lobes us. longer than the pet- als, 2-4 mm wide, + foliaceous, narrowly triangular, + deeply glandular-ser- rate except in rare aberrant forms, thinly pubescent on both sides; petals * cir- cular, white; stamens 20, anthers white to cream, except in rare, possible interserial hybids; styles 5. Fruit 8-10(-12) mm tall, + orbicular, tomentose, yel- low-green, yellow, or yellow orange to ruddy; flesh firm, dry or mealy; calyx- lobes to ca. 7 mm long, persistent, conspicuous, patent; nutlets 4-5, dorsally furrowed, laterally smooth. Common name.—One-flowered hawthorn. Distribution. —This species is widespread in the southeastern part of the United States (Fig. 7) from Long Island (New York) to eastern Texas with an outlier in Tamaulipas (Mexico). Inland it reaches the line Oklahoma, Missouri, southern Ohio, West Virginia. There is a remarkable disjunction in the Missis- sippi embayment between trans-Mississippian and cis-Mississipian populations that mirrors a similar pattern in C. spathulata (Phipps 1998). Crataegus uniflora is usually found in sand barrens and open scrubby woodland. Crataegus uniflora should be easy to identify on account of its small size, unusually large sepals (relative to the petals), uniflorous or few-flowered inflo- rescences and yellowish to ruddy fruit. However, it has sometimes been con- fused with some members of series Lacrimatae which include similar sized xeromorphs, also of sand-plains of the southeastern United States as is ampli- fied under the series discussion. Sterile plants can most easily be distinguished from small, sterile plants of C. berberifolia Torrey & A. Gray, a superficially similar member of series Crus-galli, by the smaller number of lateral veins on the leaves. An interesting feature of this species is the occasional deep lobing of extension-shoot leaves, even narrow ones, with veins to sinuses. The United States National Champion of C. uniflora in 1998, from Gainesville, Florida, measured 18 ft tall in 1992, an unusually great stature for this species. Crataegus uniflora is somewhat variable as befits a common species, in plant size, precise leaf shape (when strikingly obtrullate with acute tip or when suborbiculate presenting a quite different facies), number of flowers to an in- florescence (usually one), and anther color (usually ivory to cream). However, 15 specific names listed under series Parvifoliae (Palmer 1925), 11 of which are accounted for in this paper, attest to this variation and in some cases may repre- sent valid, perhay tic ies or, as we shall see, perhaps interserial L L[ a°e 432 BRIT.ORG/SIDA 22(1 = hybrids. Crataegus uniflora had never hitherto been typified perhaps partly because the protologue is unusually diagnostic for one of the period on Crataegus. The protologue clearly represents C. uniflora as construed here. I have thus neotypified on what I regard as the widespread and common form:a small shrub, with coriaceous, + spatulate to narrow obovate leaves, usually numer- ous fine straight thorns, 1-2 flowered inflorescences, glandular-pectinate se- pals, anthers white to cream and yellow-green to yellow-orange fruit. Among this group are some particularly dwarf forms of C. uniflora from laterites in Georgia that have been observed with over one hundred flowers when less than 0.3 m tall. There are two main kinds of deviation from the above norm, firstly, a group where multi (3-6)-flowered inflorescences occur and second, odd forms very like the putatively typical group but with often small and unlobed sepals. We will deal with the multiflowered types first. The most striking of those that can confidently be included in C. uniflora is C. parvifolia Aiton, lectotypified above, its type with 2-6 flowered inflorescences. A similar form is C. trianthophora, nicely illustrated by Faxon in Sargent’s protologue, scattered through the range of the species and being the main form of C. uniflora in Mis- souri. Multiflowered forms such as parvifolia and trianthophora represent one extreme of variation in the uniflora complex but the existence of two- and three- flowered inflorescences on some plants makes any varietal recogntion dubi- ous at this point. Generally, forms with multiflowered inflorescences have larger and relatively broader leaves than C. uniflora but this is not the case with the parvifolia complex. Multiflowered specimens with broad and slightly lobed leaves and unpitted nutlets are however, assigned to C. brittonii. The other main kind of variation is exemplified by C. croomiana (Figs. 4, 9), whose type comes from Tallahassee, Florida, and which has the distinction of pink anthers and entire to merely glandular-serrate calyx-lobes. Out of the approximately 1200 specimens of the uniflora complex that in the UWO data- base, ten have + entire calyx-lobes and come from the northern Florida coun- ties of Alachua, Gadsden, Leon and Liberty. The foliage and non-geniculate twigs are, nevertheless, quite typical for C. uniflora. These specimens may be part of the variation that Murrill (1942) noticed. Further investigation is re- quired to see whether C. croomiana, plus perhaps some other names, should be upheld, possibly as sporadic interserial hybrids with members of series Lacrimatae. Crataegus bisulcata from North Carolina has similar calices though ‘rounder’ leaves while the striking C. grossiserrata (Fig. 4) with a type from ‘Florida’ (no location) also has similar calyx-lobes, though often orbicular (un- usual in this series) extension shoot leaves and particularly strong and sharp leaf-teeth. A feature of these broad, extension-shoot leaves is a tendency to deep lobing, sometimes with veins to sinuses. If populations of such forms could be 433 located today, studied and shown to be worthy of name recognition, they might prove to be the same as the untypified var. grossulariifolia of Loudon. Names listed in the synonymy but so far undiscussed all belong to entities more like typical C. uniflora as defined earlier and we present below a table illustrating some of the differences among these and other forms (Table 1). Such is the homonymous C. uniflora Du Roi, judging by the protologue obviously the same species as the Muenchhausen one, and is not worth neotypifying. The differences among the remainder are small and apparently not sharp, but it is possible that they represent local, perhaps apomictic, races. The information in this table is taken from protologues, amended by personal observation. Uncom- mon forms with suborbiculate leaves may be the same as the untypified var. florida of Loudon but, as these grade into typical forms, are not considered worthy of recognition. 2. Crataegus pet Egel. Bull. Torrey Bot. Club 36:640. 1909. (Figs. 5, 8). Tyrr: U.S.A. NORTH CAROLINA. Buncombe Co.: Biltmore, 15-18 Sep 1908, W. W. Eggleston 4134 (HOLO- TYPE: NY; ISOTYPE: i ? Crataegus arenicola Ashe, J. Elisha Mitchell Sci. Soc. 17:5. 1900 ? Crataegus rhodella Ashe, J. Elisha Mitchell Sci. Soc. 18:19. 1902. Shrub, slender, to 3 or 4 m tall; thorns 3-5 cm long, often numerous, + slender, + straight to slightly recurved, blackish; extending shoots pubescent, at one year glabrous, somewhat shiny gray-brown, older becoming gray. Leaves decidu- ous; petioles 3-6 mm long, flared into the blade, pubescent; blades 2.0-3.5 cm long, elliptic-rhombic to broad elliptic-rhombic in general shape, generally widest near the middle, apex rounded to subacute, bases broad-cuneate; lobes O-1(-2) per side, if present, shallow, rounded: margins crenate-serrate, the teeth with caducous gland-tips; venation + craspedodromous, 3-4 main lateral veins per side; somewhat coriaceous, appressed-pubescent and dark green adaxially, soft pubescent and paler green abaxially; indumentum variably persistent with age. Inflorescences 2-6 flowered; branches dense-pubescent, bearing narrow, gland-bordered bracteoles. Flowers 14-18 mm diam.; hypanthium dense-pu- bescent; calyx lobes adaxially pubescent, narrow-triangular, margins + deeply glandular-serrate; stamens 20, anthers ivory; styles 3-5. Fruit subglobose, ca. 14 mm high, red or ruddy, sometimes ripening to dull brown, pubescent; calyx- lobes reflexed; nutlets 3-5, lateral faces plane, dorsally ridged. ommon name.—Britton hawthorn. Distribution. —Crataegus brittoniiis known from North Carolina, Georgia, Tennessee and Alabama. It is found in brushy places. Ashe’s protologue of C. rhodella, for which no specimens were actually cited, states that this species comes from western North Carolina, along the Cullasagee River, near Franklin, Macon Co. and the Little Tennessee River, where, although said to be common, it has not been found since that time. Although we have Tasie 1. Diagnostic features of named forms in Crataegus series Parvifoliae, from protologues. bePy Stature leaf shape infl., fl. no. styleno. —fruitcolour — other C.arenicola shrub ovate-rhomb. 1-5 ? ruddy C.armentalis 0.3-1m cuneate-spatul. 1-2(-3) 3-5 d C. bisulcata ? rounde ? ? ry. C. brittonii 3-4m ovate to br. ovate 1-6 5 brown C.choriophylla — arborescent obov.-elliptic 3-5 4-5 or-red thornless croomiana 1.5- bov.-rhomb. 1 5 ? C. earlei shrub oblanc., obl.-cun. 1 ? ? / C. gregali 1-3m cun., obov.-obl. 1-2(-3) 3-5 ruddy pyriform fruit C. grossiserrata small obov.-spatul. ] - ? coarse leaf teeth C.raleighensis shrub obov.-orbic. 1-(2-3) ? yell-gr. C. rhodell 3m rhomb., ovate 3-6 3-5 uddy diam. fls. 18mm C. smithii 1-2m V 1 5-6 or.to or-gr. flowers 7-10 days before uniflora C.trianthophora 0.6-1.2m obl-obov. 3 3-4 ruddy abbreviations use br.= broad gr.= green pk. = pink cr= cream lacin.= laciniate ur. = purple cun.= cuneate obl. = oblong Serr. = serrate ent.= entire oblanc. = oblanceolate Spat. = spatulate fls. = flowers obov.= obovate wh. = white gl. = glandular or.= orange y.= yellow (L)ZZ VIS/DYO'LINE r n Fic. 5 Biltmore Herb. s.n. ( si eahil ans, Oe eR + sie DOV). S. Laurie-Bourque del.; scale bars = 1cm. ting from Small s.n. = NY); flowering from 436 BRIT.ORG/SIDA 22(1) located none of this material, the protologue is sufficiently detailed that C. rhodella is likely conspecific with C. brittonti, the only significant difference appearing to be anther color (rose in C. rhodella) and, if conspecific, C. rhodella would become the correct name for the species. Because neither the type of C. rhodella, nor any authentic material annotated by Ashe, can be located, however, neotypifying this species is problematical. Two flowering specimens collected by Harbison (Harbison 11589 and 14999) from Dillsboro, Jackson Co., North Carolina are labelled Crataegus rhodella and with such a prominent collector there is necessarily some presumption that he was correct and that their anthers were pink. However, the label data is silent on anther color and the leaf form resembles C. x vailiae. Pink-anthered brittonii-like forms should nevertheless be carefully sought out by those that have the resources to do so. Crategus arenicola appears to be a smaller form of C. rhodella but in the absence of authentic ma- terial and similar difficulties surrounding neotypification one cannot be sure. A final group of larger plants (also 3-4 m tall) with longer petioles, ovate to broad-ovate leaves with usually sharply toothed margins, also deeply glandu- lar-serrate sepals and somewhat eroded nutlets is sufficiently different that they are referred to the putative interserial hybrid, C. x vailiae Britton, below. 3. Crataegus Xvailiae Britton, Bull. Torrey Bot. Club 24:53. 1897. (Figs. 6, a TYPE: U.S.A. VIRGINIA: banks of Roanoke River, 29 May 1890, A. Brown, s.n., T. Hoge, A.M. \ M. Timmerman, N.L. Britton and E.G. Britton (LECTOTYPE designated here: NY). Crataegus missouriensis Ashe, Bull. North Carolina Exp. Stn. 175:100. 1900. TyPE: U.S.A. MISSOURI. Shanon Co.: Montier (NEOTYPE to be designated by Phipps et al. in press Crataegus conjungens Sarg., J. Arnold Arbor. 3:200. 1922. Typr: U.S.A. VIRGINI A. Alleghany Co. Clifton Forge, 3 May 1919, T.G. Harbison15078 (-9) (HOLOTYPE: A; ISOTYPE: NCU). Bushes to 2-4(-5) m tall; thorns to 2.5 cm long; extending twigs densely pubes- cent, at | yr thinly pubescent; at 2 yrs glabrous, dark, shiny blackish-brown to dark gray; older dark gray; mature bark not recorded; thorns absent or 4-6 cm long, few, straight to slightly recurved, somewhat narrow, black at two years. Leaves deciduous; petioles 3-6 mm long, pubescent, occasionally glandular; blades 4-5(-6) cm long, elliptic to rhomb-elliptic in general shape, symmetri- cal about the middle but sometimes either end widest; subacute at the tip, ta- pered toa cuneate base; with up to 3 extremely shallow lobes per side; margins with large acute or subacute teeth except near the cuneate base; margins eglandular (on material seen by me), teeth normally sharp; venation craspedo- dromous with 4-6 veins per side; appressed pubescent above, more obscurely so at maturity, very thin pubescent (glabrescent) on the surface below but veins below persistently dense pubescent; somewhat coriaceous at maturity. Inflo- rescences 2-8(-12) flowered; branches very dense appressed-pubescent, bear- ing a few caducous, linear, membranous, gland-margined bracteoles or larger (to 15 x 3mm) more herbaceous bracteoles in the lower parts; late-flowering. Fic. 6 Crataegus fees er eee ply 1885 (A); fl ing from Harbison 15071 (NCU). S. Laurie-Bourque del.; scale bars= 1cm. 438 BRIT.ORG/SIDA 22(1) rie —— fe ae 4 7 a fy SY ) Oe ae Number of Records i 0 Fic.7 | distrit f C. uniflora in tt I United $ based on 1160 records. Not mapped north of Ohio River or Maryland and Delaware. Flowers 15-20 mm diam.; hypanthium densely spreading pubescent; calyx- lobes 6-8 mm long, narrowly triangular, somewhat foliaceous, abaxially slightly pubescent, margins deeply glandular-serrate; petals + circular, white; stamens 20, anthers cream, pink or red; styles 3-5. Fruit 7-9 mm thick, subglobose, hairy, straw yellow to ruddy at maturity in cultivation at Kew, recorded as deep red and succulent in Missouri (Bush); calyx-lobes long, margins + pectinate, re- flexed in fruit; nutlets 3-5, dorsally grooved, laterally usually shallowly pitted. Common name.—Vail’s hawthorn. TERS POE par ne ApS igen | Sa Se Oy Mumiber of Records iti a 3 [ * | Literature Record Fic. 8.C level distributi p of C. brittonii based on 17 records seen and including literature references for C. sedelle faa Macon Co., NC. Distribution.—Vail’s hawthorn has a sparse distribution in the United States and is primarily a species of southern Appalachia in Virginia, Tennessee, North Carolina and Georgia. It is also known from Missouri and Arkansas. It occurs in brushy places or on riverbanks. With its similarities to C. uniflora, C. x vailiae may be a hybrid between that species and an no in series Macracanthae, most likely C. calpodendron, on account of its laterally eroded nutlets and late anthesis. Crataegus conjungens appears to be a smaller form of C. x vailiae. Vail’s hawthorn is an attractive plant in flower and fruit when well grown, especially in forms with reddish fruit, the color nicely set off by the glossy foli- age. It is named for Anna Murray Vail, librarian at the New York Botanic Gar- den, who was one of the collectors of the type material. 440 BRIT.ORG/SIDA 22(1) Number of Records 0 Fic. 9. County level distributi fC jana based on 16 records. APPENDIX: FURTHER CITED SPECIMENS Crataegus Xvailiae Britton ARKANSAS. Hempstead Co.: 1905/04/21, Bush, B.F 14 (HUH).GEORGIA. Baldwin Co.: 1914/05/06. Harbison, 1.G. ey (HUH). Morgan Co.: 1978/05/06, Hill J.W.406 (FLAS). MISSOURI. R Aliebes 1892/ 07/23, MacKenzie, K.K. 400 (NYBG). Shannon Co.: Hee 7, Bush, B.F. 3F (CM). NORTH CAROLINA Burke Co.: 1957/06/11, Bell, C.R.8759 (NC). Guilford Co.: 1927/06/29, Manning, WE.and a Ga 1383 (HUH). Jackson Co.: 1901/05/23, Ashe WW. 32 ae Lincoln Co.: 1957/06/11, Bell, CR. 8891 (GA).Macon Co.: 1919/05/10, Harbison, TG. (A). Richmond Co.: 1954/05/12, Melvin, L. (NCU).Rowan Co.: 1904/05/17, Beadle C.D. 10103 (NYBG). Swain Co.: a eee ne i I.G.1591 (A). Vance Co.: 1874/04, Faxon, C.E. (MO). Wake Co.: 1938/04/25, ee : ‘: 3758 (MQ). VIRGINIA. Alleghany Co.: 1909/05/19, Eggleston, W.W. 4378 (NYBG). Amelia Co.: /07/24, me vee (VCU). Bucha- nan Co.: 1909/10/21, Eggleston, WW. 5594 (NYBG), Sti Co.: oo Brown,H.A.et al (NYBG). WEST VIRGINIA. Summers Co.: 1933/05/13, Boone W. 564 (WVU). ee 441 ry Number of Records 0 =a OF CO 1- ro 9- 4 Fic. 10. County level distribution map of C. X vailiae based on 39 records. Crataegus brittonii Egg). GEORGIA. Rabun Co.: 1893/08/04, Small, .K. (NY). NORTH CAROLINA. Buncombe Co.: 1934/05/ 26, Palmer, E.J.42550 (NY).Madison Co.: 1908/09/21 -22, Eggleston, W.W.4162 (NY).Stanley Co.: 1909/ 10/24, Eggleston, W.W. 5615 (NY). TENNESSEE. Cocke Co.: 1897/08/24, Kearney, -H., Jr.596 (NY). Crataegus uniflora Muenchh. ALABAMA. Bibb Co.: 1977/04/14, Sessler, A.H. 1004 (ALU). Blount Co.: 1966/06/28, Clark, R.C. and Hare, T.B. 4077 (NCU). Bullock Co.: 1967/07/19, Clark, R.C. 16433 (NCU). Chambers Co.: 1967/06/02, Clark, R.C. 13470 (NCU). Choctaw Co.: 1964/04/20, McDaniel, S. 4222 (IBE). Coffee Co.: 1967/07/04, Clark,R.C. 15618 (NCU). Covington Co.: 1967/06/20, Clark, R.C. 14402 (NCU).Crenshaw Co.: 1931/04/ 16, Palmer, J.P. 38686 (MO). Cullman Co.: 1891/06/13, Sudworth, G.B. (US). Dallas Co.: 1920/04/19, Harbison, T.G. 15275 (NCU). DeKalb Co.: 1955/05/06, Churchill, LA. (MSC). Escambia Co.: 1969/04/18, Harmon, C. 046 (NCU). Greene Co.: 1966/08/06, Clark, R.C. 6954 (NCU). Hale Co.: 1966/06/07, Will- 442 BRIT.ORG/SIDA 22(1) iams,N.337(ALU).Henry Co.: 1966/07/28, Clark, R.C.6141 (NCU). Jackson Co.: 1899/06/18, Harbison, G. (NCU). Jefferson Co.: 1899/06/02, oe ).Lee Co.: 1897/08/11, Farle, FS. (MO). ewe Co.: 1982/10/03, Gunn. S.C. 1366 (AUA).M n Co.: 1969 ss Freeman, - 695 (AUA). Marne Co.: 1967/06/05, Clark, R.C. 13644 (NCU). Sie Co.: 1977/04/23, Ross, L. 107 (ALU). Mobile Co. 1898/04/15, Mohr, C. (ALU). Monroe Co.: 1959/06/1 es en,H. ne rnenee Co.: 188 0/07/ 15, Mohr, C. (US). Perry Co.: 1967/08/10, Clark, R.C. 17835 (NCU). Pike Co.: 1931/04/15, Palmer, EJ. 38656 (MO).Randolph Co.: 1967/04/16, Clark, 2.C. 11365 (NCU). St. Clair Co.: 1951/06/24, Hollis, C.G. (ALU). Sumpter Co.: 1966/06/06, Drapalik, D. 1362 (NCU). Talladega Co.: 1966/06/20, C oe R. 5 Heard, 1A, 2895 (NCU). Tallapoosa Co.: 1973/10/06, Pringle G. ie aa A). Tuscaloosa Co.: 1982/1 07, Burckhalter, R.015 (UNA).Walker Co.: 1967/05/08, nee A.C. 12564 (NCU).Winston Co.: ie /05 10, Clark, R.C. 12995 (NCU), ARKANSAS. Cleburne Co.: 1967/0 nos Demaree, D. 56271 (IBE). Dallas Co.: 1975/07/05, Sparkman 69902 (IBE). ast Co.: a 0/13, Palmer, EJ. 26535 (A). Fulton Co 1909/05/21, Bush, B.F. 5690 (A). Garland Co.: 1925/04/23, Palmer, EJ. 26885 (A). Howard Co.: 1909/ 10/07, Kellogg, JH. (A). Independence Co.: | on Thomas, D, 9093 (NLU). lzard Co.: 1929/04/ 27, Palmer, EJ. 35547 (A). Marion Co.: 1914/06/14, Palmer, EJ. 5974 (A), Quachita Co.: 1975/08/06, Demaree, D. 69628 (\BE). Pope Co.: 1932/09/20, Merrill, G. 36 (A). Saline Co.: 1972/07/15, Demaree, D. 6524] (IBE). Sharp Co.: 1970/07/24, ali D. 15255 (NLU). Stone Co.: 1971/09/18, Demaree, D.64280 (IBE). White Co.: 1887/07/23, Coville FV. 171 (US). DELAWARE. Kent Co.: 1941/05/09, Tatnall, R.R. » (DOV). New Castle Co.: 1930/05/30, - nall, R.R. 896 (DOV). Sussex Co.: 1902/09/12, Norton, J.B.S. (MARY). District of Columbia: 1889/05/18, Burgess, E.S. (MARY).FLORIDA. Alachua Co.: 1982/ 03/14, Easterday, J.C. 821 (USF). Clay Co.: 1869/03, Canby, WM. (US). Columbia Co.: 1895/07/11-19, Nash, V. (A). Duval Co.: 1894/07/09, Curtiss, A.H.4553 (US), Escambia Co.: 1949/04/15, Hood, S.C. 1915 (FLAS). Franklin Co.: 1923/09/27, Harbison, 1.G.6046 (A).Gads 1966/04/30, McDaniel, S. 7407 (IBE). Hamilton Co.: 1900/04/17, Hugar, A. pl Jackson Co.: 1929/04/12, Palmer, E.J. 35299 (US). Jefferson Co.: 1950/09/30, Kruz, H. (FSU). Leon ioe 1920/10/06, re nison, 1.G.5710 (A). Liberty Co.: 1957/05/10, Uttal, JL. 5382 (LYN). Marion a 85/07/18, Hansen, B.F, Van Etten, M. and Wunderlin R.P. 9981 (USF). Polk Co.: 1985/06/12, Hopkins, : 3410 (FLAS). Union Co.: 1939/08/18, Murrill, WA. (FLAS).Wakulla Co.: 1964/06/21, Godfrey, R.K.64178 (FSU).GEORGIA. Bartow Co.: 1971/08/01, Wa- ters, J.C. 136 (AUA). Berrien Co.: 1973/05/06, McCarty, I. (MO). Bibb Co.: 1901/05/26, No Collector (MO). Brooks Co.: 1966/07/13, Faircloth, W.R. 3465 (MO). Burke Co.: 1938/04/10, McVaugh, R. and Byron, J.H. 2470 (GA). Chatham Co.: 195 ie Duncan, WH. 21113 (DHL). Clarke Co.: 1929/03/12, Dunnaway, JM, E7433 (GA). ee Cox 07/26, Bozeman, J.R. 6614 (NCU). Cobb Co.: 1943/05/02, Wurdack, J. Saale Co.: 1900/0 es Sargent, C.S. (A). DeKalb Co.: 1966/( eae, Wood, A. (GA). oughert « 1900/08/13 pane W.R.and baie .505 (US). Echols Co.: 1965/07/16, Dean, R. and Faircloth, te 2297 (GA). Elbert Co.: 1977/09/10, Coile, N.C. 1424 (GA). Flo on Co.: 1956/08/09, Lipps, L. 957 (TENN). Glascock Co.: 1966/07/15, Faircloth, WR. 3499 (MO). Greene Co.: need 8/13, Duncan, WH. 1585 (GA). Gwinnett Co.: 1908/04/12, Allard, H.A. 106 (US). Habersham Co.: 1914/05/ 06, Ashe, W.W. (NCU). Hancock Co.: 1966/08/09, Bozeman, LR. 7314 (NCU). Hart Co.: 1942/04/26, Duncan, W. 4846 (GAM). Houston Co.: 1979/06/01, Dixon, D. ee (NLU). Jackson Co.: 1948/04/30, Steen A. 5085 (US). es Co.: 1966/08/08, Bozeman, ee 8 (NCU). Lanier Co.: 1965/07/30, Faircloth, WR. and Norsworthy, J. 2603 (NCU). Liberty Co.: a Harbison, I.G. (A). Lowndes a 1973/05/17,McCarty, I. (MO).Macon Co.: 1898/1 0/02 aie WM., Muir, J.and Sargent, C.S. (DOV). Meriwether Co.: 1945/04/17, Henry, M.G. 3982 (GA). Morgan Co.: 1979/04/22, Hill, L. W848 (GA). sgh Co.: LA ae Montgomery, F. 1025 (GA). pet Co.: 1966/07/02, Clark, LV. (GA). Put- «1942/04/30, Duncan, WH. 4913 (GA).Rabun Co.: 1893/08/07, Small, .K. (MO). Richmond oe 1900/04/25, psa 1 (DOV ). Screven Co.: Peer 7,Cron nee 5004 (GA). Stephens Co.: 1975/07/21, Boufford, D.E. and Wood, E.W. 2096 Meseaaans Co.: 1940/05/11, Duncan, WH. 2342 se Troup Co.: 1829/06/12, Sargent, C.S. (A). Turner Co.: 1947/1 7 Thorne, R.F. 7509 (GA). Washington Co.: 1970/06/26, Hobbs, G.B. and ons " 52 (NCU). ILLINOIS. Wabash Co.: 1875, Schneck, i (MQ). KENTUCKY. Boyd Co.: 1923/05/20, Horsey, R.E. 1662 (A). Pulaski Co.: 1939/09/08, pate tN Co wo [o> — =~ Brown, E.L.2637 (A). LOUISIANA. Bi ille Parish: 1967/06/07, Thieret, /. W. 26568 (LAF). Caddo Par- ish: 1969/06/15, Thieret, J.W. 31277 (LAF). Caldwell Parish: 1973/06/02, Marx, Pet al 1207 (NLU). Clai- borne Parish: 1975/04/26, Haynes, R.R.4807 (ALU). Desoto Parish: 1979/11/10, Dixon, D. 2526 (NLU). Morehouse Parish: 1972/05/04, Thomas, D.R. 28684 (NLU).Orleans Parish: 1916/04, Cocks, R.S. (NO), Rapides Parish: 1976/06/18, Shultz, CJ). and S.£. 142 (NLU). Tangipohoa Parish: 1901/09/01, Cocks, R.S. (NO). Union Parish: 1983/07/23, Moore, D.C. 1670 (NLU). MARYLAND. Allegany Co.: 1940/06/ 12, Norton, J.B.S. (MARY). Caroline Co.: 1928/06/03, Tatnall, R.R. 78 (DOV). Montgomery Co.: 1918/ 09/29, Hunnewell, FW. 5661 (GH). Washington Co.: 1906/07/11, Jones, WR. and Shreve, F. 689 (US). Wicomico Co.: 1903/06/06, Norton, J.B.5. (MARY).Worcester Co.: 1940/06/24, Jeffery, W.F.and Norton, J.B.S. (MARY).MISSISSIPPI. Clarke Co.: 1967/09/09, Jones, S.B.and Jones, C. 15241 (NCU). pee Co.: 1961/08/20, McDaniel, S. 2689 (IBE). Jones 1978/06/03, Morgan, D.678 (IBE). Kemper Co.: 1859/ 05/01, Holgand, £. (MO).Lauderdale Co.: 1922/04/18, Harbison, T.G.6043 (A).Lowndes Co.: 1 ee 02, Holmes, W.C. 1701 (NATC). Newton Co.: 1961/07/22, McDaniel, 5.2627 (IBE). Wayne Co.: 1967/04/ 12, Bass, C, Jones, 5.B., Prescott, V. and Reynolds, J.D. 11631 (GA). MISSOURI. Carter Co.: 1893/05/16 Eggert, H. (MO). Dent Co.: 1927/09/06, Kellogg, J. (MO). Douglas Co.: 1934/08/19, Steyermark, J. 14705 MOQ). Ozark Co.: 1928/06/28, Palmer, E.J. 34812 (GH). Reynolds Co.: 1934/09/28, Steyermark, J. A. 15577 (MO). Shannon Co.: 1978/05/18, Summers, B. 396 (MO). St. Louis Co.: 1879/09/16, eee G. (MO). Texas Co.: 1934/08/15, Steyermark, J.A. 14530 (MO). Wayne Co.: 1893/06/22, ae >} (CM). NEW JERSEY. Burlington Co.: 1955/08/18, Churchill, A. (MSC). Cape May Co.: 1 i 5, Killip, E.P 2406 (US). Cumberland Co.: 1964/07/27, Gillis, W.T. 5646 (MSC). Monmouth Co.: 1976/06/ 24, Schwenk, J. (MO). Salem Co.: Vanco ip d,J.H. 1902 (MO).NEW YORK. Cattaraugus Co.: 1895/07/17, Andrews, E.F (AUA). Richmond C 79/08/22, Schrenk, J. (MO). Suffolk Co.: 1879/06/ 03,LSK (MO). NORTH CAROLINA. Alamance a ee Hammond,A.E.and Ramseur, G.$.2254 (NCU). Alexander Co.: 1935/05/08, Verrhoof, O. (NCSC).Anson Co.: 1956/09/21, Ahles, H.E.and Leisner, R.S. 19469 (NCU). Bladen Co.: 1948/06/10, Fox, WB. et al 1705 (NCSU). Brunswick Co.: 1954/08/01, Melvin, L. (NCU). Buncombe Co.: 1908/09/15, Eggleston Denote ieee Co.: 1958/09/09, Bell, R.C. 15054 (NCU). Carteret Co.: 1923/06/30, Holmes, J.S. (NCU). Caswell Co.: 1958/05/22, Bell, RC 17893 (NCU). Catawba Co.: 1891/06/25, Heller, AA and Small, J.K. (US). Chatham Co.: 1988/05/19 Swab, £.C. 162 (USCH). Chowan Co.: 1898/07/29, Kearney, T. Jr. 1903 (US). Cleveland Co.: 1956/06/22, Ahles, H.E. and Leisner, R.S. 15253 (NCU). Craven Co.: 1919/05/26, Harbison, T.G. 15090 (NCU). Cumberland Co.: 1957/05/04, Ahles, H.E.and Hammond, A.24389 (NCU). Davidson Co.: 1916/06/07, Totten, H.R. (NCU). Davie Co.: 1916/06/08, Totten, H.R. (NCU). Durham Co.: 1932/08/02, Blomquist, H.L. 83 (US). Forsyth Co.: 1948/09/07, Correll, D.S. and Correll, H.B. 14478 (DUKE). Franklin Co.: 1956/07/ 10, Ahles, H.E. and Leisner, B.S. 16508 (NCU), Gaston Co.: 1956/09/17, Ahles, H.E. and Leisner, R.S. 18810 (NCU). Granville Co.: 1956/05/22, Ahles, H.E.and Bell, C.R. 12756 (NCU). Greene Co.: 1934/05/12, Tot- ten, H.R. (NCU). Guildford Co.: 1953/05/12, Melvin, L. (NCU). Halifax Co.: 1956/07/19, Ahles, H.E. and Leisner,R.S. 16975 (NCU). Harnett Co.: 1956/09/29, Laing, H.349 (NCU). Henderson Co.: 1962/08/21, Pittillo, D.546 (GA). Hertford Co.: 1958/07/08, Ahles, H.E.and Duke, J.A.46018 (NCU). Iredell Co.: 1958/ 05/18, Ahles, H.E.and Britt, R.41050 (NCU). Jackson Co.: 1940/07/1 tere H.R. (NCU). Johnston Co.: 1919/04/26, Harbison, I.G. 14967 (NCU). Jones Co.: 1954/08/08, Corbin and pee (NCU). Lee Co.: 1955/06/27, Kessler, P-A. 350 (NCU). Lenoir Co.: 1908/10/12 see WW. 4279 (US). Lincoln Co.: 1957/06/11, Bell, R.C.8907 (NCU). Madison Co.: 1908/09/21, Eggleston ie: (CM).Mecklenburg Co.: 1958/10/02, Ahles, H.E.and Duke, .A.49968 (NCU).Montgomery Co.: 1970/05/18, Wells, E.F 2888 (NCU). Moore Co.: 1900/08/20, Ashe W.W. (CM). New Hanover Co.: 1963/06/30, Ahles, H.E. and Mc- Crary, A. 537 (NCU). Northampton Co.: 1958/07/07, Ahles, H.E. and Duke, JA. 45726 (NCU). Orange Co.: 1969/08/08, Whigham, D. 1251 (NCU). en Co.: 1951/07/31, Batson, WT. Jr. 749 (DUKE). Pitt Co.: 1956/05/09, Eason, M. 20 (NCU : ae Co.: 1921/08/11, Peattie, D.C. 1163 (NCU). Randolph Co.: 1965/10/10, Harper, FH.and Harpe een Rockingham Co.: 1956/07/07, abi F.13628 (NCU). Rowan Co.: 1908/09/1 2 oe eston, WW. 4105 (CM). Rutherford Co.: 1956/08/30, Melvin, L. 5007 (NCU). Scotland Co.: 1937/05/03, Coker, W.C. (NCU). Stanly Co.: 1891/08/18, Heller, A.A. and — 444 BRIT.ORG/SIDA 22(1) Small, .K.49 (MO).Stokes Co.: 1958/07/26, Radford, A.E. 37727 (NCU). Transylvania Co.: 1961/07/05, Rodgers, C.L.61328 (DUKE). Vance Co.: 1956/10/01, Ahles, H.E. and Leisner, R.S.20430 (NCU). Wake Co.: 1964/07/14, Sawyer, G.P Jr. 1256 (USCH). Warren Co.: 1968/06/12, O'Briant, S. and Radford, A.£. 45482 (ALU).Washington Co.: 1958/06/15, Radford, A.F. 35225 (NCU). Wayne Co.: 1957/05/03, Radford, A.E. 22009 (NCU). Wilkes Co.: 1938/06/13, Stewart, S.C. and Stewart, L. (NCU). OKLAHOMA. Choctaw Co.: 1850s-1900s, Edwards, J.E. (GH). Garrett Co.: 1905/06/08, Shreve, F483 (US).Marshall Co.: 1963/06/ 19, Ware, G. (NATC). Pusmataha Co.: 1931/05/26, Palmer, EJ. 39406 (MO). PENNSYLVANIA. Clinton Co.: 1895/06/10, Glatfelter, N.M. (MO), Delaware Co.: 1903/05/20, Graves, J.A. (MO).North Hampton Co.: 1891/08/03, Porter, T.C. (US). SOUTH CAROLINA. Abbeville Co.: 1957/06/29, Radford, AE. 25986 (NCU). Aiken Co.: 1898/08/06, Eggert, H. (MO). Allendale Co.: 1953/04/06, Batson, W.T.and Kelley, WR (USCH). Anderson Co.: 1809/07/24, Davis, J. (CM). Barnwell Co.: 1952/06/13, Batson, W.T.and kelley, WER. (USCH). Beaufort Co.: 1917/04/29, Churchill, LR. (MO).Calhoun Co.: 1957/05/19, Ahles, H.E.and Haesloop, J.G. 25520 (NCU). Charleston Co.: 1957/05/20, Ahles, H.E. and Haesloop, J.G. 25769 (NCU). herokee Co.: 1957/09/18, Ahles, H.E. 34428 (TENN). Chester Co.: 1957/06/27, Bell, R.C. 9536 (NCU). Clarendon Co.: 1914/05/22, Stone, W. 271 (A). Colleton Co.: 1983/04/04, Rayner, DA. 1593 (USCH). Darlington Co.: 1941/05/15, Smith, B.£. 947 (USCH). Edgefield Co.: 1957/05/12, Radford, A.E. 22648 (NCU). Fairfield Co.: 1983/06/22, Gould, B., LaBorde, O.and Pugh, A.E. (NLU). Florence Co.: 1982/04/ 24, McFarlane, 8.1191 (USCH). Greenville Co.: 1934/09/20, Wherry, E.7. (A). Horry Co.: 1957/09/12 Totten, H.R. 1576 (DHL). Jasper Co.: 1954/04/19, Ahles, H.F. 7807 (NCU). Kershaw Co.: 1960/05/25, Herring, WA. 3 (USCH). Lancaster Co.: 1975/05/10, Boufford, D.E. 15999 (CM). Laurens Co.: 1991/07/ 09, Horn, C.N. and Mytko, D.E.4564 (USCH). Lee Co.: 1957/06/06, Radford, A.E. 24320 (NCU). Lexington Co.: 1934/05/19, Palmer, E.J.42423 (MO).McCormick Co.: 1960/10/14, Mc Comb, C.65 (GA).Newberry Co.: 1957/06/25, Bell, R.C. 9186 (NCU). Oconee Co.: 1942/05/09, Prince, A.E. 4001 (A). Orangeburg Co.: 1983/05/23, Aulbach-Smith, CA, Long, G.and Tallevast, T. 2476 (USCH). Pickens Co.: 1975/07/18, oe . and Wood, E.W.2077 (NCU).Richland Co.: 1976/08/26, Ne/son, /.B.68] (IBE). Spartanburg 06/04, Bell, R.C.8169 (NCU). Sumpter Co.: 1933/04/17, Harbison, I.G.and Totten, H.R. (NCU). aioe ea 993/05/09, Horn, C.N. 6627 (USCH). Williamsburg Co.: 1957/06/12, Radford, A.£. 24830 (NCU). York Co.: 1957/06/05, Ahles, H.E.and Haesloop, .G. 27098 (NCU). eee aera Co.: 1961/06/30, Ellis, WH. 28599 (TENN). Cocke Co.: 1898/07, Ruth, A. 286 (US). Hamilton Co.: 1921/04, eet 1024 (US).Knox Co.: 1938/06/29, Sharp, AJ. 232 (TENN).Lawrence Co.: 18 ea 8, Ruth, 4 (MO). Union Co.: 1934/07/10, Kelly 2473 (TENN). TEXAS. Bell Co.: 1902/05/08, Reverchon, J. a Jasper Co.: 1963/04/14, Correll, D.S. a Correll, H.B.27229 (MO).Smith Co.: 1949/05/15, Shinners, JH.11183 (GA). VIRGINIA. Alleghany Co.: 1909/05/19, Eggleston, W.W.4374 (MO).Amelia Co.: 1937/ 05/25, Lewis, J.B. 555 (VPI). Amherst Co.: eas 1, Crandell, D. 9669 (LYN). Appomatox Co.: 1967/ 07/07, James, FC. 7231 (NCU). Bedford Co.: 1871/08, Curtiss, AH. (GH). Botetourt Co.: 1967/05/06, James, F.C. (NCU). Brunswick Co.: 1968/04/21, James, F.C. 13252 (NCU). Buckingham Co.: 1951/06/ 29, Jarrett, G.P and Little R.L. 14622 (US). Campbell Co.: 1893/07/20, Hel pone 099 (MO). Caroline Co.: 1968/04/28, James, FC. 13895 (NCU). Chesterfield Co.: 1971/06/10, Johnson, M.F.23220 (NCU). Craig Co.: 1967/07/19, James, EC. 7630 (NCU). Dinwiddie Co.: 1965/08/18, James, FC. 2375 (NCU). Fairfax Co.: 1970/07/31, Kiltz, BF. 57 (NLU). Fluvanna Co.: 1921/06/03, Eggleston, WW. 17740 (US). Franklin Co.: 1966/07/15, Ramsey, G.W.et al 4938 (LYN). Frederick Co.: 1941/06/19, Palmer, E.J.45271 (A). sas Co.: 1935/09/07, Adams, .W.and Wherry E. 2323 (A). Goochland Co.: 1970/05/28, Johnson, M.F. 320 Snes ae Co.: 1944/06/22, foe M.L. and Lewis, J.B. 14609 (US). Hanover Co.: 1 se 26, James, F.C. 13446 (NCU). Henry Co.: 1968/05/28, James, F.C. 10443 (NCU). Lee Co.: a James, FC.9791 ae Louisa Co.: ieee Sharp, AJ. 1102 (TENN).Lunenburg Co.: 1965/09/19, James, FC. 3071 (NCU). Mathews Co.: 1852/05, Bischoff, G.G. (MO). Mecklenburg Co.: poner Seaman, a (NCU). Middlesex Co.: 1966/07/14, James, F.C. (NCU). Montgomery Co.: 1962/ 07/02, Kral,R 8 (FSU).New Kent Co.: 1977/05/12, Bims, S. (VCU). Patrick Co.: 1950/07/24, van B.6813 ae ae Co.: 1893/07/30, Heller, A. 1099 (A). Powhatan Co.: 1967/06/08, jeniee F.C.6279 (NCU). Prince George Co.: 1936/06/19, Fernald, M.L.etal 5791 (GH).Roanoke Co.: 1969/07/ — 445 29, Uttal, L.J.6648 (VPI). Rockbridge Co.: 1881/05, /.8.C. (MO).Rockingham Co.: 1956/07/01, Allard H.A. 21957 (US). Shenandoah Co.: 1970/05/19, Churchill, A. (MSC). Smyth Co.: 1892/06/20, Small, JK. (US). Suffolk City, 1980/09/17, Wieboldt, TF. 590 (WILLI). Surry Co.: 1934/08/17, Totten, H.R. (NCU). Zuni Co.: 1908/08/19, Render, A. (A). WEST VIRGINIA. Grant Co.: 1979/05/15, Duppstadt, H.029 (WVA). Greenbrier Co.: 1959/07/29, Clarkson, R.B. 2992 (NCU). Hampshire Co.: 1929/05/30, Walker, EH. 303 (GA). Hardy Co.: 1955/06/16, Bartley, F. and Hicks, L.E.2122 (NY). Mineral Co.: 1928/10/06, Saat W.W. & Kelly, J.W. 22859 (US).Morgan Co.: 1969/05/10, Downs, R.M. 6349 (NCU). Wyoming Co.: 1900/ 08/13, Morris, E.L. 7189 (US). ACKNOWLEDGMENTS The curators of the following herbaria are thanked for the loan of specimens: A, ALU, AUA, BALT, CM, DHL, DOV, DUKE, FLAS, FSU, GA, GH, IBE, JSU, LAF LSU, LYN, MARY, MO, MSC, NATC, NC, NCC, NCSC, NLU, NO, NY, TENN, UCC, UMO, UNA, US, USCH, USF, VCU, VPI, VSC, WCU, WILLI, WVA. Vicky Papworth (BM) went to great lengths to help with the type of C. parvifolia. Bob O’Kennon of the Botanical Research Institute of Texas, Fort Worth and Ron Lance, now of Chimney Rock Park, North Carolina, are thanked for their par- ticipation in some of the first author's field trips to the area. Ms. Alison Lacon, who was an undergraduate at the time, is thanked for compiling the large speci- men database of C. uniflora without which the detail of mapping used here could not have been attained. Susan Laurie-Bourque is thanked for the beauti- ful line illustrations. The National Sciences and Engineering Research Council of Canada funded this work via a discovery grant to the first author. REFERENCES Cronauist, A. 1980. Vascular flora of the southeastern United States, vol.1, Asteraceae. Uni- versity of North Carolina Press, Chapel Hill. Murritt, W.A. 1942. New Florida hawthorns. Castanea 7:19-30. Pavmer, E.J. 1925. Synopsis of North American Crataegus. J. Arnold Arbor. 6:5—128. Puipps, J.B. 1988. Crataegus (Maloideae, Rosaceae) of the southeastern United States, |. In- troduction and series Aestivales. J. Arnold Arbor. 69:401-431. Puipps, J.B. 1998. Synopsis of Crataegus ser. Apiifoliae, Cordatae, Microcarpae and Brevispinae (Rosaceae subfam. Maloideae). Ann. Missouri Bot. Gard. 85:475-491. Pipes, J.B., R.J. O’KeENNoN, and R.W. Lance. 2003. Hawthorns and Medlars. Timber Press, Port- land, OR. Pripps, J.B.,G. Yarskievycn, and K.Woop.(in press). Typification of Crataegus (Rosaceae) names from the Missouri flora. Harvard Pap. Bot. Sarcent, C.S.1922.Notes on North American trees, X:new species and varieties of Crataegus. J. Arnold Arbor. 3:182—207. [Crataegus choriophylia, p.201] BRIT.ORG/SIDA 22(1) Book REVIEWS SHatolyA Dr La Tour and RichArp De LA Tour. 2001. The Herbalist’s Garden: A Guided Tour of 10 Exceptional Herb Gardens: The People Who Grow Them and the Plants That Inspire Them, (ISBN 1-58017-410-8, hbk.). Storey Pub- lishing, 210 MASS MoCA Way, North Adams, MA, 01247. U.S.A. (Orders: # 67410, 1-800-441-5700; wwwstorey.com). $27.50, 229 pp. color photos, color illustrations, 8 3/8" x 10" Herbalists and gardeners (or both) will enjoy this very readable coffee ele book. The authors vho also run an herb store are a husband-and-wife team of passionate medicinal plant gardeners This book is a tour of the gardens of ten well-known herbal medicine experts ai tas une Rosemary Gladstar, and others. Each section includes - Ey shies of the BarccnErs and the h | OLLIE LO really Step inside thes Se Calill tory behind their gardens. The copious color retreats. A particular gem is the section on James | Duke’ s “farmette” in Maryland. A renowned ethno- botanist and researcher, Dr. Duke's six-acre herb garden includes 80 plots, each centered ona specific health condition, such as cancer or respiratory conditions. Included in each section are lists of the gardeners’ favorite plants, watercolor garden plans, medicinal tea recipes, and tips for growing your nnician, Botanical Research Institute of own medicinal plants.—Marissa Oppel, MS, Herbarium Tec Texas, Fort Worth, Texas 76102-4060, U.S.A. LONDA SCHIEBINGER. 2004. Plants and Empire: Colonial Bioprospecting in the At- lantic World (ISBN 0-674-01487-1, hbk.). Harvard University Press, 79 Garden Street, Cambridge, MA 02138, U.S.A. (Orders: www.hup.harvard.edu/ index.html). $39.95, 320 pp., 20 black-and-white illustrations, 6 1/2" x 91/2" This page-turner is a fascinating glimpse into the mone of colonial! pa eats duri ane the’ ae ] LATS eighteenth century. ” When botany was young, voyaging from the New World ina mad race to discover the most economically and medicinally useful ae Colonial bioprospecting was a cut throat business involving botanical spies and pirates. The author presents an overview of this rough-and-tumble era, focusing on the voyaging botanists and those armchair botanists who stayed in Europe and had their plants collected by assistants, ship captains, and others. A botanical artist and scientist, Maria Sibylla Merian, was one of very female voyaging botanists during this time period. While in the rae bean, Merian described the “peacock flower” and its use by slave women as an abortifascient. Schiebinger’s ¢ arefully constructed history of this discovery and the reasons it never reached mainstream Europeans is an interesting portrait of Euro- pean reproductive mores and the business of botany in the 1700s.—Marissa Oppel, MS, Herbarium Technician, Botanical Research Institute of Texas, Fort Worth, Texas 76102-4060, U.S.A. SIDA 22(1): 446. 2006 ANEW SPECIES OF SIENANTAIUM (MELANTHIACEAE) FROM TENNESSEE, U.S.A. B. ie Wofford Dept. of Fc Biolog The Uni iyeisiy of nee SA Knoxville Tennessee 37996 ewofford@utk.edu ABSTRACT Stenanthium diffusum Wofford, sp. nov, is described as a new species from Fentress, Morgan, Pickett, and Scott counties on the northern Cumberland Plateau of Tennessee. It is most closely related to S. gramineum but differs in having (1) a more diffuse inflorescence, (2) flowers on the mid-portion of lateral panicle branches spaced =8 mm apart and with distinct pedicels (1.6-4 mm long), and (3) the uppermost non-bracteal stem leaf less than 15 cm above ground level. It also flowers later and has ecological requirements differing significant y from S gramineum Stenanthitum liffus a4) 11S an ex- tremely rare rockhouse endemic in need of consideration for state and federal ee species status. RESUMEN Se describe una nueva especie Stenanthium diffusum Wofford, sp. nov., de los condados de Fentress, Be Hee y peek del norte ne Gumbel and Plateau de Tennessee. Esta estrechamente tener (1) una eee mas difusa, (2) flores en la parte media lateral de las ramas eidel paniculo espaciadas 28 mm y con aoa otis (1.6-4 mim de longitud), is (3) la ba superior del tallo no Practeal a menos ede cm sobre el nivel del suelo. También f] y = tante S.gramineum Stenanthium diffusum es un eaaone ea raro de sale que requiere el estatus de especie amenazada estatal y federalmente. TAXONOMIC/NOMENCLATURAL PERSPECTIVE In the traditional sense, the genus Stenanthium (Liliaceae sl.) is represented by one species in Asia, S. sachalinensis F Schmidt, and from two to four taxa in North America. Stenanthium occidentale A. Gray, western featherbell, occurs in California, Oregon, Washington, Idaho, Montana, British Columbia, and Al- berta (USDA, NRCS). In the eastern United States, depending upon taxonomic interpretation, Stenanthium may be composed of the typical variety, S. gramineum (Ker Gawler) Morong var. gramineum (eastern featherbell), and two additional varieties, S.gramineum var. micranthum Fernald and S.gramineum var. robustum (S. Watson) Fernald. Stenanthium gramineum sl. ranges from Florida to Texas and northward to Indiana and Pennsylvania. Local established waifs are known from upper Michigan (Voss 1972). Fernald (1946) provided a treatment of the Stenanthium gramineum com- plex in the eastern United States and concluded that there is a a confluent series SIDA 22(1): 447 — 459. 2006 BRIT.ORG/SIDA 22 — = ) of variation and that...“As the best Ican now do propose the following defini- tion of the three varieties.” Although it is my impression that he had little faith in their distinctiveness, he did provide the following key for others to compare with their collections: a. Stem (dry) 4-10 mm thick at lowest exposed internode; leaves rather crowded be- low, rapidly cnn below paniegyl firm to coriaceous, ly opaque, the larger one wide, t ell prominently raised {ribs ee ce urface; panicle lax, the ae distant, the flowers mostly chine ee the often flexuous branches; perianth 3-8(-10) mm long; capsules ovoid-urceolate, 6-9 mm long, on spreading to reflexed pedicels; seeds 5-5.5 mm ne Stem 0.5-1.9 m high, 4-10 mm thick at base; perianth 5-10 mm long S.gramineum var. typicum (= var. gramineum) Stem 0.25—1 m high, 1.5-5 mm. thick at the base; perianth 3-4(-5) mm long var. micranthum a. Stem (dry) 7-15 mm thick at lowest exposed internode, up to 1.8 m high; leaves crowded and numerous nearly up to panicle, thin and membranaceous, translu- cent, the larger ones 1-3 cm wide, their ribs mostly immersed in the tissue; panicle usually dense, with flowers crowded along the stiffly ascending branches; perianth 5-10 mm long;capsules oblong-subcylindric, 9-10 mm long, crowded and ascend- ing to horizontally spreading; seeds 5-8 mm long Since Fernald’s (1946) assessment, the two varieties of typical S. gramineum have been largely unrecognized as distinct taxa in local and regional floras of the eastern United States as well as in the recent Flora of North America treat- ment by Utech (2002). Utech also cited and agreed with Gates (1918) and Johnson (1969) that the varieties of S.gramineum are indistinct and sympatric. lagree that significant morphological variation and overlap of characters exist but the extremes, however, are strikingly distinct and warrant consideration for additional study. Recently, Zomlefer and Judd (2002) made the following significant new combinations in the tribe Melanthieae based on parsimony analysis of ITS and trnL-F DNA sequence data, distributions, chromosome numbers, and morpho- logical data: Stenanthium sachalinensis = Anticlea sachalinensis (F. Schmidt) Zomlefer & Judd, Stenanthium occidentale = Anticlea occidentalis (A. Gray) Zomlefer & Judd, Zigadenus densus (Desr.) Fernald = Stenanthium densum (Desr.) Zomlefer & Judd, and Zigadenus leimanthoides A. Gray = Stenanthium leimanthoides (A. Gray) Zomlefer & Judd. These former Zigadenus taxa share significant morphological and geographical distinctions from the S.gramineum complex and their transfer to Stenanthium may be subject to debate by bota- nists emphasizing morphology and biogeography. Both Zigadenus densus an the poorly defined and taxonomically ambiguous Z. leimanthoides have a small gland at the base of the obovate, round-tipped tepals. They are primarily re- stricted to the southeastern U.S. Coastal Plain with the exception of a few out- liers in the mountains of West Virginia, Virginia, and North Carolina, and on the Eastern Highland Rim of Tennessee. Stenanthium gramineum and S. var. robustum WOFFORD, A NEW SPECIES OF STENANTHIUM FROM TENNESSEE 449 diffusum lack a gland at the base of the lanceolate, tortuose-tipped tepals. Stenanthium gramineum is widespread throughout most of the eastern U.S. while S. diffusum is restricted to the northern Cumberland Plateau of Tennes- see. Conversely, adoption of the new combinations of Zomlefer and Judd would leave the genus Stenanthiumss. restricted to the eastern United States. BIOLOGICAL/ ECOLOGICAL / FLORISTIC RELATIONSHIPS The Cumberland Plateau is perhaps the most floristically diverse physiographic province in Tennessee, as shown in a recent treatment of the woody plants of Tennessee (Wofford & Chester 2002). This treatment exemplifies that the Cumberland Plateau has both more native genera and native species (and lesser taxa) than any other physiographic province in the state and surprisingly a few more than the Blue Ridge which includes the Tennessee portion of the Great Smoky Mountains National Park and numerous other high mountain peaks. The highly dissected northern portion of the Cumberland Plateau in Ten- nessee (Fentress, Morgan, Scott, and Pickett counties) and Kentucky (McCreary Co.) has a unique habitat commonly referred to as rockhouses. These are am- phitheater-like semicircular recessions under overhangs and ledges of Penn- sylvanian sandstone. In addition to providing moist, sandy soil, other physical features unique to this habitat include lower light quality, lower summer and higher winter temperatures, higher relative humidity, and lower evaporation rates. Rare and/or unique taxa occur here at or behind the drip line and are only rarely in direct sunlight or rainfall. The flowering plant taxa of interest often have paler, more delicate leaves that probably would not withstand direct sunlight or heavy rainfall [see Walck et al. (1986) for an excellent review of sand- stone rockhouses with emphasis on ecology and evolution of endemic taxal. Several taxa closely related (with an asterisk) to Tennessee rockhouse endemics are often common in adjacent forests or outcrops. Examples of these sister species pairs are: Thalictrumclavatum DC.* and T. mirable Small, Arenaria glabra Michx.* and A. cumberlandensis Wofford & Kral, Ageratina altissima (L.) King & HE. Robins.* and A. luciae-brauniae (Fern.) King & HE. Robins., Silene viginica L* and S. rotundifolia Nutt., and Heuchera villosa Michx.* and H. parviflora Bartl. In addition, three pteridophyte taxa endemic to sandstone rockhouses in the eastern U.S. occur on both the Cumberland Plateau and Blue Ridge provinces of Tennessee (Walck 1986); these are: Vittaria appalachiana Mickel & Farrar, Trichomanes intricatum Farrar, and Trichomanes boschianum Sturm. The only rockhouse endemic that is also a Tennessee endemic is the herein described new species of Stenanthium. SPECIES DESCRIPTION Field observations for numerous seasons from Cumberland Plateau rockhouses have resulted in the discovery of yet another undescribed rockhouse endemic. 450 BRIT.ORG/SIDA 22(1) This new taxon is most closely related to S.gramineum but the distinct differ- ences in morphology, restricted distribution, phenology, associated taxa, and habitat requirements warrant recognition and description of a new species of Stenanthium, to wit: Stenanthium diffusum Wofford, sp. nov. (Figs. 1, 2, 4). Tyee: TENNESSEE. Pickett Co. Pickett State Park, beneath and at the margin of rockhouse ledges and overhangs on Ladder Trail at crossing of Thompson's Creek, 36°33'18"N, 84°47'55"W, elev. 1597 {t, 23 Sep 2005, BE. Wofford 20051, with D.K. Smith, G. Bresowar, & S. Huskins (HOLOTYPE: TENN; ISOTYPES: BRIT, GH, MO, NCU, NY, US) St hiogrami {fini | differt intlorescentia m; aay floril i ly i ramis later ali ae i yniciula ca. ee cel dis spos yeITIS 34¢ in | : C 3-4 mm longis portatis, et folio cauli i inus q 15cm supra es Plants glabrous perennial herbs witha single stem up to | m tall, mostly shorter, arising from a shallow seated tunicate bulb 2-8 cm deep; the bulb 2 cm high, 1 cm wide on reproductive individuals and surrounded by a dark, fibrous net- work of remains of previous years’ leaf bases. Roots fibrous, numerous, up to | mm wide. Leaves 7-10, linear, entire, up to 3 dm long, 1-3(-4) cm wide, apex acute to slightly rounded, sheathing, predominately basal, only | or 2 above the basal rosette, conduplicate basally, reflexed and flattened distally, lowermost leaves often reclining; basal and cauline leaves (non-bracteal leaves) 15 cm or less above ground level, bracteal leaves becoming progressively reduced toward the inflorescence axes. Inflorescence a terminal, bracteate, diffuse panicle up to 3.dm wide of nodding, simple or compound racemes, the terminal portion race- mose, erect, and upto 1.5dm long or mostly shorter or absent, at least not longer than the lateral racemes; lateral branches 6-10, up to 15 in larger plants, each subtended by a bracteal leaf that becomes smaller toward the stem apex, axis of lateral branches up to 1.5 dm long. Flowers numerous, mostly bisexual or with the lowermost functionally staminate and the uppermost functionally pistillate, distinctly pedicellate, those on the mid-portion of panicle branches (5-)8-15(-18) mm apart and on slender pedicels (_-)1.6-4(-5) mm long, those of the terminal raceme, if present, of similar length; ultimate pedicel bracts 13 mm long, pale margined, clasping at the base. Tepals 6, spreading to slightly campanulate, scarcely connate basally, imbricate, narrowly lanceolate, apex acute to acuminate, tortuous, white at anthesis with a pale green midrib, be- coming green and white margined with fruit development, persistent; sepals 4.5mm long, 1.2 mm wide; petals 5 mm long, | mm wide, slightly narrower and longer than the sepals, scarcely tumid along the basal margins but not glandular or nectariferous. Stamens 6, in two series, antitepalous, scarcely epitepalous; fila- ments white, 0.7 mm long, recurved; anthers L-locular, yellow, 0.4 mm long and wide, reniform, basifixed, extrorse; dehiscing intoa peltate, dumbbell shaped disc; pollen yellow, ellipsoid. Gynoecium 3-carpellate, syncarpous, placentation axile; ovary about one-third inferior, white, 1 mm long, becoming green and glaucous WOFFORD, A NEW SPECIES OF STENANTHIUM FROM TENNESSEE 451 Fic 1 uel typ Lee ah? ALL. (We ££, {INNCA , TENN) 452 BRIT.ORG/SIDA 22(1) with fruit development, ovules up to 12, most aborting; styles 3, recurved, 0.4 mm long, persistent. Fruit a glaucous capsule 7-9 mm long, 3-4 mm wide; styles per- sistent and recurved: dehiscence septicidal; seeds fusiform, sometimes flattened on one side due to compression, 1-6/locule, 3-3.5 mm long, | mm wide, brown, with irregular wrinkles, flattened and slightly arcuate at the tip. Habitat, distribution, and phenology.—Stenanthium diffusum is restricted to sandy, moist soil on ledges and below rockhouse overhangs on the northern Cumberland Plateau of Tennessee (Fig. 5). It is presently documented from only five populations in Morgan, Fentress, Pickett, and Scott counties (Fig. 6). The largest are found in Pickett State Park where they are associated with rockhouse endemics Silene rotundifolia, Heuchera parviflora, Vittaria appalachiana and the federally endangered Arenaria cumberlandensis. lt flowers from mid-Sep- tember to early October with fruits maturing in mid-November. Additional collections examined. TENNESSEE. Fentress Co.:ca.6 mi S Pickett Park, shi eras ledges, sandstone, 6 Sep 1978, R. Kral 62661 (MO & VDB, as photocopy!); Twin Arches, 13 Oct _D. Eagar s.n.(VDB, as photocopy!)*. Morgan Co.: beneath sandstone ledges at crossing of Tennessee en ley Authority transmission line above tributary to Clear Fork River, 36°18'59'N, 84°46'46'W, elev. ca. 1300 ft, 3 Oct 2005, B.E. Wofford 20053, with D.K. Smith & G. Sexton (TENN). Pickett Co.: acid sandy oak-pine woods above Thompsons Creek ca. | mi N Park Headquarters, 7 Jul 1973, R. Kral 50563; in SISNINT: p] sand of rockhouse, | lidden Passage Pickett State Park, IBA ug 1973, A. Sharp sn C1 ENN): Shai p Place Quad., Pickett State Park, Ladder Trail, series of sandstone rockhouses near Hwy. 154, in damp soil ol sandstone rockhouse overhangs, with hemlock, near Arenaria cumberlandensis, 07 Aug 1993, Milo Pyne 93-204(TENN, VDB, as photocopy!); moist sanc Ladders Trail, Pickett Park, 22 Sep 1974, R. Kral cece VDB, as photocopy); Pickett State Park, rockhouse on Hidden Passage Trail, ca. 100 yards E of its jet. with Group Camp Trail, 36°33'35'N, 84°47'26'W, elev. 1538 ft, 26 Oct 2005, B.E. Wofford ce with G. Sexton, D. Estes, & J. Beck (TENN). Scott Co.: in moist, sandy soil at base of sandstone bluffs at Twin Arches, ca. 0.2 mi. w of Charit Creek. 36°32°30"N, 84°44'09"W. 26 Oct 2005, BE. Wofford 20052, with D.K. Smith, G. Bresowar, & S. Huskins (1 ane *The correct county for this specimen is Scott Co. stone ledges in hemlock-white pine forest above DISCUSSION Stenanthium diffusum is easily distinguished from S. gramineum by morphol- ogy, geography, phenology, and habitat (Table 1). Morphologically, the most ap- parent difference between these two taxa is the nature of the inflorescence. The inflorescence of S. diffusum is broader, up to 3 dm wide, more diffuse, and the terminal, wand-like raceme is absent or nearly absent (Fig. 2). The diffuse na- ture of the inflorescence of S. diffusum is due to the more numerous and longer secondary panicle branches that usually exceed 3cm in length. In S.gramineum the inflorescence is less diffuse and rarely exceeds 1.5 dm in width except in some individuals perhaps referable to var. robustum; in addition, the terminal raceme is always present and may comprise up to 20-70% of the total inflores- cence length (Fig. 3). Furthermore, in S. gramineum, secondary branches are infrequent and generally less than 2 cm long. WOFFORD, A NEW SPECIES OF STENANTHIUM FROM TENNESSEE 453 Fic. 2. Stenanthium diffusum habit at anthesis. Image taken at type locality, Ladder Trail, Pickett State Park, Pickett County, Tennessee. 454 BRIT.ORG/SIDA 22(1) Taste 1.Morphological and other comparisons of Stenanthium diffusum and S.gramineum. Quanti- tative values are based on measurements from specimens deposited in the University of Tennes- see Herbarium (TENN). S. diffusum S.gramineum Pedicel length at mid-portion of lateral (1-)1.6-4(-5) (0-)0.3-1.1(-1.7) inflorescence branches (mm) Distance between pedicels at mid-portion (5—)8-15(-18) (1-)3-7(-8.5) of lateral inflorescence branches (mm) Distance from ground to uppermost (3-)4-14(-15) (15-)22-66(-76) cauline leaf (cm) Distribution northern Cumberland Plateau of TN FL to TX,n to IN and PA Flowering period (at ca.same latitude) mid a to mid-Oct May to late Aug abitat moist, shady,sandstone — dry to mesic woodlands rockhouses and open areas Stenanthium diffusum and S.gramineum also may be separated by quanti- tative morphology, phenology, distribution, and habitat (Table 1). The flowers of the mid-portion of the lateral inflorescence branches of S. diffusum are sepa- rated from each other by =8 mm and are borne on distinct pedicels (=1.6 mm long) that often equal or exceed the subtending bracts. In comparison, the [low- ers of the lateral inflorescence branches of S.gramineumare sessile or subsessile on pedicels (<1.1 mm long) that are equal to or shorter than the subtending bracts and are separated from each other by <7 mm, giving the inflorescence a less diffuse appearance. Interestingly, although the pedicels on the lateral branches are sessile or subsessile, those of the terminal wand-like raceme are distinctly pedicellate and may be up to 5 mm long. Another reliable character- istic that readily separates S. diffusum and S. gramineum is the distance from the ground to the node of the uppermost cauline leaf (not bracteal inflores- cence leaf). The distal-most cauline leaf of S.diffusum is <14 cm above ground level whereas the distal-most cauline leaf of S. gramineum is >22 cm above ground level. Although somewhat difficult to quantify from herbarium mate- rial, there also appears to be a difference in leaf length/width ratio between these taxa, with the tendency toward a lower leaf length/width ratio in S. diffusum. Other non-morphological differences include: (1) phenological differences, ie., S.diffusum flowers from mid-September to mid-October and S.gramineum, at this approximate latitude, flowers from May to late August, (2) S. diffusum is a narrow endemic to the northern Cumberland Plateau of Tennessee in sand- stone rockhouse habitats characterized by siliceous soils, low light intensity, high relative humidity, lower evaporation rates, and lower summer and higher winter temperatures than the surrounding forests. The more widespread S. WOFFORD, A NEW SPECIES OF STENANTHIUM FROM TENNESSEE Fic.3. Stenanthium gramineum habit at anthesis. gt fl BRIT.ORG/SIDA 22(1) h, from holotype Fic. 4. Stenanthium diffusum, image of flowers at mid- WOFFORD, A NEW SPECIES OF STENANTHIUM FROM TENNESSEE 457 Fic. 5. Habitat of Stenanthium diffusum. Sandstone rockhouse formations at type locality, Ladder Trail, Pickett State Park, Pickett County, Tennessee. gramineum occurs in a much broader range of ecological conditions in numer- ous habitats including rich woods, moist to dry woods, meadows, flood plains, prairies, and grassy balds. The following key is provided for ease of identitica- tion: 1. Inflorescence a diffuse panicle up to 3 dm wide, the terminal racemose portion reduced or absent;flowers on mid-portion of lateral branches with distinct pedicels 1.6-4 mm long and 8-15 mm apart; uppermost non-bracteal stem leaf 4-14 cm above ground level S. diffusum Inflorescence not diffuse, typically up to 1.5 dm wide, the terminal racemose por- tion present and up to 3 dm long; flowers on mid-portion of lateral branches with pedicels 0.3-1.1 mm long and averaging 3-7 mm apart; uppermost non-bracteal stem leaf 22-66 cm above ground level S.gramineum (including var. micranthum and var. robustum) CONSERVATION CONCERNS The northern Cumberland Plateau of Tennessee and adjacent Kentucky har- bors a suite of species commonly referred to as rockhouse plants. This unique flora is restricted to sandstone floors, ledges, and overhangs. There are several 458 BRIT.ORG/SIDA 22(1) ad em aul an eee @epmeemen eae ar > oa] VA NC sc Legend \ ® Populations \ \ 100 ‘ TT Kilometers Fic 6 r + J Ly + Es | - £ Cy oy AT; critical factors required, in concert, to maintain these unique habitats: moist sandy soil, low light intensity, high relative humidity, low evaporation rate, and ower summer and higher winter temperatures than adjacent forests (Wofford & Smith 1980). The environmental factor most likely to have an immediate and negative impact would be removal or severe thinning of adjacent woodlands. This would increase light intensity and temperature while reducing relative humidity and moisture. An aggressive management plan should be placed into motion on public lands (especially at Pickett State Park and Forest and the Big South Fork National River and Recreation Area) aimed at protecting a part of this critical habitat and its associated flora. In addition to protecting a unique part of Tennessee's natural heritage, protection would provide opportunity for study of biological/evolutionary problems yet to be resolved, i. e., speciation and sis- ter pair relationships, micro and macroclimates, endemism, rarity, etc. There- fore, itis highly recommended that the Tennessee Department of Environment and Conservation and the U.S. Fish and Wildlife Service, in coordination with public land managers, conservation groups, and interested individuals, propose — _ WOFFORD, A NEW SPECIES OF STENANTHIUM FROM TENNESSEE 459 both state and federal endangered status for Stenanthium diffusum. Hopefully, these efforts can, in concert, accomplish the necessary management and mis- sion plans to preserve this unique habitat and flora. ACKNOWLEDGMENTS I thank the directors/curators of MO and BRIT/VDB for providing images of specimens. Special thanks goes to the ey and graduate students at the Uni- versity of T for their « numerous field trips; to Dwayne Estes who carefully reviewed early drafts of the manuscript; to Guy Nesom who provided the Latin diagnosis; to Chris Fleming for constructing the distribu- tion map; to anonymous reviewers for their helpful comments, to Jerry Drown for providing the image of Stenanthium gramineum, and to the Hesler Fund, University of Tennessee, for funding field trips. REFERENCES FerNALD, M.L. 1946. Stenanthium in the eastern United States. Rhodora 48:148-152. Gates, R.R.1918.A systematic study of the North American Melanthiaceae from a genetic standpoint. J. Linn. Soc. Bot. 44:131-172. JOHNSON, R.G. 1969. A taxonomic and floristic study of the Liliaceae and allied families in the southeastern United States. Ph.D. dissertation. West Virginia University. USDA, NRCS. 2005.The PLANTS Database, Version 3.5 (http://plants.usda.gov). Data com- piled from various sources by Mark W. Skinner. National Plant Data Center, Baton Rouge, LA. 70874-4490 USA. Utec, F.H.2002. Stenanthium.|n:Flora of North America Editorial Committee. Flora of North America north of Mexico. Magnoliophyta: Liliidae: Liliales and Orchidales. Oxford Univ. Press, New York, New York. Vol. 26. Pp. 88-89. Voss, E.G. 1972-1996. Michigan flora, 3 parts. Part 1,Gymnosperms and monocots. Bloom- field Hills and Ann Arbor.P.418 Watck, J.L., JM. Baskin, and C.C. Baskin. 1986. Sandstone rock houses of the Eastern United States, with particular reference to the ecology and evolution of the endemic plant taxa. Bot. Rev. 62:311-362 Worrord, B.E.and R.Krat.1979.A new Arenaria from the Cumberlands of Tennessee. Brittonia 31:257-260. Worrorb, B.E. and E.W. CHester. 2002. Guide to the trees, shrubs, and woody vines of Ten- nessee. The University of Tennessee Press, Knoxville. Worrorb, B.E. and D.K. SmitH. 1980. Status report on Arenaria cumberlandensis Wofford & Kral. Report submitted to U.S. Fish & Wildlife Service, Atlanta. Zomterer, W.B. and W.S. Jupp. 2002. Resurrection of segregates of the polyphyletic genus Zigadenus 5.l. (Liliales: Melanthiaceae) and resulting new combinations. Novon 12: 460 BRIT.ORG/SIDA 22(1) Book REVIEW RiA Loonuizen. 2005. The Elder in History, Myth and Cookery. (ISBN 1-903018- 31-5, pbk.). Prospect Books, Allaleigh House, Blackawton, Totnes, Devon TQ9 7DL, United Kingdom. (Orders: The David Brown Book Co., PO. Box 51], 28 Main St., Oakville, CT 06779, U.S.A., 860-945-9329, 860-945-9468 fax, Email: david.brown.bk.co.snet.net). $17.95, 140 pp., 51/2" x 71/4" This petite book is an affectionate ode to the European elder (Sambucus nigra). It includes chapters on elder botany, mythology, and cuisine, as well as medicinal and other uses for the plant. Ina chap: of ter titled “The Healing E Ider,” the author reviews traditional medicinal uses of the elder. The proper harvest and drying of eldenlowst>: e uSeHOET IES and leaves is ite Piesnone io making. oint- } ments, syrups pouivs es that the E | tiviral and ad ic eff pporting i liti easa treat- ment for ee A chapter named “The C acme! “Ider” describes Bidens recipes, which sneha e such delicacies as elderf ia tuack ureter wine Lula jelly, CEA sour soup, and recipes for the Jew’s Ear mushroom (Auric ie-judae), which grow r wood. These reci- J pes can be prepared using the American elder na us ioe nsis). However, caution should be can | rt ff t. Pep used because large amounts of elderberries and bar red t ee the book are illustrations, photographs, and quotes ae to the elder. The author es vith a quote by the poet Seamus Heany: “I love its blooms like saucers brimmed with meal, its berries a pine of shot, a buoyant spawn, a light bruised out of purple. Elderberry? Itis shires dreaming wine.’—Marissa N. Oppel, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, Texas 76102-4060, U.S.A. SIDA 22(1): 468. 2006 A NEW SPECIES OF DODECATHEON (PRIMULACEAE) FROM THE NORTHERN COAST RANGE OF OREGON AND WASHINGTON Kenton L. Chambers Department of Botany and Plant Pathology Oregon State University 2082 Cordley Hal Corvallis, Oregon 97331-2902, U.S.A. ABSTRACT Dodecatheon austrofrigidum is described as a species occupying scattered sites in the Coast Ranges of northwestern Oregon and adjacent W, ere USA. It is similar in floral morphology to, but widely disjunct from, D. frigidum, a species of the interior mountains and arctic tundra of Alaska and northern British Columbia. It also is ecologic cally distinct, being associated with streams, water- falls, cliffs, rocky river banks, and high elevation moist grassland and basalt talus slopes in a mari- time climatic zone. Its relationships are with members of Sect. Dodecatheon, but its seedling mor- phology is different from any described thus far in the genus. Morphology of the seed testa is illustrated by scanning electron microscopy. RESUMEN Dodecatheon austrofrigidum se describe como una espec upa lugares espaciados de la dillera Costera del Noroeste de Oregon y de W. ashington, U SA Es en en 2 puorteloe al i pero muy disyunta de D.frigidum le Alaska y norte de British Colum bia. imapien es ecologicamente diferente, estando asociada a arroyos, cascadas, acantilados, lechos rocosos de rios, y praderas humedas a gran altitud y laderas basalticas n una zona climatica marina. Esta relacionada con los miembros de la Sect. Dodecatheon, pero la morfologia de sus plantula liferente de cualquiera de las hasta gee descritas en el género. Se Ha ilustra la moriolose de testa seminal al microscopio electrénico de barr INTRODUCTION The northern Coast Ranges of Oregon and immediately adjacent Washington, especially on the higher peaks, are known to harbor a small suite of endemic plant species. Prominent in this list are Cardamine pattersonii LF Hend., Castilleja chambersii M. Eggers & R. Meinke, Erythronium elegans P Hammond & K.L. Chambers, Filipendula occidentalis (Wats.) Howell, and Saxifraga hitchcockiana Elvander. A floristic affinity to the Olympic Mountains and coastal Alaska is suggested by more northern taxa whose ranges extend south to this region. These include Carex macrochaeta C.A. Mey., Cladothamnus pyroliflorus Bong., Erigeron peregrinus (Pursh) Greene var. peregrinus, Lewisia columbiana var. rupicola (English) CL. Hitchc., Prenanthes alata (Hook.) Dietr, Saxifraga caespitosa var. emarginata (Small) Rosend., Senecio flettii Wieg.. and SIDA 22(1): 461 — 467. 2006 pas 462 BRIT.ORG/SIDA 22(1) Synthyris schizantha Piper (Chambers 1973, 1974). Saddle Mountain, Clatsop County, Oregon, is a well known botanical site where many of these species have been found (Detling 1954). Also known from Saddle Mountain, near the 1000m summit, is a dwarfed form of Dodecatheon, which the monographer, H,J. Thompson (1953, p. 118) re- ferred, with some hesitation, to D. pulchellum (Raf.) Merrill (his D. radicatum Greene). A number of years ago! was directed to populations of vigorously grow- ing examples of this plant along the Trask River, Tillamook County, Oregon, which I brought into cultivation. This tentative new species was then made known to interested botanists under the name Dodecatheon austrofrigidum, which has now come into general use for the plant. [here provide a description and further documentation of this species. Dodecatheon austrofrigidum K.L. Chambers, sp. nov. (Figs. 1, 2) Type: US.A. Or- EGON. Tillamook Co. Trask River Rd., 6.1 mi E of junction with Long Prairie Rd., TIS, R8W, §28, elev. ca. 200 ft, in patches of moss on bare, cea: rocky bank of the ees River, be- tween low and high water marks, with Saxif llii, S. mertensiana, Mi 13 May 1989, K.L Chambers ae OTYPE: OSC: ISOTYPES: FSU, G, MARY, MO, NY, OSC, UC, US, WS, WTU a Planta ad Dodecatheon frigidum et Y aes — Sea variis ase ede ci seis glabro dentibus triangularibus acutis lobis corollae ibe ] eris connectivis antherarum purpureis laevibus stigmate non capitato ab D. fr an radicibus numerosis fibrosis validis caudice plerumque per annos paucos persistenti differt, ab D. dentato corolla lavandula non alba differt; chromosomatum numerus 2n = 88 Perennial herb; roots numerous, stout, fibrous, often producing adventitious buds; caudex usually short-lived, with subsequent years’ vegetative growth aris- ing from a terminal bud or from small buds on the roots; leaves in a basal ro- sette, glabrous, variable, 2.5-30 cm long, 0.7-7 cm wide, ovate to broadly or nar- rowly elliptic, gradually or abruptly tapering to the petiole, margins entire to irregularly sinuate-dentate or denticulate; flowering stems scapose, 5.5-45 cm igh; inflorescence a bracteate umbel of 1-7 pedicelled flowers, bracts 2.5-10 mim long, pedicels 4-55 mm long, glabrous to glandular-puberulent, especially distally; ealyx 5-11 mm, tube 1-2.5 mm, lobes 3-9 mm, triangular, acute; corolla lobes 9-23 mm long, 2-6 mm wide, lavender, tube 1-2 mm long, white with a sinuous purple line where reflexed; anthers purple, +.5-8 mm long, filaments 0.5-1 mm, free or slightly united, purple, connective smooth, purple; style ex- serted, stigma not capitate; capsule 6-16 mm long, 3.5-5.5(-7) mm wide, teeth ca. 1/5 of total length, dehiscence operculate or valvate; seeds 1-2 mm, irregu- larly rounded or prismatic, foveolate (Fig. 2); seedlings with petiolate cotyle- dons, the first principal adult root produced adventitiously from the apex of the hypocotyl, the first true leaf petiolate, arising between the cotyledons (Fig. . The chromosome number is tetraploid on the base of x = 22 characteristic of the genus. CHAMBERS, A NEW SPECIES OF DODECATHEON FROM OREGON 463 Fic. 1.Dodecath trofrigidum K.L. Chambers. A. Habit. Scale=2 cm. B. Flower. Scale=7 mm.C. Capsule with opercu- lum. Scale=7 mm. 464 BRIT.ORG/SIDA 22(1) Fic. 2. Seed. Scale=0.1 mm. Distribution.-Ranging from 30-1200 m, at scattered sites from the south- ern Olympic Mountains, WA (Mt. Colonel Bob) to Tillamook County, OR, both at high elevations in moist, grassy turf (Saddle Mountain and Onion Peak, Clatsop Co., OR) and in forested regions, where the habitats include moist ba- saltic slopes, talus, and cliff-faces, especially near waterfalls and streamlets. Further populations occur at low elevations in Tillamook Co.,on bare or m Ossy basalt rock banks of rivers, between the low- and high-water marks, in the win- ter flood-scour zone (Raven 1995, 1996). Plants in this peculiar habitat probably become established by seeds washed down from the mountain populations, and their tenacious mass of strong roots, penetrating crevices in the rock, anchors them from being washed away during the period of winter flooding. The spe- cies is allopatric in range to all other members of the genus. Flowering. —April-July, depending on habitat and elevation. Flowers are self-fertile and capable of self-pollination in cultivation. Relationship.—Thompson (1953) recognized three sections within the ge- nus, differentiated by characteristics of the stigmas, capsules, seeds, and seed- lings. The new species is best assigned to Sect. Dodecatheon, with the proviso that D. frigidum Cham. & Schltdl. should also be placed in this section, rather than Sect. Purpureo-tubulosa of Thompson. The corrected placement is strongly supported by molecular evidence provided by Mast et al. (2004) and by the — CHAMBERS, A NEW SPECIES OF DODECATHEON FROM OREGON 465 marked floral similarity between D.frigidum and D. austrofrigidum. Mast et al. (2004) show a strict consensus tree based on seven cpDNA regions, in which D. frigidum is sister to the group of other taxa in Sect. Dodecatheon, rather than forming a clade with D. conjugens Greene in Sect. Purpureo-tubulosa, as pro- posed by Thompson. In this DNA-based phylogeny, D. austrofrigidum is paired in Sect. Dodecatheon with D. dentatum Hook. ssp. utahense (N. Holmgren) Kartesz (perhaps itself a distinct species). Dodecatheon austrofrigidum differs in two significant features from the morphological characterization of Sect. Dodecatheon given by Thompson (1953). These, and an improved knowledge of the seed testa based on SEM stud- ies (Fig. 2), will be discussed in turn. Most important, I believe, is that seedling morphology and development are distinctly different than had been described for this section. In D. austrofrigidum, the seed begins with two petiolate cotyle- dons (as Thompson, 1953, Fig. lg, illustrates for D. pulchellum). The first true leaf, itself petiolate, next arises from between these cotyledons. Finally, a strong, major adult root forms adventitiously at the base of this first true leaf, between the cotyledons, replacing the seedling’s initial but ephemeral root system (Fig, 3). In the seedling of D. pulchellum (D. radicatum) illustrated by Thompson (loc. cit.), the first true leaf is linear-spatulate, not petiolate, and arises adventi- tiously from the hypocotyl well below the cotyledons; the new adventitious major root grows downward from that same point on the hypocotyl. Neither of the other two seedling types illustrated by Thompson (loc. cit.) resembles that of D. austrofrigidum. The mature capsules of D. austrofrigidum differ in their dehiscence mode from that described for Sect. Dodecatheon (Thompson, op. cit., p. 79). The sec- tion is characterized as having valvate capsules, but in D. dustrofrigidum, they are most often operculate instead. Capsules are thin-walled below and have an indurate apex where the cells are thick-walled and isodiametric. As these cells dry at maturity and shrink, tensions develop that rupture the tissue and open the capsule for seed dispersal (Fig. 1C). A transverse rupture through the tip region causes an operculum to free itself, carrying with it the dried style. Usu- ally five lengthwise splits also occur at the same time, but some of the five teeth thus formed may in turn split in half, so that dehisced capsules may be found with up to ten teeth. In a sample of 181 capsules, the average number of teeth was 6.28, the numbers of capsules with 5, 6, and 7 teeth being approximately equal. The initial five splits can often be observed in the operculum (Fig. 1C). Valvate dehiscence, where the lengthwise splits reach the apex, is usually noted in smaller, less robust capsules. Seeds of D. austrofrigidum (Fig. 2) conform to the description of Sect. Dodecatheon (Thompson op. cit., p.75-76). They are irregular in shape but of - ten flattened on two or more sides, becoming even more prismatic than in Thompson's drawing (op. cit., Fig. lf). They do not resemble the membranous- 466 BRIT.ORG/SIDA 22(1) t ¢.3. Seedling, showing petiol yled 1 first leaf. A major adventiti tight, repl he initial, short- lived seedling hypocotyl and root at the left. Scale=5 mm. margined seeds of Sect. Capitulum (Thompson loc. cit.). The testa is formed ini- tially of tightly packed bulbous cells, which at maturity deflate into irregular, rimmed, dish-shaped foveae (Fig. 2), best visualized by SEM. In view of these newly published observations on D. austrofrigidum, it is likely that a survey of CHAMBERS, A NEW SPECIES OF DODECATHEON FROM OREGON 467 seed, seedling, and capsular morphology in all the species will contribute toan improved understanding of intrageneric relationships. Cited collections: U.S.A. OREGON. Clatsop Co.: Onion Peak, elev. 3064 ft, T4N, RIOW, $22, 29 Jun 1971, aus a 3169 (CAS, OSC); same, 21 Jul 1971, Chambers 3258 (OSC); Saddle Mtn, T6N, R8W, $33, 29 972, D. Jaques 973 (OSC); same, elev. 2600 ft, TON, R8W, S19, 24 May 1950, Detling 6550 (CAS, ae "v0 madals Mi elev. 3200 ft, 10 Jun 1928, G. R Patterson s.n. (ORE). Tillamook Co.: Trask River Rd.a ,on pasa pani ef Trask R. ieee 1980, Chambers 4694 (OSC); same, 10 Jun 1979.6. Lewiss.n (OSC. S d.aboy k Ck., elev. ca. 3200 ft, TZN, R7W, S13 1, photo, Jun 1981, G. Lewis s.n.(OSC); near Diamond Falls, he State Forest, elev. 1480 ft, T2N, ROW, S32, 13 Jun 2001, Chambers 6300 (OSC), Kilchis Falls, Tillamook State Forest, elev. 1540 ft, TIN, R8W, $3, 13 Jun 2001, Chambers 6299 (OSC). WASHINGTON. Grays Harbor Co.: Mt Colonel Bob, above Moon- shine =e elev. 3800 ft, T23N, R8W, S18, 20 Jul 1983, E. Alverson 590 (OSC); same, 6 Jul 1994, D. Davis cific Co.: Willapa Hills, headwaters of West Fork of Grays R., elev. 1500 ft, TILN, R7W, S10, : yee eee Powell 1324 (OSC): same, elev. 1800 ft, 23 Apr 1995, J. Powell 1410 (OSC). ACKNOWLEDGMENTS The author gratefully acknowledges the help of the following colleagues in this study: Dave Buchholz, Dan Clough, Thea Cook, Aaron Liston, Cathy Maxwell, Barbara Moore, Andrea Raven, James Reveal, Larry Scofield, Rena Schlachter, Clint Smith, and Al Soeldner. John Megahan prepared the illustrations. Special thanks are extended to Gerald Carr, who made the cytological preparations, and George Lewis, who first directed me to populations of Dodecatheon austrofrigidum on the Trask River. Austin Mast and Harry Thompson provided prompt and helpful reviews. REFERENCES CHAMBERS, K.L. 1973. Floristic relationships of Onion Peak with Saddle Mountain, Clatsop County, Oregon. Madrono 22:105-114. CHameers, K.L. 1974. Notes on the flora of Clatsop Co., Oregon. Madrono 22:278-279. Detune, L.E. 1954. Significant features of the flora of Saddle Mountain, Clatsop County, Oregon. Northwest Sci. 28:52-60. Mast, A.R., D.M.S. Feuer, S. Ketso, and E. Conti. 2004. Buzz-pollinated Dodecatheon originated from within the heterostylous Primula subgenus Auriculastrum (Primulaceae):a seven- region coDNA phylogeny and its implications for floral evolution. Amer. J. Bot. 91: 926-942 Raven, A.N. 1995. Dodecatheon austrofrigidum monitoring on the Trask River Peninsula, Tillamook County, Oregon. Berry Botanic Garden, Portland, OR. Raven, A.N. 1996. Summary of six years of research on Dodecatheon austrofrigidum, the Frigid Shootingstar, at the Trask River Peninsula, Tillamook County, Oregon. Berry Botanic Garden, Portland, OR. THompson, H.J. 1953. The biosystematics of Dodecatheon. Contr. Dudley Herb. 4:73-154. 468 BRIT.ORG/SIDA 22(1) Book REVIEW WILLIAM C. BurGer. 2006. Flowers: How they Changed the World. (SBN 1-59102- 407-2, hbk.). Prometheus Books, 59 John Glenn Drive, Amherst, NY 14228- 2197, U.S.A. (Orders: 716-691-0133, 716-691-0137 fax, http://www. prometheusbooks.com/). $23.00, 337 pp., b&w figures, color photographs, glossary, and notes, 5" X 7" William Burger has written a book that is hard to put down. Flowers: How they Changed the World is a fantastic introduction to the angiosperms, their evolutionar y idv ces, and the roles plat gh play in the evolutionary history of man. This book is written in such a way ah at it will appeal to those read- ers looking for an introduction to flowering plants. However, it also provides the seasoned readers much food for thought. The writing style is very ss and conversational with a good deal of wit, k yet includes an abundance of information. Th s many topics including: what is a flower, the purpose of flowers, flower friends, ne enemies, how to distinguish flowering plants, what makes flowering plants special. It also discusses how primates, people and flowering plants relate to rer, which each other and in the end how flowers changed the world. Topics flow well into each ot greatly adds to the readability of the text. The author is clever about ke ing new subjects and making these topics non-threatening. For exam ie the book starts with introductions to four plants, a geranium, rose and two lilies. The author hints at the importance of distinguishing traits between monocotyledons and dicotyledons, but doesn’t delve into more detail until later chapters; smartly building on these unassuming intro- ductions made in the first chapter. This book contains well-written discussions and a wonderful introduction to major evolutionary advances in plant reproductive biology. Among these are how pollen helped flowering plants do well on land, the value of deciduous leaves, the biology related to losing leaves from a plant, and the evolutionary advances in plant defenses of chemicals that help prevent peEovory and disease. The author includes many other interesting discussions on subjects such as the importance of out- sea in eee and how plants have adapted ways to 2 out- nrere bree ding, as well as the diversity in fo re of [lowerin ng pla ants. The i : c ay 5 _ cover information on how plants have been oa tats to people, focusing on mankind's develo ment of agriculture and the importance of biodiversity to our lives and the Earth. There is ales. a section that summarizes the important role plants have played in creating our world and the modern lives of man in ten major stages. lowers: How they Changed the World by William Burger is an extraordinary and interesting kgrounds and inter- jon book to read; full of engaging information for readers of all ages, educational bac ests. The text is written in an easygoing style and covers a wide range of topics related to the evolu- tion and biology of flowering plants. The author has also gone above and beyond typical notes for the rea der; he eeiauceh both ee mation < | refer rences s for each chapte1 topic accented by his own edu- cated val Lucke ydoo, Herbarium, Botanical Research Institute of Texas, 509 Pe- can Street, Fort Worth, TX a 4060, U.S SIDA 22(1): 468. 2006 THE LECTOTYPIFICATION AND 197" CENTURY HISTORY OF CROTON ALABAMENSIS (EUPHORBIACEAE SS.) Kenneth J.Wurdack Department of Botany Smithsonian Institution PO. Box 37012, NUNH MRC-166 Washington, DC 20013-7012, U.S.A. wurdackk@si.edu ABSTRACT A detailed account is presented of the historical | he di of Croton alabamensis var. Ts one eco E.A. Smith in 1877 avd its subsea ent study by 19th century American botanists. A lectotype is designated and its RESUMEN Este articulo presenta una descripcion detallada de los eventos relacionados con el descubrimiento de Croton alabamensis var. alabamensis (Euphorbiaceae ss.) por E.A. Smith en 1877 y su estudio subsiguiente por botanicos en el siglo XIX. Ademas, se designa un lectotipo para esta especie y se discute su origen. INTRODUCTION The progress of botany in the southern United States during the latter half of the 19th century has generally not been critically studied. Most of the major figures have yet to be treated by more than brief biographical sketches. Histori- cal narrative is usually traced from the perspective of a particular person or institution, but a few rare southern plants (e.g., Croton alabamensis E.A. Sm. ex Chapm., Dionaea muscipula Ellis, Elliottia racemosa Muhl. ex EIL., Franklinia alatamaha Bartr. ex Marsh., Shortia galacifolia Torr. & A. Gray, and Torreya taxifolia Arn.) have attracted sustained botanical interest from discovery to the present day and are therefore of noteworthy historical interest. Croton alabamensis contains an especially rich and complete historical record that is chronicled here to recover lost details about the type collection and reveal ex- tensive and collegial interactions among the network of botanists of the day. Croton alabamensis is, in habit, the largest native United States representa- tive of the tropical-centered family Euphorbiaceae s.s. The species has a disjunct distribution between Alabama and Texas and contains two morphologically well-differentiated, geographically-named varieties (vars. alabamensis and texensis Ginzbarg, respectively). Croton alabamensis is readily distinguished from other North American species of Croton L. by its shrubby habit, petaliferous female flowers, and covering of lepidote scales that give the abaxial leaf sur- — SIDA 22(1): 469 - 483. 2006 470 BRIT.ORG/SIDA 22(1) faces a silvery appearance. Webster initially suggested its affinities were with section Andrichnia Baill, but later referred it to section Lamprocroton Mull. Arg. and then section Argyrocroton (Mull. Arg.) G.L. Webster (Webster 1967, 1996, Webster et al. 1996). Recent molecular phylogenetic studies have affirmed a sis- ter relationship between the two varieties and suggested new affinities with a small “basal” Croton group (Moacroton clade) containing Cuban endemics Moacroton Croizat and Cubacroton Alain and other Croton species (mostly sec- tion Corylocroton G. L. Webster) ranging from the Caribbean to Central and South America (Berry et al. 2005; van Ee & Berry 2005, Wurdack et al. 2005; van Ee et al. in press). Croton alabamensis var. alabamensis is known from central Alabama and a single probably erroneous report from central Tennessee (see below; Farmer & Thomas 1969; Wofford & Chester 2002). In Alabama the typical variety oc- curs in Tuscaloosa County along the Black Warrior River and in adjacent Bibb County from the Cahaba River drainage. It was first discovered in Bibb County by Eugene A. Smith in 1877. Ironically, Smith was apparently unaware of the populations along the Warrior River in his home county, Tuscaloosa, which were found by Roland M. Harper in 1905 (Harper 1906b). Croton alabamensis var. texenesis was an unexpected discovery in 1989 from Texas, 1000 km away from the typical variety (Ginzbarg 1991, 1992; Aplet et al. 1994). The Alabama plants from the Black Warrior and Cahaba river systems are separated by ca. 40 air km but show morphological, cytological, genetic, and ecological differences (Farmer 1962; Farmer & Thomas 1969; van Ee et al., in press). The plants are semi-evergreen and undergo meiosis and bloom in early spring (February and March) from overwintering flower buds formed the prior season. In Bibb County the Croton forms dense understory thickets from 1-2.5 m tall at 76-91 m of elevation in thin soils over limestone or dolomite above waterways within an 8 km radius of Pratt’s Ferry, an area roughly bounded by Schultz Creek on the west and Six Mile Creek to the east. The Cahaba River and its nearby tributaries cut steeply through the bedrock to form numerous dry rocky bluffs on which the Croton is especially abundant. Pratt’s Ferry has been reported to be the type locality of Croton alabamensis var. alabamensisand, as one of the most easily accessible localities for the plant, has been the source of numerous recollections. Pratt’s Ferry was a public ferry crossing of the Cahaba River just south of the community of River Bend and was probably named (although the Pratt family had many members in the area) for Mary Pratt (1800-1882), who ran the ferry from 1854 through at least the Civil War (Ellison 1984). Pratt’s Ferry and aptly named River Bend are ata large bend in the river where a ferry crossing could be established with relative ease. An iron bridge was built at the crossing in 1902 (Harper 1906a; the old roadbed is evident on the south side of the river and an iron pylon for this bridge re- mains on the north shore, pers. obs.) that was superseded by the present bridge WURDACK, LECTOTYPIFICATION OF CROTON ALABAMENSIS 471 crossing of County Road 26 just downstream from (west of) the original bridge. The exact nature of the ferry landings has been obscured. However, based on topography (ie., limitations imposed by adjacent bluffs), the ferry route appears to have diagonally spanned the river with the northern landing upstream and the southern landing downstream of the current bridge. In the vicinity of Pratt’s Ferry, the Croton presently occurs on the north side of the river ina small popu- lation immediately west of the current bridge and on the south side as a more or less continuous band east of the bridge for at least 1.6 km (pers. obs.). Despite a sustained interest in Croton alabamensis since its discovery in 1877 and concomitant visits to Bibb County by many botanists, the flora in the area has only recently been documented and found to be rich in endemic and disjunct taxa (Allison & Stevens 2001). Most of the endemics occur on ecologi- cally sensitive patchy glades (Ketona Glades) developed over Ketona Dolomite. Populations of the Croton in the Cahaba River drainage are often found around, but not restricted to, the fringes of the Ketona Glades (pers. obs.). Despite occa- sional logging activities, the region appears relatively unspoiled, remote, and undeveloped today and similar toa vintage 19" century riverbank photograph (Davenport 1988:40). A very different picture ges when looking at the his- torical record of regional industrialization. Taken in that light, Charles Mohr’s dire prediction over 100 years ago that the survival of the Croton in its native habitat “will in all probability be of short duration” may not have been un- founded (Mohr 1889). The limited distribution of both Croton varieties, as well as the Ketona Glade flora, has caused conservation concerns. Some habitat in Bibb County has been recently preserved (ie., The Nature Conservancy's Bibb County Glades Preserve). Discovery of Croton alabamensis Eugene Allen Smith (1841-1927), professor of geology and mineralogy at the University of Alabama, was appointed State Geologist in 1873 of the newly re- established Alabama Geological and Agricultural Survey. He began surveying the geology of the state during summer expeditions in a custom-built field wagon (modeled after a horse-drawn ambulance and preserved at the Univer- sity of Alabama) in a manner documented in photographs (see Wolfe 1983:99) and in his copious geological field notes (Smith 1870-1889). Although the origi- nal field notebooks were lost, transcriptions made during a 1930's Depression- era public works project (probably WPA or preceding agencies) remain and al- low a detailed picture of Smith’s field activities to be reconstructed here. His recurring interest in Bibb County was economically oriented toward an inves- tigation of coal and iron deposits that had originally made the region a leader in antebellum iron production (Ellison 1984). In the mid-19" century the Bibb County countryside was dotted with ironworks, mill dams, stone quarries, mines for ore (limonite) and coal, and associated communities. Today few traces 472 BRIT.ORG/SIDA 22(1) remain of this industrialization and the environmental degradation it caused is scarcely evident. Once thriving industrial communities are now extinct (ie., Tionis) or severely depopulated (i.e., River Bend and Six Mile). Smith first explored Bibb County in 1873 and in more detail in 1875. On August 17, 1875 he collected ore samples and explored associated Brighthope ironworks or “bloomery” (originally called Little Cahaba Furnace and in op- eration by 1850; Ellison 1984), the first blast furnace in the county and about 140 m above a wooden dam (Browne’s Dam) on the Little Cahaba River that provided its power. The seats for the dam timbers appear today as a series of steps cut into a riverbank rock outcrop at the Bibb County Glades Preserve near Bulldog Bend (pers. observ; Ellison 1993:51). The apparent present day natural state of this area, which contains some of the finest Ketona Glades, is remark- able considering the past destructive activities engendered by the adjacent iron- works, although it is also possible that disturbance supported or enlarged the glade community. The following day (ie., August 18,1875) Smith continued from the Sinks (sec. 26, T24N, RIOE) on lower Six Mile Creek, where the creek flows underground except at flood stage, southwestward to Pratt’s Ferry, where he noted fossiliferous “limestone” containing the gastropod Maclurites magnus LeSueur. It seems improbable that the Croton escaped visual detection, even if not considered noteworthy at the time. The trip credited with the discovery of Croton alabamensis was from July 18 to August 4, 1877. On July 18, Smith left Tuscaloosa and on July 24 his geo- logical party set up camp on the north side of the Cahaba River near Pratt’s Ferry. On the way, Smith described an area near Schultz Creek “where the Coal Measures are cut olf by a fault bringing up Knox Dolomite which ina series of glades or bare hills, with crumbling and much decomposed Dolomite stretches N.E. for a considerable distance” (entry for July 21; Smith 1870-1889). This is on the presently depauperate western end of the Ketona Glades zone and suggests that an extensive loss of glade habitat has since occurred. “Knox Dolomite” as used by Smith would have broadly included Ketona Dolomite which was named in 1910 (see Geological Survey of Alabama 1894) The label with Smith’s first collection of the Croton notes “Privet or Privy, Pratt’s Ferry. 26. 7. 77” (see Fig. 1). | interpret the date notation as July 26, 1877. Privet is probably derived from the vernacular for Eleagnus spp., which Croton alabamensis vegetatively closely resemble. Smith rarely recorded botanical observations in his geological field notes, and July 26 is no exception. On that date, Smith spent the morning west of Pratt’s Ferry on the north side of the river and then after lunch crossed over the river. On July 28, 1877, however, he makes a record of the Croton from a spot | estimate to be near the head of Little Schultz Creek, 3 km downstream from Pratt's Ferry. His field notes for that date read: _ {4 WURDACK, LECTOTYPIFICATION OF CROTON ALABAMENSIS 473 2( 774058 ) a RS SOHAL HERS! ; 1UM OF CHARLES MOHR Presented ta the Smithsonian Institution, 1901 terlom. Ulaba KemendS ty se 38 xX STAT. EF HABITAT, ee -_ PML Fer HAE li are | iis 2 es : ee A Ce tra i TU eee ee Fic. 1. Crot, Lak G Dy (Pay ME aN EY gt i ra | * acclacedt hin Ila-ti fk pie ay soci- Il label in bottom left ( | li ) read “Type S Spm: pecan’ [hand of C. Mohr} "Prive or Privy. Pratts ay re iE i velonals) to Ala. Hoel "hand of E A. Smith] f Alak cant cate I y. Discovered by or E. ‘ Smith, August 27" 1879. — forms the banks of the little Cahaba ri bl ilM y” [hand of C. Mohr]. 474 BRIT.ORG/SIDA 22(1) “Went with Mr, Mat Wallace in his skiff from opposite the point of the mountain [eastern end of Big Mountain] down nearly opposite the ore bank. The limestone bluffs furnished exactly the proper ground for the greatest variety of ferns. Amongst those I had not seen before were Adliantum|,. capil- lus-veneris, Cheilanthes alabamensis and one other which resembles it, but has lobed pinnules in- stead of entire ones. After climbing through an almost impenetrable thicket of what the natives call privet [Croton pasa and another shrub now in flower [probably Forestiera ligustrina Poir] and extremely fragran x thousands of bees we came to our ore bed” (Smith 1870-1889). Smith later recalled “the ie ocality of Croton alabamense, is Pratts Ferry in Bibb County, on [the] Cahaba River and upon limestone. Date of discovery, Au- gust 1877” (Smith 188ld; confirmed in Smith 1907). Smith probably encoun- tered the Crotonat other times during this 1877 trip, and assuredly so (based on present distribution) on August | while on the Cahaba River bluffs, 0.4 km north of where Cottingham Creek joins the river, while examining where “Knox Do- lomite” crosses. Smith appears not to have revisited the Pratt’s Ferry area until August 1889, based on field note chronology and referring to his “observations of 12 years ago” (ie., 1877). Mohr variously credits the discovery to 1874 (Mohr 1898, 1901), 1877 (Mohr 1889), or 1879 (Mohr 1887; see Mohr’s label notation in Fig. 1). Smith (881b) recalled, “I think | must be the discoverer as I know of no one else who has ever collected any of the material... It makes almost inpenetrable “Privet Thickets” as they are called, and the peculiar aroma of the leaves as you feel your way through, is exceedingly characteristic. | have not seen the plant anywhere else, than along the banks of the Cahaba River where it grows upon limestone much like Cedar.” From press to print In January 1878, Smith approached Charles Theodore Mohr (1824-1901) about preparing a mounted, organized herbarium for the Geological Survey from Smith’s collections. Mohr’s poor health and business obligations put the project on hold until May 1879, when he received the plants (Mohr 1879a). By July the Croton had come to Mohr’s attention and he directed an inquiry to Smith: “Amongst some other interesting plants collected at Pratts Ferry (vi. Forrestonia [sic], Forrestiera ligastrina [sic] et al.) | find a shrub marked “Privet.” Is it a truly indigenous plant or escaped from cultivation[?] It is very near Eleagnus Canadensis (the specimens being without flowers or fruit | cannot decide positively). It should be very a indeed to find that Northern shrub at once com- ing up afar in Central Ala{[bama].” (Mohr 187¢ The specimen (Fig. |) described by Mohr in this letter and annotated by him “Type Spm. [specimen] is apparently Smith’s earliest collection previously noted; it is in young bud. Mohr later recalled, “I found first a sprigg [sic] of it amongst the plants of the Geol. Survey of Ala. collected by the Professor [Smith] not- withstanding the poverty of the specimens bearing only a male spike with the flowers yet in the bud. I recognized as plant entirely new” (Mohr 1882a). After examination of a single male flower, Mohr tentative oe referred the Croton to section Hendecandra Eschw. and sent the specimens off to George WURDACK, LECTOTYPIFICATION OF CROTON ALABAMENSIS 475 Engelmann (1809-1884) with a cover letter dated February 24, 1880, noting that “perhaps the plant is known to you from Texas or N. Mexico” (Mohr 1880b). Mohr wrote to Smith, “I was somewhat astonished to find the Croton (Privet) by Dr. Engelmann referred to Cr. argyranthemum [Croton argyranthemus Michx,], a straggly herb not infrequent in our dry pine barrens here... It will be neces- sary to secure large specimens with a portion of the lower stem to establish its character as a true shrub and in full bloom with male and female flowers, the plants appear to be dioecious” (Mohr 1880c). Little progress had been made by October 11, 1880, when Mohr reported to Smith, “the Croton continues to be [a] riddle to all who get hold of it” and lamented the lack of adequate fertile mate- rial. The lack of fertile material explains his belief that the plant was dioecious (it isactually moneocious, with typical bisexual Croton inflorescences), though by this time he had the remnants of female flowers (Mohr 1880d). Mohr recalled, “Some time later I was so fortunate to find a unripe coped and a female flower, from which it was evident that it has its pl [sic] the section Eleuteria” (Mohr 1882a). There is no mention of the Croton in the Alabama floristic check- list Mohr and Smith published in 1880 (Mohr 1880a). Mohr turned to Alvan Wentworth Chapman (1809-1899), who was then involved in the preparation of a second edition of his Flora of the Southern United States (ie. sau 1883a). Mohr (1881la) wrote to Smith: ] ] for the * roton.Can you not geta slip for me to plantin my gar iene I have sent at his request all of the material I had received of yours (except that what I oad sent before to Prof. Engelmann who certainly took a wrong view of it) to Dr Chapman. He was anxious to describe the plant in the new edition of his Flora, he agreeing with me at first sight that [it] is see In the pursuit of more material, Chapman wrote directly to Smith, who replied April 8, 1881: “The croton you mention is a shrub he height of 8 to 10 feet. [have seen some ae three puenes in amiclee ie nae an aa ee thicket, and the aromatic odor i ility [probably Jacob S. Hansberger, see below] who has nronsed to get specimens du po oa of the year so that | may be able to get sufficient material for eee Dr. Mohr WG now all my material. When | get other specimens, I will remember to send you some” (Smith 1881la). Chapman requested all the material Mohr had “on hand,” and Mohr reported to Smith that the request had been honored including a specimen with “ripened fruit in perfect state” (possibly this collection is US-956957, see below) (Mohr 188la, 1881b). In a letter dated July 10, Smith (1881b) reported that he was ex- pecting new material in a “week or two,” presumably from his “correspondent.” He also approved Chapman's proposed name “Croton alabamense” (Lc.). On July 17, Smith was able to send Chapman “a small box containing specimens of Cro- ton alabamense, in different stages of its growth.” He continued, “From these, I hope you will be able to get all the information heretofore lacking. You might plant some of the seeds and perhaps raise it. | have several flourishing plants in 476 BRIT.ORG/SIDA 22(1) flower pots, which I have lately received from Pratt’s Ferry” (Smith 1881c). These additional specimens proved sufficient for the complete description. The “18 inch seedlings” received from the “correspondent” in July 1881, Smith planted in his yard and they attained a height of “7 feet” by 1889 when Mohr reported on their development (Mohr | Chapman traveled north, ne with Charles Mohr while passing though Washington (Mohr 1882b), and delivered the manuscript for his Flora to the printer in the last week of July 1882 (Chapman in Oakes 1932). Proof correc- tions continuing into the fall and the preface was dated December 26, 1882 (Chapman 1883a). Copies of the Flora were circulating by February 1883 (Chap- man 1883b). The main body of the Flora was merely a corrected version of the first edition (Chapman 1860) and new taxa, including the Croton, were con- tained in an appended supplement. Chapman (Lc.) credited Smith as the au- thority for the species but added “ined.,” an amendment finally removed in the entirely revised third addition of the Flora (Chapman 1897). Mohr was preoccupied with work for Charles Sprague Sargent (1841-1927) on forest trees for the Tenth Census of the United States and could not visit Bibb County until November 1882 (vouchers at A,GH, MO and US variously give the date as November 11 or 12). Mohr located the Croton while collecting wood samples of Quercus durandii Buckley on his final trip for Sargent. Of the trip Mohr notes: “About the middle of last month I was over on the little Cahaba river near Tionus [sic]. 1 went by way of Montevallo; having to return the same day I made 32 miles on horseback, arriving at my quarters Mr. Aldriches at 10 o'clock pm... At the time of my visit to the ‘Privet’ thickets near Tionus the plant was just putting forth the flower buds of the staminate spiklellets, which as Mr. Hansberger informs me open with the first warm days of le spring. Nota ve a of evenarudimentary pistillate [lower could be found and the fruit of this season was entirely gone... Mr. Ha tl received me with the genuine warm hospitality of the es n gentlemen, went oe me through the woods and gave me all assistance in his power.. 1am bound to visit again this locality so aes on [sic] botanical trea- sures” (Mohr 1882c). Tionis was a community located along lower Six Mile Creek (sec. 26, T24N, RLOE; fide Smith 1870-1889) and readily evident on period maps (see online histori- cal map archive at the University of Alabama, http://alabamamaps.ua.edu/ index.html). A post office was established in 1870 and Jacob S. Hansberger (d. 1887) served as postmaster from 1874 until 1887 when it was discontinued prior to his death (Stewart 1983). Bibb County deed books show that he owned over 200 ha of land in the vicinity (accrued as many parcels over a period of years). Smith knew Hansberger from his Bibb County trips and they had explored Six Mile Creek together shortly after Smith first discovered the Croton (fide entry for August 1, 1877; Smith 1870-1889). It is probable that Smith instilled in Hansberger an interest in the Croton and that he served as Smith’s “correspon- dent” and local expert. The exact locations of Tionis and Hansberger’s house are presently uncertain and | have not been able to assess the patchy Croton WURDACK, LECTOTYPIFICATION OF CROTON ALABAMENSIS 477 distribution in relation to them. Even today it is clear that Croton is readily avail- able in the general area without having to travel all the way to Pratt’s Ferry (ca. 6.4 km to the west). In the fall of 1882 Frederick Hoffmann, editor of Pharmaceutische Rundschau, requested Mohr to submit an article on Croton alabamensis for that journal (Mohr 1882b). Mohr had noted that the Croton bark was of interest to “pharmacognosy” as a possible native substitute for cascarilla that came from the bark of the West Indian Croton eluteria (L.) Sw. Mohr decided to defer writ- ing the article until he could obtain flowering material to clear up the “point of the sexual relation of their flowers” and have a photograph of the plant taken. In March 1883 Smith sent Asa Gray a print of the photograph (annotation on back of print says the photo was taken February 28, 1883; Smith 1883) and Mohr received a copy in May (Mohr 1883a). In June 1883, while collecting wood for the Louisville Cotton Exposition, Mohr returned to Pratt’s Ferry, ostensibly for Quercus durandii wood, but also unsuccessfully for Croton seeds and flower- ing specimens. (Mohr 1883p). Mohr’s labors finally bore fruit. In a paper read in January 1884 before the Mississippi Valley Horticultural Society, he presented his views on the possible use of Rhus cotinoides (= Cotinus obovatus Raf.), Neviusia alabamensis A. Gray, and Croton alabamensis as horticultural subjects (Mohr 1884). The same three species were treated in an expanded form by Mohr for the January 1887 issue of Pharmaceutische Rundschau (Mohr 1887). An etching of a cultivated seedling was substituted for the originally intended photograph. Preparation for this article included the examination of additional blooming material that was col- lected by Smith in his garden and received by Mohr in April 1886 (Mohr 1886). This material probably included a collection in the Mohr Herbarium dated April 16,1886 from “my yard (Tuscaloosa), transplanted from Pratt’s Ferry” (E.A. Smith s.n., UNA-00020895, annotated by Mohr “Type Spm.”). In 1888, Mohr’s prepara- tion of an article for Garden and Forest brought renewed questions to Smith on the life history of the Croton and requests for specimens (Mohr 1888). Mohr hoped to have the article sent off to press by the end of August 1888 and it ap- peared in the December 11, 1889 issue of Garden and Forest. In March 1889 Smith sent seedlings of the Croton to Sargent and Mohr (Smith 1889). Typification and historical collections Chapman gave no collection data beyond “central Alabama’ in his original de- scription, therefore a lectotype is selected here: Croton alabamensis E.A. Sm. ex Chapm., Flora of the Southern United States, ed. 2:648. 1883. Lectotype: UNITED STATES. ALABAMA: E.A. Smith s.n.(US-935923, ex John Donnell Smith herbarium). Other original material: Alabama, 4 Apr 1881 (date given on supplemental tag attached to stem), E.A. Smith s.n. (MO-1904803); Alabama, E.A. Smith s.n. (US-956957, ex Biltmore Herbarium). Alabama, Tuscaloosa, E.A. Smith s.n. (F-99491, ex H.N. Patterson herbarium; MO-784596, ex Chapman estate from 1899 purchase by MO). 478 BRIT.ORG/SIDA 22(1) Herbaria (GH, MO, NY, UNA, and US) containing the best representation of collections from Chapman, Mohr, and Smith were searched for historical Cro- ton collections. Five Croton specimens were located that bear evidence of hav- ing been seen by Chapman (ie., bear writing in Chapman’s hand), although additional specimens may turn up in his widely scattered exciccatae (see Stafleu and Mennega [1997] for a listing of the 27 herbaria with Chapman material). It is unclear whether the two sheets labeled “Tuscaloosa” by Chapman should be considered original material. If taken asa literal exact locality then the collec- tions came from cultivated material and most likely post-date the preparation of the original description (i.e., post 1881). Smith did cultivate plants in Tusca- loosa that were eventually a source of specimens, but these were probably the small, valuable seedlings also received in the summer of 1881, and not likely to become herbarium fodder at that time. On the other hand, label locality data secondarily penned by Chapman and not the collector might be distorted or incomplete. Chapman did generalize that Pratt’s Ferry was “near Tuscaloosa” (see below) or he may have referred to Smith being based in Tuscaloosa. John Donnell Smith (1829-1928) corresponded with and received speci- mens from Chapman during at least 1884-1886, based on specimen provenance notations in Donnell Smith’s hand and fragments of Chapman letters attached to herbarium specimens at US. Donnell Smith’s personal herbarium (including the lectotype, see Figure 2 with embossed stamp) and library were donated to US in 1905. The lectotype has an attached undated fragment of a Chapman let- ter (consistent in the distinctive paper with other letter fragments established as dating from 1884-1886), presumably to Donnell Smith, that reads: “Croton Alabamensis is not found along R{ailJRoads and being a shrub will remain pretty quiet in its secluded home near Tuskaloosa [sic]. Strange that it should be in flower every month from Sept to May through the coldest weather and yet they sent me specimens collected in all these months!” Based on the detailed historical account given above, it appears that Chapman had material from two collectors (E.A. Smith and J. Hansberger), multiple dates and perhaps several Bibb County localities on hand for his original diagnosis. The circumstantial evidence (ie., Smith did not visit the area between the time of the Croton discovery and publication, and he also refers to receiving speci- mens from a “correspondent”) suggests that much of the material was collected by Jacob Hansberger near Tionis, and the lectotype may have been collected by him in April 1881 and sent to Smith, who forwarded the specimens to Chap- man for description. The two branches on the lectotype sheet have young fruits at slightly different developmental stages and may come from separate late spring collections. Of the five sheets of original material, only the selected lec- totype contains an additional printed label that indicates the specimen was associated with the “Flora of the Southern United States and Supplement’ (ie, Chapman 1883a). In addition to sending duplicates to correspondents, Chapman sold sev- WURDACK, LECTOTYPIFICATION OF CROTON ALABAMENSIS U.S. National Herbequm. det 00730774 pea Mee, TOTYPE: Croton alabamensis E. A. Smith ex Chapman Fl. S. U.S. ed. 2: 648 Ky Wolk 17-2.0904 TT ne , 1883 Cee rea Pig ins, EX HERS. / United Stat " and Supplement, A. W. Chapman, M.D.” : sae how ar Sane -- Brett: Af ‘ nee ~ tte bs af V2 yore Pad yb ry rieee JA Loulin Mabivrrrecrdit, F Kidugy Bias ate VE Dop Mir Slat ri aeate alqhamensic(F A S, Fic. 2.Crot I “ Chapman. See text for transcription of letter. 480 BRIT.ORG/SIDA 22(1) eral sets of specimens that are massive enough to have been described as “her- baria.” However, he always retained a personal working collection, the residue of which was purchased from his estate by MO in 1899. In 1896 Chapman se- lected and sold the best set of material on which his 1883 Flora was based, to the Biltmore Herbarium in Asheville, North Carolina. The Biltmore Herbarium, modeled after the Royal Botanic Gardens, Kew, was established ca. 1894 for George Vanderbilt (1862-1914) asa scientific adjunct to his lavish Biltmore Es- tate. At the time of the Chapman purchase, the herbarium was trying to estab- lish itself as one of the foremost botanical institutions in the southern United States through purchasing and exchanging collections, hiring of collectors, and publishing the eponymous scientific journal Biltmore Botanical Studies. The enterprise withered after about 10 years of activity (peaking ca. 1901-1903) due to Vanderbilt’s financial downturn and untimely death. A 1916 flood destroyed three-quarters of the collection and the residue of ca. 25,000 sheets was given to US in 1917. The Chapman collection was reportedly stored in a “vault” and largely salvaged (Maxon 1917; Boynton 1936). Only one flood-damaged fruit- ing specimen (US-956957, stamped ex Biltmore Herbarium) of Croton alabamensis clearly has this provenance and was used at the Biltmore in an illustration (see Lounsberry 1901:306). It is possible that other C. alabamensis specimens, including other type material at Biltmore, were destroyed. An anomalous Tennessee record (Tullahoma, Coffee Co., 10 Aug 1899, TG. Harbison 725, NCU-9391) has connections with Biltmore. Thomas Grant Harbison (1862-1936) was a member of the Biltmore Herbarium staff and made extensive collections of the southern flora. His incomplete Biltmore field notes deposited at US (partial number series of duplicate field notes bracketing, but not includ- ing, the Croton collection in question) place him in Tullahoma, but suggest errors on the specimen label which was made long afterwards (ie., Harbison 725 would have been collected on 12 Aug 1899 and not 10 Aug as reported with the speci- men). A Croton alabamensis collection with the same collection number and from a well-established locality “On sandstone bluffs along the Warrior River, Tusca- loosa, Alabama,” T. G. Harbison 725, 11 Oct 1911, A) may have been a source of confusion. The credibility of the Tennessee record is doubtful. ACKNOWLEDGMENTS I thank the curators and staff of the noted herbaria and archives for access to their collections and permission to quote archival materials. | am grateful to LJ. Davenport (Samford University) for the Smith to Chapman typescripts and to B. van Ee for unpublished information on his recent genetic studies. J. Alli- son, LJ. Davenport, LJ. Dorr (US), and J. Reveal (emeritus MARY) provided help- ful comments on the manuscript. At the Geological Survey of Alabama, A. Sartwell provided information on Smith and alerted me to the archives formerly in his care and LS. Dean clarified Smith’s geological terminology. WURDACK, LECTOTYPIFICATION OF CROTON ALABAMENSIS 481 REFERENCES A.uison, J.R.and T.E. Stevens. 2001.Vascular flora of Ketona Dolomite outcrops in Bibb County, Alabama. Castanea 66:154-205 Aptet, G.H.,R.D. Laven, M.B. FALKNer, and R.B. SHaw. 1994. Population and site characteristics of a recently discovered disjunct population of Croton alabamensis (Euphorbiaceee). Sida 16:37-55. Berry, PE., A.L. Hipp, K.J. Wuroack, B. Van Ee, and R. Rina. 2005. Molecular phylogenetics of the giant genus Croton and tribe Crotoneae (Euphorbiaceae sensu stricto) using ITS and trnL-trnF DNA sequence data. Amer. J. Bot. 92:1520-1534. Boynton, F.E. 1936. Letter to C. D. Beadle, 28 October, from Old Fort. 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Trans. Mississippi Valley Hort. Soc. 2:216-29, Mone, C.T. 1886. Letter to E.A. Smith, 22 April, from Mobile. W.S. Hoole Special Collections Library, University of Alabama. Mone, C.T. 1887.Ueber drei vereinzelte Burger des Floren-Gebietes der nordamerikanischen Sudstaaten, Pharm. Rundschau (Berlin and New York) 5:8—11. Mone, C.T. 1888. Letter to E.A. Smith, 25 August, from Mobile. W.S. Hoole Special Collec- tions Library, University of Alabama. Mone, C.T. 1889. The la[te]st addition to the shrubs of eastern North America. Gard. & Forest 2:592, 594. Mour, C.T. 1898. Letter to A.M. Ferguson, 19 April, from Mobile. Missouri Botanical Garden Herbarium (attached to specimen of Croton alabamensis). WURDACK, LECTOTYPIFICATION OF CROTON ALABAMENSIS 483 Mour, C.T. 1901. Plant life of Alabama: an account of the distribution, modes of associa- tion,and adaptations of the flora of Alabama, together with a systematic catalogue of the plants growing in the state. Contr. U.S. Natl. Herb, 6:1-921. Smith, E.A. 1870-1889. [WPA? typescripts of 1936]. Field notes of E.A.Smith.v. 1:1870-1875, Vv. 2:1876-1882, v. 3:1883-1889. Library, Geological Survey of Alabama. Smith, E.A. 1881a.Letter to A.W. Chapman, 8 April, University of Alabama. [WPA? typescript] Library, Geological Survey of Alabama. SmitH, E.A. 1881b. Letter to A.W. Chapman, 10 July. [WPA? typescript] Library, Geological Survey of Alabama. Smith, E.A. 1881¢. Letter to AW. Chapman, 17 July. [WPA? typescript] Library, Geological Survey of Alabama. Smitu, E.A. 1881d.Letter to A.W. Chapman, 7 September, from Prattville. [WPA? typescript] Library, Geological Survey of Alabama. Smito, E.A. 1883. Letter and photograph to A. Gray, 11 March, from Tuscaloosa. Gray Her- barium (filed with Croton alabamensis), Harvard University. Smith, E.A. 1889. Letter to C.S. Sargent, 26 March, from University, Alabama.Gray Herbarium (filed with Croton alabamensis), Harvard University. Smith, E.A. 1907. Letter to R.M. Harper, 8 September, from University, Alabama. Harper Pa- pers. W.S. Hoole Special Collections Library, University of Alabama. Starteu, FA. and E.A. MenNeca. 1997. Taxonomic literature. Supplement IV: Ce-Cz. Koeltz Scientific Books, Germany. Stewart, F.R. 1983. Alabama's Bibb County. Unpublished Doctor of Education dissertation, University of Alabama. vAN Ee, B. and PE. Berry. 2005. The Caribbean-centered Moacroton clade within Croton (Euphorbiaceae sensu stricto).Abstract. Available at:http://www.2005.botanyconference.org VAN Ee, B., N. Jevinski, PE. Berry, and A.L.Hiee.In press. Phylogeny and biogeography of Croton abamensis (Euphorbiaceae), a rare shrub from Texas and Alabama, using DNA se- quence and AFLP data. Molec. Ecol Weester, G.L. 1967.The genera of Euphorbiaceae in the southeastern United States. J. Ar- nold Arbor. 48:303-430. Weester, G.L.M.J. pet Arco Acuitar, and B.A. SmitH.1996. Systematic distribution of foliar tri- chome types in Croton (Euphorbiaceae). Bot. J. Linn. Soc. 121:41-57 Weesster, G.L. 1993. A provisional synopsis of the sections of the genus Croton (Euphorbiaceae). Taxon 42:793-823. Worrorp, B.E. and E.W. CHester. 2002. Guide to the trees, shrubs, and woody vines of Ten- nessee. The University of Tennessee Press. Wotre, S.R. 1983. The University of Alabama. A Pictorial History. The University of Alabama Press. Wuroack, K.J., P. HorFMANN, and M.W. ee 2005. Molecular phylogenetic analysis of uniovulate Euphorbiaceae (Euph ceae sensu stricto) using plastid rbcLl and trnL-F DNA sequences. Amer. J. Bot. 92:1397-1420. =e 484 BRIT.ORG/SIDA 22(1) Book REVIEW Mark Turner and Puytis Gustarson. 2006. Wildflowers of the Pacific Northwest. (ISBN 0-88192-745-7; 978-0-88192-745-0, flexibind). Timber Press Inc, 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.tim berpress.com, mail@timberpress.com, 503-227-2878, |-800-327- 5680, 503-227-3070 fax.). $27.95, 512 pp., 1248 color photos, 1221 maps, 31 line drawings, 5 3/4" x 81/4". More than 1200 native and non-native ee are described and illustrated, including 1247 color photos and 1219 ran ( distribution). The region covered is all of Washing- ton and Oregon, a: Seurhiwectern Br itish Columbia and the northern third of California. An ecoregions map and excellent ey intro een nae “C sas Geography, and Plant Habitats,” set the stage. The photos lowers as well as critical vegetative fea- tures, and the morphological and meee notes are ae and to the point. Six categories of flower color (white, yellow, orange, pink to red or red-purple, violet to oe or blue- purple, brown and green) provide the first division in the arrangement of species, followed wit} ‘h color category by ma- jor structural features (e.g, number of petals, petals forming a a or not, ovary inferior or superio composite flowers). My perennial response to “color-coded” taxonomy is that so many people ene this book will wish the blue- ene Ww Bite: flowered (etc.) species of a single genus were together, but apart from this personal bias, this ! “third-generation” field guide, about as good as they get, useful to everyone interested in the flora of this region.—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A. SIDA 22(1): 484. 2006 TWO NEW SPECIES OF ELYMUS (POACEAE) IN THE SOUTHERN U.S.A. AND OTHER NOTES ON NORTH AMERICAN ELYMUS SPECIES Julian J.N. Campbell The Nature Conservanc cy ( (} entucky Chapter) 42 leas ae Street Lexington, b 40508-2018, U.S.A. ieamipbel@tne org ABSTRACT wo new species of Elymus L. are described, E. churchii JJ.N. Campb. and E. texensis J.J.N. Campb. Elymus churchii grows in the Ouachita and Ozark Mountains of Arkansas, Missouri and Oklahoma. It is similar to E. svensonii G.L. Church, but differs in its lemmas, which are usually pubescent, and onger awns; its spikelets, which have fewer florets; its ienicno des, which are longer, wit teed dorsal ridges and green longitudinal bands; and its foliage, which is typically less pubescent and less glaucous. Elymus texensis grows on the Edwards Aue in southwest Texas. It is similar to E. pringlei Scribn. & Merr, but differs in its larger anthers; its glabrous lemmas; its larger spikelets; its longer spikes, with ee internodes that are esas with oe longitudinal bands, and gener- ally glabrous; and its foliage, which is ly pilo e. The new succes are sais from yi ] £ few collections, and need much more cee to assess their bution. They are published here to allow inclusion in the Flora of North America, pone 24, and to stimulate further attention to them. Some additional notes and corrections are appended regarding the author's previous publications on Elymus. RESUMEN Se describen dos nuevas especies de Elymus L., E. churchii J.J.N. Campb. y E. texensis ne Campb. Elymus churchii vive en las montafias Ouachita y Ozark de Arkansas, Missouri y Oklahoma. Es simi- lt a E. svensonii — L. Church, p ncia por sus es que son penal pubes ntes, y ill u is, que son mas poe con eostillac derales Rigpidas y bandas verdes longitudinales; y su - llaje, que es tipicamente menos pubescente y menos glauco. Elymus ene vive en el Edw a Plateau en el sureste de Texas. Es saaues a E. ae oO ] ] . ll a a aq n as fe con Bande verdes longitudinale y pene erie glabras; y su follaje, ae es a veces ee piloso. Las nuevas especies se conocen de pocas colecciones, y necesitan mucho mas estudio para evaluar su estatus gent y distribucion geografica. Se publican aqui para que nee incluirse en mila Flora de Norte América, volumen 24, y para estimular la atencion sobre ellas. Se anaden les y correcciones a Sree previas del autor sobre Elymus INTRODUCTION In 2002, I provided descriptions, specimen data, and distribution maps for two taxa of Elymus that seemed different from other members of the genus with paired spikelets and reduced or vestigial glumes (Campbell 2002a). I refrained SIDA 22(1): 485 — 494. 2006 486 BRIT.ORG/SIDA 22(1) then from formal taxonomic recognition, but after further revison | am now more confident. To allow inclusion in the Flora of North America, volume 24 (M.E. Barkworth et al. in prep.), | present below the two new names, anda key to the species of Elymusthat have paired spikelets and reduced or vestigial glumes. There has been much uncertainty in the circumscription and phylogeny of these species, which would have been grouped together under the broad con- cept of Elymus interruptus Buckley by Hitchcock and Chase (1951). Elymus churchii J.).N. Camp., sp. nov. (Fig. 1). Type: US.A. ARKANSAS: Conway Co: Petit Jean State Park, rocky bluffs, PO. Morrilton, 1500 ft, 3 Jul 1957, D. Demaree 37234 (HOLOTYPE: UARK:; IsoTyPES: OKL, SMU-BRIT). Affine Elymo svensonii G.L. Church sed differt lemmibus plerumaque pubescentibus aristis (10-)20- 30(-35) mm longis, spiculis flosculis 3(-5), internodis spicarum (5-)7-13(-18) mm longis viridi-vittatis dorsaliter hispidis, foliis paginis adaxialibus glabris vel pilosis, et plantis non glaucissimis. Plants cespitose, of ten somewhat, but not strongly, glaucous. Culms 50-120 cm, erect; nodes usually 4-8, exposed or covered, often reddish-brown or blackish, glabrous. Leaves evenly distributed; sheaths usually glabrous, or sometimes pubescent at the summit; auricles 1-2 mm, often reddish brown or blackish; ligules up to 1 mm, often reddish-brown; blades 3-11 mm wide, lax, adaxial sur- faces glabrous or short-pilose. Spikes 10-18 cm long, 3-5 cm wide, slightly nod- ding, with 2 spikelets per node; internodes (5-)7-13(-18) mm, about 0.2 mm thick at the thinnest sections, flexuous, with green longitudinal bands along the con- cave sides, glabrous except for the two hispid dorsal ridges; disarticulation be- low each floret, but not below the glumes. Spikelets usually appressed, 10-15 mm (excluding awns), with 3(-5) florets. Glumes often unequal (differing in length by more than 5 mm), or sometimes vestigial to absent from the upper spikelets or throughout, 0-15(-20) mm long including the undifferentiated awns, indurate at base, 0.1-0.3 mm wide, setaceous to subulate, with 0-1 dis- tinct vein, glabrous, awns often outcurving; lemmas 8-10 mm, pubescent, or occasionally glabrous, awns (10-)20-30(-35) mm, slightly to (at maturity) strongly outcurving; paleas 7-9 mm, obtuse to truncate, or emarginate; anthers about 2.5-3 mm, evident in June. 2n = unknown. Elymus churchii grows in dry rocky soils, often base-rich, in open woods on ridges, bluffs and river banks of the central Ouachita Mountains and the western Ozark Mountains, in Arkansas, Oklahoma and Missouri. It was previ- ously included within a broad concept of E. interruptus (Steyermark 1963; Smith 1991). It is similar to E. svensonii, which is disjunct in Kentucky and Tennessee. It might reasonably be combined as a subspecies or variety of F. svensonii, but differs in its lemmas, which are usually pubescent, and have longer awns; its spikelets, which have fewer florets; its rachis internodes, which are longer, with hispid dorsal ridges and green longitudinal bands; and its foliage, which is typi- cally less pubescent and less glaucous. It may have originated, like E. svensonii, CAMPBELL, tii | MG ck. Sah calek he D ea ZN \ eee ~~, os eo ~~ aw, ith v7 % a ———_ =. Fao even Stl Op haa a oo Fic. 1 Ely hurchii (d f Demaree 29337 unless noted). A. Habit. B. Upper | i f cul itt pil viewed on plane with alternating spread of spikelets (B2 from Palmer 57957). C. Sheath summit and blade bases. D. hdaviallesfeurtace- showing Weer ey . ‘al (showing variation in size), viewed in pl pikelet spread (with abaxial view of Igl pikelet, and largely side view of lateral glume). F. Spikelet, with lateral view of florets (F2 from Palmer 57957). G. Mature floret in abaxial view (| ft) and adaxial vi (right). H. Cross-secti f mature, indurate gl I (showi internode. J ize). I. Cross-section of central rachis 488 BRIT.ORG/SIDA 22(1) from introgression of E. canadensis L. and E. hystrix L., and there are occasional specimens suggesting transitions to both species (Campbell 2002a). This spe- cies is named after George L. Church, who conducted several useful studies of Elymus in eastern North America, focusing on problems of hybridization and the species with reduced glumes (Church 1967). ParatyPes: ARKANSAS. Baxter Co.: rocky shaded blulf, White Rv. PO. Lakeview, Bull Shoals Dam Reservoir, 600 ft, 1 Jul 1950, D. Demaree 29337 (OKL, TEX). Logan Co.: Magazine Mt, about 45 mi east of Fort Smith, 8 Aug 1942, D.M. Moore 420118 (TEX), Sep 1947, D.M. Moore 470639/642 (UARK, NCU, US), and 30/31 Jul 1949, D.M. Moore 490422/441 (UARK). Newton Co.: Big Bluff above Buffalo Rv, 5 mi below pes: 5 Jun 1953, D.M. Moore 53259 (UARK). OKLAHOMA. Le Flore Co.: Rich Mt., roadside. oak-] nant, 16 Jun 1940, H. Taylor 110 (OKL); see also See ener ihe salves collection known from Missouri has relatively short rachis intenodes (ca. 5-7 mm): Christian Co., 3 miles west of Nixa, rocky wooded banks, 24 Jun 1954, E.J. Palmer 57957 (SMU). I previously suspected that all of Steyermark’s (1963) records of Elymus diversiglumis Scribn. & CR. Ball or E. ee aa in Missouri could be grouped with the plants described here as E. churchii(Camp- bell 2002a). However, some of these records were based on misidentifications: EJ. Palmer 66416 (UMO) is E. glabriflorus var. australis (Scribn.&@ CR. Ball) JJ.N. Campb.; EJ. Palmer 55939 (UMO) is E. hystrix var. bigelovianus (Fernald) Bowden. Elymus texensis J.J.N. Camp, sp. nov. (Figs. 2a, b). Type: US.A. TEXAS. Gillespie Co: Ser- pentine Mounds, about 9 mi N of Willow City, hilly area vegetated mainly with grasses, 18 May 1906, E.S. Nixon 531 (HOLOTYPE: TEX 5322) Affine Elymo pringlei Scribn. & Merr. sed differt antheris 4.5-6 mm longis, lemmil s glabris, spiculis 25-40 mm longis flosculis 5-8, 3 icis 9-20 cm longis phauaue (5- ce 15(- ens mim longis an vittatis glabris praeter margi _foliis pag hirsutis vel dense brevipilosis. Plants cespitose, glaucous. Culms 70-110 cm, erect; nedes usually 4-6, mostly exposed, glabrous. Leaves evenly distributed; sheaths glabrous; auricles about 0-1 mm, sometimes adherent to sheath summit, pale to purplish brown; ligules about | mm, erose; blades 2-9 mm wide, lax, or somewhat involute, adaxial sur- faces thinly scabrous-hirsute or densely short-pilose. Spikes 9-20 cm long, 2- 25 cm wide, erect to slightly nodding, with 2 spikelets per node; internodes (5-)7-15(-22) mm, about 0.1-0.3 mm thick at the thinnest sections, with slight dorsal ridges, with green longitudinal bands along the concave sides, glabrous except for the ciliolate margins; disarticulation below each floret, but not be- low the glumes. Spikelets appressed, 25-40 mm (excluding awns), with 5-8 flo- rets (including terminal rudiment). Glumes subequal, 14-24 mm long includ- ing the undifferentiated awns, 0.1-0.3 mm wide, setaceous, with O-1 distinct vein, glabrous, awns more or less straight; lemmas 8-12 mm, glabrous, awns 8- 25mm, straight, flexuous or slightly curving; paleas about 7-11 mm, obtuse or narrowly truncate; anthers 4.5-6 mm, evident in May. 2n = unknown. CAMPBELL A 489 Elymus texensis is known only from three collections that were made on calcareous bluffs and hills, in juniper woods and grassy areas on the Edwards Plateau of southwest Texas (Campbell 2002). It is similar to E. pringlei, but dif- fers in its larger anthers: its glabrous lemmas; its lar ger spikelets; its longer spikes, with rachis internodes that are longer, with green longitudinal bands, and gen- erally glabrous; and its foliage, which is sometimes densely pilose. It is notable that the northernmost known collection of E. pringlei appears somewhat atypi- cal and may be transitional to E. interruptus or E. texensis: MEXICO, Coahuila, 54 mi SE of Big Bend National Park, south end of Sierra Maderas del Carmen, Canyon de la Fronteriza, ryolite area, J. Henrickson 15045 (TEX). That collec- tion has relatively robust glumes, long rachis internodes, spikelets with up to 6 florets (including the terminal rudiment), and lemmas that are virtually gla- brous (Campbell 2002a). This new species is named after the great state of Texas. ParATYPES: TEXAS. Burnet Co.: Inks Lake State Park, limestone bluffs, juniper woods along creek just east of HQ, 19 May 1983, R. & G. Kral 70066 (VDB, SMU-BRIT). Uvalde Co.: chalk bluff on Nueces River, 12 May 1938, V.L. Cory 29073 (US 3039432). KEY TO-FLYMUS FYS ER EAGCAND ITS: ALLIES These two new species appear to belong ina natural group with Elymus hystrix and other allies. The following key distinguishes the members of this group. Brief notes on ranges are inserted, but Campbell (2002a) should be consulted for maps and further details. This group of species is characterized by their reduced or vestigial glumes, with O-1(-2) veins, tapering from the base, 0-24 mm long including awns, of - ten differing in length by at least 4 mm, 0.1-0.5(-0.7) mm wide, persistent after florets disarticulate; and by their rachis internodes usually 0.1-0.3 mm thick at the thinnest sections, often with green longitudinal bands along the concave sides. This group should be contrasted with the more heterogeneous group that consists of E. virginicus L., E. canadensis L., E. glaucus Buckley and their allies. The latter species have glumes with 2-5(-8) veins, widening or linear above the base, 4-45 mm long including awns, subequal, 0.2-2.3 mm wide, persistent or disarticulating: and rachis internodes usually 0.2-0.8 mm thick at the thinnest sections, usually lacking green bands. Elymus interruptus remains a somewhat problematic taxon, with specimens that can fit within either of these outlines. Moreover, various hybrids of E. hystrix with other species will fit within either outline, and will require deeper analysis. It is likely that some allies of E. hystrix in this group originated from Peeauar (Church 1967; ee 2002a). 1. Spikelets widely divergent to perpendicular at traight (rarely slightly curving); glumes vestigial or 1-3 mm long, ea some ee glumes up to 10(-20) mm long and 0.1-0.2 mm wide but with no distinct vein; spikes more or less erect [widespread in eastern North America, but unknown in Texas] E. hystrix 490 BRIT.ORG/SIDA 22(1 Sie Se Soa a L — Fic. 2a Fly is(2ad fi E.S. Nixon 537). A. Habit. B Upy I ti f cul it pil , Viewed on 1 ey I j £ 1 L « ILI J L ry) a Tl rf £. 2 J hairs EF AAA ee J | 1 _ ans 1 £ *1 | rs “aL L oe * £ = ~ i F Fr t J spikelet, and | ly si ). F. Spikelet. with lateral vi f fl G. Mature fl (left) and adaxial view (right). H. Cross-sections of mature, indurate gl base(s). I. Cross-section of central rachis internode. a= CAMPBELL, A 491 Fic. 2b. Ely is(2bd fi VL. Cory 29073). A. Habit. B. Upper porti f cul it t pil viewed on f g I | D Al £ _ g hairs E.M hisi | 1gl ; i di pl f pil | I i g spikelet, and | gely sid Igl ). F. Spikelet, with | lvi f fl G. Mature floret in abaxial vi (left) and adaxial view (right). H. Cross-sections of mature, indurate gl base(s). I. Cross-section of central rachis internode. 492 BRIT.ORG/SIDA 22(1) 1. Spikelets usually appressed, never perpendicular; lemma awns straight or curving; glumes sometimes vestigial, but usually 1-24 mm long, 0.1-0.5(-0.7) mm wide, often with a distinct vein; spikes erect, nodding or penden 2. ‘ ic well-developed, at least 12 mm long, Sieetel lemma awns straight moderately curving; spikes erect to slightly nodding. aes (6-)9-15(—22) mm long (excluding awns), each with 2-5 florets; lemma awns moderately outcurving at maturity; glumes 0.2-0.5(-0.7) mm wide [in the southern Rocky Mountains and adjacent regions, from southern California to ea Texas to northern Coahuila E. interruptus 3. Spikelets 18-40 mm long (excluding awns), each with 3-8 florets; lemma awns straight to a curving at maturity; glumes 0.1-0.3(-0.6) mm 4, Anthers 2.5-4 mm long;lemmas eae hispid to thinly strigose soils cent; spikelets 18-25 mm long, with 3-6 florets; spikes 4-12 cm long, ra- chis internodes 3-6 mm lo eter ee longitudinal bands, hispid on dorsal ridges; blades thinly scabridulous, hispidulous or pilose on veins [along the Sierra Madre Orientale, in eastern Mexico from Coahuila to Veracruz] E. pringlei 4. Anthers 4.5-6 mm long;lemmas glabrous; spikelets 25-40 mm long, with sieaa alee ae cm lenig co internodes (5-)7-15(-22) mm long, pt for ciliolate margins; blades thinly scabrous -hirsutet to dense ely pilose [on the Edwards Plateau, in south Tex E. texensis 2. Some glumes less than 12 ae pair usually differing in length by at least 4 mm when developed, or one or both vestigial: lemma awns [eon at ma- a often oe SO; ve aaa nodding to penden 5. Rachis inter —6(-9) mm long;glumes (0.1-)0.2—-0.5(—0.6) mm wide; lem mas eee to strigose, at least near margins; sheaths aeane plants not glaucous to moderately glaucous [mostly in the northern Great Plains, from Saskatchewan to Ontario to lowa] E. diversiglumis . Rachis internodes (4—)6-13(-18) mm long; glumes 0.1—-0.3 mm wide; lemmas glabrous or pubescent; sheaths glabrous or villous; plants usually glaucous, sometimes strongly so 6b usually pubescent, the awns (10-)20-30(-35) mm long; spikelets with 3(5) florets; rachis internodes (5-)7—13(-—18) mm long, with green lon- gitudinal bands and hispid dorsal ridges; blades glabrous or short-pilose; plants not strongly glaucous [central Ouachita and western Ozark moun- ns in Arkansas, Missouri and Oklahoma] E. churchii glabrous, or occasionally arate near apex, the awns (8-)10—20 (-25) am ng; spikelets with (3—)4—5 florets; oo ~)6-10(-12) mm long, without green longitudinal bands, glabrous; blades usually vil- lous; plants strongly glaucous [central Interior Low Plateaus, in Kentucky and Tennessee] E. svensonii n OTHER NOTES The taxonomy of Elymusin North America hasa long tortuous history. In draft- ing a treatment of the species with paired spikelets, | have made some sugges- tions for nomenclature, which will hopefully guide botanists in their interpre- tation of these taxa (Campbell 1995, 1996, 2000, 2002a, 2002b, 2002c). However, it is clear that much deeper analysis is needed, including a proper focus at the CAMPBELL A 493 genetic level, instead of relying just on the rather inadequate morphological differences. There has been much taxonomic confusion, misidentification, and erroneous reporting of distributions. Many mysteries remain. An Elymus hystrix-like collection from New Mexico Ananomalous collection that may have similarities to both Elymus hystrix and E. interruptus is of interest here: NEW MEXICO [Colfax Co.], Cimarron, wooded bank, 6 Jul 1939, W.A. Silveus 4928 (TEX). This specimen was listed by Church (1967) under his “atypical” E. hystrix group with filiform glumes. Unfortunately all the florets are lost, but the glumes are narrower (ca. 0.2 mm) and more widely spreading than typical E. interruptus, and rachis internodes are shorter (ca. 5 mm). This specimen suggests that plants closely related to E. hystrix may still be found in northern New Mexico or nearby. The closest documented E. hystrix is in eastern Oklahoma. Does this New Mexico plant have a distinct origin, per- haps from some isolated introgressed population? Elymus interruptus in California [recently discovered the following specimen of Elymus interruptus: CALIFOR- NIA, Fresno Co., Pine Ridge, altitude 5300 ft, “Plants of the Sierra Nevada Moun- tains,” 15-25 Jul 1900, H.M. Hall & H.P Chandler 317 DOV). This collection was initially annotated as E. canadensis. It appears to be the first record of E. inter- ruptus from California. The few other records of E. canadensis from the state should be checked. Need to clarify typification of Elymus virginicus Some of the Linnaean material of Elymus virginicus does not clearly match our current typical concept of that species, and should probably be included with E. glabriflorus (Vasey ex L.H. Dewey) Scribn. & CR. Ball. | proposed to the Inter- national Association for Plant Taxonomy that the name E. virginicus be retained for the typical plants of Hitchcock and Chase (1951) and most other authors, with a new type established for that species (Campbell 1996). However, more careful analysis of the several sheets of Linnaean material is needed, and the Nomenclature Committee remains undecided (D. Nicholson & R. Soreng, pers. comm.). After the treatment in Flora of North America is published, there will undoubtedly be further consideration of this problem, and I hope to revive the proposal with more evidence. Corrections to Campbell (2002b) (a) In the key, leads (number 13 & 14) to E. glaucus, E. hirsutus J. Presl. and E. dahuricus Turcz. ex Griseb. will be modified substantially for the Flora of North America. E.dahuricus isa rare Asian introduction that appears close to E. hirsutus; its lemmas can be glabrous, not just scabrous or hispid, as stated in the key. (b) In the key, the lead (number 15) to E. interruptus versus E. canadensis and E. wiegandii Fernald should omit the spike internode thickness character, which reflected some erroneous data. 494 BRIT.ORG/SIDA 22(1) (c) By priority, the correct name for Elymus submuticus (Hook.) Smyth & Smyth is E. curvatus Piper. ACKNOWLEDGMENTS lam deeply grateful to Mary E. Bark worth, who has provided continual encour- agement and constructive advice during the 20 years that I have dabbled ec- centrically in Elymus. Without her invitation to contribute to the Manual of Grasses, and then the Flora of North of America, my efforts would not have progressed. She provided insightful comments on an earlier draft of this manu- script. [am grateful also to Kathleen M. Capels, who corrected several errors and inconsistencies in the key and pointed out the priority of E. curvatus. REFERENCES Campsett, JJ.N. 1995. New combinations in eastern North American Elymus (Poaceae). Novon 5:128. Campeett, J.J.N. 1996. Proposal to conserve Elymus virginicus (Poaceae) with a conserved type. Taxon 45:128-129. Campeett, J.J.N.2000.Notes on North American Elymus species (Poaceae) with paired spike- lets. |.E. macgregorii sp.nov.and E.glaucus ssp. mackenzii comb. nov. J. Kentucky Acad. Sci. 61:88-98. Campeett, JJ.N. 2002a. Notes on North American Elymus species (Poaceae) with paired spikelets. Il. The interruptus group. J. Kentucky Acad. Sci. 63:19-38. Campsett, J.J.N. 2002b, Notes on North American Elymus species (Poaceae) with paired spikelets. Ill. A synoptic key. J. Kentucky Acad. Sci. 63:39-46. Camesett, J.J.N. 2002c. Notes on North American Elymus species (Poaceae) with paired spikelets. IV. A key to the species and varieties in Kentucky. J. Kentucky Acad. Sci. 63: 47-52. Cuurcn, G.L. 1967. Taxonomic and genetic relationships of eastern North American spe- cies of Elymus with setaceous glumes. Rhodora 69:121-162. HitcHcock, A.S., and A. CHase. 1951, Manual of the grasses of the United States. 2nd ed. U.S.D.A. Misc. Pub. 200. Smith, E.B. 1991. An atlas and annotated list of the vascular plants of Arkansas. 2nd ed. Published by the author, University of Arkansas, Fayetteville. STeveRMARK, J.A. 1963. Flora of Missouri. lowa State University Press, Ames. A NEW HYBRID GENUS AND 12 NEW COMBINATIONS IN NORTH AMERICAN GRASSES Mary E. Barkworth Intermountain Herbarium tah S Logan, Utah 84322-5305, U.S.A. mary@bi are ABSTRACT One new hybrid genus, x Pascoelymus = Pascopyrum X Elymus, and 12 new combinations are pre- sented for North American grasses. Six of the new combinations are for hybrids in the Das Stipa arnowiae is transferred to Achnatherum and Hystrix californica (= Elymus ) to edie sg Egret is aan as a subspecies of Elymus ceoiaiue Avi mophila Jon n the findings of others, and the previously recog- nea subspecies of Peeudormenneria spicata are reduced to forms. Reasons for the changes are provided. RESUMEN Se Diecut un nuevo género hibrido, sbascoelymiis = = mascop uD x A oF V2 ee Snes ticede. Stipa arnowiae se transfiere a Achnatherum y Hystrix californica (=Elymus cries a ae Agropyron riparium se reconoce como una subespecie de Elymus lanceolatus, Ammophila s = oS champlainensis se reduce a una subespecie, basandonos en los hallazgos de otros, y la subespecie | 7 eal. p ] } } J £ ie ; mT I i f los cambios. INTRODUCTION Volume 24 in the Flora of North America series will be the second of the two volumes on grasses. Its completion requires publishing the following 12 names. Eleven of the 12 new combinations are for previously recognized taxa; one is a new hybrid genus. Six of the new combinations, and the new genus, are for hybrids in the Triticeae. Identifying the parents of lees without knowing which species were present in the vicinity is al fficult. In most instances, I have accepted the parentage suggested by previous workers when naming these taxa; in one instance, I have felt compelled to disagree. The known distribution of many of these hybrids is limited. The remaining six names affect non-hybrid taxa. Two reflect a generic change and four a change in rank. I have attempted to examine type material of all the taxa treated. As indi- cated below, however, this has not always been possible. All the taxa mentioned will be described and illustrated in the Flora of North America volume 24. The SIDA 22(1): 495 — 501. 2006 = 496 BRIT.ORG/SIDA 22(1) illustrations will also be available at hetp://herbarium.usu.edu/webmanual/. To the extent permitted by the institutions that own them, photographic im- ages of the type specimens examined are also available via this site and will be available via TROPICOS. The images include close ups of individual parts in addition to images of the whole specimen. NOMENCLATURAL TREATMENT POEAE Ammophila breviligulata subsp. champlainensis (F Seym.) Walker, Paris & Barrington ex Barkworth, comb. nov. Basionym: Ammophila champlainensis F Seym., Sida 2:349. 1966. TyPF: U.S.A. NEW York: Lake Cham plain, Au Sable Point, in sand, 3 Jul 1902, Nellie F. Flynn s.n. (HOLOTYPE: VT). — Walker, Paris, and Barrington (1998) reported that Ammophila breviligulata sensu stricto Fern.and A. champlainensis differ completely in glume length and flowering time, and tend to differ in inflorescence length. They noted, however, that the morphological and molecular uniformity of A. champlainensis, com- bined with the prevalence of vegetative reproduction in both taxa, is consis- tent with the hypothesis that all populations of A. champlainensis are derived from a single genetic individual. For this reason, they recommended acknowl- edging the distinction between the two taxa at the subspecific, rather than spe- cific level. 1 present the new combination here so that it may be used in volume 24 of the Flora of North America. = STIPEAE rece arnowiae a L. Welsh @ N.D. Atwood) Barkworth, comb. nov. BASION S.L. Welsh & N.D. Atwood, Utah FI. (ed. 3):799. 2003. U.S.A. UTAH. Kane Co. T4+ 35, R4W,S 13,ca 19 mi Eof Johnson Canyon Jct; pinyot liper-sagebrush-com munity at 1740 m, on white, gypsiferous member of the vane For mation, 30 May 2001, S.L. Welsh & T. O’Dell 28062 (HOLOTYPE: BRY; ISOTYPE UTC-245001). = This taxon is very similar to Achnatherum hymenoides (Roem. & Schult.) Bark worth, differing in having loosely contracted panicles with non-divaricate branches. TRITICEAE encontraron vi I triples doblesya p forma } 1 ] ] ie basi As alto (x = 9)—Heterotheca, Noticastrum "Pipi y problemen +] Tome ntaurum—los sleue ates de) se una n primitiva, similar pero con menos seas! interiores ae dé le ] que se ae en nee y Eucephalus Astereae. Los cuatro géneros tienen un vilano cuadruple que consta de cuatro series, a veces . 1) : fa [ees | q . i ee SIDA 22(1): 503 — 531. 2006 504 BRIT.ORG/SIDA 22(1) de sedas afiladas de longitud media; una serie primaria externa de sedas afiladas del 80-95% de la longitud de la serie interna; y una serie primaria interna de sedas normalmente clavadas. Tanto las flores radiadas como las del disco tienen una ordenacion similar, excepto los frutos sin vilano de H. sect. Heterotheca. El alg | t ries son distintas y en otras se intergr oa entre ellas. ea | ies del vilano d reduccion _ de x = En los géneros con numero base inferior Bradburia, pie y Oshertia, se ha ea una reduccion en el numero de series, con vilanos cuadruples, triples, do bles /a veces simples. En Chrysopsis, la serie secundaria externa esta bien desarrollada, ae : serie secundaria interna esta ausente o reducida a unas pocas sedas. En Osbertia, El vil s simple o incluye una cea a serie ] de 1-2 sedas cortas; todas las sedas largas son afiladas 0 com ho solo débi ‘la y una pequena muestra por especie, el numero de sedas pot fruto vario entre 20-125: Noticastru m, 46- 125con una media de 87 sedas; eed 80-85; Heterotheca, 23-80 con una see de 43 sedas; Pityopsis, 25-46, con una media de 37; Bradburia, flores radiadas, 30-36, del d 5; Chrysopsis and Croptilon, 18-36 con una media i: 28; Oshe >rtid, 20-35, con una media de 27. | ae menor numero de sedas tuvieron O- lew en la serie secundaria interna de longitud media. INTRODUCTION Goldenaster genera in the subtribe Chrysopsidinae Nesom (Asteraceae: Astereae) have long been reported to have a double pappus consisting of a short outer whorl of narrow to broad scales or linear bristles (e.g., Gray 1884; Fernald 1950; Cronquist 1968, 1980; Semple 1981, 1996; Semple & Bowers 1985; Semple et al. 1988). Nesom (1994a) in his protologue to subtribe Chrysopsidinae described the pappus as “2-3(-4)-seriate, persistent, the inner of 1-2 series of generally flattened bristles, outer of much shorter setae, bristles, or scales,” but did not elaborate as to the number of series in each genus. Nesom (2000) described the pappi of the North American genera as follows: Croptilon, “pappus a single se- ries of persistent, thick and rigid, equal-length, tawny to reddish-brown capil- lary bristles”; Chrysopsis (including Bradburia), “pappus in 2 series, the outer very short bristles, the inner of long capillary bristles”; Heterotheca, “‘pappus of tawny to whitish barbellate bristles somewhat uneven in length, witha shorter, outer series of lanceolate scales or bristle-like squamellae” and noted the epappose to few bristled ray floret cypselae of sect. Heterotheca; Osbertia, “pap- pus uniseriate, with numerous, nearly non-barbellate bristles, a short, outer series commonly present in O. chihuahuana’, Pityopsis, “pappus 2-seriate, the inner of barbellate bristles, the outer of much shorter setiform bristles or se- tae”, and Tomentaurum, “pappus of 45-60 white, barbellate bristles in several series, with a few inconspicuous setae or very slightly widened bristles, 0.5-L5 mim long.” Nesom (1994a, 2000) did not indicate which taxa had the flattened inner bristles. Noticastrum is native to South America and was not discussed in Nesom (2000). Zardini (1985) monographed the genus and described the pap- pus as being in two series, the external morphologically similar to the internal but of short bristles, normally straw-colored or reddish [five species] to pur- plish in N. diffusum and the bristles barbellate especially distally. Zardini also noted that the cypsela body was glandular in some taxa. fou SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 505 Many other North American Astereae have been reported to have a simple pappus. Some asters often treated in Aster sensu lato have been reported to have a “double” pappus (Eucephalus Nutt., Sericocarpus Nees: Gray 1884; Cronquist 1955) or a “triple” pappus (Doellingeria Nees: Cronquist 1968, 1980; Nesom 1994a,b; Semple et al. 2002). The “triple” pappus reportedly had two inner whorls, the outer slightly shorter and tapering and the inner bristles clavate. However, Hood and Semple (2003) demonstrated that nearly all species of gold- enrods, which had been reported to have a simple pappus, in fact had a double pappus with two primary series of long bristles (the outer shorter and taper- ing, the inner clavate) and at least some species had a secondary outer series of a few very short bristles, ie. Solidago has a double or triple pappus. Semple and Hood (2005) demonstrated that many North American aster genera have a double, triple, or quadruple pappus and concluded that the likely primitive pappus of the North American Astereae consists of four series: the primary in- ner series of long, clavate (flattened) bristles, a slightly shorter primary outer series of long tapering bristles,a secondary inner series of tapering bristles 40- 80% the length of the primary inner bristles, and a secondary outer series of short bristles or narrow to broad squamellae usually only 10-15% the length of the inner primary bristles. Hood and Semple (2003) proposed a terminology for a triple pappus, ie. secondary outer whorl, primary outer whorl, and pri- mary inner whorl bristles, which Semple and Hood (2005) modified to cover the discovery of a fourth series of bristles found in asters but not seen in gold- enrods. They also noted that “series” was a more accurate label than “whorl” based on observations of both North American and Old World aster genera. They concluded that due to the subtlety of the differences between the longer bristle series in many species, it would be practical to treat the pappus of Symphyotrichum as “appearing simple” in keys to identification. The details of the “double pappus” of the Chrysopsidinae were re-examined in light of the recent discoveries on pappus traits of goldenrods and asters and the results are presented in this paper. MATERIALS AND METHODS A preliminary survey was undertaken to examine the pappus of one or two specimens of representative species of the genera of the Chrysopsidinae sensu Nesom (2000). Subsequently, a more detailed survey was conducted involving eight genera, 58 species, and 94 taxa of subtribe Chrysopsidinae listed in Table 1. Observations were made using a dissecting scope (10-70) or a compound light microscope (20-400). The degree of the clavateness of bristle tips was determined using the 0-4 rankings described in detail in Hood and Semple (2003). At least five different fruits from each species were observed under the dissecting microscope at a maximum of 70x; in some cases many fruits from ie ] many individuals examined to investigate the frequency of variation. The 506 BRIT.ORG/SIDA 22(1) majority of observations were made on specimens in the WAT Herbarium, ad- ditional specimens on loan from MEXU, MO, OS, SD, TEX-LL and UC (Holmgren et al. 1990) were also ined. In addition to the several methods used in evalu- ating pappus features listed by Hood and Semple (2003), assessment of the char- acteristics of the short secondary outer scales or bristles was also systemati- cally undertaken with three traits recorded. The large difference in numbers of pappus bristles per fruit noted in the preliminary survey and the literature led to recording a rough estimate of bristle number for each taxon by counting the number of bristles on three to five representative fruits per taxon. These were compared to counts of numbers of bristles in (Smith 1965; Nesom 1991b; Semple & Bowers 1995; Semple 1996). Thus, Table | has eight columns of observational data plus a column for additional comments, while only four were reported by Hood and Semple (2003) and five in Semple and Hood (2005). Observations on the compound microscope were made from both un- mounted fruits and from slides prepared following Semple and Hood (2005). For each species, usually two to three disc floret cypselae with or without co- rollas were mounted in Cytoseal-60'™ mounting medium under a cover slip; immature ray floret fruits were also often mounted on the same slide for com- parison. Observations at 20-400 on the compound microscope were made similarly to the observations under the dissecting scope at 30-70. Observa- tions made on the two kinds of scopes were compared and any discrepancies were resolved by re-examining specimens. Digital photomicrographs were taken using a Nikon CoolPix 990 camera manually held against the ocular lens of either the dissecting or compound mi- croscope. Pictures were taken of specimens under the compound light micro- scope with either below stage or above stage lighting. Scanning Electron Micro- scope (SEM) photomicrographs were made using Kodak FX or Ilford Pan F Plus film commercially developed and subsequently digitized by scanning the nega- tives. Final digital illustrations were made using CorelDraw 12® from digital im- ages edited with Corel PhotoPaintl2® (Corel Corp.). In some illustrations, the con- trast was manipulated to increase the difference between pappus bristles and background for illustrative purposes; backgrounds were darkened considerably and sometimes bristles were lightened somewhat to correct for uneven lighting and exposure; the converse was done in bristle tip silhouette illustrations RESULTS AND DISCUSSION The pappus of most species of the Chrysopsidinae consists of 3-4 series (Figs. 1-7) while a minority of taxa has a pappus of 1-2 series (Fig. 8). The quadruple pappus consists of the following series: |) a well develop secondary outer series usually of many, short, linear-bristly to broad scales; 2) a secondary inner se- ries of mid-length, tapering, non-clavate bristles that were generally 40-80% the length of the primary inner series (Figs. LE J, 2F 2N, 4K, 5A, 5M, 6A); 3) a SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS eile onion sone nee Fic. 1. Fruit and ts of Heteroth lisc fruit pt A; scal Tmmin A and F,=0.5 mm in C-D and H—- | and = 10 min E.A-B.H. 9 liflora (Semple & Semple 5575 WAT). A. Mature epapp yand papr i (long) and bristles. CE. H. inuloid ( & Woodland 1928 WAT), SEMs. C. Tip of mid lengt ths Jary inner bristle. D. Secondary outer series scales and overlapping bases of longer bee peal of barbs on me person of inner series PBUE: F- a H, mexicana Uppne el mall: F. paawe fruit ( ).G. Upper p \ )and f y ou LH. isessiliflara subsp bolanderioides (Semple & Chmiel ki 8918a WAT: SEMs) H Weakl f pri series bristles.1.5 lary les.J.H. (H 671WAl), tip pemg length secondary inner iste Hee en ee (Semple 9902 WAT), c tips of primary i primary outer bristles. L- Lt . oregona var pacta (Semple & Heard 5693 WAT).L. Silhouette of tion of mid lengtt 1 I 1 long primary outer and inner bristles. M. Linear secondary auter bristle (arrow). TCTIQUIT SCCUNUaTy TIC 508 BRIT.ORG/SIDA 22(1) pH primary outer series of tapering bristles that were 85-95% the length of the primary inner series (Figs. 1B, 1], lJ, 3D, 4D, 4G, 4K, 5G, 5K, 6KC, 7B, 7E, 71, 7K, 8A, 8C): and 4)a primary inner series of very subtly to moderately clavate tipped bristles (Figs. 1B, HI, 4D, 5B, 0D, 7E, 7J). A four-series pappus is plesiomorphic in the North American Clade (sensu Noyes & Rieseburg 1999) of the tribe Astereae (Semple & Hood 2005). Heterotheca, Noticastrum, and Pityopsis, with the plesiomorphic chromosomal base number of x = 9, has a quadruple pappus in all or most species, as does the monotypic Tomentaurum, whose chromosome number is not yet known but is hypothesized here to be x=9 based on indument and fruit similarities with x = 9 Noticastrum. In the genera with lower chromosomal base numbers, Croptilon (synonym: Haplopappus sect. Isopappus: Hall 1928), Chrysopsis, and Bradburia, the pap- pus is usually reduced to three pappus series or less but sometimes had I-few bristles of one or both of the missing series. Osbertia, which also was included in Haplopappus by Hall (1928), has a vestigial secondary series and a possible biseriate primary series. Thus, in all genera of the Chrysopsidinae, some evi- dence was found for the presence of at least 2-3 pappus series and usually more. A large range in pappus lengths and numbers of bristles was observed among genera and within some genera of Chrysopsidinae. The smallest fruits with the lowest numbers of bristles were seen in Bradburia, Chrysopsis, Croptilon, and Osbertia. These traits vary considerably in Heterot Pityopsis. The majority of species of Noticastrum examined had the longest pappus bristles and the highest number of bristles observed. Pappus and fruit traits are discussed in detail below by genus with higher base number genera presented first. Data for all taxa examined are presented in Table 1. Heterotheca (x = 9; all 24 species, all 48 taxa examined).—Cypselae straw colored to brown, obconic, compressed laterally or not, those of ray florets of - ten 3-angled, 1-4.3 mm long, sparsely to moderately strigose, dimorphic in sect. Heterotheca (Fig. 1A), ray florets usually lacking a pappus and glabrous, rarely pappose and intermediate; ribs 5-10, sometimes dark, resin filled; pappi qua- druple or sometimes triple through loss of the secondary inner series, series often intergrading; 23-80 mid to long bristles, averaging about 43 per fruit for the genus, barbs usually evenly arranged around the round shaft for most or all the length of the bristle; secondary outer series obvious to inconspicuous, of linear to narrowly triangular or oblong scales (Figs. LD, I) or linear bristles (Fig. IM), these usually 5-15(-30)% the length of the longest primary bristles; rarely up to 40% (some specimens of H. mucronata var. harmsiana), secondary inner bristles many (H. oregona, Fig. lL) to few or absent (e.g., H. villosa var. nana), 35- 85% the length of the longest primary bristles, sometimes grading in length from linear secondary outer bristles into the primary outer bristles (e.g., H. brandegei, H. stenophylla, H. thiniicola), primary inner bristles tapering dis- tally, 85-95% the length of the primary inner bristles, fewer than the primary — heca and Taste 1. Pappus variation in subtribe Chrysopsidinae; disc floret cypselae (ray fruit for B hirtella). Clv, degree of clavateness of primary inner bristles (0 = not clavate to4= svongy clavate), Clv- -Tap, clavate alternating with tapered bristles (primary inner and outer whorls, y = yes, - not obviously so); Lgth, primary outer bristles shorter than primary inner bristles; Ovrlp, degree of overlapping of bristles at the base (0 = not observed: 1 = San overlap; 2 = definite overlap); 2"9-0, evidence for a secondary outer whorl of short scaly-bristles; 2ne- Oo pata. mie of the secondary outer mites (0 = linear, 1 = narrowly triangular, 2 = lanceolate); % = length compared to 1° inner bristles; No. Br= ap Jate number of mid to long bristles (2"¢ inner, 1° outer and inner whorls; small sample size); percents listed are bristle lengths compared to the length bathe: primary inner series. Taxon Clv Clv- Lgth Ovrlp 2rd. 2"4_9 24.9 No. Comments Tap 0 wid % Br Heterotheca (x=9) sect. Heterotheca (di phic; ray cypselae nearly always lack pappus) grandiflora 1 y y ] y 0-1 5-15% 50-60 2°¢ inner bristles few, 40-70% H. inuloides var. inuloides 1-2 y y 1 y 1 5-15% 60-65 2°? inner bristles many, 35-65% var. roseum ] y y 1 y O-1 5-10% 60-65 2° inner bristles few, 50-70% var. viridis Q-1 y y ] y 1 5-10% 60-65 2” inner bristles few, 70-80% H.leptoglossa 2 y y 1 y 1 5-15% 35-45 2” inner bristles few, 70-80% grading into 1° outer bristles H. subaxillaris subsp. latifolia ] y y 2 y 0-1 5-10% 25-30 2” outer bristles 5-10%; 2™ inner bristles O-very few, 60-70% subsp. subaxillaris ] y y ] y O-1 5-15% 30-45 2™ outer bristles 5~10%; 2" inner bristles few, 40-70% sect. Ammodia (rayless) H. oregona var. compacta ] y y ] y @ 5-8% 60-75 2™ outer bristles obscure; 2" inner bristles few, 50% var. oregona ] y y 2 y 0 5-10% 75-85 2” outer bristles obscure: 2° inner bristles few, 50% SLIVUL Sfidd¥d JVNIGISdOSYAHD ‘J1dW3S 605 TABLE 1. cont. OLS Taxon Clv Clv- Lgth Ovrlp 2nd. 24.9 24.9 No. Comments Tap 0 wid % Br var. rudis 1 y y | y 0 5-10% 50-55 2°° outer bristles few; 2°? inner bristles few, 40-70% var. scaberrima 1 y y | y 0 15-30% 60-70 2" inner bristles few, 40-70%, grade into 1° outer bristles sect. Phyllotheca H. brandegei (rayless) 1 y y 1 y 0-1 10-15% 35-55 2™ inner 70-80%, grade into 1° outer bristles H. canescens 1 y y 1 y 1 5-10% 25-40 2.4 inner bristles few, 50-70% H.camporum var. camporum 1 y y 1 y 0-1 10-15% 30-45 2" inner bristles O-few, 70% var. glandulissimum 2 y y 1 y 0-1 5-10% 30-45 2° inner bristles few, 50-70% H. fulcrata var.amplifolia O-1 y y 1 y O-1 10-15% 35-45 2™ inner bristles few, 55-75% var. arizonica ] y y ] y O-1 10-20% 35-45 2” inner bristles very few, 60-65% var. fulcrata 0-1 y y 1 y 1-1+ 10-15% 25-45 2° inner bristles O-few, 70-80% var. senilis 1 y y 2 y 1 5-15% 28-45 2°4 inner bristles few, 70-85% grade into 1° outer bristles H. gypsophila 1-2 y y ] y 0 5-10% 30-40 2" inner bristles few, 70-85%, grade into 1° outer bristles H. jonesil 1 y y 2 y 1 5-15% 25-35 2” inner bristles O-few, 70% H.marginata 0-1 y y 1 y 0-1 10-15% 40-52 24 inner bristles very few, 70-80% H. mexicana 0-1 y y 1 y 0-1 10-15% 40-60 2° inner bristles few, 40-60% H. monarchensis 1 y y 1 y 1+ 10-15% 25-40 2° inner bristles O-very few, 70% H. mucronata var. mucronata 1 y y 1 y 0-1 5-10% 35-50 2° inner bristles few, 80-85% var. harmsiana ] y y 1 y 0-1 10-40% 35-40 2™ outer bristles can be long; 2” inner bristles few, 70-80% (L)@2 WaIs/9Y¥O' LY TABLE 1.cont. Taxon H. pumila H. rutteri H. sessiliflora subsp. bolanderi subsp. echioides var. bolanderioides var. echioides var. camphorata subsp. fastigiata var. fastigiata var. sanjacintensis subsp. sessiliflora H. shevockil H. stenophylla var. angustifolia var. stenophylla H. thiniicola Clv Clv- Lgth Ovrlp 2nd 24.9 24.0 No. Comments Tap 0 wid % Br 0) - y 2 y 0 10-20% 35-45 2™ inner bristles very few, 75-80% 0 = y 1 y 0-1 20-25% 35-46 2™ inner bristles very few, 75-80% 1 y y 1 y 1 5-15% 50-70 2” inner bristles few-many, 55-85% 2 y y 1 y 1 X-X% 35-60 2°? inner bristles few, 55-70% 1 y y 1 y 1 5-15% 25-50 2" inner bristles 0 or few—many, 60-70%, some fruits have significantly few bristles and lack 2" inner whorl 2 y y 1 y 1 5-15% 35-45 2°? inner bristles few, 60-70% 0-1 y y 2 y 0 5-30% 50-70 2" inner bristles many, 45-85%, grade into 2° outer and 1° outer whorls ] y y 2 y 0 5-30% 50-80 2” inner bristles Lo) 45- ica grade into 2" outer inner whorls 1-2 y y 2 y 1+ 5-10% 45-50 2™¢ inner bristles few, 50-70% 1 y y 0) y O-1+ 5-10% 35-45 2™ inner bristles few, 60-65% ] y y 1 y 0-1 5-15% 35-45 2°? inner bristles few, 50-70%, grade into 1° inner bristles 1 y y 1 y 0) 15-10% 25-42 2°? inner bristles few, 60-70%, grade into 1° inner bristles 1 y y 2 y 0 10-25% 25-35 2°? inner bristles many, 30-60%, grade into 2"° outer and 1° inner SLIVUL SddWd JVNIGISdOSYAH) ‘J1dWAS LLS TABLE 1. cont. Taxon H. villosa var. ballardii var. depressa var. foliosa var. minor var.nana var. pedunculata var. scabra var. sierrablancensis var. villosa H. viscida H. zionensis Noticastrum (x=9) acuminatum N.calvatum N. diffusum N. gnaphaloi oides N. sericeum Clv Clv- Lgth Ovrlp 2nd. 24.9 2nd_Q No. Comments Tap 0 wid % Br ] y y ] y O-1 5-10% 35-45 27 inner bristles few, 60-70%% 1 y y 1 y 0) 10-15% 5-45 2° inner bristles few, 50-65% 1-2 y y 0 y 0-1 8-15% 35-45 2” inner bristles few, 60-65% ] y y 2 y 0-1 5-15% 30-45 2” inner bristles few, 60-70% 1-2 y y 0) y 0-1 10-15% 28-40 2” inner bristles O-very few, 65% 2 y y 1 y 1 10-25% 25-45 2°° inner bristles O-very few, 80% | y y 0) y 0-1 8-15% 35-45 2°° inner bristles very few, 60-70%, grade into 1° inner bristles 1-2 y y 1 y 0-1 10-15% 35-45 2° inner bristles 70-80%, grade into 1° inner bristles 1 y y 0 y 0) 5-15 30-50 2” inner bristles 60-70%, grade into 1° inner bristles 1-2 y y 0 y 1 5-15% 23-40 2° inner bristles very few, 50-60% 1 y y 1 y ) 10-15% 30-45 2™ inner bristles few, 50% 0-1 y y 2 y 0 5-20% 75-90 2™ inner bristles many, 65-75% 0-1 y y 2 y 0 5-10% 90-100 2” inner bristles few, 80% 0-1 y y 2 y 0 10-30% 5-80 barbs of bristles tinted red; 2"° inner bristles few, 60-80% 0-1 y y 2 y 0 5-10% 100-115 2” inner bristles few, 80% 0-1 y y 2 y 0 10-20% 45-50 2™ inner bristles few, 40-80% 0-1 y y 2 y 0 10-30% 80-90 2” inner bristles few, 80-85% 1 y y 2 y 0 5-15% 75-80 2" inner bristles few, 70-85% 0-1 y y 2 y 0 10-20% 120-125 2°? inner bristles few, 70-80% ZLS (L)@Z VOIS/DYO"LINa TABLE 1.cont. Taxon (lv Clv- Lgth Ovrlp 2nd. 24.0 24. No. Comments Tomentaurum (unknown) T.nivea 0-1 y y 2 y 0 10-20% 75-85 2” inner bristles few to many, 60-70% Pityopsis (x=9) sect. Pityopsis P falcata 0-1 y y 1 y 0 10-15% 30-40 2° inner bristles O-few, 45-80% P flexuosa 1 y y 1 y ) 5-10% 30-50 2°° inner bristles 0O-few, 45-85% P pinifolia 1 y y 2 y 1 10-15% 25-30 2” inner bristles not seen thii 0-1 y y ] y 1 5-15% 25-35 2™ inner bristles, few, 40-85% sect. Graminifoliae P. r var. adenolepis 0-1 y y 2 O-1 10-20% 25-35 2™ inner bristles O-few, 70-80% var. aspera 0-1 y y ] y 0-1 5-15% 30-35 2” inner bristles few, 70-80% P. graminifolia var. aequilifolia 0 y y 0) y 0-1 10-15% 40-45 2° inner bristles 0O-few, 70-80% var. graminifolia 1 y y 2 y 0 10-20% 32-40 2° inner bristles few, 75-85% var. latifolia 0 y y 1 y 0 5-15% 35-45 2°¢ inner bristles 0O-few, 60-85% var. tenuifolia 0 y y 1 y 0-1 5-15% 35-45 2°° inner bristles O-very few, 70- 85%, grading into 1° outer bristles var. tracyi 0-1 y y 0 y 0-1 5-10% 35-45 2 inner bristles few, 50-85% P oligantha 0-1 - y 1 0-1 5-10% 25-40 2” inner bristles O-very few, 75-85% Croptilon (x=7,6,5,4) C. hookerianu var. graniticum (x=7) 0 = y 0 y 0 5-20% 30-35 2°? whorls absent or very rarely a short linear bristle present SLIVYL SMdd¥d JVNIGISCOSYAHD ‘31dW3S ELS Tasle 1. cont. Taxon var. hookeri (x=6) var. validum (x=5) C.rigidifolium (x=6, 5) C. divaricatum (x=4) Bradburia (x=4,3) irtella B.p ilosa (lv Clv- Lgth Ovrip ana 24.9 2n4_() No. Comments Tap 0 wid % Br ) ~ y 0 y ) 5-10% 25-30 ° outer vityetes a 2” inner oe 5 O-few 0 7 y 0 y 0 (5-10%) 25-30 2°° outer vac Ae rarely vestigial (or abnormal short 1° bristle) ) - y ) n 0) 5-10% 25-30 2” out bristles O-few; 2" outer bristles few 35-75%, grade into 1° bristles 0 - y 1 n - = 25-30 2” outer bristles not seen; 1° outer bristles 70-80%, sometimes grade nto 1° in O0-1(-2) y y 2 y 2 10-15% 30-36 Ey flo ae inner bristles few, 0-60%; 1 r 70-95%, grade into 1° nae see text for comments on pigmentation: disc florets—re- duced pappus, 0-5 short scales to long broad-based bristles O(-1) y y 0 y 2 8-20% 18-25 2™ outer whorl of large, obvious to naked eye, lanceolate-oblon scales 0.1-0.2 mm wide; 2°? inner bristles O-few, 75-85%, usually absent, grade into 1° bristles; 1° comments on pigmentation. Pls (L)@Z VOIS/9YO' LYS TABLE 1.cont. Taxon Clv Clv- Lgth Ovrip 2nd. 2n4.9 24.9 Comments Tap 0 wid % Br Chrysopsis (x=5, 4 and x, = 9; most species lack 2" inner bristles and have few primary outer bristles) C.de 2 y y y 0 15-25% laneyi C.floridana dfreyi var. godfreyi var. viridis C. gossypina (x, = 9) subsp. cru/seana subsp. gossypina subsp. hyssopitolia C. highlandsensis C. lanuginosa C. latisquamea C. linearifolia subsp. dressii subsp. linearifolia C.mariana (x = 4) C. scabrella C. subulata Osbertia (x=5) O. bartlettii ] y << Me. Oe eS y <— os ain ai ai a a ] —- Oo —- 000 © y Se Ss Ss OS 0 10-15% 10% 5-10% 10-15% 10-15% 10-15% 15-30% —15% 25-30 30-35 30-35 25-35 20-25 20-25 2 inner bristles not seen 2™ inner bristles not seen 2” inner bristles not seen 2” inner bristles not seen 2° inner bristles not seen 2° inner bristles not seen 2° inner bristles not seen 2™ inner bristles very few, 55-60% 2™ inner bristles not seen 2™ inner bristles not seen 2™ inner bristles not seen 2™ inner bristles not seen 2” outer bristles no wider than in some other species of genus (sample includes 2x, 4x, 6x, 8x); 2” outer bristles absent; 1° outer bristles 0O-few 2” inner bristles not seen 2™ inner bristles not seen 1° vary in length (80-100%); one fruit had 1 fine 2°° bristle 25% the SLIVHL SNdd¥d JVNIGISdOSYAHD ‘1d WS SLs T l.cont Taxon Clv Clv- Lgth Ovrip 2 24.9 274.9 No. Comments Tap 0 wid % Br length of the longest 1° bristles O. chihuahuana 1-2 y y 0-1 y 0 12% 25-30 2° outer bristles few; 2"° inne bristles 45-65% of the length of the longest 1° bristles; 1° outer bristles 85-95%, grade into 1° inner bristles O. stolonifera 0-1 - y 1 y 0 5-40% 25-30 2™ outer bristles grade into 2 inner bristles; 2" inner bristles 50— 70% of the length of the longest 1° bristles; 1° outer bristles 90-95% grade into weakly clavate 1° inner bristles (L)@2 VaIs/9¥O' LIYE SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 517 inner bristles; primary inner bristles very weakly (Fig. 1B) to moderately clav- ate (Fig. LK), 3.4-10 mm long. Noticastrum (x = 9; eight of 19 species examined).—Cypselae straw col- ored to brown, fusiform, 4-6 mm long, sparsely to moderately strigose, some- times stipitate glandular distally (Fig. 2B-C) or over entire surface (Fig. 2H); ribs, 16-22, surface between ribs usually golden-brown, translucent, resin filled; pappi quadruple, each series usually grading into next, 45-125 mid to long bristles, averaging about 87 per fruit for the genus, straw colored, orange-rust or purplish (e.g., N. diffusum; Fig. 2A), yellow-rust pigment in shaft over entire length, the red pigment concentrated in the barb tips (Fig. 2F), barbs evenly arranged around the round shaft along the entire length of the bristles; sec- ondary outer series obvious to inconspicuous, linear triangular scales (Figs. 2N) to linear bristles (Fig. 2B), usually 5-20(-30)% the length of the longest pri- mary bristles; secondary inner bristles many, 40-85% the length of the longest primary bristles (Fig. 2E); primary inner bristles many, 85-95% the length of the primary inner bristles; primary inner bristles very weakly (Fig. 2D) to some- times moderately clavate, 7-14 mm long, much exceeding the disc corolla lobes. Tomentaurum (x = unknown; monotypic, IT. nivea examined).—Cypscelae fusiform, 4-5 mm long, densely strigose (Figs. 3B-C); ribs 10-18, shallow, ob- scured by hairs; pappi quadruple, each series usually grading into next, 75-85 mid to long bristles, averaging about 80 per fruit, straw colored, barbs evenly arranged around the round shaft along the entire length of the bristles; sec- ondary outer series obvious to inconspicuous, very linear triangular scales to tapering bristles (Figs. 3H-D, 10-20% the length of the primary inner bristles; secondary inner bristles few to many, 40-85% the length of the longest pri- mary bristles (Fig. 3G); primary inner bristles many, 85-95% the length of the primary inner bristles (Fig. 3D); primary inner bristles very weakly to weakly clavate (Fig. 3E-F), 10-11 mm long. Pityopsis (x = 9; all seven species, all 11 taxa examined).—Cypselae straw colored to often dark reddish-brown, narrowly to broadly fusiform, 3-4 mm long, ribs 8-10, sparsely to moderately densely strigose; pappi quadruple or sometimes triple through loss of the secondary inner series, 25-46 mid to long bristles, averaging about 37 per fruit for the genus, light tan to straw colored, sometime pale rust colored proximally, barbs evenly arranged around the round shaft along the entire length of the bristles; secondary outer series of linear (Figs. +E, G arrows) to broad (Figs. 4+K-L), barbed scales, 5-15(-20)% the length of the primary inner bristles; secondary inner series of tapering bristles, few, 45-85% the length of primary inner bristles; primary outer series of tapering bristles 90-95% of primary inner bristles (Figs. 4C-D, E H; arrows); primary inner series of weakly clavate bristles (Fig. 4D), 4-9 mm long. Croptilon (x = 7, 6, 5, 4; all three species, all five taxa examined).—Cy pselae straw colored to brown or reddish-brown, sometimes mottled, fusiform to nar- 518 BRIT.ORG/SIDA 22(1) Fic. 2 papy i ticast : (Zardini 1094 TEX). A. Fruit with rusty reddish antl secondary outer bristles (arrow) and the bases longer seri istl leb mm. €. SEM of stipitate glands (arrows) on upper fruit body; scale il = 100 um. D. lips of primary inner series srs, E. ee Ui seeunealy inner t ? Peet (H hbach 52879 MO). H. Fruit t | I istle ( ).1 Detail of multi- ribbed ues mony with stipttate ganas kart): - N. macrocephalum (Montes 14942 MO). I. Maule fruit are Im- f K.T clavate to very v p of primary r Pa tert tl 1 PP fal y rina tle f 1 y bristle and Uppel mid portions (open arrow). SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 519 Fic. 2.5 . 1 Pema re £T. ° 1° Leisite (Dre, ln TIZAL RAN) lah Tmmin A-B =0.5 mm in H. A. Nearly fruit with lary outer series (1°o), secondary inner series (2°i), primary outer series (1°0) J id . id (1 O°V bL 7-4] . J* 7 J R Eruit hndv C€ AA A 4 £f, Hy | J nm TT: £| - 1 car fl | r é co r é f vf a few are broken off. E-F. Tips of primary inner bristles. G. Tip of dary i bristle ( ) and mid porti f four primary bristles. H-I. Secondary outer series bristles. rowly obconic, 2-3.2 mm long, ribs 10-20, sometimes either the ribs or faces between them golden-brown to brown translucent, resin filled, sparsely to moderately densely strigose (Figs. 4A, G, K); pappi quadruple, triple, double, or sometimes single, 25-35 mid to long bristles, averaging about 28 per fruit for the genus, somewhat to dark rust colored in shaft, darkest proximally, becom- ing paler to whitish distally, barbs not pigmented, barbs evenly arranged around the round shaft along the entire length of the bristles; secondary outer series of linear bristle-like scales, O-few (Figs. 5D-E, I-J, N-O), 5-10(-20)% the length of the primary inner bristles (Figs. 5D-F, I-J, O); secondary inner series of taper- ing bristles, absent or rarely 1-2, 50-70% the length of primary inner bristles (Fig. 5A, M; arrow) or these just atypically short primary outer series bristles; primary outer series of tapering bristles 80-95% of primary inner bristles, grad- ing into primary inner bristles (Figs. 5A, 5G, 5K-L); primary inner whorl of non- clavate/very weakly clavate (Fig. 5B) to weakly clavate bristles (Fig. 5L), 4-9 mim long. Croptilon has been treated as Haplopappus sect. Isopappus (Hall 1928; Smith 1965) and described as having a pappus with “a single series of equal (or nearly 520 BRIT.ORG/SIDA 22(1) Fic. 4. Frui its of Pityopsis, disc frui let 1mm in A-B and I,= 100 Lim inJ, = 0.5 mm in K, and = 0.1 mmin L.A. P graminifolia var. - aequilifolia, mature fruit (Semple et al. 3992 WAT). B—E. P. falcata (Semple 3366 WAT). B. au aay: C- 2 Tips of weakly caval primary inner bristles and attenuate, shorter primary outer bristles (arrows).E y | Es sels ead a re F. Tips of primary inner and outer bristles G. Bri ly secondary outer i | 0537 WAT, tips of primary inner and outer (arrows) series bristles. \- L. ane var. mdenelepD (empleet a wor, lL. Maullte murchad ys = oe NS J. Clavate tips of primary inner series bristles. y highligl K) and proximal portions primary series ae SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS Fic. 5. Fruit and pappus traits of Croptifon, disc fruits; scale bars = 1 mm. A-D. C. hookerianum var. hookerianum. A Mature fruit (Semple & Brouillet 3363 WAT).B—D. Turner et al. 46 TEX-LL.B. Tips of primary bristles. C. Tips of secondary inner bristle and upper mid portions of two longer Hee D. Tip of suis outer arisile, E.C, negkeanum var. graniticum, secondary outer bristle (E.B. Smith 622 TEX). F. 4259 TEX). G—J. C. rigidifolium (G and |, Cory 11886 TEX; ee Nesom 5216 TEX, SEMs). G. Mature fruit. H. tine of pri- mary bristles. I-J. Bases of primary bristles and secondary outer buses fartaws). K- iM C iver Cemnle Cimiclowsse0n) WAT). K. Mature uth a iy of panany bristles. M > 522 BRIT.ORG/SIDA 22(1) equal) capillary bristles, more or less ferruginous” (Smith 1965). Nesom (2000) as noted in the introduction also described the pappus as being ina single se- ries. The results presented here indicate that the pappus neither consists of a single series nor are the bristles equal or nearly equal. The higher base number taxa (e.g., C. hookerianum var. graniticum with x=7 ) are more likely to have three or four series of bristles, while the lowest base number taxon, C. divaricatum with x = 4, may have only 1-2 series. The gradation in lengths of longer bristles and the lack of clearly clavate bristle tips results in uncertainty in determining the number of series present. The conclusion that multiple se- ries are present is based on observations of many fruits, some of which have longer bristles of 2-3 distinct lengths. The much shorter outer bristles clearly exterior to the long bristles are essentially the same as the secondary outer se- ries scales/bristles of some species of genera long recognized to have a short outer series, e.g. Heterotheca, Noticastrum, Pityopsis and Chrysopsis. The small fruit size generally correlates with a lower number of bristles and very few bristles in some series. Bradburia (x = 4, 3; two species with very different pappi traits).—1) B. hirtella: ray florets fertile, disc florets functionally staminate, ovary aborts; ray floret cypselae straw colored to brown or reddish-brown, obconic, + triangular in cross-section, ca. 2mm long, 5-10 shallow ribs per face, moderately strigose, more densely so along angles (Fig. 6A); pappi quadruple or triple, the series grad- ing into each other, 18-26 mid to long, long-barbellate bristles (Fig. 6A), densely pigmented red proximally to only the barbs pigmented one third to one half the length, pale yellow to pale rust grading to white distally; secondary outer bristles linear, 5-20% the length of the longest inner bristles (Fig. 6B), barbs evenly arranged around the round shaft along the entire length of the bristles, uneven on the distal portion of the longer bristles; secondary inner bristles grad- ing from secondary outer into primary outer, tapering, 25-85% the length of primary inner bristles; primary inner bristles round proximally, flattened dis- tally and weakly to moderately clavate (Fig. 6C), 2.5-3.5 mm; mid to longer bristles pigmented proximally, the red pigment concentrated in the barbs (Figs. OB, D). Disc floret pappi reduced to 0-5 scales and/or broad-based proximally flattened bristles (Figs. 0G-J), the longest weakly clavate (Fig. 6H). — 2) B. pilosa: ray and disc pappi similar, cypselae obconic, slightly compressed, sparsely stri- gose, 10-20 shallow ribs; pappi triple or double (Fig. 7K); secondary outer scales linear to lanceolate or oblong (Fig. 6L), 0.05-0.2 mm wide, barbellate proximally, margins jagged, especially distally; secondary outer bristles absent, primary outer bristles tapering, 0-2, 75-85% the length of the inner bristles; primary inner bristles tapering or only very remotely clavate; all longer bristles pig- mented proximally with the concentration decreasing distally, yellow to rust, pigment in shaft, not barbs. The combination of broad, whitish, light-reflecting scales and proximally SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 523 Saganaeier* 4 of - : Fic. 6. Fruit and papy tae ds Bigapuna, scale bars = ao mm. oe J. B. hirtella (Nesom 7470 WAT). A-E. Ray fruit. B. g b g g outer brist tle caivow); e Tip of Ae! fattened Hong pet L 4 1 D MAtd Ld at ¥ 1 aL 4; TL L iT devel J. Functionally inate disc floret. F. Floret with Jary le ( ") 1 two long broad-b ad hrictles G.B f f] ith | j | H. Tit ra he GA. Tiy oe in G. J. Details of scales bases. K-L. B. pilosa, disc fruit (Semple & Brammall 2753 WAT). K. Mature fruit. L. Secondary outer series scales. 524 BRIT.ORG/SIDA 22(1) darker-pigmented, long bristles as background makes the secondary outer se- ries of Brad buria pilosa obvious to the naked eye. The narrower to mid-width scales are similar to broader scales in some species of Chrysopsis and Heterotheca. The widths of the secondary outer scales/bristles in Heterotheca and Chrysopsis form a continuum that overlaps at the wide end with the nar- rower width scales of B. pilosa. The broader scales of B. pilosa are unique in the subtribe. The broad scaly bases of the few, long bristles attached to the aborted ovary of the disc florets of B. hirtella are also unique in the subtribe. Possibly, the genes controlling normal scale and bristle development have been expressed together in single structures rather than separately in different series of struc- tures in disc floret pappus development. Therefore, but in different ways, the two species of Bradburia have broader pappi members than found in all other genera in the subtribe. The phylogenetic position of Bradburia is uncertain. Nesom (199la, d) in- cluded the two species of Brad buriain Chrysopsis sect. Brad buria. Semple (1996) transferred C. pilosa to Bradburia as defined here. In the cladistic study by Semple and Tebby (1999), Bradburia grouped with Croptilon and Heterotheca, while Chrysopsis grouped with Pityopsis, Noticastrum, and Tomentaurum. Based on pappus traits, Bradburia is no more similar to Croptilon than to Chrysopsis. Pigmentation of bristles is similar in Bradburia and Croptilon, but a well developed secondary outer series is shared by Brad buria and Chrysopsis, although the scales are much broader on average in B. pilosa than in any spe- cies of Chrysopsis. A DNA sequence based phylogeny of the subtribe has not been published. Until such a study is completed, the phylogenetic position of Bradburia within the subtribe remains uncertain, although it is likely derived from either the Croptilon or Chrysopsis lineages, unless a separate dysploid se- ries from x = 9 is hypothesized. Chrysopsis (x = 5, 4, x2 = 9; all taxa examined, 11 species, 15 taxa).— Cypselae straw colored or rarely dark red-purple, compressed obconic, smooth (Fig. 7G) or with 1-10 shallow ribs and sometimes 1-5, yellow to red brown, clavate, translucent ridges per side (Fig. 7D), sparsely to densely strigose, 1.5-3 mm, pappi triple or rarely quadruple, sometimes double in C. mariana, barbs evenly arranged around the round shaft along the entire length of the bristles; secondary outer series of linear (Fig. 7F) to narrowly triangular, jagged-edged scales (Figs. 7C, 7L), 0.4-1.4 mm, 5-15(25)% the length of the primary inner bristles, 20-40 mid to long barbellate bristles in 1-2(-3) series, whitish to straw colored; secondary inner series usually absent, rarely 1-3 tapering bristles 45- 85% the length of primary inner bristles; primary outer series of tapering bristles 85-95% of primary inner bristles (Figs. 7B, E, larrow, K); primary inner series of weakly to moderately clavate bristles (Figs. 7E, I-J), 4-7 mm long. Some fruits of C. mariana (x = 4) have a true double pappus of just short secondary outer series scales and long, clavate primary inner series bristles (Fig. — SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 525 Fic. 7. Frui J its of Ct is, disc frui let Sl 0.5 mm in L.A-C. C. latisquamea BW aah 2753 WAT) A. Mature fruit B. Tips of bri C.5 y De ms scales D. Fruit epady jish-brown resin- filled rib Co (Semple et al. 312 pap pus, C. delanevi (Semple & Semple 7476 WAT).F. Li bristle-li le ( ) ifolia (Godfi 75759 WAT). . at ioe une Cis godireyi (Semple & Godfrey 3149 wal): ae C. mariana Cemple & Chmielewski 6214 WA I uter series; 0—2 primary outer bristles on other fruits SOU l. ips of primary inner bristles and one primary outer bristle (arrow). J. Linear t narrowly trian gular y 526 BRIT.ORG/SIDA 22(1) 7H, 7L); others fruits produced by the same individual have a few tapering pri- mary outer bristles (Fig. 71. This is true for fruits at all four ploidy levels 2n=8, 16, 24, 32: Semple & Chinnappa 1986). In his cladistic study, Nesom (1991a) scored the secondary outer scales of both Bradburia pilosa and C. mariana as “(1) scales”, but the latter has scales that are very narrowly triangular to only narrowly triangular on the same fruit like some other species of Chrysopsis, not like those of B. pilosa. Osbertia (x = 5; three o to brown, cylindrical-fusiform, 1.0-1.5 mm long, sparsely strigose, ribs 8-16, su- perticial (Figs. 8N, T), sometimes either the ribs or faces between them golden- brown to brown translucent; pappi quadruple, triple, double (Fig. 8M) or single and rarely vestigially triple, the series distinct or grading into each other (Fig. 8A), 20-35 bristles, barbs evenly arranged around the round shaft along the entire length of the bristles, barbs sometimes small and less obvious; second- ary outer bristles linear, 3-10, sometimes grading into secondary inner bristles, absent in O. bartlettii or very rarely one linear, fine, short bristle (Figs. 8R-S), 5- 40% the length of the inner long bristles; secondary inner bristles tapering, fine, linear, 5-10, 45-70% the length of primary inner bristles, absent in O. bartlettii; primary outer bristles fewer than and often grading into the primary inner bristles, 85-95% the length of the primary inner; primary inner bristles weakly tosometimes moderately clavate tapering (Figs. 8C, I-]), or in O. bartletii tapering and subequal or the shortest 80% if all long bristles are included ina single series, 2.5-7.5 mm long. The pappus of Osbertia differs in the three species, but in general all have rather linear and short-barbed bristles. The four series are most distinct length- wise in O. chihuahuana (Figs. H-L) and least distinct or reduced to two or one series in O. bartlettii. In the type species C. stolonifera, the secondary outer and inner series grade together, as do the primary outer and inner series, but the secondary and primary series are clearly of different lengths (Figs. A-B). The two secondary outer series could be interpreted as a single series highly vari- able in length, and the primary outer and inner could be interpreted asa single series grading from non-clavate shorter bristles to weakly clavate longer bristles. Overall there is a spiral arrangement from very short, fine, outer bristles to long coarser inner bristles, with the outer members of the series clearly placed ex- ternally to and overlapping the longer inner members of the series. Previous authors have considered the pappus of Osbertia to be simpler than described above. Turner and Sundberg (1986) described the pappus of Osbertia as having “numerous setae ina single series” 4+-6 mm long. Nesom (2000) noted that O. chihuahuana Turner & Sundberg had “a short, outer series commonly present” (i.e.,a double pappus) but did not describe the individual bristles or scales. Nesom (1991c) stated that pappus of O. bartlettii was “uniseriate, of 15- 20 barbellate bristles without an outer series.” The single short outer bristle Ss three species examined) — Cypselae straw-colored SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 527 Fic. 8. Frui i Osbertia, disc frui le bar= 1mm. A-G. 0. stolonifera (Ton 1027 TEX-LL). A. Imma- ture fruit. B. Distal portion oflonder iste. hs au A) ELLA vowen inner bristles. D-E. Tips of mid length econdary yl a G.S pectively. H— te Me chihu ag 7 sare edo 2232 TEX Distal half of ( indi possibl ).1 tal portion of primary bristles. J. a tay ier K. Tip of possibl lary bristles (center in s L. Linear sec- ondary outer bristle (arrow). M—T. 296 WAT; SEM images, Muller 2854 UC.M. Immature fruit with disc corolla attached. N. Fruit body sam oe Q. Tips of long primary bristles. R-S.Tip and base of rarely seen short outer bristle, respective 528 BRIT.ORG/SIDA 22(1 = reported above was found on one fruit of the few present on the single collec- tion of the species at WAT (Nesom 6296), but none were visible in the material from UC used for the SEM examination (Fig. 8H). The phylogenetic position of Osbertia is uncertain and the genus only ten- tatively has been included here in the Chrysopsidinae. It was included in Haplopappus sensu lato by Hall (1928), which sensu stricto is part of the Machaerantherinae Nesom. Nesom (1991c) placed Osbertia in the Chrysopsidinae, and Nesom (2000) placed it close to Chrysopsis. In the RFLP DNA study by Lane et al. (1996), Osbertia stolonifera was placed basal to the clade including Ericameria/Macronema, Xylothamia, and Tonestus but was not included in the ITS DNA sequence study by Beck et al. (2004). In the latter study, Ericameria, Xylothamia, and Tonestus were each placed in different clades sepa- rate from the clade including Heterotheca villosa and Chrysopsis gossypina. The phylogenetic position of Osbertia within the North American clade needs ad- ditional study. On the basis of pappus traits, Osbertia fits into the subtribe Chrysopsidinae as a derived lower base number taxon with a reduced number of bristles, sometimes reduced number of series, and derived non-clavate to weakly clavate longer bristles. Further study is needed to determine if such traits would exclude it from other proposed subtribes. SYSTEMATICS IMPLICATIONS There is considerable variation in the pappi traits of taxa within the Chrysopsidinae, which can not be defined in part on the basis of usually hav- ing a double pappus. Even within individual genera there is variation in the number of series, how distinct the series are from each other, and the number of long bristles per fruit. There are patterns to the variation such that a “norm” for each genus can be given, but these are not diagnostic due to with-genus varia- tion and overlapping of ranges of variation between genera. For example, the number of bristles in Noticastrum, with a quadruple pappus, is higher com- pared to other genera; the bristles in all series are linear; and the tips of the longest secondary inner bristles differ little from those of the shorter bristles in the primary outer series and secondary inner series. In Heterotheca with a usually quadruple pappus, the number of bristles averages less than Noticastrum but more than occur on the fruits of the lower base number taxa; and there are usually only a few mid-length bristles making up the secondary outer whorl. However, in H. oregona the number of bristles is within the range found in Noticastrum, but the series are generally discontinuous in length and thus do not grade into each other. In other species of Heterotheca, the bristle series grade into each other, but there are fewer bristles, e.g., H. thiniicola. In Chrysopsis with a usually triple pappus, the number of bristles and the absence of secondary inner bristles is fairly consistent; the primary inner bristles usu- ally are more obviously clavate than in most species in other genera of the SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS 529 subtribe; and fruits with relatively broad secondary outer scales have some lin- ear scales. Chrysopsis mariana with x = 4 has a double pappus or “slightly” triple pappus (1-2 primary inner bristles), while the secondary outer scales are like other species in the genus and the primary inner bristles are obviously clavate (especially when examined at 50-100H). Although pappus descriptions of the genera of the Chrysopsidinae need to be longer to encompass the details and range of variation in the series of bristles, the individual genera themselves can not be diagnosed on the basis of pappus traits alone. Therefore, the details of pappus features presented here are likely to be more significant for cladistic/ phylogenetic studies then for identifying specimens. The genera with the higher base numbers have more pappus elements in more series than do the lower base number genera with more derived and simpler pappi. Lastly, when the pappi of members of other subtribes have been examined in detail, general trends in differences of derived features may emerge. At this point, however, more data are needed. ACKNOWLEDGMENTS This research was supported by a Natural Sciences and Engineering Council of Canada Discovery Grant to J.C.S. A preliminary analysis of pappi traits was included in a cladistic study of the subtribe done by Laura Woodworth as part of a BIOL499 senior research project at the University of. Waterloo. Dwayne Estes (TENN) kindly provided data on the pappus of Pityopsis ruthii. The tech- nical assistance of Jennifer Hood and Naomi Steenhof is gratefully acknowl- edged. REFERENCES Beck, J.B.,G. Nesom, PJ. Catie, R.K. SMALL, and E.S. ScHILLING. 2004. Is Solidagininae monophyl- etic? Taxon 53:691-698. Cronaquist, A. 1955.Vascular plants of the Pacific Northwest. Part 5: Compositae. University of Washington Press. Seattle. Cronaquist, A. 1968. Aster, Chrysopsis, and Heterotheca. In: H.A. Gleason, ed. The new Britton and Brown illustrated flora of the northeastern United States and adjacent Canada. Hafner Pub. Co., New York. Cronauist, A. 1980.Vascular flora of the southeastern United States — Vol. 1 Asteraceae. The University of North Carolina Press, Chapel Hill. FerNato, M.L. 1950. Gray’s manual of botany 8" Ed. D.Van Nostrand Company, New York. Gray, A. 1884. Synoptical flora of North America. Vol. 1, part 2. vison, Blakemean, Taylor & Co., New York. Hatt, H.M. 1928. The genus fe da a phylogenetic study in the Compositae. Publ. Carnegie Inst. Wash. 389:1 Hoimaren, PK., N.H. Houmeren, and eo BARNETT. 1990. Index herbariorum. Part |. The herbaria of the world, ed. 8. Regnum Veg. 120:1-693. 530 BRIT.ORG/SIDA 22(1) Hoop, J.L.A.and J.C. Sempte. 2003. Pappus variation in Solidago (Asteraceae: Astereae). Sida 20:1617-1630. Lane, M.A.,D.R. Morcan, Y. Sux, B.B. Simpson, and R.K. Jansen. 1996. Relationships of North Ameri- can genera of Astereae, based on chloroplast DNA restriction site data. /n DJ.N. Hind (Editor-in-Chief). Proceeding of the International Compositae Conference, Kew, 1994. Vol. 1. Systematics (vol.eds DJ.N. Hind and H. Beentje). Royal Botanic Gardens, Kew. Nesom, G.L.1991a. Union of Bradburia with Chrysopsis (Asteraceae: Astereae), with a phy- logenetic hypothesis for Chrysopsis. Phytologia 71:109-121. Nesom, G.L. 1991b.Tomentaurum (Asteraceae: Astereae), a new genus of goldenaster from Chihuahua, México. Phytologia 71:128-1 Nesom, G.L. 1991c. Transfer of Heterotheca bartlettii to Osbertia (Asteraceae: Astereae). Phytologia 71:132-135. Nesom, G.L.1991d.A phylogenetic hypothesis for the goldenasters (Asteraceae: Astereae). Phytologia 71:136-151 Nesom, G.L. 1994a. Subtribal classification of the Astereae (Asteraceae). Phytologia 76: 193-274 Nesom, G.L. 1994b. Review of the taxonomy of Aster sensu lato (Asteraceae: Astereae), emphasizing the New World species. Phytologia 77:141-297. Nesom, G.L.2000. Generic conspectus of the tribe Astereae (Asteraceae) in North America, Central America, the Antilles and Hawaii. Sida, Bot. Misc. No. 20. Botanical Research Institute of Texas, Ft.Worth. Noyes, R. D. and LH. Rieseperc. 1999. ITS sequence data support a single origin for North American Astereae (Asteraceae) and reflect deep geographic divisions in Aster S.L. Amer. J. Bot. 86:398-41 2. Semete, J.C. 1981. A revision of the goldenaster genus Chrysopsis (Nutt.) Ell. nom. cons. (Compositae-Astereae). Rhodora 83:323-384. Sempte, J.C. and F.D. Bowers. 1985. A revision of the goldenaster genus Pityopsis Nutt. (Compositae: Astereae). Univ. Waterloo Biol. Ser. 29:1—34. es J.C.and C.C. CHINNaPPA. 1986. The cytogeography of Chrysopsis mariana (Compositae: reae): Survey over the range of the species. Rhodora 88:261-266. Sani J.C. and J.L.A. Hoop. 2005. Pappus variation in North American Asters. |. Double, triple and quadruple pappus in Symphyotrichum and related aster genera (Asteraceae: Astereae). Sida 21:2141-2159. Sempte, J.C. and L. Teesy. 1999. A cladistic analysis of subtribe Chrysopsidinae (Asteraceae: Astereae). XVI International Botanical Congress, August 1-7, 1999. St. Louis, MO. Ab- Stract No. 2852; Poster No. 401. Semete, J.C., C. Leeper, C. Leury, and L. Gray. 1988. Heterotheca sect. Ammodia (Compositae: Astereae): a multivariate study of H.oregona and specimens of Brewer's (golden)aster. Syst. Bot. 13:547-558 Sempte, J.C. 1996. A revision of Heterotheca sect. Phyllotheca (Nutt.) Harms (Compositae: Astereae): the prairie and montane goldenasters of North America. Univ.Waterloo Biol. Ser. No. 37:1-164 SEMPLE, CHYRSOPSIDINAE PAPPUS TRAITS Semete, J.C., 5.B. Hearo, and L. Brouitet. 2002. Cultivated and native asters of Ontario Compositae: Astereae):Aster L. (including Asteromoea Blume, Diplactis Raf.and Kalimeris (Cass.) Cass.), Callistephus Cass., Galatella Cass., Doellingeria Nees, Oclemena E.L.Greene, Eurybia (Cass.) S.F. Gray, Canadanthus Nesom, and Symphyotrichum Nees Virgulus Raf.). Univ.Waterloo Biol. Ser.41:1-134. Smith, E.B. 1965. Taxonomy of Haplopappus section Isopappus (Compositae). Rhodora 67:21 7-238. — =alt —— — including Turner, B.L. and S. Sunosera. 1986. Systematic study of Osbertia (Asteraceae-Astereae). PI. Syst. Evol. 151:229-239. ZAROINI, EM. 1985. Revision del genero Noticastrum (Compositae-Astereae). Rev. Mus. La Plata, n.s. 13.83:313-424. 532 BRIT.ORG/SIDA 22(1) Book REVIEW Bossy J. Warp. 2004. The Plant Hunter’s Garden. (ISBN 0-88192-696-5, hbk.). Timber Press Inc. 133 S.W. Second Ave, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@tim| a he com, 503-227-2878, l- 800-327-5680, 503-227-3070 fax). $39.95, 340 pp, b&w drawings, color photos, bibliography, index, 7 3/8" x 10 3/8". If you get excited at the sight of the season's first seed and plant catalogs, you should pick up a copy of the new book The Plant Hunters Cares n. Author popey Ward includes the profiles of 32 modern } day plant hunters many of the plant offerings in those catalogs. The people included are from finotighout hie oat and are associated with an assortment of nurseries, horti- cultural research associations, and/or small companies offering plants or seeds. The focus of the in- cluded plant collectors is very ae native plants, regional plants, ‘ aa plants, rock and alpine gardens, plants/seeds from inaccessible locales, unusual or hard to find species or varieties, and even collectors with very specific interests, such as bulbs, shade plants, and single genera. Each chapter focuses ne plant hunter/company. First off, the person or company is pro- filed g nee some oe ios mation on the collector's history and plant interests. Then some inl non travels and exploration for new plants is discussed. A conclusion for each includes a picture of the collector and company contact information. The bulk of each chapter concentrates on the plants collected by, introduced by, grown by and promoted by the various plant hunters. The chapters are filled with colorful photos of oa as well as some origin descriptions and growing zone boy atien for various plants. Throughout the chapters are brief stories about how some plants ] how the person met someone who introduced or passed along the plant to them. The descriptions of the discovery of these plants were a little less adventurous than this reader ex- pected, but these discoveries are still interesting to read about. An example of sucha discovery is that of the small, white flowered, bronze leaved plant Jabrosa volkmannii by John Watson and Anita Flores, found growing in pumice on the side a volcano. There is also interesting information on how these hunters cultivate their finds. For instance the book ue a few “rock gardeners” growing species in various rock crevices including Panayoti Kelaidis of the Denver Botanical Gardens who designed and manages a 450,000 kg, 0.4 ha rock garden with 4000+ plant species. The book concludes with a chapter on invasive plants that gives an honest look at w > a] at weeds and invasive species are and the variety of vectors of ial AnEFOGEC ion. If or | li ho thrill i icipati f new species and vari- eties of plants, on will thoroughly enjoy reading this Geek and will benefit [om aie contact cs mation supplied for each collector. Travel with these plant I naeult of discovery, excitement of their plant fin vithin the pages of Ward’s The Plant Hunter’s Garden.—Lee eon Herbarium, figvantcat Re nee Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A SIDA 22(1): 532. 2006 NOTES ON TYPES IN APOCYNACEA EAASCEEPIADOIDEAE IN CUBAN HERBARIA AND FOUR LECTOTYPIFICATIONS IN WEST INDIAN GONOLOBINAE Alexander Krings Paul R. Fantz sie elt Department of Botany Department of Horticultural Science orth Carolina State University North Carolina State University Ralei a North Carolina 27695-7612, U.S.A. Raleigh, North Carolina 27695-7609, U.S.A. Alexander_Krings@ncsu.edu Paul_Fantz@ncsu.edu ABSTRACT Four asclepiadaceous types not listed in a previ f the collections of the Instituto de Egologia) vy SIStetnanA, Elabaiia< Cuba (H AC) are prov rided, alone witha giscussion of typuication t I 1 Charles V 1] [ for Gonolobus stephanotrichus Griseb., Orthosia acuminata Griseb., Orthosia oblongata coy and b Poicilla ovatifolia Grise RESUMEN a, t ] A ] ] 4 ] ] =| : -j : ae | ] oO C L de igen de Feit Sse a, Habana, Cuba (HAC), junt di ion de] bl tipificacion asociados con las Colecciones Charles Wright. 7 eapouan lectotipificaciones de ibn stephanotrichus ee 2 iseb,, y Poicilla ovatifolia Griseb. A revision in progress by Krings of subtribe Gonolobinae (Apocynaceae: Asclepiadoideae) in the West Indies has led to the discovery of four additional types at the Instituto de Ecologia y Sistematica, Habana, Cuba (HAC). These specimens were not listed ina previous catalogue of types at HAC (Cardenas & Herrera 1991). Types in subtribe Gonolobinae were not found at the Jardin Botanico Nacional, Habana (HAJB; for a complete list see Gutiérrez et al. 1997). The expanded list of Apocynaceae: Asclepiadoideae types at HAC is presented in Table 1. The asclepiadaceous original material at HAC belongs to two groups: (1) species with unproblematic typification due to designations by the original author(s) and (2) species requiring lectotypification due to complications in collecting practice and lack of holotype designations. Typification is unproblematic for Marsdenia micrantha Alain, Matelea alainii Woodson, and Marsdenia bicolor Britton & P. Wilson, as these species were described relatively recently and as each original author designated holo- types. Thus, corresponding material at HAC (Table 1), was identified easily as either holotype, isotype, or paratype based on the respective protologues. An SIDA 22(1): 533 — 537. 2006 = 534 BRIT.ORG/SIDA 22(1) Taet E 1.List of Apocynaceae: Asclepiadoideae types at the Instituto de Ecologia y Sistematica, Habana, Cuba (HAC). Previously unlisted types (Cardenas & Herrera 1991) are marked by an asterisk. (1) Species with unproblematic typification “Marsdenia bicolor Britton & P. Wilson, Bulletin of the Tor bok Botanical Club 50:47. 1923. Type: CUBA: Léon 10787 (isotype: not seen); Léon 10788 (paral SS Marsdenia micrantha Alain, Revista de la Sociedad Cubana de Botanica 13:59. 1956, Type: CUBA: Al ain, Acuna & Lopez 5678 (HOLOTYPE; IsoTyPE: NOt seen) a aia inii San ee Contribuciones Ocasionales del Museo de Historia Natural del Colegio Salle’ 4.1956 ener 5029 (8 TYPE!) (2) Species lectotypified herein (see text) sae stephanotrichus Griseb., Catalogus plantarum cubensium 177.1866. E: CUBA: Wright 2969 (syntyee!) *Orthosia acuminata Griseb., Catalogus plantarum cubensium 175, 1866. Tyee: CUBA: an (ISOLECTOTYPE!) hs ‘thosia ot Griseb., Catalogus plantarum cubensium 176. 1866. Type: CUBA: Wi ight 2967 (sc LECTOTYPE!) *Poicilla ovatifolia Griseb., Catalogus plantarum cubensium 177.1866. Type: CUBA: Wright 2965 (syntyee!) isotype of Marsdenia bicolor was reported previously by Cardenas and Her- rera (1991), but not seen as part of this study. A paratype (Léon 10788) at HAC was not listed by Cardenas and Herrera (1991). Grisebach (1866) described a number of taxa based on material collected by Charles Wright in the Antilles. These included the following four species for which original material was found at HAC and for which lectotypification is necessary: Gonolobus stephanotrichus Griseb., Orthosia acuminata Griseb., Orthosia oblongata Griseb., and Poicilla ovatifolia Griseb. (Table 1). Grisebach studied Wright material distributed to him by Asa Gray (GH). However, these speci were not necessarily true duplicates of a single gath- ering. Indeed, there have been instances of mixed material joined by a single number, as Gray appears to have provided the same number to specimens Wright may have collected from different locations on different dates (Howard 1986, 1988). Thus, Howard (1986) stated [brackets ours]: “The determination of the type collection, therefore, depends on examin- ing the sheet Grisebach saw and named, which is preserved in Géttingen [GOET, Universitat Gottingen]. This should match one of the fragments preserved by Gray. The GOET specimens however, rarely have field tickets, so the date and sored of the type collection can only be determined, if at all, from the GH she oe (1988) noted that Grisebach likely did not see the Wright collec- KRINGS AND FANTZ, 535 tions sent to the Sauvalle herbarium in Habana (currently deposited in HAC), as Asa Gray distributed this material from GH. In contrast to Howard (1988), however, it should be noted that the fact that Grisebach may not have seen the Wright specimens in the Sauvalle herbarium, has no bearing on their status as original material and availability for lectotypification in the future. If judged only by this fact, according to Div. 2, Ch.2, Sect. 2, Art. 9 of the International Code of Botanical Nomenclature or ICBN (Greuter et al. 2000), they would none- theless be considered part of the original material and, as duplicates of one cited collection, would be syntypes as articulated by Fantz (1993). Gonolobus stephanotrichus.—Two specimens of Wright 2969 reside at GOETI. No field tickets accompany the GOET specimens. Both sheets include flowers and one also includes a fruit. Fruits are not described in Grisebach’s protologue. Duplicates of this number are found at BREM!, G!, GH!, HAC), K!, NY!, P!, UC), US!, and reported for MO, although this specimen could not be ob- tained on loan. The collection labels of the sheets at GOET are blue and bear the dates 1856-7 crossed out. No additional dates are given. The collection la- bels for duplicate numbers housed everywhere else, but HAC, are white and bear the dates 1860-64 (including the MO specimen, an image of which could be seen from the Missouri Botanical Garden website: http://www.mobot.org). The collection label for the HAC specimen is white and bears the date 1865. This specimen is sterile and quite poor due to insect damage. Field tickets re- maining with the sheet at GH seem to indicate that the three mounted frag- ments (all in flower) were collected at different times Jan., Mar, Apr.). However, it isimpossible to tell which fragment is associated with which field ticket and furthermore, none can be correlated with the GOET sheets, as these lack field tickets altogether. All material under the number 2969 (incl. the duplicate at HAC) does appear to belong to the same species. Thus, in light of the available facts, the Wright 2969 sheet at GOET that bears flowers, but lacks fruit, is herein designated lectotype for Gonolobus stephanotrichus Griseb. This action agrees witha 1984 annotation by R.A. Howard whose lectotypification (1988) appeared ina microfiche appendix, not in print, and thus is not effectively published and has no standing in nomenclature. Rankin @ Greuter (2000) reported a similar case in Antillean Aristolochia. The remaining extant sheets distributed under Wright 2969 retain their status as syntypes. An additional fruiting specimen of Gonolobus stephanotrichus is housed at S, bearing a Wright label of 1861 and the preliminary number 164. Although a determination on the label is provided in Grisebach’s hand, we do not consider the specimen original material as fruits were not mentioned in the protologue. Orthosia.—The only known type material for O. acuminata (Wright 2966) is deposited at BM, G, GH, HAC (2 sheets), and K. These specimens bear white labels with the dates 1860-64. The mounted field ticket on the GH sheet reads: 536 BRIT.ORG/SIDA 22(1) “Asclepias—Fl. (except the white stigma) green. Farallones San Andre Oct 27.” As Wright 2966 (GH) contains fifteen inflorescences and is in very good condi- tion; itis here designated as the lectotype for Orthosia acuminata Griseb. Wright 2966 (G!) contains four inflorescences and is in superior condition to the HAC material. Wright 2966 (BMI, G!, HAC!, K!) should be considered isolectotypes. Original material of Orthosia oblongata was located at BM!, G!, GH!, HAC, K!, and S!, and reported for MO, although the specimen could not be obtained on loan. These specimens all bear white labels with the dates 1860-64 (includ- ing the MO specimen, an image of which could be seen from the Missouri Bo- tanical Garden website). The mounted field ticket of Wright 2967 (GH) reads: “Asclepias—Fl. green—a white speck at the tips of the segments. Stigma white. Loma de Ranjel June 17.” Field tickets do not accompany the other specimens. Wright 2967 (GH) is herein designated lectotype for Orthosia oblongata Griseb., considering the duplicates at G, HAC, K, MO (provided that the specimen does not constitute another species), and S isolectotypes. Wright 2967 (GH) is in good condition, with numerous inflorescences. Lectotypification decisions for both Orthosia names were based on the in- terpretation that sheets of both Wright 2966 & 2967 are original material, as they were respectively cited in the according protologues. There is no evidence that they were collected from different localities, although this cannot be ruled out altogether, given the notorious problems with Wright collections. However, in our opinion, when there is no specific evidence to the contrary, if collection number and identification match, the protocol ought to hold that the speci- mens be considered duplicates of a single gathering. We recognize that “isolectotype” is not an ICBN type designation, but assign it herein should lec- totypes become lost or destroyed or additional syntypes are located that we have not examined. Poicilla ovatifolia.—Syntype material (Wright 2965) was located at BMI, BREMI, G!, GH!, GOET!, HAC), K, NY!, P!, S!. and UC! No field tickets accompany the GOET specimen or any syntype, except the GH specimen. Accompanying field tickets of the GH specimen suggest that the sheet is comprised of at least two collections made at different times (Mar, June), although three fragments are mounted. The two fragments mounted on the right contain inflorescences; the fragment mounted on the left contains infructescences. Both field tickets refer to flowers with neither one mentioning fruits. Fruits are not described in Grisebach’s protologue and are not present on any other syntype beside the GH specimen. Except for the GOET specimen, collection labels of all other known specimens are white and bear the dates 1860-64. The GOET specimen bears a tan label with a printed date of 1860, although the zero appears to have been crossed out. It is heavily written on in Grisebach’s hand and is herein desig- nated lectotype for Poicilla ovatifolia Griseb. The studied (and matching) duplicately-numbered material in other herbaria remain syntypes. KRINGS AND FANTZ, 537 ACKNOWLEDGMENTS We thank the curators and staff of the following herbaria for searching, or pro- viding access or loans of their collections: B, BG, BH, BKL, BM, BOLO, BR, BREM, BsG RUE GC, CGE, COLO, CReDUKE 2 ETL PEAS ER PIG .G GHaGOEr1,-H, HAC, HAJB, HBG, IA, JJ, ISC, JBSD, JE, K, LD, M, MICH, MIN, MO, MSU, NEU, NSW, NY, O, OXE P, PH, RSA, S, U, UC, UPS, US, USF TUR, WILLI, WU, Z. AK also thanks Eldis Bécquer and Lutgarda Gonzalez for stimulating conversation regarding typitication and extends grateful appreciation for the kind hospital- ity afforded by the Jardin Botanico Nacional, Habana, and especially Fabiola Areces and Julio Lazcano. Thoughtful reviews of a previous version of the manu- script were provided by Carlos Sanchez and Alain Loigier. REFERENCES Cardenas Atvarez, A.and P. Herrera Ouiver. 1991. Catdlogo de las colecciones histdricas y de tipos del Herbario de la Academia de Ciencias de Cuba. Editorial Academia, La Habana. FANTZ, PR. 1993. Notes on Clitoria (Leguminosae) in Southeast Asia. Novon 3:352-355. GreuTer, W., J. McNelt, FR. Barrie, H.-M. Buroet, V. DEMOULIN, TS. Fitcueiras, DH. Nicotson, P.-C. Sitva, J.E. Skoe, P. TREHANE, N.J. TuRLAND, and D.L. Hawkswortu. 2000. International code of botani- cal nomenclature (St Louis Code).Regnum Veg. 138. Koeltz Scientific Books, Konigstein GriseBach, A.H.R. 1866. Catalogus plantarum cubensium. Wilhelm Engelmann, Leipzig. Gutiérrez Amaro, J., |. ARIAS GRANDA, H, Manitz,and E. ALGuiLar CeLpa. 1997.Los tipos del herbario “Prof. Dr. Johannes Bisse” del Jardin Botanico Nacional (HAJB): |. Revista Jard. Bot. Nac. 18:21-50. Howard, R.A. 1986. Notes on Quiina (Quiinaceae) and Ilex species (Aquifoliaceae) in Cuba. Brittonia 38:13-16. Howarp, R.A. 1988. Charles Wright in Cuba, 1856-1867. Chadwick-Healy, Alexandria Rankin Rooricuez, R.and W.Greuter. 2000. Notes on Aristolochia linearifolia and A. stenophylla (Aristolochiaceae), a vicarious species pair from the Greater Antilles (Cuba and Hispaniola). Willdenowia 30:131-139. 538 BRIT.ORG/SIDA 22(1) Book REVIEW Joun Dawson and Ros Lucas. 2005. The Nature of Plants: Habitats, Challenges & Adaptations. (ISBN 0-88192-675-2, hbk.). Timber Press Inc. 133S.W. Second Ave, Suite 450, Portland, OR 97204-3527, US.A. (Orders: wwwtimberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $39.95, 314 pp., 205 color photos, 5 color drawings, glossary, and index, 7.38" x 10.38". The new book called The Nature of Plants is a plant lover's dream. Have you ever wondered how it’s possible for any plants grow in the tundra? Or how underwater plants get their flowers pollinated? The authors have provided readers with a wonderful resource to answer these and many of other questions about plants surviving in extreme environments. ue textis peppered with rich color pho- de tographic examples of the plants and | | in the tex In each chapter, the reader is introduced to unique environments, conditions, and climates; and then told about all the special challenges faced by plants that live there. For example, we're told about parasitic plants, desert plants (both hot and cold), fire effects, salt ene plants on windy terrain an interactions with animals, fungal and even bacterial interactions between the plants. e authors provide a good informational background for the reader regarding the environ- mental i climatic situations of the plants. For example, in one chapter titled “Not Enough Water” = : the authors include an explanation of why deserts even exist and where they're located. This book covers a lot more than just North America when they go into detail on the deserts of the world in- ee temperate deserts and the special adaptations that plants make for each environment. eading about the pee adaptations | to these challenging environments makes this book par- aie interesting. After a | lescript { climates or special conditions, the author includes ormation about common plant surv ‘val strategies. For example, the chapter about fire shows how in some plants use fire as a regeneration process; after being damaged, they release seeds or flower. Very often, the strategies are given with specific plant examples along with beautiful color photographs that help the reader visualize such ceaned fel sicaseas These images are mace wondertul because many of the images shov nts i tic locations. In addition, one chapter focuses on the various methods of reproduction ne the different vectors for pollen transfer. This sn included a wonderlul discussion on the benefits and problems of self{-pollination and out-crossing in plants. The book The Nature of Plants: Habitats, Challe nig s & Adaptations is recommended for anyone with an interest in plants and their ability t urvi he harshest of habitats. This book does not require much botanical background to enjoy, although an understanding of sane family traits will be helpful. A glossary is included for the less common terminology. All topics are explained with diverse examples and fantastic color photographs. Read The Nature of P “e Habitats, Chal- lenges & Adaptations and take a peek into the fascinating survival strategies of plants in extreme situations and environmental conditions.—Lee Luckeydoo, Herbarium, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A SIDA 22(1): 538, 2006 COMMENTS ON THE GERBERA-COMPLEX (ASTERACEAE: MUTISIEAE) Hans V. Hansen Botanic Garden Ae 2 of Copenhagen er Farimagsgade Dk . fee oes K, DENMARK ABSTRACT The recent transfer of Chaptalia hintonii to Gerbera is doubted, as is any transfer of New World members to the Old World part of the Gerbera-complex. On a biogeographic basis it seems more probable that Gerbera is an Old World clade, while Chaptalia, Trichocline, and T. hieracioides(some- times considered to be a Ger per are New Word groups. The geographic distribution of Leibnitzia rth America, Asia) and Aml lata (W. Australia) is enigmatic. Additional input of ee is needed for a full Spaces of the phylogeny in the Gerbera-complex RESUMEN f. a Py cal | a baa aad: es Pees ] Ur ansfe rencia de elementos del BEKO Mundo ala Datte del compl Gerbera del Viejo Mando: En base a la ] biogeografia l Ae: Mune mientras que og eae evi L t Thies ioide ( | J ipachogene Mundo. La distribucion geogrfica de Leibnitzia (Norte iva ee. Asia) y Ambbsper ma a (Oeste de A tica. Se necesitan datos adicionales para una completa comprension b de la nee del somiplejs Gerbera. In recent papers, Katinas (1998, 2004b) and Nesom (2004a,b) discussed the ge- neric position of Chaptalia hintonii Bullock within the Gerbera-complex (the scapose complex of Mutisieae subtribe Mutisiinae sensu Cabrera 1977). Their discussion continues the long-standing debate regarding OTUs within this com- plex. To recapitulate briefly, Jeffrey 967) used LM to point out characters of taxonomic value in the group, mainly pertaining to cypselar pubescence and pappus-hairs. When [ took up similar studies (e.g., Hansen 1985, 1990) SEM had been introduced, but my conclusions deviated little from those of Jeffrey. Nesom (1983) contributed with his first study of the Gerbera-complex (American Leibnitzia) and next focused on Chaptalia Vent. (Nesom 1984a,b, 1995, 2004a,b; Cabrera & Nesom 2003), while other workers have also published studies dealing with the scapose group (Katinas 1998, 2004a,b; Sancho & Katinas 2002; Moraes 1998; Hind 1999, 2001; Roque 2005). All these studies, with their general access to living material, have amplified our knowledge about the Gerbera-complex. The complex includes about 100 species, and evidently it takes subtle char- SIDA 22(1): 539 — 543. 2006 540 BRIT.ORG/SIDA 22(1) acters to delimit monophyletic groups within it. This is noteworthy in view of its wide distribution: Trichocline Cass. Lulia Zardini, and Chaptalia Vent. in the New World, Leibnitzia Cass. in North America and Asia, Gerbera L. in Africa, Madagascar, and Asia (i.e., Gerbera sensu Hansen 1990), Perdicium L. in W. Cape, Uechtritzia Freyn in Asia,and Amblysperma spathulata (A.Cunn.ex DC.) DJ.N. Hind in W. Australia. It has been disputed whether Amblysperma falls within the limits of Trichocline (Hind 2001, and implicitly Hansen 1990, contra Zardini 1975 and Katinas 2004a), and whether the transfer of Trichocline hieracioides (Kunth) Ferreyra (Ecuador, Peru) to Gerbera by Zardini (1974) is correct. The most significant problem, however, is with Chaptalia, for which no complete revisionary treatment exists; Burkart (1944) only considered part of the genus in detail. Nesom (2004a) and Katinas (2004b) remarked that while Jeffrey would split the complex into smaller genera, | suggested the recognition of one large genus (hence Gerbera). This calls for an explanation. The relatively ancestral position within Asteraceae of Mutisieae (however circumscribed, but at least excluding subfamily Barnadesioideae (Benth. G Hook.f.) K. Bremer & R.K. Jansen) is now supported by morphologic and molecular evidence. Of relevance here is that part of the classic Mutisiinae in all studies is indicated to be mono- phyletic, with the Gerbera-complex placed in a relatively advanced position. Actually, the Brazilian monotypic genus Lulia (Zardini 1980) (ie., Trichocline nervosa Less.), by its monocephalous, albeit non-scapose habit, stands between the Gerbera-complex and its sister (a group including Mutisia Lf, Chaetanthera Ruiz & Pav, Duidaea S.F Blake, and Pachylaena D. Don in Kim et al. 2002; Lulia was not considered). In Hansen (1991), | anticipated these aspects and then reasoned that if Mutisia and Chaetanthera are not split into minor taxonomic units, it would not be logical to split the Gerbera-complex either, hence all species should fall within Gerbera. This explains my way of reasoning which was not versus Jef- [rey (1967), but simply an alternative provided by cladistic reasoning. Katinas (2004b:938-939) presented a key to the scapose group with seven genera (including Lulia, but with Amblysperma sunk into Trichocline and awaiting a complete revision of Chaptalia). In this key there were autapomorphies for each genus (if we accept that Trichocline hasa special type of cypsela hairs). However, there is a complication in Katinas’s key couplet 6, since Gerbera p.p. (sects. Gerbera, Parva, and Isanthus) do not have the apomorphic state ‘trimorphic rays. This is precisely where the problems with Gerbera (in its current sense) arise. Nesom (2004a, cl. Nesom 1995; Cabrera & Nesom 2003) has maintained that Chaptalia hintonii belongs to Chaptalia sect. Chaptalia. The traits which led Katinas to move it to Gerbera (all florets bilabiate, outer ray florets long, ray florets with staminodes) can be found in various Chaptalia, and the cypsela HANSEN, COMMENTS ON THE GERBERA-COMPLEX 541 hairs in C. hintonii do not deviate in morphology from those in Chaptalia sect. Chaptalia p.p. (the C. lyratifolia group). I believe that Nesom has the strongest point, namely that the ‘deviating traits in C. hintonii do not disrupt its alliance with Chaptalia sect. Chaptalia. Rather, C. hintonii possesses states which are parallelisms/reversals to those in Gerbera. As Nesom indirectly argues, it seems more probable on a biogeographic basis that Gerbera is an Old World clade, while Chaptalia, Trichocline, and the enigmatic T. hieracioides are New World groups (not overlooking problems with Amblysperma in W. Australia and Leibnitzia, which is divided between Asia and North America). Evidently, various authors during their close survey of taxa detect excep- tions to the currently used taxonomy and thus find reason to allocate this or that species. But the question is whether we push the case too far. We know more today than we did in Hansen (1990) and hardly disagree on the polariza- tion of states. Hence, a matrix with all relevant placeholders in the Gerbera- complex treated as OTUs ultimately should be prepared, because only then it will be revealed, if the addition of new characters can disrupt the unity of (some of) the currently recognized taxonomic groups. Presumably, however, the con- sensus tree will still be burdened with polytomies, since the number of known synapomorphies is so small. Additional input of data is therefore much wel- come. However, along with the detection of amplified variation in ray floret morphology, cypselar vestiture, etc. the selection of relevant characters may cause more and more debate. A good example is the presence of trichomes on corollas in 13 species of Trichocline, three of Onoseris, and one investigated spe- cies of Uechtritzia (Sancho & Katinas 2002). Is this information of phyloge- netic relevance or not? Nesom (2004a:932) has now suggested that Chaptalia sects. Lieberkuhna (Cass.) Burkart and Loxodon (Cass.) Burkart possibly may be separated at ge- neric rank, among other things due to their distinctive cypselar vestiture. Katinas (2004b) likewise predicted taxonomic alterations at generic rank. The discussion between the two authors reveals considerable disagreement, both with respect to observation of characters (e.g., whether ray flower staminodes are present or not) and how to interpret their phylogenetic relevance. | must stay neutral, since I never studied Chaptalia in detail, but clearly the main is- sue is to decide whether Chaptalia is monophyletic (even if split into two gen- era), with evolution of character states parallel to those in Gerbera, or whether some species of Chaptalia really should be moved to Gerbera or even some- thing else. ? ACKNOWLEDGMENTS I thank Guy Nesom and an anonymous reviewer for their most insightful com- ments which improved much on this paper. 542 BRIT.ORG/SIDA 22(1) REFERENCES Burkart, A. 1944. Estudio del género de Compuestas Chaptalia con especial referencia a las especies argentinas. Darwiniana 6:505-594, Caprera, A.L.1977.Mutisieae-systematic review.|n: Heywood,V.H.,J. Harborne, and B.L.Turner, eds. The biology and chemistry of the Compositae I]. Academic Press, London. Pp. 1039-1066. Caprera, L.and G.L. Nesom. 2003. A new species of Chaptalia (Asteraceae: Mutisieae) from Mexico and rediscovery of Chaptalia mexicana. Sida 20:1363-1369. Hansen, H.V. 1985. A taxonomic revision of the genus Gerbera (Compositae-Mutisieae) sections Gerbera, Parva, Piloselloides (in Africa), and Lasiopus. Opera Bot. 78:1-36 Hansen, H.V. 1990. Phylogenetic studies in the Gerbera-complex (Compositae, tribe Mutisieae, subtribe Mutisiinae). Nordic J. Bot. 9:469-485. Hansen, H.V. 1991. Phylogenetic studies in Compositae tribe Mutisieae. Opera Bot. 109: Hino, DJ.N. 1999. Notes on Chaptalia (Compositae: Mutisieae) in Brazil. Kew Bull. 54: 933-939, Hino, D.J.N.2001.A new combination in Amblysperma (Compositae: Mutisieae). Kew Bull. 56:711-713. JerFrey, C. 1967. Notes on Compositae: II. The Mutisieae in east tropical Africa. Kew Bull. 21:177-233. Karinas, L. 1998. The Mexican Chaptalia hintonii is a Gerbera (Asteraceae, Mutisieae). Novon 8:380-385. Karinas, L.2004a.Amblysperma should be retained under Trichocline (Asteraceae, Mutisieae) Taxon 53:108-112. Katinas, L. 2004b. The Gerbera complex (Asteraceae: Mutisieae): to split or not to split. Sida 21:935-940. Kim, H.G., D.J. LoockerMAN, and R.K. Jansen. 2002. Systematic implications of ndhF sequence variation in the Mutisieae (Asteraceae). Syst.Bot. 27:598-609. Moraes, M.D. be. 1998. Chaptalia hermogenis (Asteraceae: Mutisieae), a new species from the Brazilian Atlantic Rain Forest. Novon 8:173-175. Nesom, G.L. 1983. Biology and taxonomy of American Leibnitzia (Asteraceae, Mutisieae). Brittonia 35:126-139. Nesom, G.L. 1984a. A new, widespread species of Chaptalia (Asteraceae: Mutisieae) from Mexico. Rhodora 86:127-130, Nesom, G.L. 1984b. Taxonomy and distribution of Chaptalia dentata and C. albicans (Asteraceae: Mutisieae). Brittonia 36:396-401. Nesom, G.L. 1995. Revision of Chaptalia (Asteraceae: Mutisieae) from North America and continental Central America. Phytologia 78:153-188. Nesom, G.L. 2004a. Generic placement of Chaptalia hintonii (Asteraceae: Mutisieae). Sida 21:929-933. Nesom, G.L. 2004b. Response to “The Gerbera complex (Asteraceae: Mutisieae): to split or not to split” by Liliana Katinas. Sida 21:941-942. — HANSEN, COMMENTS ON THE GERBERA-COMPLEX 543 Roque, N. 2005. A new species of Chaptalia (Compositae, Mutisieae) from Minas Gerais, Brazil. Kew Bull.60:133-135. SancHo, G. and L. Katinas, 2002. Are the trichomes in corollas of Mutisieae (Asteraceae) really twin hairs? Bot. J. Linn. Soc. 140:427-433. ZaARDINI, EM. 1974. Sobre la presencia del género Gerbera en America. Bol.Soc. Argent. Bot. 16:103-108. ZARDINI, E.M. 1975. Revisi6n del género Trichocline (Compositae). Darwiniana 19:618-733. ZARDINI, E.M. 1980. Lulia - un nuevo genero de Compuestas. Bol. Soc. Argent. Bot. 19: 255-258. BRIT.ORG/SIDA 22(1) Book REVIEW He_en Heipert. 2000. Papermaking with Garden Plants and Common Weeds. (Paperback edition). (SBN 1-58017-622-4, pbk.). Storey Publishing, 210 MASS MoCA Way, North Adams, MA, 01247, U.S.A. (Orders: # 67622, 1-800-441- 5700, www.storey.com ). $14.95, 112 pp., color photos, drawings, appendix, glossary, resource guide, index, 91/4" x 10" Papermaking With Garden Plants And Common Weeds isa good resource for crafters and profession- als interested in making their own paper from natural materials. Author Helen Heibert has put to- gether a wonderful reference with ee instructions on three methods of papermaking with a variety of natural materials. These instructions include practical advice on keeping records of re- sults with different materials, detailed fiber processing instructions, and sidebars of personal expe- riences and successes of the author. he book begins with preliminary information on responsible and safe plant fiber and mate- rial collecting. This eee information is followed by descriptions of the types of possible fibers: bast, leaves, grass and ees materials oe uae ad and recycle fibers. Next she pre- f king four mould iia cee a tep-by- sents the with the different ee techniques. a the different alkali recipes, she presents ssing instructions covering cooking, beating, retting, and safety information. In a S Wit step proce to that, she includes information on the additives for protecting and coloring fiber materia > =) artificial and natural dyes. Along with the Co instructions, the author has ae included great color images that highlight the step-by | the three Western style with a mould and deckle, Japanese paper eels with a sugeta and formation aid (recipes included), and t There is a helpful troubleshooting section for potential sian os e Ne- palese method using a deckle bo that may be encountered Pee the paper making process. Multiple pressing techniques are also presented with color images and include instructions on how to make your own paper press as well as wet paper handling suggestions and drying technique e author has included prone with instructions ol ten different artists who used ten differ- .A small personal story box is included with ent plant materials and pa each artist's project agattionahy, seven step- ry oe projects are presented including: flower petal envelope, lampshade, and accordion book within paper, | ealowing the Section on projects is important iaigr mation including lists of papermaking plants, with common and Latin names, glossary, reading list, and a resource guide Author Helen Heibert has provided adventurous crafters with well-guided instructions on paper- making techniques and projects. The et are recommended for adults and, due to the hemical king involved, children with dervision. The paper types and projects sug- gested and included in the book Papermaking with Garden Plants and Common Weeds are promising for scra BOSanE a making, craft groups, interior design, education, youth projects and man 8 8 y _uckeydoo, Herbarium, Botanical Research Institute of Texas, 509 Pe- othe can cee Fort Worth, TX 161 “ 4060, U.S.A. SIDA 22(1): 544, 2006 VALIDATION OF THE NAME DAHLIA SUBLIGNOSA (ASTERACEAE) Dayle E. Saar Paul D.S@rensen Department of Biological Sciences Department of Biological Sciences Murray State University Northern Illinois University Aurray, Kentucky 42071, U.S.A Dekalb, Illinois 60115, U.S.A. Article 33.3 of the International Code of Botanical Nomenclature (ICBN) (St. Louis Code 2000) states that a new combination based ona previously and val- idly published name is not itself validly published unless its basionym is clearly indicated and fully referenced. While the original work was cited regarding the new combination of Dahlia dissecta S. Watson var. sublignosa P.D. Sorensen to Dahlia sublignosa (PD. Sorensen) D.E. Saar & PD. Sorensen, the basionym was not “clearly identified” (Saar & Sorensen 2005). Therefore, the combination is validated here in accordance with the ICBN. Dahlia sublignosa (PD. Sorensen) D.E. Saar & PD. Sorensen, comb. & stat. nov. BASIONYM: Dahlia dissecta var. sublignosa PD. Sorensen, Rhodora 71:35]. 1969. TyPE: MEXICO. Tamaulipas: 4.8 km N of Miquihuana in forest dominated by Pinus (99°47' N Lat; 23°36' W Long.), elev. ca. 2100 m, 14 Jul 1949, Stanford, Taylor, & Lauber 2436 (HOLOTYPE: NY; ISOTYPES: GH, MICH, TEX, UC, US-2, WTU). Asan explanatory note, the holotype specimen collected by Stanford, Taylor, & Lauber 2436 cited in our paper is a correction of the original basionym, which was incorrectly stated as collected by Stanford, Rutherford, and Northcroft 2436. ACKNOWLEDGMENTS We thank Teri Bilsborrow for bringing this oversight to out attention and Kanchi Gandhi for helpful comments regarding this validation notice. REFERENCES Saar, D.E.and PD. Sorensen. 2005. Dahlia subli pecies in its own right. Sida 21:2161- 2167. SIDA 22(1): 545. 2006 546 BRIT.ORG/SIDA 22 — 1) Book REVIEW FREDERICK R. TROFH and Louis M. THompson. 2005. Soils and Soil Fertility. 6th edition. (ISBN 0-8138-0995-X, hbk.). Blackwell Publishing. 2121 State Ave., Ames, TA 50014-8300, U.S.A. and 9600 Garsington Road, Oxford, OX4 2DQ, UK. one. 515-292-0140, 515-292-3348 fax 1-800-862-6657, www.blackwell » Vix | (7) LI] ] lly 1 ae lee Pee g.com). $89.95, 498 pp., 231 ‘iiietaciona. 7 210" understood lan- Soil and Soil Fertility is a well-illustrated book that presents soil t guage. It would be suitable as a textbook for classes on general soils or soil fertility. Additionally, this title could also be a beneficial reference text for classes in agronomy, hydrology and agricultural engineering. Within the preface, the authors mention that this book has been “designed for use as a textbook for the introductory course in soils for students in agriculture and related sciences.” Chapter topics include 1) Soil; 2) Soil Formation; 3) Physical Properties of Soils; 4) Soil Mechanics; 5) Soil Water; 6) Soil Organic Matter; 7) Soil Mineralogy; 8) Soil Chemistry; 9) Amending the Soil; 10) Fertilizers; 11) Nitrogen; 12) Phosphorus; 13) Potassium; 14) Calcium, Magnesium, and Sulfur; 15) The Micronutri- ents, 16) Variations in Plant Composition; 17) Soil Classification; 18) Land Use and Soil Manage- ment; 19) Water Management, 20) Soil Erosion and Its Control; and 21) Soil Pollution. The order of chapters How well into one ae pune on previous chapters’ materials, starting with soils basics and mineralogy, d g and fertility and ending with environmental topics and land stewandehip The text is very readable and is accompanied by many supportive illustrations, cule and charts. These diagrams and een on Lae ize ee ene white wee mans ne other e easily dis- tinguishable designs. TI I lly offer an example problem with éeplonation: Basic soil tests/analyses are iced hg the text in- — cluding items such as particle size analysis using Stoke’s law, a hydrometer, and a RUSLE soil loss equation example. At the end of each chapter is an im et notes section, and a ” of questions that if answered, would demonstrate an understanding of the text In addition to all of the discussion of general soils information, this book other pn tant souls Subiccts:> soil cea and soil classification. The section on soil rie isa ehel il fertility fllowee Hace daunaaors nou s in plants, the ] Jp] einr ee micronutrients. These chapters include chemical use and deficiency symp nutrient cycle and availability, fertilizer types for that nutrient, application info “e more. The chap- ter on soil classification includes a number of different systems with background and explanation on each system, additionally covering land use and land practices to reduce soil loss. In the end, there isa measurement and con Nees summary as well as a glossary. Soil and Soil Fertility by Troeh and Thompson would be an excellent textbook for soils and related classes. oS isa —— intelligent book os l of s sup por tive black and w pied illustrations within xt. Althou Sa the : gh g textbook g ng this text for =e mation and a better understanding of soil properties and soil nese ee oa: keydoo, Her- SIDA 22(1): 546. 2006 A NEW COMBINATION IN POLYTRICHASTRUM (BRYOPHYTA: POLYTRICHACEAE) G.L.Smith Merrill Department of Botany The Field Museum 1400 South Lake Shore Drive res ey linois 60655-2496, U.S.A. ABSTRACT (NA : \ The new combination Polytrichastrum alpinum var. sylvaticum L. Merrill is provided. RESUMEN L Se aporta la nueva combinacion Polytrichastrum alpinum var. sylvaticum (Menzies) GL. Merrill. The following transfer is proposed for use in a treatment of the moss family Polytrichaceae in the Flora of North America North of Mexico, vol. 27, the first of the projected three bryophyte volumes. Additional discussion will be found in the flora treatment. = Polytrichastrum alpinum var. sylvaticum (Menzies) G.L. Merrill, comb. nov. BASIONYM: Polytrichum sylvaticum Menzies, Bot. Zeit. (Regensburg) 1:74. 1802. TyPE: U.S.A: Northwest Coast of North America, Menzies (HOLOTYPE: E!). Pogonatum alpinum var. aye Scien Lindberg, Bot. Not. 1867: 130. 1867. Polytrichum alpinum var. sylvaticun ndberg, Acta Soc. Sci. Fenn. 10:6. 1872, “silvaticum.” Polytrichum macounti iment in ene Bull. Torrey Bot. a 16:96. 1889; Pogonatum macounii (Kindberg) Kindberg & Macoun, Cat. Canad. Pl. 6:152. 1892. Pogonatum alpinum var. macounii (Kindberg) Cardot & Thériot, Proc. Wash. Aon Sci. 4:328. 1902. Polytrichum alpinum var. macounii (Kindberg) Saito, J. Jap. Bot. 31:75. 1956. Pogonatum alpinum subsp. macounii (Kindberg) Persson in Persson & Viereck, Lindbergia 9:16. 1983 Habitat and distri onen in North America.—On soil in moist woods, common in the lowlands, iddl British Columbia, southern Alaska, Oregon, Washington. Polytrichastrum alpinum i is a widespread, polymorphic species, common at middle to upper elevations throughout the North Temperate zone, in south temperate South America, the Antarctic, and Australasia. The var. sylvaticum isa giant form of the Pacific Northwest with stems as muchas 14 cm high, simple to laxly branched, sharply serrate leaves to 19 mm long, and large, plump, cy- lindric capsules, often somewhat zygomorphic and widest below the middle. REFERENCES Frye, T.C.1937. Polytrichaceae. In: Grout, AJ. Moss flora of North America north of Mexico. Pp. 99-128 Lawton, E.1971.Moss flora of the Pacific Northwest. Hattori Botanical Laboratory, Nichinan,Japan. SIDA 22(1): 547, 2006 548 BRIT.ORG/SIDA 22(1) Book REVIEW STEVEN CARROLL and STEVEN SALT. 2004. Ecology for Gardeners. (ISBN: 0-88192- 611-6, hbk.). Timber Press Inc. 133 S.W. Second Ave, Suite 450, Portland, OR Oi 204-5021. US.A.(Orders: www.tim berpress.com, il@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $29.95, 420 pp., ll baw figures, 170 color photos, glossary, and common, scientific and subject indices, 6" x 9" =) With Ecology For Gardeners, authors Steven Carroll and Steven Salt have written a book that is es- = sential for any gardener’s shelf. Their book keeps ecological principles in mind while dealing with the subject of gardening. Ecology for Gansenirs provide x ii many? ints to pau 1 think about. It contains many cause & effect situations anc between abiotic and biotic factors in rs your garden. The text includes discussion of seemingly Go topics that one may not instinc- tively link together at first; such as how soil temperature relates to nutrient availability for plants, intraspecific competition and plant spacing, pruning as herbivory, energy roles in a garden, the abil- ity of the wind to alter the shape a garden, the importance of microbes to the nutrient cycling & avai a ity in oe puch etc. neler topics include plant biology, non-plant aan ai heir environment ts,an oo gi — as applied a (stewardship and gar i management) Each chapter begins with a basic and easy to understand intro dueion about the topic. After the introduction ios each ¢ Baptett ane fopies is div ute into were organized es For example, the chapter i etition, herbivory, pre- oO pa Se) dation, parasitism and mautualism, Another example is the garden environment chapter that is di- Vv ee into sun, air, water sell energy, food isha and nutrient Se within the alee: - stem. The tyle is the chapter on appliec eee in etn ane isa subtopic of ateee management. I con- - tail cussion on mulching varieties, benef each mulch type. The chapter on applied ecology in the garden really helps th the reader bring together all the previous information of other chapters and may lead to many “Eureka!” moments. Clear color ee phs exemplify many ach of the various plants, items, and occurrences referenced within e Ap The book ee logy, for Garde ners is full many a provoking topics; especially for com- i nd E monly used gical ba hind them. The authors have done a fan- tastic job of ever ts eedlonics! part of gardening very accessible to gardeners. Readers will defi- nitely understand more of the ecology that occurs in, uae and pada the plants in their garden. More importantly, gardeners will be able to make more informed garden management decisions af- ter reading Carroll and Salt’s Ecology for Gardeners —Lee Lucke ydoo, Herbarium, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. SIDA 22(1):, ): 548. 2006 A NEW COMBINATION IN TETRAPHIS (BRYOPHYTA: TETRAPHIDACEAE) Judith A. Harpel Herbarium, Burke Museum Box 35325, University of Washington Seattle Washington 98195-5325, U.S.A. wjharpel@earthlink.net ABSTRACT n Tetraphis pellucida Hedw. var trachypoda 1 | larify the taxonomic The mbinat : ane - ae moss Reais trachypoda and to recognize the morphological difference of having a straight, slightly papillose seta. A revised key to the genus, a diagnosis of var. trachypoda, and speci- mens examined are provided. RESUMEN Se propone la nueva combinacion Tetraphis pellucida Hedw. var. trachypoda, para aclarar el rango taxonomico del musgo Tetraphis ee y para reconocer las diferencias morfologicas de tener una seda derecha ligeramente papilosa. Se ofrece una clave revisada del género, una ee de la var. trachypoda, y los especimenes os Kindberg (1893) described Georgia trachypoda from specimens collected by John Macoun in British Columbia as follows: “leaves subovate or ovate-oblong, short-acuminate and acute. Perichaetial leaves with long, sublinear, acute acu- men; costa not excurrent. Capsule straight; pedicel straight, rough in the upper part.” In 1900 Paris accepted Hedwig’s genus Tetraphis and Georgia trachypoda Kindb. became T. trachypoda (Kindb.) Par. Currently Tetraphis consists of two recognized species, neemicwar Cue ex Milde and T. pellucida Hedw. Because the gametophytes are undisti ble, these two species are separated by se- tae characteristics. Since its description, Tetraphis trachypoda has been recog- nized at various taxonomic ranks. In 1936 Grout combined G. trachypoda under T. geniculata noting the seta roughness and dismissing the straight seta. He comments that “I: geniculata is not materially different in macroscopic appearance from T. pellucida, as the seta is not always noticeably bent at first glance and the bend is sometimes absent as in Kindberg’s Georgia trachypoda (the only difference of importance that he notes in his description). Specimens in the herbarium of the National Museum of Canada, labeled Georgia trachypoda and collected at same date and place as the type and probably a part of it, have the seta only slightly rough- ened and not geniculate, otherwise normal.” Van Der Wijk et al (1969) and Savicz-Ljubitzkaja and Smirnova (1970) fol- lowed Grout’s lead and also placed T. trachypoda under T.geniculata. In Russia, SIDA 22(1): 549 — 554. 2006 550 BRIT.ORG/SIDA 22(1) Abramova et al. (1954) regarded T. trachypoda as having only a weakly mamillose stalk, and therefore combined it under T. pellucida. They did not comment on its distribution in Russia. Crum and Anderson (1981) in their treat- ment of Tetraphis, comment, “Rare specimens with some setae more or less roughened above but not at all geniculate qualify as Georgia trachypoda Kindb., which Grout placed in synonymy of Tetraphis geniculata. The sporadic occur- rence of such plants (in Ontario, Minnesota, Montana and British Columbia at least) in close association with plants with smooth setae seems to indicate that they are anomalous forms of T. pellucida.” Ignatov and Afonina (1992) in their check-list of mosses of the former USSR also treated T. trachypoda asa synomen of T. pellucida. Forman (1962) in his discussion of the family Tetraphidaceae makes no mention of T. trachypoda but does comment that “separating the two species of Tetraphis is easy when mature sporophytes are present because very few colo- nies show intermediate sporphytic characters. In such intermediate cases the tuberculate character of the seta is more constant than the geniculate one.” Be- tween 1936 and 1992 T. trachypoda has been synonymized under both T. geniculata and T. pellucida. While examining specimens of Tetraphis from North America for the Bryo- phyte Flora of North America project, it became apparent that an additional morphological feature of the seta could be used to help clarify the confusion surrounding [. trachypoda. Although Weber and Simone (1977) determined that the papillose or tuberculate surface of the seta in T. geniculata was the result of protruding end-walls of the cells, they did not mention the spiral vs. straight arrangement of the cells. Their paper focuses on the nature of the papillae and does not comment on the fact that the papillae are also the result of the seta cells becoming straight rather than spirally twisted. Tetraphis pellucida hasa straight smooth seta, with all of the cells strongly spirally twisted throughout its length and it is never papillose. The cells of the seta in Tetraphis geniculata are spirally twisted up to the point where the bend occurs. Above the bend the cells are straight and conspicuously papillose by projecting cell-end walls. Tetraphis trachypoda on the other hand hasa straight seta, in which the cells are spirally arranged and interspersed with short sec- tions of cells (or sometimes only a few cells) that are straight, which results ina few papillae on a normally smooth seta. When the type specimen of T. trachypoda was examined, this new characteristic was verified. The change of cell arrangement correlated with papillae and a bend in the seta of T’ geniculata, and straight, consistently spirally arranged, smooth seta cells in T. pellucida appear to be characteristics sufficient to maintain both of these taxa as individual species. Tetraphis trachypoda on the other hand ap- pears to be intermediate between T. geniculata and T. pellucida, but its charac- HARPEL, A NEW COMBINATION IN TETRAPHIS 551 teristics are not sufficient to maintain it as a separate species. It is suggested, therefore, that T. trachypoda be treated as a variety of T. pellucida based on the straight, slightly papillose setae, the outermost cells of which have alternating zones of cells with either spiraled or straight side walls. Tetraphis pellucida Hedw. var. trachypoda (Kindb. ex Paris) Harpel, comb. nov. Figs. 1, 2). Georgia trachypoda Kindb., Rev. Bryol. 20:93. 1893. TyPE: CANADA. BRITISH COLUMBIA: Sicamous, 7 Apr 1889, J. Macoun s.n. (HOLOTYPE: S!). Plants in dense turfs or scattered patches, green above, reddish-brown below. Stems 8-15 mm uptolcm tall. Leaves erect-spreading, ovate to ovate-lanceolate, 1-2 mm long, keeled, costa subpercurrent, margins entire, plane to broadly re- curved, upper medial cells smooth, irregularly rounded-hexagonal, lower basal cells oblong-linear. Stalked, discoid gemmae present in a rosette of rounded bracts (occasionally weakly developed) formed on top of an attenuated stem. Seta 6-14 mm long, erect + flexuose, cells lightly papillose usually near the base, cells spirally twisted but interspersed with sections of straight cells, twisted when dry. Capsule narrowly cylindrical, 2-3 mm long, peristome teeth 4. Spores 10-17 um, green to yellowish-green, papillose. Capsules mature in spring to early summer, rare on well rotted wood, stumps and logs. The distribution of T: pellucida var. trachypoda is interesting because it occurs in North America and Russia, although at the present time, the geo- graphic range in Russia is unknown. Based on the North American material examined, it appears to only be found in areas where both T. pellucida and T. geniculata occur, which suggests that it would occur in the Russian Far East where both of these species also overlap. Examination of material frorn this area and additional fieldwork would =e to validate this hypothesis. ANADA. British Columbi 7 Apr 1889, Macoun s.n.(S). LABRADOR: eras. Bay, 10 Jun 1949, Schofield 32 (NY, UBC). NEW BRUNSWICK. Restigouche Co.: Mount Carleton Provincial Park, 12 Aug 1988, Bristow s.n. (MICH). Northwest Territories. Nahanni National Park, 7 Jul 1974, Scotter 22554: 15 Jul 1976, Steere 76-786, 76-822: 25 Jul 1974, Marsh 4550 (NY). UNITED STATES. COLORADO. Gunnison Co.: 3 Aug 1969, Weber B-34769 (COLO, NY). MONTANA. Lake Co.: Flathead Lake, 26 Jun 1960, Schofield 11632 (NY, UBC). WYOMING. Yellowstone National Park, 17 Aug 1953, Welch 16435 (NY). Teton Co.: 29 Aug 1973, Hermann 25571 (NY) KEY TO SPECIES OF TETRAPHIS . Seta geniculate, cells smooth, and spirally twisted below the bend, cells straight and papillose by projecting cell walls above the bend 1.T. geniculata . Seta not geniculate, outer cells smooth, and spirally twisted throughout the entire length of the seta, not papillose, or sometimes lightly papillose near the 2. Seta smooth, superficial cells spirally twisted throughout the entire ae gth 2a.T. salted: var. pellucida 2. Seta usually lightly papillose near the base, superficial cells spill twisted but interspersed with sections of straight cells 2b.T. pellucida var. trachypoda — — Vow @ene Piura sy _ at Ff i Trak spats ABE areas es Ob rvs, Cy, Lthiroker v9 F9 eA ff SRK CK dope {cr ] gy 7 - PP « _J.Macoun s.n. (S). Size marker in lower right corner equals 2.5 cm. Re oat Oh. mr "3 ye, A a Fb Lite gewigh ma ed L wcoitth : : ft sarha , de - BRIT.ORG/SIDA 22(1) HARPEL, A NEW COMBINATION IN TETRAPHIS 553 AMOS var.trachypoda \ AS) 5 NP © CE e) CO STR ZO JO " 4 ea we C PD © CY oO or wn rT ~ ion p Q Oo ist) far) Q oO i=} ine) te oOo wn a] oO (@ oO n =| o a: | a 3 lean oOo wn usadas y u seguida por se Asteraceae, Lamiaceae, Solanaceae (6 especies cada una), Cucurbitaceae, Malvaceae, y Poaceae (5 especies cada una). El analisis de las formas vitales de estas plantas medicinales indica que estan representadas todas con predominio de las herbaceas. Para conservar esta riqueza Batra se ak ere una esttaicely ion tanto poe janes ie Src como Bal a abordar | | I I ] ] p> ies a }; 1 auc ir ncluir la for mulaci la produccion sostenible de plantas medicinales eden su cultivo. INTRODUCTION Pakistan has the distinction of having the largest contiguous gravity-flow irri- gation system in the world. The Pakistani irrigation system has been greatly expanded since independence in 1947. This irrigation system serves as a life- line for sustaining agriculture and is the main source of livelihood for the 70% population. Punjab has a well-developed irrigation system with a net work of In; Disclaimer: The views expressed in this articles are ee of authors and do not necessarily aes nt those of their Bene departments. The publication o endorse- ment of the use of these plants as herbal medicine. The authors or their departments make no warranty, ex- ee or implied, and assume no legal liability for the use of these plants for medical purposes SIDA 22(1): 593 — 606. 2006 594 BRIT.ORG/SIDA 22(1) — canals, water channels and distributaries. The canal-irrigated areas account for 90% of agricultural production and employ 54% of the labor force (Khan 2002). Pakistan is rich in plant resources with more than 6,000 plant species o higher plants (Nasir & Rafiq 1995). The irrigated areas of Punjab are rich in phyto-diversity, containing a wealth of cultivated crops, natural vegetation and wild flora (Ahmad 2003). This includes plants used in traditional herbal medi- cine. Research on medicinal plants in the irrigated areas of Punjab has been generally ignored in respect to developing a scientific understanding of the dynamics of natural resources, organization and evolution of human systems. This apparent lack of attention to irrigated, cropped areas has resulted ina pau- city of scientific data about the natural vegetation of these areas. There are a few studies listing the medicinal plants from various regions of Pakistan (Ah- mad & Waseem 2004; Athar & Ahamd 2004; Athar & Siddiqi 2004; Gill 2003). However, the irrigated areas of Punjab have not been explored adequately in floristic surveys, and a comprehensive documentation of the medicinal flora of the region also is lacking. Pressure on the wild plants of these areas is ex- pected to increase with increasing agricultural, urban and industrial activities. The natural vegetation, including medicinal plants, of these areas is severely degrading, and there is an immediate need to protect and preserve these plant resources. This study is aimed at compiling a list of medicinal plants in canal- irrigated areas of Punjab with information about their taxonomic position, life form and pharmaceutical utilization of different plant parts. The study also explores the reasons for the decline of medicinal plants in the irrigated areas and suggests measures for their conservation. rH je MATERIALS AND METHODS A literature search was conducted to determine the medicinal plants used for various disorders or diseases in canal-irrigated areas of Punjab, Pakistan (Baquar 1989, 1995; Joshi, 2003; Maheshwari 2003; Nasir & Ali 1972; Nasir & Rafiq 1995; Palaniswamy 2003; Rahim 1996; Rizvi 1998: Shaheen et al. 2003; Tivari & Tandon 2004; William & Ahmad 1999). Their taxonomic position, life form and pharmaceutical utilization of different plant parts were also determined. The plants included herbs, shrubs and trees. The genera were arranged alphabeti- cally within each family. The English and local names that are commonly as- sociated with these plants are provided. The nomenclature and classification followed Nasir and Ali (1972), and author citations followed Brummitt and Pow- ell (1992). RESULTS AND DISCUSSION This study represents the first comprehensive survey of medicinal plants in the irrigated areas of Punjab, Pakistan with suggested strategies to conserve this natural wealth by both saving the medicinal plants and meeting their increased TaBLE 1.Some important medicinal plants in irrigated areas of Punjab. Species Local name English name Life form Part used Pharmaceutical uses MONOCOTYLEDONS Agavaceae Agave americana L. Banskeora Agave Herb Leaves, roots Astringent Alliaceae Allium cepa L. Onion Herb Bulbs, leaves, seeds Anti-ulcera Allium sativum L Lehson Garlic Herb Bulbs, leaves eae a pressure Asparagaceae Asparagus adscendens Roxb. Musli sufaid Garden asparagus = Shrub Seeds, roots Antidiarrhoeal, demulcent galactogogue Asphodelaceae Aloe vera (L.) Burm. f. Gheekawar Aloe Herb Whole plant Cosmetic agent, febrifuge, anthelmintic Cyperaceae Cyperus rotundus L. Dela Sweet cyperus Herb Roots Diaphoretic, lithontriptic, liver tonic Poaceae age lias (L ee Tabasheer Bamboo Tree Leaves, roots Tonic, astringent antidiarrhoeal Ousa 5 (L.)Vo Bamboo Tree Leaves, stem Styptic, tonic, desiccative ymbopogon citratu Izkhir Russ grass Herb Stem Antiseptic, stomachic, styptic (DC.) Stapf. Cymbopogon jwarancusa Khowi ghas Russ grass Herb Flowers Detoxifier, astringent and tonic (Jones) Schult. Zea mays L. Makai Maize, corn Herb Flowers Astringent, chloretic, diuretic and remedy for urinary infection DICOTYLEDONS Amaranthaceae Achyranthes aspera L. Pathkanda Prickly chaff Herb Whole plant Astringent, diuretic NVLSINVd ‘ IVNIDIGUW GVAly ONY avadny 965 Tasce 1. continued Species Local name English name Life form Part used Pharmaceutical uses Apiaceae Apium g lens L. Soya Dill Herb Roots, seeds, Rheumatism, arthritis, indigestion stem flatulence, urinary t inflammation and on neurasthenia, sleeplessness, anxiety and nervous breakdown Centella asiatica (L.) Urb. Brahmi booti Pennywort Herb es Anti-inflammatory, diuretic Coriandrum sativum L. Dhania riander Herb Leaves, stem Antispasmodic, appetizer, aromatic Daucus carota L Gajar Carrot Herb Roots, stem Aphrodiasic, exhilarant, diuretic Foeniculum vu inate Mill. Saunf Fennel Herb Leaves, stem Digestive, lactagogue, masticatory, flavoring agent Trachyspermum ammi (L.) Ajowan Omum Herb Leaves, stem Stimulant, antispasmodic Sprague ex Turrill Apocynaceae Alstonia scholaris (L.) R. Br. Shaitan-ka- Devil's tree Tree Bark Fever treatment Jhad Nerium oleander L. Kaner Oleander Shrub Leaves, roots Abortifacient, attenuant anti-inflammatory Tabernaemontana divaricata Chandani Nero's crown Herb Flowers, roots Anthelmintic (L.) R.Br.ex Roem. & Schult. Asclepiadaceae Calotropis gigantea (L.) Madar Gigantic Shrub Whole plant Expectorant, alterative W.T. Aiton swallowort Calotropis procera (Aiton) Ak Swallowort Shrub Whole plant Diaphoretic, emetic, diuretic W.T. Aiton 96S (L)@2 VaIS/D¥O' LIYE Taste 1.continued = => = Species Local name English name Life form Part used Pharmaceutical uses S = Asteraceae ai Ageratu nyzoides L. Ajganda Herb Flowers, leaves Emetic, digestive tonic = lendula officinalis L. Zergul Calendula Herb Flowers Diaphoretic, antiemetic = Carthamus tinctorius L. Kusumba Safflower Herb Flowers, stem mmenagogue, ae sedative, = stimulant Eclipta prostrata (L.) L. Bhangra Eclipta Herb Whole plant Deobstruent, antiasthmatic atricaria recutita Babuna Chamimila Herb Carminative, stimulant Xanthium strumarium L. Banukra Cocklebur Herb Whole plant Diaphoretic, emollient Boraginaceae k Cordia dichotoma G. Forst. Lasora Sebestan plum Tree Fruits Demulcent, laxative = Heliotropi Oont chara Herb Whole plant Skin disorders = Brassicaceae Brassica juncea (L.) Czern. Rai Brown mustard Herb Seeds Emetic Lepidiu tivu Chandrasur Common cress Herb Leaves, roots Stimulant, anti-inflammatory Sisymbrium irio L. Khub kalan Hedge mustard Herb Leaves, seeds Skin disorders actaceae Opuntia dillenii (Ker Gawl,) Chappal Prickly pear Shrub Whole plant Demulcent, expectorant Cannaceae Canna ind Aqeeq Indian Shot Herb Roots, seeds Diaphoretic, diuretic, demulcent annabinaceae Cannabis sativa L. Bhang Indian hemp Herb Flowers, leaves Sedative, narcotic Capparidaceae Capparis spinosa L. Karir Capper plant Shrub Roots Anti-inflammatory, anti-flatulent, resolvent Cleorne gynandra L. Huinul Caravalia seed Shrub Leaves, seeds Rubefacient, anthelmintic Cleome viscosa L. Hurhur Wild mustard Herb Leaves, seeds Rubefacient, anthelmintic £6S Tage 1.continued Species Local name English name Life form Part used Pharmaceutical uses Chenopodiaceae Chenopodium alb Bathu White goose foot Herb Seeds Anthelmintic phania amb d Sak Mexican tea Herb Seeds Vermifuge Mosyakin & Clements Combretaceae Terminalia arjuna (Roxb. ex Arjan Arjun tree Tree Bark Stomachic, anti-ischemic and oe ) vee & a dla abcae iv a (Gaertn.) Bahira Beleric myrobalan Tree Fruits Bron suilstany antispasmodic poe ot gmatic, expectorant and sedative activities Terminalia chebula Retz. Harir Bidda nut Tree Fruits Stomachic, alterative Convolvulaceae Cuscuta reflexa Roxb. Akas bel Dodder Herb Seeds Carminative, alterative, anodyne Crassulaceae siete oe (Lam.) Pers. Zakhame hayat Life plant Herb Leaves Antiseptic, styptic urbitacea as co ee (L.) Tumma Colocynth Herb Fruit, roots Anthelmintic, antipyretic Schrad. Citrullus lanatus (Thunb.) Tarbooz Water melon Herb Fruits, seeds Laxative, brain tonic Matsum. & Nakai var. /anatus Lagenaria siceraria (Molina) Kaddo Gourd Herb Seeds Brain tonic andl. ita vetnangula (L ’ ‘a Ghia tori Herb Fruits Bitter tonic, diuretic Karela Bitter gourd Herb Fruits, a seeds Anthelmintic, purgative Euphorbiaceae Chamaesyce hirta (L.) Millsp. Dhodak Snake weed Herb Leaves, stem Purgative, rubefacient Euphorbia neriifolia L. Thanda thor Milk bush Tree Latex, roots Purgative, expectorant, antispasmodi 865 (L)@Z VaIsS/OYO'LINA TaBLe 1. continued Species Local name English name Life form Part used Pharmaceutical uses Phyllanthus niruri L. Bhoin avail Myroblan Herb Leaves, roots Diuretic, astringent, bitter tonic Ricinus communis L. Arind Castor oil Shrub Leaves, seeds Emetic, anti-inflammatory Fabaceae (Caesalpinioideae) Bauhinia purpurea L. Kaliar Purple Bauhinia Tree Bark, flowers, Purgative, indigestion, body pain flower buds Bauhinia semla Wunderlin Kandla Bauhinia Tree Bark, leaves, Astringent, diarrhea, dysentry flowers Bauhinia variegata L. Kachnar Mountain ebony Tree Bark, flower Blood purifier, anthelmintic 5, roots Cassia fistula L. Amaltas Golden shower Tree Fruits, leaves, roots Emollient, febrifuge ista absus (L.) H.S. Chaksu Chaksu Herb Leaves, seeds Astringent, detersive, heamotonic Irwin & Barneby opthalmic Senna alata (L.) Roxb. Dadmurdan Ringworm shrub Tree Leaves Purgative, expectorant, aperient Senna alexandrina Mill n Senna Herb Leaves Purgati Tamarindus indica L. Imli Tamarind Tree Bark, leaves, fruits ooling, antibilious Fabaceaae (Mimosoideae) cacia modesta Wall. Phulai Tree Gum Emollient Acacia nilotica (L.) Delile Kikar Acacia Tree Flowers, leaves Emollient, astringent, styptic Albizia iter .) Benth. Siris Lebbek tree Tree Bark, leaves Astringent, oral therapy Mimosa p Chhui-mooi Touch me not Herb Leaves Emetic, renal diseases pene pee Alhagi maurorum Medik. Jawansa Camel's thorn Shrub Whole plant Laxative, demulcent, expectorant Clitoria ternatea L. Gokarni Butterfly pea Herb Roots, seeds Cathartic, dem Cullen plicata (Delile) Bakuchi Babchi Herb Seeds Anthelmintic, ee C.H. Stirt Erythrina stricta Roxb. var. Pangra Coral tree Tree Bark Cathartic, anthelmintic suberosa (Roxb.) Niyomdham WNDIGIW aVAly ONY dVaAy NVISDIVd 665 Tasle 1. continued Species Local name English name Life form Part used Pharmaceutical uses Indigofera tinctoria L Nee| Indigo plant Herb Whole plant Deobstruent, alterative Sesbania grandiflora (L.) Bansa Sesbania Tree Roots, flowers Rheumatism, fever, diabetes Per eee sesban (L.) Merr. Janter Sesbania Shrub Leaves, flowers Suppuration, anti-fertility Trigonella foenum-graecum L Methi Fenugreek Herb Leaves, seeds Aperient, nutritive, expectorant Vigna unguiculata (L.) Walp. Kulatha Horse gram Herb Seeds Astringent, aiere lithontriptic subsp. cylindrica (L.) Verdc. Lamiaceae Lallemantia royleana (Benth. Tukham-i- Herb Seeds Cooling, mucilaginous Benth. balanga anti-inflammatory Mentha arvensis L. din Mint Herb Whole plant Carminative Mentha xpiperita L. Podina Peppermint Herb Leaves Aromatic oil, digestive aa Ocimum basilicum L Niazb Sweet basil Herb Flowers, seeds Carminative, expectoran Ocimum tenuiflorum L. Tulsi Holy basil Herb Flowers, leaves, Stimulant, antiseptic roots Salvia plebeia R. Br. Salbia Sage Herb Seeds Treating boils and wounds Linaceae Linum usitatissimum L Alsi Linseed Herb Seeds Diuretic, emollient Lythraceae Lawsonia inermis L. Mehndi Henna plant Shrub Whole plant Hair tonic, hepatic disorders lvaceae Abutilon bidentatum A. Rich. Pattaka Country mallow Herb Leaves, seeds Demulcent, diuretic Abutilon indicum (L.) Sweet Peeli booti Country mallow Shrub Leaves, roots, seeds Demulcent, laxative, sedative Gossypium herbaceum L Kapas Cotton erb eeds Detersive, emollient, expectorant Malva sylvestris L. ee Common mallow Herb Fruits, leaves Throat infections Sida cordifolia L. mak Country aa Shrub Leaves, roots Astringent, diaphoretic 009 (L)@Z ¥QIS/9YO LYS Table 1.continued Species Local name English name Life form Part used Pharmaceutical uses Meliaceae Azadirachta indica A. Juss. Neem Margosa tree Tree Whole plant Adrenalgic stimulant, dyspepsia, skin ae insecticidal Melia azedarach L. Drek Persian lilac Tree Flowers, leaves Anthelm antispasmodic Toona ciliata M.Roem. Tun Indian mahogany Tree Bark ern Moraceae Ficus benghalensis L. Bar Banyan Tree Leaves, latex Astringent, viscous, resolvent desiccative Ficus religiosa L. Pipal Pipal tree Tree Bark, fruits Astringent, antiasthmatic Ficus virens Aiton Pilkhan Tree Fruits, stem Astringent, stomachic ringaceae Moringa oleifera Lam. Soanjan Horse radish tree Tree Flowers, leaves Anthelmintic, antipyretic Myrtaceae Barringtonia acutangula (L. Jugar Indian Oak Tree Bark, roots, seeds Astringent, emetic Gaertn Mine communis L. Wilayti Myrtle Shrub Fruits, leaves Antiseptic, disinfectant mehndi Psidium guajava L. Amrood Guava Tree Fruits, seeds Astringent, febrifuge, antiseptic, laxative taginaceae Se diffusa L. Itsit Hog weed Herb Leaves, roots Diuretic, diaphoretic, anthelmintic Mirabilis jalapa L. Gul-i-abbas Four o'clock plant Shrub Leaves, roots Demulcent, antispasmodic Nymphaeaceae Nelumbo nucifera Gaertn. Kanwal Water lily Herb Fruits, leaves Cardiac tonic, demulcent Nymphaea lotus L. Nilofar White lotus Herb Flowers Cardiac tonic, hepatic disorders IWNDIGIW UVALY ONY ava NVLSIAVd L09 TasB_e 1. continued 709 Species Local name English name Life form Part used Pharmaceutical uses Oleaceae lasmi inale L. Chambeli Jasmine Shrub Flowers, leaves Anthelmintic, diuretic Nyctanthes arbor-tristis L. Kuri Night jasmine Shrub Leaves, seeds Antibilious, laxative Oxalidaceae Oxalis corniculata L. Surchi Indian sorrel Herb Leaves Anti-scorbutic, refrigerant Papaveraceae Argemone mexicana L. Stayanasa Yellow thistle Herb Whole plant Alterative, stimulant Pedaliaceae Sesamum indi Til Sesame Herb Seeds Aphrodiasic, fattening, nourishing Plantaginaceae Plantago ovata Forssk. lsabaghol Plantain Herb Bark, seeds Cooling, mucilaginous anti-inflammatory Plumbaginace Plumbago ee L. Chitrak Lead wort Herb Roots Stimulant, antiseptic, detersive refrigerant Portulacaceae Portulaca oleracea L. Dholica Purslane Herb Seeds Demulcent, diuretic, astringent Ranunuculaceae Nigella sativa L. Kalonji Black cumin Herb Seeds Diuretic, galactagogue, stimulant Rl amnaceae phus jujuba Mill. Beri Jujube Tree Leaves, seeds Anti-inflammatory Rutaceae Aegle marmelos (L.) Corréa Bel Bil Tree Flowers, leaves Laxative, expectorant Salvadoraceae lvad leiodes Decne. Pil Tooth brush tree Tree Bark, leaves, seeds Vesicant, stimulant Salvadora persica L. Pilu Mustard tree Tree Fruits, leaves Diuretic, anthelmintic (L)@Z ¥OIS/9YO'LIYS Taste 1. continued Species Local name English name Part used Pharmaceutical uses Sapindaceae Cardiospermum hali. Kanphuti Baloon vine Whole plant Demulcent, laxative Scrophulariaceae Bacopa monnieri (L.) Pennell. Brahmi Thyme Gratiola Whole plant Febrifuge, nervine Simaroubaceae Ailanthus excelsa Roxb. Maharuk Talbot Bark, leaves Astringent, febrifuge baaanee Mirch Red pepper Leaves, seeds Pungent, sialgogque Datura metel L. Datura Apple of Peru Leaves, seeds Narcotic, toxic, antispasmodic Tamatar Tomato Fruits Antioxidant, carminative Sol anum dete L. Mako Black nightshade Roots, seeds Cardiac tonic, diaphoretic Solanum virginianun Kandiari Indian Solanum Whole plant Expectorant, diuretic Withania somni ae Asgandh Winter cherry Whole plant nthelmintic, stimulant Tiliaceae Corchorus olitorius L. Kost Jute plant Seeds, leaves Carminative, astringent Grewia asiatica L Phalsa Grewia Fruits Carminative Zingiberaceae Curcuma longa L. Haldi Turmeric Seeds Antimutagenic, antioxidant y hyllaceae Fagonia cretica L. Prickly cover eaves Refrigerant, febrifuge Peganum harmala Harmal Harmal Roots, seeds Narcotic, hypnotic, emetic Tribulus terrestris L. Pakhra Puncture vine Aphrodisiac, diuretic TWNIDIGIW UVALY ONY avaaAy NVLSTAWd £09 604 BRIT.ORG/SIDA 22(1 demands in the traditional health care system. A total of 131 medicinal plant species in 112 genera belonging to 52 plant families are reported from the ca- nal-irrigated areas of Punjab (Table 1). It is interesting to note that most of the medicinal plants were Dicots (48 families), while Monocots constitute only a mi- nor fraction of them (3 families). The most important family was Fabaceae con- taining 21 medicinal species. This was followed by Apiaceae, Asteraceae, Lamiaceae, Sol (6 species each), Cucurbitaceae, Malvaceae, and Poaceae (5 species each). Life form analysis of these medicinal plants indicated that all life forms were represented, but the majority belonged to herbaceous forms (Table 1). Herbs (80 species) were found to be the most common life form fol- lowed by trees (32 species) and shrubs (19 species). Most frequently utilized plants parts included leaves (54 species), seeds (44 species) and roots (27 spe- cies). Whole plants of 18 species were utilized in herbal medicine. The bark, fruits and flowers (16 species each) and stem and branches (10 species) also served as medicines. Onion and garlic bulbs and gums and latexes from some of the plants were also used as medicines. The biological and physical changes associated with fragmentation and loss of habitats and canal irrigation have many residual and secondary effects that aggravate the problem of phyto-diversity loss. These include pollution, decline of soil fertility, water logging, the salinization or acidification of soil and water, the siltation of water channels and increased desertification. Among these, salinity and water logging of soils in Punjab are the most important prob- lems (Davidson 2000). Saline soils, high temperatures and increased rate of evaporation, and limited water supply pose a serious limit back on the growth and distribution of native plants (Hussain et al. 2003). Desertification is an- other threat to native plants of Punjab because it particularly affects the biota of areas with 200-800 mm of average rainfall (Khan 2000). The activities that lead to or enhance desertification include cultivation of unsuitable terrains and soils, over-grazing and lack of proper livestock management practices, and ex- cessive removal of shrubs and trees for fuel wood (Ahmad & Waseem 2004). This has resulted in the loss of many herbaceous species, including medicinal plants, in many areas of Punjab (Khan 2000). The irrigated areas of Punjab are as rich in phyto-diversity and deserve all possible protection and conservation measures to prevent loss of wild plants. The conservation and sustainable management of medicinal plants requires a strategic action at different levels. As part of the conservation of medicinal plants, the management and use of agricultural lands need major revamping. Farmers should realize that their methods of land use and farming practices affect the surrounding environment and the organisms in that environment. Asa first step towards better land management, clearing and fragmentation of native vegetation should be stopped and the extent of native plant reserves should be increased. os may AKBAR AND ATHAR, MEDICINAL PAKISTAN 605 Medicinal plants are one of the few natural products of developing coun- tries that could be marketed at higher prices (Athar @ Ahmad 2004; Shaheen et al. 2003). The ever growing market and demand for herbal ingredients cre- ates possibilities for cultivation of medicinal plants as crops (Hussain et al. 2003). Such efforts could provide a substantial supply of herbal plants and help increase the income of the farmers. With continuous increase in use and de- mand for herbal products, the gap between demand and supply is widening (Crockett & Khan 2003). It is estimated that world trade in medicinal plants and related products may reach to US$ 5 trillion by 2050 (Purohit & Vyas 2005). Most of the demand so far has been met from wild sources, which cannot last much longer. Since medicinal plants are a source of additional earnings and traditional treatments for a major part of rural population, it is recommended that these plants may be propagated and domesticated as cash crops. Thus, cul- tivation of medicinal plants could be a potential source of economic develop- ment that would also meet conservation objectives. ACKNOWLEDGMENTS Special gratitude is expressed to Joseph H. Kirkbride, USDA, Agricultural Re- search Service, Beltsville, MD, for his help in checking the nomenclature and taxonomy of the plants and M. Akmal Siddiqi, Marshfield Medical Research Foundation, Marshfield, WI, USA for valuable suggestions and helpful criticism on the manuscript. The authors are also grateful to M. Iqbal Choudhary, H.EJ. Research Institute of Chemistry, University of Karachi, Karachi, Pakistan for providing many original reprints used in the preparation of this paper. REFERENCES AHMAD, H. 2003. Capacity building for cultivation and sustainable harvesting of medicinal and aromatic plants. In: H. Anmad and A.A. Khan, eds. Conservation and sustainable uses of medicinal and aromatic plants of Pakistan. WWF-Pakistan. pp.32-37. Auman, H. and M. Waseem. 2004. Conservation status of some medicinal plants of the salt range. Zonas Aridas 8:1-8. AtHar, M.and Z. AHmab. 2004. Taxonomy, distribution and medicinal uses of medicinal le- gume trees of Pakistan. Sida 21:951-962. AtHar, M.and M.A. Sippial. 2004. Some reflections on the taxonomy of medicinal flowers of Pakistan. Sida 21:357-369. Baauar, S.R. 1989. Medicinal plants and poisonous plants of Pakistan. Printas, Karachi, Pakistan. Baauar, S.R. 1995. Trees of Pakistan. Royal Book Company, Karachi, Pakistan. Brummit, K.R. and C.E. Powe t (eds.). 1992. Authors of plant names. Royal Botanic Gardens, Kew, UK. Crockett, S.L. and |.A. KHan. 2003. Challenges of standardization: Marker compounds in plants species related and unrelated to top-selling herbs. J. Herbs, Spices & Med. Plants 10:13-24. 606 BRIT.ORG/SIDA 22(1) Davioson, A.P, 2000. Soil salinity, a major constraint to irrigated agriculture in the Punjab region of Pakistan:Contributing factors and strategies for amelioration. Amer.J.Alternat. Agric.15:154-159 Git, M.A. 2003. Cultivation of medicinal and aromatic herbs: experiences of IMHC. In: H. Ahmad and A.A. Khan, eds. Conservation and sustainable uses of medicinal and aro- matic plants of Pakistan. WWF-Pakistan. Pp.23-31. Hussain, N.,A.R. Naseem, G. Sarwar, F. Muuees, and M.Jamit 2003.Domestication and cultivation of medicinal crops on salt-affected soil. In: H. Ahmad and A.A. Khan, eds. Conservation and sustainable uses of medicinal and aromatic plants of Pakistan. WWF-Pakistan. Pp 38-44. JosHi, S.J. 2003. Medicinal plants. Oxford & IBH Publishing Company, New Delhi, India. KHAN, A.U. 2000. History of decline and present status of natural tropical thorn forest in Punjab, Pakistan. Biol. Conserv. 63:250-260. KHAN, F.K. 2002. A geography of Pakistan. Oxford University Press, Karachi, Pakistan. ManesHwarl, J.K. 2003, Ethnobotany and medicinal plants of Indian Sub-continent. Scien- tific Publishers, Jodhpur, India Nasir, E.and S.|. Au. 1972.An annotated flora of West Pakistan. Stewart Herbarium, Gordon College, Rawalpindi, and Department of Botany, University of Karachi, Pakistan. Nasir, YJ. and R.A. Raria. 1995. Wild flowers of Pakistan. Oxford University Press, Karachi, Pakistan. PaLaniswamy, U.R.2003.A guide to medicinal plants of Asian origin and culture.CPL Scien- tific Publishing Services Limited, Newbury, UK. PUROHIT, S.S.and S.P. Vyas 2005.Medicinal plant cultivation: A scientific approach. Agrobios, India. Rani, A. 1996. Our useful plants. Polymer publishers, Lahore, Pakistan, Rizvi, M.A. 1998. Medicinal flowers of Pakistan, Part-Ill. Horticultural Society of Pakistan, Karachi, Pakistan. Pp.48-52. SHAHEEN, F., M. ATTAUR RAHMAN, K.VASISHT, and M. lagat CHoubHaRY. 2003. The status of medicinal and aromatic plants in Pakistan, In: Medicinal plants and their utilization, UNIDO. Pp. 7 Vy os. Tivari, M.and V. Tanbon. 2004. Medicinal plants. sha Books, New Delhi, India. Wiuam, J.T.and Z. AHmab. 1999, Priorities for medicinal plants research and development in Pakistan. Published by MAPPA, India, and IDRC, Canada. NOTES ON THE LIFE AND WORK OF JAMES BRIGHAM MCFARLIN, FLORIDA BOTANIST Tom Palmer 1805 26" St.NW Winter Haven, Florida 33881, U.S.A. ABSTRACT James Brigham McFarlin contributed to Florida botany and his work provided a rare glimpse into the diversity that once existed in the central part of the state three-quarters of a century ago. RESUMEN ae ee Mean aaah a la botanica de Florida y su obra dio una ojeada rara a la Florida central hace 75 anos. James Brigham McFarlin (1901-1969) was among a small group of botanists whose work contributed to a better understanding of Florida’s flora through explorations and collections during the first half of the 20th Century. Unlike J.K.Small and other prominent botanists who visited Florida to collect and ex- plore, Jim McFarlin (Fig. 1) was a Florida resident. He used his familiarity with the area—particularly the Lake Wales Ridge in the central peninsula—his en- thusiasm and a keen eye to collect and record specimens that he regarded as new species or new forms of known species as well as species that were previ- ously undocumented in the state. Apart from a brief mention in Wunderlin et al. (2003), nothing appears to have been written about McFarlin’s life and work. This paper is an attempt to fill that gap. Jim McFarlin was born November 24, 1901, in New York City. He attended Mercersburg Preparatory School in Mercersburg, Pennsylvania and went on to attend Syracuse University, where he received a degree from the New York State College of Forestry in 1925. He then joined his parents, Robert Roy and Grace B. McFarlin, in Winter Haven, Florida. The elder McFarlins had moved to Winter Haven in 1918 (Burr 1974), where the elder McFarlin had purchased land and operated a citrus grove. After moving to Winter Haven, Jim McFarlin was presi- dent and manager of Winter Haven Ornamental Nurseries east of Winter Ha- ven (City Directory 1928). However, he was not a particularly successful busi- nessman and survived at times only through his father’s financial assistance (M. Bryant, pers. comm.). His primary interest at that time was botany, not the nursery business. He followed that interest from 1930 to 1933 with an assistantship from the Univer- sity of Michigan, where he pursued a doctorate. His graduate assistant stipend was $600 a year (Cherie Peterson, pers. comm.). His thesis work involved exten- SIDA 22(1): 607 - 613. 2006 608 BRIT.ORG/SIDA 22(1) Fic. 1. James Brigham McFarlin (front left). sive collecting in Polk County and some surrounding areas in connection with the preparation of his dissertation (McFarlin 1935). No degree was awarded and his assistantship was not renewed after 1933. By 1933 McFarlin was married with one child, unemployed, and in the PALMER, JAMES B. MCFARLIN, FLORIDA BOTANIST 609 depths of the Depression. He found temporary work with the U.S. Department of the Interior's Civilian Conservation Corps. He was employed initially in the Chippewa National Forest, Cass Lake, Minnesota. In 1934 he was transferred to the CCC camp near Sebring, Florida, where work was underway to develop Highlands Hammock State Park, one of Florida’s first state parks. McFarlin be- came the botanist at the Highlands Hammock Botanical Garden and Arbore- tum and served on the Executive Committee of its Botanical Garden Associa- tion (Altvater 1966). He was recommended for that position by Harley Harris Bartlett, then chairman of the Department of Botany and director of the Bo- tanical Garden at the University of Michigan (letter, 17 February 1934, to Harry Lee Baker, Florida state forester). Dr. Bartlett had been a member of McFarlin’s graduate committee at Michigan. McFarlin was responsible for preparing some of the early interpretative materials for Highlands Hammock State Park (McFarlin 1939). Some of his field notebooks containing notes on specimens he collected during his field work in Florida were found around 2000 during an inventory of historical artifacts at the park, but their importance was not recognized until 2005. The pages in the notebooks are being digitally copied for use in a planned exhibit at the park’s CCC museum (D. Smith, pers. comm.). McFarlin lived in the Sebring area until at least 1942, but later moved back to Winter Haven (Burr 1974) and lived there until moving to Bradenton in 1949, shortly after the death of his father in October 1948. In Bradenton, probably because of the inheritance he received from his father’s estate, he was able to resume his career in the nursery business. He operated Begonia Gardens there until his death on March 10, 1969 (Anonymous 1969). According to his obituary, he and his wife Marie, who he married in 1927, had a son, James R. McFarlin, a daughter, Sarah Wells, and four grandchildren. McFarlin began his botanical collection as early as 1923, according to cor- respondence between 1924 and 1935 with J.K. Small (Florida State Archives), from whom he sought information and advice on specimens he collected and for whom he collected specimens. McFarlin’s early collections were deposited with the herbarium at Syracuse University. Later collections involving about 5,000 specimens were sent to the herbaria at the University of Michigan and the University of Florida (McFarlin 1935). Some of his specimens have been deposited in the U.S. National Arboretum (Gann et al. 2002) and other locations. In addition to vascular plants, McFarlin was also interested in mosses and liverworts. He at one time had prepared displays of some of his specimens in mahogany cases (M. Bryant, pers.comm.). He eventually donated his bryological collections to the University of Florida and to the National Museum of Canada in Ottawa (K. Perkins, pers. comm.). That collection supported a publication (McFarlin 1937) in which he concluded that the local moss population in Polk County, was “relatively depauperate of species.” McFarlin collected in south 610 BRIT.ORG/SIDA 22(1) Florida as well and was noted as having collected the first specimens for a hand- ful of species in that region of Florida (Gann et al. 2002). McFarlin’s major botanical exploration, which resulted in the collection of the bulk of the specimens, occurred in 1930 and 1931 while a graduate student. McFarlin collected throughout Polk County, a large (5,300 sq. km.) county lo- cated in the central part of Florida. At the time he was collecting the area was still relatively undeveloped. He made repeated collecting trips to several loca- tions, including Marion Creek near Haines City, Lake Deer near Winter Haven and Kissengen Spring near the Peace River south of Bartow. McFarlin never received his doctorate, but the reasons are unclear. Accord- ing to the University of Michigan's Transcript and Certification Office, he was enrolled as a graduate student from 1930 to 1933 (M. Henderson, pers. comm.). In a September 1, 1935, letter to J.K. Small he wrote of having completed his dissertation and informed. Small he was ready to submit it to his doctoral com- mittee. He wrote on April 29, 1935, to E B. Mains, one of his professors at the university and was told that. McFarlin should first submit his thesis to John H. Ehlers, chairman of his committee. Yet there is no record of his dissertation’s ever having been received at the University of Michigan (E. Voss, pers. comm.). Although McFarlin’s university transcript is unavailable because of claimed privacy concerns, the Certification and Transcript Office agreed to examine the transcript and said it does not indicate why Mr. McFarlin did not complete his degree. (M. Henderson, pers. comm.). McFarlin’s 1935 dissertation, however, has survived. A copy of the original document was completely retyped by Margaret L. Gilbert, a biology professor at Florida Southern College in Lakeland, Florida, in 1961, with duplicates given to several individuals and institutions within the state. The thesis is a tightly writ- ten work of 228 pages (as retyped), the bulk consisting of an enumeration of the vascular flora arranged by family. Each species is reported by scientific name, synonyms, and at times common names, with a brief statement of habitat and location. Nearly all entries also carry McFarlin’s field collections numbers for the species. A key is given to the species of genera with more thana single species. Asa supplement to the main body of the thesis, McFarlin provided a sta- tistical tabulation of its contents: 159 families, 560 genera, and 1131 species. He then recorded 131 species he regarded as weeds, and 53 species as introductions He gave further complete tabulations for the species of “high pine land,” “scrub,” “high hammock,” “flatwoods,” “prairie,” “low hammock,” “swamp,” “bayhead,” “cypress head,” “marsh,” “lakes,” “streams,” “clay pits,” “bogs,” “shallow ponds and drainage ditches,” “shores,” and “sandy beaches.” He followed this with lists of “epiphytes,” “parasites,” 11 and “saprophytes.” McFarlin recorded in the last pages of his thesis what he believed to be 2 species new to the United States, 7 new to Florida, 7 species new to science, 7 new varieties, 14 new forms, and 3 new combinations bya s3 boas PALMER, JAMES B. MCFARLIN, FLORIDA BOTANIST 611 McFarlin’s 1935 thesis and his supporting specimens (mostly those at the University of Florida) have provided clues for further botanical exploration. His work, though unpublished and little known, offers a rare glimpse of the former distribution and abundance of many species that are now much less common in central Florida. A few examples are worth mentioning. McFarlin reported collecting the first U.S. specimen of Hairlike Spikerush (Eleocharis nana),a species that is still considered rare and has been collected from only two other Florida counties (Wunderlin & Hansen 2000). However, the collection of this specimen under- lines one of the problems with reviewing McFarlin’s work. The reported collec- tion site (Lake Lynch near Lake Alfred) is unknown and appears neither on modern maps nor on maps from that period, such as the 1927 Polk County Soil Survey that he cited in his dissertation. Several other sites where McFarlin re- ported collecting regularly, such as Highland Gully near Lakeland and Faulkner Hammock near Bartow, are similarly undocumented as to their locations other than being near some present-day city. In an undated letter to E.B. Maines, Mc- Farlin refers toa map “showing all of my collecting locations,” but no such map was included in any of the known copies of his doctoral dissertation. In some cases, McFarlin found species beyond their known ranges in Florida at the time, suchas the terrestrial orchid Wild Coco (Eulophia alta). Until then the species was known only from the Big Cypress Swamp area, according to Small (1933). McFarlin claimed (McFarlin 1935) to have collected a number of specimens that either represented species previously unknown to Florida or entirely new species. In the case of the new species for Florida, it appears possible that some of the claimed discoveries, such as Nash’s blue-eyed grass (Sisyrinchium brownei) or Ludwigiana brevipes, may have been misidentified since modern summaries of Florida’s botanical life, notably Wunderlin and Hansen (2000), do not include them. However, others suchas Verbena bonariensis and Commelina benghalis are known to exist in Florida and are actually quite common. McFarlin’s only published new species was a small- leaved sand holly that he named Ilex pygmaea (McFarlin 1932). He was unaware that the tree had been named previously, by Small in 1924, and his name is now considered a syn- onym of the scrub holly, [lex opaca var. arenicola. All but one of the six other new species mentioned in his unpublished thesis had no nomenclatural stand- ing and were largely overlooked. The exception, which has recently received attention, is the pink-flowered Scrub (McFarlin’s) Lupine, which McFarlin named Lupinus aridorum. This plant was collected by other workers as early as 1900 (U.S. Fish and Wildlife Service 1993) but McFarlin was the first to con- sider it a separate species. McFarlin did not describe the species. It was not de- scribed until years after his death by Beckner (1982), who also knew the plant in the field, recognized its distinct features, and gave it proper treatment. Though 612 BRIT.ORG/SIDA 22(1) related to Lupinus westidanus, a similar species of the Florida panhandle, it is quite separate in its range and is clearly an endemic of the central peninsula. McFarlin described the plant as “infrequent to locally abundant,” but with de- velopment of the area it has become quite rare, and is now classified as endan- gered by the state and federal agencies (Coile & Garland 2003). Asa result, the future survival of this species remains a concern. The only protected popula- tions of the plant occur at a federal preserve in Eagle Lake south of Winter Ha- ven and at a county park in the Orlando area (Kane 2003). Unprotected popu- lations persist in a handful of other sites in Central Florida. Establishment of other protected populations—including one site near Winter Haven where Mc- Farlin did much of his collecting—has been proposed (Kane 2003) and would seem toa fitting tribute to McFarlin’s work. McFarlin commented on other endemic plants of the central ridge. Another species he believed to be new was a small-leaved variety of grape that is wide- spread in the scrub habitat. He gave it the name Muscadina pygmea. Later ob- servers have noted the plant, but have never given it formal status by publish- ing its name and description. Recently McFarlin’s original specimens have been tracked down (in the Smithsonian Institution) and a paper is in press to give it formal recognition as a variety of the muscadine grape, Vitis rotundifolia (D.B. Ward, pers. comm.). McFarlin made interesting observations on the distribution of known en- demic species in the scrub habitat in Polk and adjacent counties, relatively little of which remains (Christman 1988). McFarlin’s comments were prescient re- garding this trend. For instance, he noted that the presence of Turkey Oak (Quercus laevis) was a general indication of land that was suitable for citrus, which is the crop that most frequently displaced scrub and sandhill habitat. Comparing his comments (McFarlin 1935) with those of present day observers helps us to un- derstand how dramatic that change has been. McFarlin described the beargrass, Nolina brittonidna, as “our most common species inhabiting the scrub and in- land sand dunes,” the crucifer, Warea amplexifolia, as “locally abundant,” the scrub plum, Prunus geniculata as “widespread,” the morning-glory, Bonamia grandiflora as “frequent in the sandy soil of the scrub,” and the milkweed, “As- clepids curtissii, as “frequent in scrub.” Now they are rare and classified as en- dangered or threatened (Coile & Garland 2003; Wunderlin & Hansen 2000). He left another legacy that was probably unintended. Although McFarlin recognized the problems of invasive exotic species—his manuscript mentions Ceasar Weed (Urena lobata), Australian Pine (Casuarina esquisetifolia) and oth- ers—he was responsible for some local introductions of other invasive plant species. Through his years in the nursery trade, McFarlin was responsible for the introduction of of the orchid tree, Bauhinia variegata, into Winter Haven (Burr 1974). This tree is classified as a Class | invasive species (Florida Exotic Pest Plant Council 2005). While operating the botanical garden at Highlands — PALMER, JAMES B. MCFARLIN, FLORIDA BOTANIST 613 ] © LA] Hammock, McFarlin was apy ly for the introduction of the Flax Lily (Dianella caerulea), which continues to be a management problem in the park today (A. Hine, pers comm.). ACKNOWLEDGMENTS Ithank Daniel Ward and John Beckner for their encouragement and for provid- ing helpful comments during the preparation of earlier drafts of this account and Ward and Ron Gilmour for their review of the final draft. lalsothank Nancy Bissett for introducing me to Mr.McFarlin’s work in Polk County, Rita Lassiter for sharing her copies of J.K. Small’s correspondence, Edward Voss for help in locating relevant correspondence from the University of Michigan and Mary Brigham Bryant for sharing her childhood recollections. REFERENCES Autvarer, A.C. 1966. Highlands Hammock. Sebring Historical Society, Sebring, i Anonymous. 1969. James Brigham McFarlin. Obituary. Bradenton Herald, March 10, 1969. Becker, J. 1982. Lupinus aridorum J.B.McFarlin ex Beckner (Fabaceae), A new species from Central Florida, Phytologia 50:209-211. Burr, J.G. 1974. History of Winter Haven, Florida. Larry Burr Printing Co., Winter Haven, FL. CHRISTMAN, S. 1988.Endemism and Florida’s Interior Sand Pine Scrub. Final Project Report, Florida Game and Freshwater Fish Commission, Tallahassee. City Directory, WINTER Haven. 1928. R.L Polk Co. Jacksonville, FL. Cote, N.C. and M.A. Gartanp. 2003. Notes on Florida’s endangered and threatened olants. Florida Dept. of Agric. and Consumer Services, Gainesville. Fioriva Exoric Pest Plant Councit. 2005. Invasive plant list. http://www.fleppc.org/list/list.hAtm. Tallahassee, FL. Gann, G.D., K.A. Braptey, and S.W. Woopmansee. 2002. Rare plants of South Florida: Their his- tory, conservation, and restoration. Institute for Regional Conservation. Miami, FL. Kane, S.2003. Historical and current distributions and population status of Lupinus aridorum McFarlin ex Beckner. Masters thesis, University of Central Florida, Orlando. McFarun, J.B. 1932. Two new evergreen holly from Central Florida. Rhodora 34:16-18. McFaruin, J.B. 1935. Flora of the central portion of the Lake Region District of Florida. McFaauin, J.B. 1937. Mosses of Polk County, Florida. Bryologist 40:49-5/7. McFaauin, J.B. 1939. Highlands Hammocks Trails. A guide to the plants along the Laurel Oak Trail. Highlands Hammock State Park, FL. Smait, J.K. 1933. Manual of the southeastern flora. Published by the author. New York. U.S. FisH AND WiLouiFe Service. 1993. Endangered and threatened status for seven Central Florida Plants. Fed. Reg. 58(79):25746-25755. WUNDERLIN, R.P.and B.F. Hansen. 2000. Flora of Florida.Vol.1.Pteridophytes and gymnosperms. Univ. Press of Florida, Gainesville. WUuNDERLIN, R.P. and B. HANseN. 2003. Vascular plants of Florida, 2nd Edition. Univ. Press of Floria, Gainesville. BRIT.ORG/SIDA 22(1) Book REVIEW WILLIAMS. Justice, C. Rircuié Bett,and ANNE H. Linpsry. 2005. Wild Flowers of North Carolina (Second Edition). (ISBN 0-8078-5597-9, pbk.). University of North Carolina Press, P.O. Box 2288, Chapel Hill, NC 27515-2288, U.S.A. (Orders: www.uncpress.unc.edu/books/T-7662.html). $29.95, 376 pp., 553 color and 6 b&w illus., 6 tables, | chart, appendices, bibliography, index, 5 1/2" x ol/2". Edition 2 supplants the original from 1968 by Justice and Bellit includes 100 more species than the With conservation and preserva- original and provides brief notes on toxicity and medicinal uses. ‘irst in mind, the authors note that “Our goal in this presentation of 500 native or naturalized tion plants in North Carolina is to open eyes, minds and hearts to the story of the state’s wild flowers, their beauty, their interesting attributes, their uses, and, in many cases, their plight.” Most pages in- clude a photo and notes for each of two species, and Appendix | gives an outline of horticultural information for each species. The Edition 2 photos are better reproduced, often sharper, sometimes cropped differently, but usually reduced in size up to 30 percent. For Edition 3, better planning of increase image sizes and decrease the large areas of white space on so many of Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102- layout surely could the pages. Guy Nesom, Botanical 4068, U.S.A, SIDA 22(1): 614. 2006 FLORISTIC DIVERSITY AND COMPOSITION OF TERRA FIRME AND SEASONALLY INUNDATED PALM SWAMP FORESTS IN THE PALMA REAL WATERSHED IN LOWER MADRE DE DIOS, PERU Fernando H. Cornejo Valverde John P Janovec Botanical iesedien Institute of Texas — Peru Botanical Research Institute of Texas SN a Ae 09 Pecan St. cae de Dios, PERU Fort Worth, Texas 102-4060, U.S.A. fcornejo@brit.org jjanovec@brit.org Mathias W.Tobler Botanical Research Institute of Texas 09 Pecan St. Fort oe si - 102-4060, U.S.A. tobler@gmx.net ABSTRACT We report the results of a tree species inventory of seven hectares of lowland subtropical moist forest in the Palma Real River watershed of lower Madre de Dios, Peru, where we discovered a type of palm swamp forest vegetation that has not previously been studied. The goal of the study was to investi- gate the forest vegetation of the Palma Real watershed with special emphasis on a comparison be- tween the wetland forest vegetation and the surrounding terra firme forests. A total of 3534 trees >10 cm DBH belonging to 442 species in 62 families were measured and identified in the seven plots. Three distinct forest types resulted from Principal Components Analysis (PCA) and WPGMA cluster analysis of both family and species abundance data, including two types of terra firme for- ests and a palm swamp forest. The five terra firme forest plots sampled, with a range of 104-138 species, were less diverse than other one-hectare forest plots reported from the region, which have a diversity ranging from 151-197 species. The two palm swamp forest eee with 126 and 167 tree spe- cies, were more diverse than surrounding terra firme forest plots sampled in this study. One of the palm swamp forest plots was extremely diverse with 167 species, ae seven species less than the average diversity across 14 one-hectare terra firme plots sampled 2 other investigators in Madre de Dios, Peru. Both palm swamp forest plots had a greater number of stems distributed across smaller size classes. Both Fabaceae and Arecaceae were ane most important plant families across all plots. The Arecaceae contributed significantly to g the palm swamp forest plots, composing 16-25% of all stems compared to only 9% of the stems in terra firme plots of the region. This high abundance of the palm family is from where we derive the name we are applying to this wetland forest type in the Palma Real watershed. Using satellite imagery we illustrate the importance of the extent of these palm swamp forests throughout the region RESUMEN Reportamos los resultados de un inventario floristico de siete Becuce en un pees SHE ADICH amedo en la cuenca de la quebrada Palma Real, en Madre de Dios, Per fue investigar la vegetacion de la Quebrada Palma Real con especial Hee ae en una comparacion SIDA 22(1): 615 — 633. 2006 616 BRIT.ORG/SIDA 22(1) entre un tipo de vegetacion de Si antano con pn panier y la uegetacion de terra lirme circundante. i n 4 AyD total de 3534 arboles mayoresa fueron medidos e identificados en siete parcelas. Tres tipos de bosque résultaron de los analisis de componentes principales y analisis de agrupamiento WPGMA mediante datos de abundancia por familia y por especie en los dos tipos de ee aoe Las cinco dices yess en terra a me, con un rango de 104-138 especies por parcela, fuer sas que otras p en la region, las cuales tienen un rango de 151-197 especies. Las dos parcelas cima en . pantanoso de palmeras, con 126 y 167 especies, fueron mas diversas que las parcelas de terra firme circundantes. Una de las parcelas del bosque pantanoso de palmeras fue extremadamente diversa con 167 especies, solo siete especies menos que el promedio reportado para |4 otras parcelas estudiadas por otros investigadores en Madre de Dios. Ambas parcelas del bosque pantanoso con palmeras tuvieron una alta censidad de tallos sania en ~ cl ases diamémetricas mas pequenas, lo cual podria ser ion de laalt i . Fabaceae y Arecaceae fueron las familias mas pee para todas las eos orn contribuy6 significativamente a distinguir el bosque pantanoso con palmeras, compre aie coen te el 16-25% Ge i ee ss uaseuiak con fs a lt solo el 9% de tallosen | DOs a de Cerra firme ade la region FE el nombre que ste tipo de bosque muy humedo en la cuenca de la Quebrada de Palma Real. Usando imerenee csielicales nosotros ilustramos la importancia por su extension, de estos bosques pantanosos de palmeras en la region. Finalmente nosotros instamos a Mowe mas estudios de largo plazo sobre - eee, pees en a region, con é ome en la importancia de estos bosques de Palma Real e trac : i, Lecythidaceae) y otro: recursos para la region y sus ee INTRODUCTION A large forest plot network has accumulated in the last decade from the Ama- zonian region. The highest tree species diversity has been reported from the western Amazon where Gentry (1988) and Valencia et al. (1994) recorded 283 and 307 species respectively in one-hectare forest plots. Oliveira and Mori (1999) found 285 species per hectare on very poor soils and under reduced rainfall in central Amazonia, which is essentially the same species diversity as found by Gentry in western Amazonia. In an analysis of data from 97 existing one-hect- are forest plots from the Amazon, including the Guiana Shield, Ter Steege et al. (2000) showed that in all regions terra firme forests harbor a greater number of tree species per unit than any other forest types. Patterns of high tree species diversity have been attributed to high rainfall and more nutrient-rich soils of the western Amazon (Gentry 1988). Nebel et al. 2001) and Kvist and Nebel (2001) emphasize the importance of wetland forests to both the ecological diversity and resource use in the Peru- vian Amazon. While many forest plot inventories in the Amazon have focused on the diversity of terra firme and floodplain forests, few studies in the region have dealt with wetland vegetation. For example, only two swamp forest plots from the Amazon were available for inclusion in the large studies by Ter Steege et al. (2000, 2003), but those were excluded from the final analyses and spatial modeling. There is clearly a lack of data from wetland habitats in the Amazon. This minimal coverage of wetland vegetation leads toa generalized view of the CORNEJO ET AL., FLORISTIC PERU 617 — complexity and diversity of these ecosystems, as discussed by Kalliola et a (1991), who argue for more focused studies. The Tambopata-Candamo region and Manu National Park are two well- studied protected areas of Madre de Dios, Peru, that hold world records in diver- sity of various organisms, such as birds, insects, and mammals. National re- serves, such as Tambopata, allow for protection and sustained use of plant resources. The Palma Real watershed is inside the Tambopata National Reserve and the Bahuaje-Sonene National Park, where management plans are needed since plant and animal extraction is carried out by indigenous and local people. Yet, no floristic treatment exists for the region and only a few small areas are represented by checklists. In the Palma Real watershed and the surrounding region, there exists a vast area of terra firme forest and patches of seasonally inundated swamp forests. This mosaic of forests extends from the Tambopata Reserved Zone of lower Madre de Dios, Peru, to the Pampas del Heath in the border region of Peru and Bolivia. The seasonally inundated forests of this region are characterized by a distinctive topography that consists of small raised islands separated by shal- ow water channels during the rainy season or heavy rains in the dry season, and they sit upon a deep, fine, heavy gray soil. Because of the abundance of palm species in our study area we refer to this wetland vegetation as palm swamp forest. We have reviewed available literature that relates to the diversity, ecology, and distribution of wetland vegetation in the area, and we have found no existing description of this specific wetland vegetation. The palm swamp forests of Palma Real are different in species and family diversity and composi- tion than the palm swamp forest communities of the northern Peruvian Ama- zon described by Kalliola et al. (1991). Although Haase (1990) described palm swamp communities in northern Bolivia, there is no description that matches the palm swamp forests of the Palma Real watershed. Nor are these palm swamp forest communities covered by Beck (1984). The goal of this study was to inves- tigate the forest vegetation of the Palma Real watershed in lower Madre de Dios, Peru, with special emphasis on a comparison between these palm swamp for- ests and the surrounding terra firme forest vegetation. METHODS Study Site The study was conducted in the Palma Real watershed, a tributary of the Madre de Dios River, approximately 28 km to the southeast of Puerto Maldonado, the capital and the principal city of the Department of Madre de Dios, Peru (Fig. 1). Nested within the Tambopata-Candamo National Reserve and the Bahuaje- Sonene National Park, this area of lowland subtropical moist tropical forest (Fos- ter et al. 1994) receives about 2208 mm of precipitation annually and the aver- age temperature is 24°C (mean over 36 years). During the driest months, May to BRIT.ORG/SIDA 22(1) 618 YP 9 Naf q C7 oe Rivet ae DY” - gate = M QS ZY, ¢ Kn rd ) Limon { ° ©) < Puerto Maldonado * fro A : bd C & Oculto . x yw? & eo fs > i QW Ka) e ) < Q& Lf @ wS NY S 3s Settlements ARN) La Viuda A Plots G, y ‘Soh ~ N ; km . 0 5 10 #15 Pd Fic. 1. Location of the three si here f pl li based, Limon, Oculto, and La Viuda. hich September, the area receives occasional cold winds from the antarctic, w decreases daily temperatures to a minimum of 7-8°C, although this phenom- enon of cold fronts, known as “friajes”, lasts for only a few days at a time. The general topography is flat, dissected by small streams, most of which are without water during the dry season. Three main types of habitats exist in the Palma Real watershed, forming an interesting mosaic of vegetation zones: (1) high terrace non-inundated forest (terra firme); (2) palm swamp forests - areas with poor drainage and seasonal inundation causing the formation of swamp-like forests; and (3) seasonally inundated floodplain forest. This study focused on terra firme and palm swamp forests, which are the most extensive forest types in the study area. The Brazil nut tree (Bertholletia excelsa, Lecythidaceae) is abundant in large areas of the terra firme forests in this re- gion but it is absent from others. Seeds of the Brazil nut tree are extracted from the Palma Real watershed by harvesters (castaneros) every year during Janu- ary-March, making this an economically important region in Madre de Dios, Peru. The forests of the Palma Real watershed maintain a healthy fauna with jaguars, ocelots, tapirs, peccaries, deer, and at least six species of monkeys. No J logging activities have been conducted in this region since twenty years ago Ss CORNEJO ET AL., FLORISTIC DIVERSITY IN LOWER MADRE DE DIOS, PERU 619 when many Spanish Cedar trees (Cedrela odorata, Meliaceae) were extracted. Then terra firme forests of Palma Real that are cl ized by high Brazil nut tree density have been divided by the Peruvian government into forest conces- sions of approximately 1000 hectares each and given to local extractors living in Puerto Maldonado. Soils.—The soils of the terra firme forests are dominated by red clay and those of palm swamp forests by gray clay. Within the Palma Real zone, three representative sites were selected for quantitative studies of floristic diversity and composition (Figs. 1, 6): (1) Limon (12°32'19"S; 68°51'40"O), close to the Madre de Dios River; (2) Oculto (12°39'03'S: 68°55'40"O), in the middle of the zone; and (3) La Viuda (12°47'43'S: 68°58'07"0), to the extreme south of the zone. Field Data Seven permanent one-hectare forest plots were established in the Palma Real watershed, divided between Limon (3 plots), Oculto (3 plots) and La Viuda (1 plot). Within each site, one plot was established in terra firme (TF) forest witha known high density of 5-7 individuals of Brazil nut trees per hectare (Limon 1, Oculto 1, La Viuda 1). One plot in each of Oculto and Lim6n was established in TF forest with known absence of Brazil nut trees (Oculto 2 and Limon 2). In Oculto and Limon one plot was also established in palm swamp (PS) forest (Oculto 3 and Limon 3). Each plot consisted of a square area of 100 x 100 m, divided into 25 sub- plots of 20 x 20 m. Each corner of the one-hectare forest plot is marked witha PVCstake labeled with the respective X,Y coordinate in red paint. All individu- als, including palms, 10 cm or greater in diameter at breast height (DBH) were measured, mapped and identified to family, genus, and species or morphospecies when definite species determination was not possible. All individuals with multiple stems were considered as one individual, and the diameters of each stem were summed for basal area calculations. Every species is represented by duplicate voucher specimen collections of one or more individuals. All identifications were made in the La Molina Her- barium (MOL )at the Agraria University in Lima, Peru, where duplicate voucher specimens were deposited. The Cronquist system of classification of flowering plants was used for the purposes of this project. Data Analysis Relative diversity, relative density, and relative dominance were calculated for each plot. The Importance Value Index (IVI) was calculated at the species and family level for each plot using the sum of the three measures, relative diversity, relative density, and relative dominance. Sorensen Similarity Index (Serensen 1948) was calculated for all species found in the plots and for the 20 most abun- dant species. The Sorensen index is 2 * S}2 / (S1 + S2), where S)2 is the number of species common to both sites and S;and $2 are the total number of species found 620 BRIT.ORG/SIDA 22(1) at site one and two. Plots were also grouped based on species and family abun- dance using Principal Components Analysis (PCA) and WPGMA cluster analy- sis (MVSP 3.12, Kovach Computing Services). Comparing the raw number of species between plots without accounting for differences in stem density can give misleading results. To avoid this prob- lem we used individual based rarefaction curves as suggested by Gotelli and Colwell 2001). Rarefaction curves were calculated using the software package EstimateS 7.5 (Colwell 2005). RESULTS Forest Structure A total of 3534 trees =10 cm DBH belonging to 442 species in 62 families were measured and identified in the seven plots (Table 1). Three distinct forest types resulted from PCA: (1) palm swamp forest (Limon 3 and Oculto 3); (2) Limon terra firme forest (Limon | and Limon 2): and (3) Oculto/La Viuda terra firme forest (Oculto 1, Oculto 2 and La Viuda) (Figs. 2, 3). PCA resulted in almost no difference between the species (Fig. 2) and family level (Fig. 3),a pattern previ- ously suggested by Terborgh and Andersen (1998). Table | shows the principle structural characteristics for each plot. The palm swamp (PS) forests (PS) in both Limon and Oculto have a higher number of species than the terra firme (TF) forests at the same sites. Limon 3 has the high- est number of individuals and species of all the plots while the two TF forest plots at Oculto are the least diverse. The rarefaction curves (Fig. +) confirm these results and show that the higher diversity of the PS forest plots are not simply due toa higher stem density. The PS plots have a lower basal area than the TF plots (215 vs. 27.3). Distribution of trunk diameter classes shows the inverse J- shaped curve, witha mean proportion of 69% of individual trees found between 10 and 20 cm DBH. = Composition Monimiaceae was the most abundant family in the TF forest plots, followed by Arecaceae and Fabaceae (Table 2). In PS forest plots Arecaceae and Fabaceae were the most abundant families. Looking at the 20 most abundant families (Table 2), Lauraceae, Moraceae, Violaceae, Lecythidaceae, Chrysobalanaceae, Sapotaceae, Burseraceae, and Cecropiaceae were found in both TF and PS for- ests, whereas Myristicaceae, Strelitziaceae, Linaceae, Bixaceae, Rubiaceae and Flacourtiaceae were found only in TF forests. Of the 20 most abundant families, Myrtaceae, Dichapetalaceae, Anacardiaceae, Clusiaceae, Ochnaceae, Sterculiaceae, and Combretaceae were restricted to the PS forests. The ten most abundant fami- lies represented 64% of all individual trees found. The most species-rich family across all plots was Fabaceae, followed by Lauraceae, Moraceae, Sapotaceae, Chrysobalanaceae, and Annonaceae. The ten richest families represented 54% CORNEJO ET AL., FLORISTIC DIVERSITY IN LOWER MADRE DE DIOS, PERU Taste 1. Structural characteristics from 1 hectare plot. Limén1 Limén2 LaViuda Oculto1 Oculto2 Oculto3 Limén3 No. of stems 499 Sop 473 422 508 492 608 No. of species 138 130 130 109 104 126 167 Basal area ( m7’) 32.23 23.77 25.23 29.25 26.22 21.86 21.07 Categories by diameter (%) 10-20 cm 3.4 70.5 69.5 64.9 65.9 74.6 74.5 20.1-40 cm 27.6 23.5 22.2 23.0 24.8 203 21.9 40.1-80 cm 74 5.1 7.0 10.2 8.3 4.3 35 > 80cm 1.6 0.9 1.3 1.9 1.0 0.8 0.2 No. of species in 50% IVI 16 16 18 13 13 14 20 No. of families in 50% IVI 39 42 4] 4] 40 42 45 4.09 —- 3.27 —— 2.45 —— La Viuda A Oculto 3 A Oculto 1 Oculto 2 0.82 -- £ | —+—_-__ SEE EEEEEEEE eee a | Ne) -3.27 -2.45 -1.63 -0.82 0.82 1.63 2.45 3.27 4.09 i672 7. Limén 3 -1.63 — Limon 2, ~2:45-- A Limén 1 A» = Axis 1 Fic. 2. Ordinati f inci r t a 622 BRIT.ORG/SIDA 22(1) Limon 1 26.7—-+- A Limon 2 c SIXW -6.7 + La Viuda A A A Oculto 1 aan Oculto 3 A Oculto 2 -20.9—- Nedinati £4 Deine Fic. 3. Ordination of a | p f y J of all species. Table 3 summarizes the Importance Value Index (IVD for the 20 most abundant families per plot, per forest type, and across all plots, emphasiz- ing the importance of Arecaceae in PS forest plots, and the Fabaceae across all forest plots. Table 4 lists the twenty most abundant species in each of the seven I-ha forest plots. There were only seven species that occurred in all seven plots. Twenty species were found to occur in all five TF forest plots while 62 species were found to occur in both of the PS forest plots. A total of 243 species were found to occur in only one plot out of the seven. Out of the total of +42 species found across all the plots, 124 occurred only in the PS forest and 212 only in the TF forest. A total of 106 species were found to occur in both forest types. About +0% of the species were represented by only one individual and 65% by | to 3 individuals. Similarity Table 5 shows the Sorensen Similarity Index for all plots. Values range from 0.25 to 0.55 for all species and from 0.10 to 0.75 when only the 20 most abundant CORNEJO ET AL., FLORISTIC PERU 623 180 9 160 74 a MSc. Limon 1 i La Viuda Lim6én 3 140 5 120 4 Te Limon 2 Oculto 3 w 1007 = ) a Oculto 2 ® 894 Oculto 1 60 4 40 4 20 4 Q T T ; T T T 1 0 400 200 300 400 500 600 700 Individuals D £. + L I L L + 1 £ j j ! J L ££ Fic.4 g I p species were included. The 20 most abundant species, on average, represent 60% of all individuals (Range: 54-69%). Similarity analysis shows the same patterns already observed in the PCA. First, plots within the same forest type (TF or PS) are more similar to each other. Considering only the 20 most abundant species in each plot, this pattern becomes even more apparent. The TF forest plots from the same site (e.g., Oculto | and 2 or Limon 1 and 2) share a higher number of species with each other than they do with plots from different sites. Figure 5 provides the results of WPGMA cluster analysis based on species abundance, which shows that PS forest plots cluster together, as do TF forest plots from the same sites. When all species are considered, PS forests are more similar to the TF forest plots at the same site than to those farther away. However, this is not true when we consider only the 20 most abundant species. In this case, Limon 3 shares the same number of species with Limon Las it does with Oculto 2, and Oculto 3 and Oculto | are the least similar plots. DISCUSSION The Palma Real watershed presents a unique seasonally inundated PS forest habitat that is very different from the TF forests of the region. Based on species abundance, the two PS forest plots are clearly distinguished from the TF forest plots as shown by results of both PCA (Figs. 2, 3) and WPGMA cluster analysis (Fig. 5), as well as similarity index analysis (Table 5). Euterpe precatoria is the 624 BRIT.ORG/SIDA 22(1) nH haldt rh +f Taste 2. The 20 most diverse families in the two main forest types. Families i those tnat are among the top 20 in both forest types. Number of individuals are average by hectare Terra firme Forest palm swamp Forest Family No. of No. of Species Family No. of No. of Species individuals individuals ceae 41.6 39 Fabaceae 78 35 Lauraceae 30.6 26 Annonaceae 12 12 26.8 22 Chrysobalanaceae 18 12 Sapotaceae 9.6 1/7 Lauraceae 75 1] Chrysobalanaceae 7.6 14 Sapotaceae 16 11 Meliaceae 12.4 12 Moraceae 33.5 10 Annonaceae 94 1] Burseraceae 15.5 8 Cecropiaceae 20.2 10 Arecaceae 128.5 7 Monimiaceae 53 9 Apocynaceae 10.5 7 Flacourtaceae 6 9 Anacardiaceae 11.5 6 pocynaceae 78 7 Lecythidaceae 24 7 Ar ea 52.8 6 Clusiacea 10.5 4 Burseraceae 19.8 6 tacea 15 4 Myristicaceae 36.8 5 Cecropiaceae Ves) 3 Rubiaceae 7.2 4 M ea 8 3 Lecythidaceae 124 4 Dichapetalaceae 11.5 2 Violaceae 26.6 2 Violaceae 28.5 2 Linaceae 9.4 2 Ochnaceae 10.5 2 Strelitziaceae 14.2 | Combretaceae 8 2 Bixaceae 9 | Sterculiaceae 10 2 most abundant species in PS forest plots, Limon 3 and Oculto 3, contributing significantly to the major differentiation between the PS and TF forest plots. Astrocaryum murumuru, Attalea butyracea, Brosimum laevis, Chimarrhis sp. 1, Dipteryx micrantha, Eriotheca globosa, Manilkara bidentata, Ouratea weber- baueri, Rheedia sp. 2, Terminalia oblonga, and Triplaris americanaare restricted to PS forest plots. Calyptranthes bipenis, Iryanthera juruensis, Lauraceae sp. 1, Lauraceae sp. 2, Pourouma minor, and Rubiaceae sp. 2 are restricted to TF forest plots. Based on our results, there are two groups of TF forest plots as demon- strated by Figures 2, 3, and 5. The first group of TF forest plots, Limon | and Limon 2, is distinguished primarily by the abundance of Brosimum dlicastrum, Pseudolmedia laevigata, and Rinoreocarpus ulei, and secondarily by Phenakospermum guyanense. The second group of TF forest plots, Oculto 1, Oculto 2, and La Viuda, is distinguished by the abundance of Minquartia guidnensis, Oenocarpus batua, Roucheria punctata, Roucheria sp. 1, Rubiaceae sp. l, Socrotea exorrhiza, and Tontolea corymbosa. Analysis of family abundance demonstrates similar results (Fig. 3, Table 3). The families Fabaceae and Arecaceae are the most abundant families across ~— CORNEJO ET AL., FLORISTIC PERU 625 Taste 3. Family Importance Value (FIV) of the 20 most abundant families for each plot, both forest and across all plots (sorted by Total column). The FIV was calculated as the sum of relative a relative density, and relative dominance. Family LaViudal Limén1 Limon2 Limén3 Ocultol Oculto2 Oculto3 PS TF Total Fabaceae 35.39 33.25 42.05 52.04 25.96 29.65 47.27 48.96 33.55 37.28 Arecaceae 16.76 1446 17.85 31.91 1865 2238 48.26 3836 16.34 22.32 Moraceae 13.14 3699 39.55 2583 918 3.01 27.71 27.37 2048 21.30 Lecythidaceae 2045 3071 490 17.16 3798 1741 949 1345 22.05 20.11 Lauraceae 22.78 21.32 1584 520 23.29 2639 684 695 2239 19.16 Monimiaceae 1735 1340 19.64 1.62 2063 16.20 426 325 1765 13.28 Sapotaceae 9.24 1275 745 1436 800 10.78 7.61 11.59 10.56 11.31 Burseraceae 6.26 8.81 10.92 11.01 1286 15.82 453 810 1036 9.52 Myristicaceae 1242 969 669 446 1280 2457 141 3.13 12.01 9.17 Cecropiaceae 8.40 11.79 16.22 362 931 17.03 425 3.51 11.63 9.07 Vidas 5.80 5.51 606 1040 9.63 573 1452 12.06 7.26 8.69 Violac 2.58 22.93 12.96 1398 3.11 240 448 940 859 852 eee 8.25 5.53 662 536 440 950 1222 848 696 7.31 Flacourtiaceae 10.37 0.97 7.12 5.91 7.30 5.65 108 385 592 5.68 Anacardiaceae 2.11 499 477 5.61 5.23 450 875 7.25 460 5,25 Clusiaceae 6.18 3.7) 3.71 566 290 3.29 362 443 436 461 all PS and TF forest plots, although they are more abundant in PS forest plots. The Lauraceae and Monimiaceae are more abundant in TF forest plots. The Violaceae and Moraceae are much more abundant in all the Limon plots in- cluding the PS forest plot Limon 3, distinguishing all the Limon plots from the other plots on axis lin the PCA (Fig. 3). The Dichapetalaceae and Polygonaceae are found only in PS forest plots, helping to distinguish them from TF forest plots. A higher abundance of the Bombacaceae in PS forest plots also helps to distinguish them from the TF forest plots. The first TF forest plot group of Limon 1 and 2 is distinguished by a higher abundance of Lauraceae species than the rest of the plots, as well as by the Staphylaceae, which only occurs in Limon 1 and 2 TF forest plots. The second TF forest plot group of Oculto l and 2 and La Viuda is distinguished by a high abundance of Myristicaceae, as well as by the Bixaceae, Hippocrateaceae, and Linaceae, which only occur in those plots. PS forest plots of Palma Real have a higher abundance of palms in com- parison to the TF forest plots, making the Arecaceae one of the most significant families distinguishing the different forest types (Table 2 & 3). This is especially apparent in Oculto 3, where palms account for the top three and sixth most abundant species sampled. Palms compose 25% of the stems in Oculto 3 and 16% in Lim6n 3. In comparison, in all TF forest plots in the region palms make up only 9% of all stems. Euterpe precatoria, the most common palm in the area, = Tas.e 4. The twenty most abundant species in each of the seven one-hectare forest plots. LaViuda1 Oculto1 Oculto2 Limon] Bixa arborea sic sp. | Siparuna decipien al chrysophylla Oenocarpus bataua Socrotea exorrhiza Euterpe precatoria Lauraceae sp. 1 Roucheria punctata Iryanthera juruensis Tachigali polyphylla Calypthrantes bipenis Tetragastris a Iryanthera lae hoo sp. 1 Phenakospermum guyanense Pourouma minor Rubiaceae sp. 2 Inga gereauana Siparuna decipiens Siparuna $ Euterpe precatoria Iryanthera laevis Lauraceae sp. 1 Ouratea sp. 1 Oenocarpus bataua Protium 5 Tetragastris panamensis Iryanthera juruensis Roucheria punctata Le polyphylla nt mb Tontolea corymbosa Hirtella giandistiotla Bertholletia excelsa Pourouma minor Trichilia quadrijuga Phenakospermum guyanense Hebepetalum humirifolium Iryanthera laevis Siparuna s Iryanthera juruensis uterpe precatoria toe oo Laurac sp. 1 oe bataua Pourouma minor Eschweilera coriacea Phenakospermum guyanense Protium $s Socrotea exorrhiza Bixa arborea Siparuna decipiens Tetragastris panamensis Tontolea corymbosa Ouratea sp. 1 Lacistema aggregatum Roucheria punctata NN TWO OO Leonia glycicarpa Rinoreocarpus ulei Euterpe precatoria Siparuna decipiens Iryanthera juruensis Lauraceae sp Pseudolmedia laevis Siparuna sp. 1 Brosimum alicastrum Lauraceae sp. 2 Guarea macrophylla Inga auristillae Lauraceae sp.9 Pseudolmedia laevigata Socrotea exorrhiza ee panamensis Turpinia sp. 1 Gall ulei Celtis schippii (2D) ON: (ONS) SS SST SN 979 (L)@Z VIS/9YO'LIYS Taste 4 (continued from left) Limon2 Limon3 Oculto3 Euterpe precatoria Siparuna decipiens Phenakospermum guyanense Rinoreocarpus ulei Siparuna Tetragastis panamensis imum alicastrum pee nner p pouroura guianensis Lauraceae Pourouma minor Cecropia eae Eschweilera c Pseudolm ci ee Tachigali s eae vargasil Geissospermum reticulatum Euterpe Leonia glycica ee eens ence Eschweilera coriacea Tachigali chrysophylla Socrotea exorrhiza Rinoreocarpus ulei Ouratea cf. weberbaueri Tetragastris panamensis Brosimun lactescens Manilkara bidentata neuen tessmannil Ingae aay iis ees see Chimarrhis aude ese Rheedia CONN WWM Pl, SSS SSP Oe (COs AONE: (NO a Euterpe precatoria Socrotea exorrhiza Astrocaryum murumuru Tachigali chrysophylla Theobroma subincanum Attalea butyracea Tapura tessmannii Terminalia oblonga Eschweilera coriacea Leonia glycicarpa Brosimun lactescens Hirtella excelsa Triplaris americana Brosimun guianensis Cecropia sciadophylla Cordia cf. scabrifolia Inga gereauana JLSGO 1a TV La OfaNaQ) nud’ £79 628 BRIT.ORG/SIDA 22(1) | Taste 5. Number of common species (above) and S@rensen Similarity In v) by one-hectare plot. The diagonal shows the total number of species for each plot. Alle species EM 20 most abundant species. La Viuda Limon 1 Lim6n 2 Limon 3 Oculto 1 Oculto 2 Oculto 3 LaViuda 130 51/8 56/10 43/5 55/13 46/13 32/4 Limon1 038/040 138 61/10 56/5 49/7 49/8 36/3 Limon2 043/050 045/050 130 50/7 48/10 52/10 32/5 Limon3 0.29/0.25 037/025 034/035 167 40/4 44/5 64/8 Oculto1 046/065 040/035 040/050 0.29/0.20 109 59/15 34/2 Oculto2 039/065 040/040 044/050 032/025 055/075 104 33/4 Oculto3 0.25/0.20 0.27/0.15 0.25/0.25 044/040 0.29/0.10 0.29/0.20 126 | al Oculto 3 Limon 3 Limon 2 Limon 1 Oculto 2 Oculto 1 La Viuda Fic 5 n A f WPGMA 1 | oe | 4 £. 1 7 . L 4 is among the five most abundant species in six of the one-hectare forest plots and it is the most abundant species sampled in both of the PS forest plots, with 88 stems in Lim6n 3 and 61 stems in Oculto 3. Socrotea exorrhiza is also an important species in PS forest plots, ranking as the sixth most abundant species in Lim6n 3 with 15 stems, and the second most abundant species in Oculto 3, with 32 stems. Astrocaryum murumuru is the third most abundant species in Oculto 3, with 31 stems. In Oculto 3, Attalea butyracea, known locally as “shebon’, is the sixth most abundant species, from which has come the local common name of this forest type, shebonal, but this is not true for Lim6n 3. One of the most important families in the TF forest plots was the Monimiaceae (Figs. 2, 3), making this the first report of high abundance and dominance of this family in any tropical forest plot study. This family was rep- CORNEJO ET AL., FLORISTIC PERU 629 2] ry , Limén, Oculto, and La Viuda, and the HG ft bates im satin image from aay 2001 showing ine bw J the border of Peru and Bolivia. resented by two species of the genus Siparuna that were common in areas of tree fall gaps. The presence and abundance of Tachigali sp.,a monocarpic spe- cies of the Palma Real watershed, might explain the presence of tree fall gaps and existence of several groups of light-demanding species (e.g., Brazil nut and Cecropiaceae). We found that the PS forest plots differ structurally from other forests by a greater number of stems, the distribution of tree diameter size, and basal area, especially the site Limon 3, which has 80 more stems than other plots (Table L). In contrast, the TF forest plots in the Palma Real watershed present the typical structure in comparison to reports from other regions in Madre de Dios, and the Amazon basin, in general (Pitman et al. 2001). The PS forests of the Palm Real watershed are characterized by several months of anaerobic conditions because of the inundation of the area during the rainy season from at least November to February. As suggested by Terborgh and Andresen (1998) and Connell and Lowman (1989), who note that swamp forests are characterized by low tree diversity and high dominance by one ora 630 BRIT.ORG/SIDA 22(1) low number of species, we expected to find a relatively low species diversity in the PS forests of the Palm Real watershed. However, the most significant result of this study was that the PS forests in the Palma Real watershed were found to have a higher tree species diversity per unit area than TF forests of the region. Limon 3,a PS forest plot, was the most diverse of all plots sampled, with 167 tree species, only seven species less than the average diversity of 175 species across l4 one-hectare TF forest plots sampled in Madre de Dios, Peru, by Pitman et al. (2001). Although Oculto 3 was not as diverse as Limon 3, with 126 species it had more or just slightly fewer species than the TF forest plots sampled, and it was more diverse than the two neighboring TF forest plots from the same site (Oculto | and 2). All of our TF forest plots from the Palma Real watershed were less diverse than TF forest plots sampled in the neighboring Tambopata region, which range between 151 and 197 species (Gentry 1988; Phillips et al. 2003). It is interesting to compare the diversity of the two PS forest plots of the Palma Real watershed with other one-hectare swamp forest plots in the Madre de Dios region. Pitman et al. (1999) and Terborgh and Nufez (unpublished data) have found a range of 61-129 tree species among five one-hectare plots in Madre de Dios, Peru. Phillips et al. (2003) report 158 tree species from a swamp forest plot in the Tambopata region. The Oculto 3 PS forest plot from Palma Real falls out at the upper limit of the first range, with 126 species total. Limon 3, with 167 species, appears to be the most diverse swamp forest plot discovered to date in Madre de Dios, Peru. And, as mentioned, Limon 3 is nearly as diverse or is more diverse than many non-wetland TF and floodplain forest plots in Madre de Dios, Peru. Our results and discoveries lead us to some important questions about the forests of the Palma Real watershed and argue for continued studies of this re- gion. First of all, why are the PS forest plots of Palma Real so high in diversity in comparison to plots in neighboring non-swamp forest plots? And second, why are the TF forest plots relatively low in diversity compared to the PS forest plots of the Palma Real watershed, especially Limon 3, and other TF forest plots from the Tambopata region and Madre de Dios, Peru, in general? The importance of the PS forest in the region can be seen in the Landsat satellite image (Fig. 6), which shows the great expanse of these forests across the landscape from the Tambopata River in Peru to the Heath River on the Bo- livian-Peruvian border and beyond to the east into Bolivia. Preliminary studies in the La Torre River watershed (Cornejo, unpublished data) show a forest with the same characteristics of the PS forests of the Palma Real watershed, with a dominance in the understory of Lycopodium sp., Selaginella sp., Clusia spp., Melastomataceae spp., and other species of non-woody plants that were not sampled in our forest plots. Interestingly, such plants are also found in the vast wetland savannahs to the east, known as the Pampas del Heath along the bor- der of Peruand Bolivia. Local people in the region, in passing conversations with peed = CORNEJO ET AL., FLORISTIC PERU 631 our team, have mentioned that the Pampas del Heath wetland savannahs can develop into dense swamp forests, especially in the absence of management by fire. We hypothesize that these diverse PS forests may represent a successional stage of vegetation derived from the Pampas del Heath savannahs that are more common to the east. This hypothesis could be tested using a combination of remote sensing and intensive ground-truthing between the Tambopata region and the Pampas del Heath savannahs to the east. The Pampas del Heath sits on a thick mass of organic matter that has accumulated over hundreds of years of deposition and slow decomposition characteristic of wetland ecosystems (i.e., bogs). In other words, perhaps the scattered PS forests of the Palma Real water- shed and surrounding region represent a preliminary stage of succession from open wetland savannah to tropical forest. Pollen cores could shed some light on the succession of these vegetation types. This leads us to our second open-ended question, which is why are the TF forest plots of the Palma Real relatively low in diversity? The forests of Madre de Dios, Peru, are referred to as subtropical moist tropical forests (Foster et al. 1994). It is possible that the forests in the Palma Real watershed receive less pre- cipitation than forests to the west, from Tambopata to Manu, in Madre de Dios, Peru. However, there are no climate data available to test this hypothesis. An- other possibility is that the soils of TF forests in the Palma Real region are nu- trient-poor. We do not have data that enable us to test this hypothesis. We rec- ommend future soil sampling in association with vegetation studies in the Palma Real region. Itis possible that with continued sampling of PS and TF forests in the Palma Real watershed, other patterns of plot diversity will be discovered, including more diverse TF forests and differences in PS forest diversity. Clearly, more stud- ies are needed of PS and TF forests of the Palma Real watershed. Considering that the Palma Real watershed is one of the most important zones of Brazil nut extraction in all of the southwestern Amazon, we conclude by arguing for more sustained and long-term studies of the botanical and ecological diversity, and the conservation and management of this region. ACKNOWLEDGMENTS We would like to thank the Gordon and Betty Moore Foundation and the Ben- eficia Foundation for their support of botanical studies in Madre de Dios, Peru. We would like to thank Adrian Forsyth and Enrique Ortiz for their support of studies of Brazil nut forests in the Palma Real watershed and for their contin- ued support of the Brazil Nut Project (Proyecto Castanales) of the Amazon Con- servation Association (ACA) and the Asociacion para la Conservacion de la Cuenca Amazonica (ACCA), through which this work was originally supported. Fieldwork would not have been possible without the logistical support provided by ACCA in Peruand specifically the staff of the Brazil Nut Project in Madre de 632 BRIT.ORG/SIDA 22(1) Dios. We thank the board, administration, and staff of the Botanical Research Institute of Texas for their constant support of the Andes to Amazon Botany Program, through which this manuscript was produced. We thank Sonia Guerra, Amanda Neill, Tiana Franklin, and Keri McNew who have supported us di- rectly and indirectly in the preparation of this manuscript. We also thank the numerous individuals, such as Piher Maceda, Pedro Centeno, Pedro Maceda, three Peruvian field assistants who assisted in many aspects of the fieldwork required to complete the study. We are grateful to the Ministry of Natural Re- sources of Peru (INRENA) for their support of this project and for providing permits for research and collections in Madre de Dios, Peru. We also thank Asuncion Cano and staff of the San Marcos Natural History Museum, and Car- los Reynel and staff of the La Molina University Forestry Department in Lima, Peru, for providing herbarium support to the project. REFERENCES Beck, S.G. 1984. Comunidades vegetales de las sabanas inundazides en el NE de Bolivia. Phytocoenologia 12(2/3):321—350. Cotwett, R.K. 2005. Estimates: Statistical estimation of species richness and shared species from samples. Version 7.5. URL: http://purl.oclc.org/estimates. Connett, J.H. and M.D. Lowman. 1989. Low-diversity tropical rain forests: some possible mechanisms for their existence. Amer. Naturalist 134:88-119. Foster, R., ef AL. 1994. The Tambopata-Candamo Reserved Zone of southeastern Pert: a biological assessment. RAP Working Papers, Vol 6. Conservation International. Gentry, A.H. 1988. Tree species richness of upper Amazonian forests. Proc, Natl. Acad. Sci. 85:156-159. Gore, NJ.,and R.K. Cowwett. 2001. Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness. Ecol. Letters 4:379-391. Haase, R. 1990. Plant communities of a savanna in northern Bolivia Il.“palm swamp'’s, dry grasslands, and shrubland. Phytocoenologia 18(2/3):343-370. KALLIOLA, R., M. PuHakka, J. Sato, H. Tuimisto, and K. RUoKOLAINEN. 1991. The dynamics, distribu- tion, and classification of swamp vegetation in Peruvian Amazonia. Ann. Bot. Fennici 28:225-239, Kvist, LP. and G. Neer. 2001. A review of Peruvian flood plain forests: ecosystems, inhabit- ants, and resource use. Forest Ecol Managem. 150:3-26. Neset, G., L.P. Kvist, J.K. VANCLAY, H. CHRISTENSEN, L. Freitas, and J. Ruiz.2001.Structure and floristic composition of flood plain forests in the Peruvian Amazon |. Overstorey. Forest Ecol. Managem. 150:27-57. Ouiveira, A.D. de and S.A. Mori. 1999. A central Amazonian terra firme forest. |. High tree species diversity on poor soils. Biodivers. & Conservation 8:1219-1244. Puitups, O.L., R. VAsquez Martinez, P. Nusiez Varcas, A. Lorenzo Monteacuoo, M, CHuspe ZANS, W. GALIANO SANCHEZ, A. PENA Cruz, M, TiMANA, M. Yu-Hatta,and S.Rose.2003. Efficient plot-based floristic assessment of tropical forests. J. Trop. Ecol. 19:629-645. =e CORNEJO ET AL., FLORISTIC PERU 633 Pitman, N.C.A,, J. TersorGH, M.R. Sitman, and P. Nunez V. 1999. Tree species distributions in an upper Amazonian forest. Ecology 80:2651-2661. Pitman, N.C.A., J.W. TerBorGH, MLR. SiLMAN, P. Nunez V., D.A. Nei, C.E. Ceron, W.A. Paacios, and M. AutestiA. 2001. Dominance and distribution of tree species in upper Amazonian “terra firme” forests. Ecology 82:2101-2117. TerBORGH, J.W. and E. ANnpresen. 1998. The composition of Amazonian forests: patterns at local and regional scales. J. Trop. Ecol. 14:645-664. Ter Steece, H., D. SABATIER, H. CASTELLANOS, T. VAN ANDEL, J. DUlveNvooRDEN, A.A. DE OLiveirA, R. Ek, R. Livwan, P. Maas, and S.A.Moari. 2000. An analysis of the floristic Composition and diversity of Amazonian forests including those of the Guiana Shield. J. Trop. Ecol. 16:801-828. Ter Steece, H., N.C.A. PITMAN, D. Saparticr, H. CastTeLLAnos, P. Van Der Hout, D.C. Daky, M. SILveiRA, O. PHILLIPS, R. Vasquez, T. Van ANDEL, J. DulveNvooRDEN, A.A. DE OLIvEIRA, R. Ek, R. LILWAH, R. THOMAS, J. Van Essen, C. BAiDer, P. Maas, S.A. Mori, J. TeR80RGH, P. Nunez Varcas, H.Mocotton, and W. Morawerz. 2003. A spatial model of tree alpha-diversity and tree density for the Amazon. Biodivers.& Conservation 12:2255-277. VALENCIA, R., H. Batstev, G. Paz, and C. Mino. 1994. High tree alpha-diversity in Amazonian Ecuador. Biodivers. & Conservation 3:21-8. BRIT.ORG/SIDA 22(1) Book REVIEW CAMPBELL and LYNN LOUGHMILLER. Updated by Damon E. Waitt. 2006. Texas Wild- flowers: A Field Guide (Revised Edition). (ISBN 0-292-71286-3, pbk.). University of Texas Press, PO. Box 7819, Austin, TX 78713-7819, U.S.A. (Orders: http:// www.utexas.edu/utpress, 512-471-4032). $19.95, 278 pp., 381 color photo, ay Tee, This is the successor to the original from 1984, which, top three best sellers for the University of Texas Press, with more tha edition includes an additional 61 species (for a total of 370), but most rai photos ae been re- 1, consistently ae ere eae notes g the 1155, favayay oy 14 he new see Ongne text is mostly pncneneee: The family order i g I iff | lated for species names, about sing © -aceae’ ‘endings for the : percent of which have been supplanted or modified since 1984. Del Damon Waitt of the Wild- flower Center in Austin has been responsible for the taxonomic revision and accuracy of the photo identifications, as both of the Loughmillers have passed on In explanation of why faites 2 I sé 1 lly appearance from the first,” the intro- duction notes that the cover stock is durable and the en narrower—changes that are “part of the University of Texas Press plan to develop a complete series of natural history field guides.” Ap- parently as part of the same revamp, margins are pee spaces expanded oa the words and | less COMIPAacl COX. The new | SAaY¥e consid- lines, and the font less bold—resulting in erably smaller and seem subsidiary to the text. Edition 2 is useful, but if you don’t have the first, try to find a copy of it also, since the original Loughmiller photos, in addition to their larger size, often here it was show views more representative of the species, and a brief statement for V taken (e.g., “Photographed near Blanco State Park in April,” “Photographed in the Big Thicket in May”).—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, USA SIDA 22(1): 634. 2006 A VASCULAR PLANT INVENTORY OF STARKEY WILDERNESS PRESERVE, PASCO COUNTY, FLORIDA Emily Ferguson Richard P Wunderlin 2531 River Plaza Dr, #52 Department of Biology Sacramento, California 95833, U.S.A. University of South Florida Tampa, Florida 33620-5200, U.S.A. ABSTRACT Starkey Wilderness Preserve d and 1 by the Southwest Florida Water Management Dis- trict, contains 7,739.5 hectares with 18 slanee communities. A floristic inventory was conducted on 449.2 hectares within the property which consists of 11 community types and is considered as rep- resentative of the entire Preserve. The objective of this study, conducted from May 2003 to October 2004, was to compile a list of the vascular plant taxa found to be used by the Southwest Florida ter Management District to help in their management regimes. A total of 483 taxa were collected, representing 104 families, and 271 genera. Of these, 444 are native taxa, 16 Florida endemic taxa, 39 non-native taxa, 12 encaneciess threatened or commercially exploited taxa, and 33 county records. Each is described and an annotated list of the vascular plant taxa is presented pi RESUMEN La reserva de Starkey Wilderness, gestionada por el Southwest Florida Water Management District, tiene 7,739.5 hectareas con 18 aeaaae vegetales. Se realiz6 un inventario floristico en 449.2 hectareas de la reserv 1 tipos de comunidades y que se consideran representativos de la Reserva entera. El cbeiv0 us este ae realizado desde mayo de 2003 hasta octubre de 2004, fue compilar una lista d I sadas por el souces Florida Water Manage- ment District para ayuda égi | ion. Se col de 483 taxa, que ce 104 familias, y 271 géneros. De slice 444 ¢ son taxa nativos, 16 taxa endémicos de Florida, 39 taxa aldctonos, 12 taxa en peligro, amenazados o explotados comercialmente, y 33 citas del ois Se describen las comunidades vegetales y se presenta una lista comentada de los taxa de plantas vasculares INTRODUCTION Florida is a unique state with a relatively flat topography and generally fast draining soils. These conditions would presumably lead toa flora with low di- versity, but that is not the case. The state is about 1000 km long and spans about 6.5 latitude resulting in a diverse flora consisting of nearly 4,200 taxa of native and naturalized non-native vascular species (Wunderlin & Hansen 2003), the third most diverse in the United States. As Florida is also the fourth most popu- lated state, it is important to work to conserve and protect the state’s flora. Several agencies in Florida work to secure large tracts of landinan effort to prevent development and to establish areas of conservation. One of these, the Southwest Florida Water Management District (6WFWMD), owns many par- cels of land for a variety of water management practices including the protection SIDA 22(1): 635 — 659. 2006 636 BRIT.ORG/SIDA 22(1) and development of potable water supplies, aquifer recharge, water quality en- hancement, restoration and protection of natural systems, and structural flood control. The property in which the inventory was conducted is owned and managed by SWFWMD. The results of this research are intended to help SWEWMD make knowledge-based management plans for its property. Site Overview Physical Location.—Starkey Wilderness Preserve is located in southwestern Pasco County, Florida, near the west coast of central Florida (Fig. 1). It is 6.4 km northeast of Seven Springs and 4.4 km northwest of Odessa. Main access to the Preserve is Wilderness Road approximately 5.6 km from State Road 54. A por- tion considered to represent the major habitats found within the entire Preserve was selected for this inventory. The Starkey Wilderness Preserve is included within Sections 1-17, 21-24, and 26-27, Township 26 South, Range 17 East and Sections 10-11, 13-15, 22-27, and 33-36, Township 25 South, Range 17 East. The entire Preserve comprises approximately 7,739.5 hectares. onsists of 18 plant communities. Of these, 11 occur within the study site. The study site, consisting of 449.2 hectares, includes portions of Sections 3-4, 8-10, and 15-17, Township 26 South, Range 17 East. The boundaries of the study site comprise a power line artery along the western edge, a paved biking trail at the north, an unpaved road running due south along the eastern edge, and the Anclote River along the south. Early Inhabitants.—The first humans who lived near the Starkey Wilder- ness Preserve were the Safety Harbor Culture or the Tocobaga, who inhabited the Florida Gulf coast (Lawson et al. 1981). The period of their inhabitance was from 900 A.D. through the early eighteenth century (Milanich 1994). The settle- ment area extended from the Withlacoochee River (north and east boundary of Citrus County) south to the Charlotte Harbor area (Charlotte County). Tocobaga arrowheads have been unearthed on the Starkey property, suggest- ing their use of the land to hunt (Lawson et al. 1981). Approximately 33 prehis- toric archaeological sites have been identified as temporary campsites for hunt- ing within the Starkey Wilderness Preserve (GSWFWMD 1990). Hunters on the Preserve could have originated from two different sites. The first, a flint work- shop located 3.2 km north of the Anclote River along the coast; and the second, Tarpon Springs at the mouth of the Anclote River (Willey 1949). In the eigh- teenth century, after the Spanish established themselves in much of the state, the remaining Tocobaga are believed to have joined the Creek tribes from fur- ther north as they moved south seeking refuge from persecution by the new settlers. The land now included in the Starkey Wilderness Preserve passed from the Federal Government to the State of Florida and private ownership beginning in 1856 (Lawson et al. 1981). Principal land uses were for turpentine production FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PERSERVE, FLORIDA Soil types oN QB Adamsville fine sand C — Basinger fine sand | Cassia fine sand — Chobee soils [3] Immokalee fine sand ___] Myakka fine sand 2 Paola fine sand Pomello fine sand MM Pomona fine sand fT Sellers mucky loamy soil f | Smyrna fine sand : (99 Wauchula fine sand 0 250 = 500 1,000 1,500 2,000 oo = J Mer Meters Fic. 1. A. Location of the Jay B. Starkey Wild P , Pasco County, Florida. B. Jay B. Starkey Wilderness Preserve ith +h f+h tod C pl re ary Jay B. Starkey Wilderness Preserve study area. 638 BRIT.ORG/SIDA 22(1) and for cypress and pine lumber. In 1937, Jay B. Starkey and his partners, Ernest, Dave, and Howard Cunningham, bought 6,070 hectares. The C-S (Cunningham- Starkey) Ranch was used to raise cattle by both families. As land values in- creased and cattle yields decreased, portions of the land were sometimes sold to developers. Jay B. Starkey, who outlived his partners, finally sold the cattle and the equipment to his son Jay. B. Starkey, Jr. The senior Starkey still lived and worked the land with his son for cattle ranching, and later during World War IL. converted about 30 hectares into an orange grove. Over the years, items such as timber, pine stumps, and resin were sold to provide extra income. In 1975, as the population increased in the county, Jay B. Starkey, Sr. decided to preserve the land from development by selling part of the ranch to SWFWMD with the understanding that SWFWMD would maintain the land in its natural state (Lawson et al. 1981). By 1980, SWFWMD had acquired 1,407.9 hectares (Wells 2004). The SWEWMD has purchased additional property since then and the Starkey Wilderness Preserve now comprises approximately 7,739.5 hectares. Climate.—The climate of Pasco County is humid and subtropical (Chen & Gerber 1990). The constant cyclical pattern consists of cool, dry winters and warm, humid summers. Annual mean temperature is 22°C (SWF WMD 1988). During the winter months, December through February, the monthly mean temperature is 16°C. The temperature will usually drop to freezing at least once a year. During the summer months, from June to September, the monthly mean temperature is about 27° C, with highs reaching about 33°C. Rainfall in Pasco County varies seasonally, oscillating between the rainy season and the dry season (SWF WMD 1988). From October to May, rainfall is less than 10.2 cm per month, while during the summer months, June through September, the area receives about 18.4 cm of rainfall per month. This accounts for about half of the rainfall for the year. In spring, central Florida, including Pasco County, experiences harsher and longer, dry drought-like conditions than either northern or southern Florida (Chen & Gerber 1990). The high amounts of rain that fall from June to September are attributed to the convective clouds, sea breezes, and tropical storms. By the beginning of October, the rainfall de- creases by about 50 percent from that of the summer months, indicating the beginning of the dry period. Geology.—The geology underlying the Starkey Wilderness Preserve is the Suwannee Formation deposited during the Oligocene epoch of the Tertiary period (34 to 24 MYBP) (Scott et al. 2001). This formation is a yellow to white, fossiliferous, fine-grained limestone (Wetterhall 1964). It is a very porous for- mation that contains the Upper Floridan Aquifer. Below the Suwannee Forma- tion are the Ocala and Avon Park Formations, both deposited during the Eocene epoch of the Tertiary period (55 to 34 MYBP). Both of the lower formations are also important parts of the Floridan aquifer. Soils.—Twelve distinct soil types occur within the study site. The domi- FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PERSERVE, FLORIDA 639 nant types are: Chobee, frequently flooded (22%), Adamsville fine sand (19%); Myakka fine sand (13%); Immokalee fine sand (11.5%); and Sellers mucky loamy fine sand (10%). Other soil types present are Pomona fine sand (8%); Pomello fine sand, 0 to 5 percent slope (6%); Wauchula fine sand, 0 to 5 percent slope (4%); Cassia fine sand, 0 to 5 percent slope (3%); Smyrna fine sand (1.5%); Paola fine sand (1%); and Basinger fine sand (1%) (Stankey 1982). Topography and Hydrology.—Starkey Wilderness Preserve is located in the Gulf Coastal Lowlands physiographic province, which is characterized by low elevations and poor drainage. Within the study site, the elevation ranges from 9.1 to 16.8 m (USGS 1974). The majority of the site has an elevation of 10.7 to 12.2 m. The swamp and river floodplain area ranges from 9.1 to 10.7 m. The Anclote River, located primarily in Pasco County, is 27.4 km long. About 8.1 km of the Anclote, excluding the south branch, is within the Starkey Wilderness Preserve (SWFWMD 1990). The southern border for the study site is an approximately 3.2 km section of the Anclote River (fig. Ll). The Anclote is characterized as a slow moving, turbid, acidic, black water stream. The chan- nel is not more than 6 m wide at any point along the southern boundary of the study site. During low flow stage, the stream is confined to the channel. As the rainy season starts, the Anclote overflows its banks inundating the floodplain forest, and remains at flood stage for two to three months. When the river over- flows its banks, the water flowing into the floodplain removes waste and pro- vides nutrients to the floodplain community. The sediment load settles out as the water moves out of the channel. Heavier sediments are deposited first closer to the channel due to their weight, and finer, lighter sediments settle out more slowly further away from the channel. Disturbances Affecting the Study Site—With the exception of the effects of cattle grazing, lumbering, and other minor events, the study area has under- gone few changes. The western boundary of the study site is the right-of-way for the Florida Power Corporation electrical transmission line. This 46 m wide artery was constructed in the 1970s (SWFWMD 1990). In order to maintain the right-of-way the area is frequently mechanically cleared. In addition, SWF WMD built 14 active production water wells within the Preserve. These wells pump water from the Preserve to provide potable water for the city of New Port Richey. Three production wells occur within the study site. An approximately 3,240- hectace portion of the Preserve, the Jay B. Starkey Wilderness Park, is open to the public for a number of recreational opportunities. In order to provide these recreational activities, SWFWMD constructed a paved road, hiking/biking trails, horse riding trails, primitive camp sites, and semi-primitive road beds. A number of these features occur crinee the study site as well. The most recent disruption within the study site was a wildfire that was believed to have started from a lightning strike June 3, 2004. After persisting through the night, the wildfire consumed approximately 162 hectares on June — 640 BRIT.ORG/SIDA 22(1) 4, primarily in a band through the center of the study site. This fire occurred towards the end of the research collection period and thus had little impact on data collection. Management Regimes.— The management plan devised and implemented by SWF WMD (1990) includes several conceptual land uses. The most impor- tant function for the District is to manage the well field withdrawals. The con- sumptive use permit renewed in 1979 allows the city of New Port Richey to withdraw 30 to 57 million liters per day from the 14 wells within the Preserve (SWFWMD, 1990). This water withdrawal has produced shorter hydroperiods, lower peak water levels, increased rates of succession into wetland areas, and unusual plant associations (SWFWMD 1990). To manage areas of historical significance, SWF WMD protects them from disturbance by keeping hiking and horse-riding trails away from them. SWFWMD has opened the Preserve for “low intensity resource-based” recre- ation by building facilities for local use (SWFWMD 1990). These recreational facilities are managed by Pasco County. They include hiking, biking, horse riding, birding, camping, and any other non-motorized activities. The hiking/ biking trails are separate from the horse riding trails to avoid safety hazards. Land management practices exercised by SWFWMD include a number of ways to improve or preserve the quality of the property. The District uses pre- scribed burning to conserve various communities and to restore habitats. Dur- ing the summer months, when there is greater thunderstorm activity, higher lightning frequency giving way to natural wildfires which SWFWMD contains and controls. The District normally burns during the winter months in order to limit and control fires (SWFWMD 1990). A population of feral hogs has disturbed the soil while rooting, causing damage to some habitats and previously preserved historical sites. Part of the management plan is to reduce or minimize their impact. METHODS Vascular plant specimen vouchers were collected from May 2003 to October 2004 using standard field collection and herbarium techniques. Notes were made for each collection regarding the habitat and frequency of occurrence. One complete set of voucher specimens was deposited at the University of South Florida Herbarium (USF). The floras of Wunderlin (1998) and Wunderlin and Hansen (2003) were utilized for identifications, followed by verification with specimens in the USF herbarium. Vascular plants of special interest include those taxa that are Federal and/ or State listed species and exotic species listed by the Florida Exotic Pest Plant Council (FLEPPC 2005). Species unreported for Pasco County as determined by Wunderlin and Hansen (2005) are noted. The plant community classification for the study site follows that of Florida FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PERSERVE, FLORIDA 641 Natural Areas Inventory (FNAI1990). The plant communities are described and include the dominant species found within each layer (overstory, understory, and herbaceous ground layer). RESULTS AND DISCUSSION A total of 483 taxa were collected within the study site within Starkey Wilder- ness Preserve. The flora is comprised of 104 families and 271 genera (Table D). The families with the greatest number of taxa are: Ast (66), Poaceae (62), Cyperaceae (39), and Fabaceae (28). The genera with the most abundant taxa are: Rhynchospora (12), Quercus (10), Hypericum (9), Andropogon (8), Ludwigia (8), and Dichanthelium (8). About 92% of the flora in the study site is native. Sixteen (3%) of the spe- cies found are endemic to Florida. Four of the 39 non-native taxa are listed by the Florida Exotic Pest Plant Council (FLEPPC 2005). Thirty-three taxa are records for Pasco County (Wunderlin & Hansen 2005). Twelve taxa are listed as endangered, threatened, or commercially exploited in Florida (Coile & Gar- land 2003). No Federally listed taxa occur on the site. Starkey Wilderness Preserve consists of 18 plant communities. Eleven of these occur within the study site. Comparing the Preserve with the study site shows the study site is representative of the Preserve (Table 2). The plant com- munities that fall outside the boundaries of the study site are not dominant ones and contribute only 4% of the total area of the Preserve. Thus the plant list com- piled for the study site is fairly representative for the majority of the Preserve. NATURAL COMMUNITIES The classification for plant communities found within the study site following the Florida Natural Areas Inventory guidelines (FNAI 1990, 2004) are discussed below from the lowest elevation type to the highest followed by the ruderal areas. Floodplain Forest.—The floodplain forest is a forested wetland that con- sists of 64.3 hectares adjacent to the River. Flooding is closely linked to the rain events that occur throughout the summer months. The soil is Chobee (Stankey 1982) The dominant tree species that make up the overstory of the floodplain forest are Acer rubrum, Carpinus caroliniana, Fraxinus caroliniana, Ilex cassine, Nyssa sylvatica var. biflora, Persea palustris, Quercus laurifolia, Quercus virginiana, Sabal palmetto, Taxodium distichum, Ulmus alata, and Ulmus americana. Some of these are buttressed due to the long hydroperiods. Note- worthy epiphytic species are Encyclia tampensis, Epidendrum conopseum, Pleopeltis polypodioides, and Tillandsia spp. The woody understory includes Berchemia scandens, Campsis radicans, Cephalanthus occidentalis, Psychotria nervosa, Psychotria sulzneri, and Toxicodendron radicans. The herbaceous ground layer consists of a number of pteridophytes including Blechnum 642 BRIT.ORG/SIDA 22(1) serrulatum, Osmunda cinnamomea, Osmunda regalis, Thelypteris interrupta, Taste 1. Synopsis of the vascular plants found in the Jay B. Starkey Wilderness Preserve study area. Taxa Genera Families Endemics Non-native Co. Records Pteridophytes 13 9 7 0 1 2 Gymnosperms 7 4 3 0 0 0) Monocotyledons 164 71 24 2 17 14 Dicotyledons 299 187 70 14 21 17 Totals 483 27) 104 16 39 33 Taste 2, Summary of the plant communities in the Jay B. Starkey Wilderness Preserve and the study area. Plant Community SWP SA Plant Community SWP SA Basin Marsh 101.2 Oak Scrub 70.4 Basin Swamp 1,422.1 61.5 Pine Plantation 22.2 Clastic Upland Lake 10.9 Ruderal and Disturbed = 888.3 0.4 Depression Marsh 48.6 2.0 i Pine Scrub 206.0 20.2 Dome Swamp 2347 10.5 dhill 275.6 344 Dry Prairie 0.8 ate Flatwoods 3314 40.5 paieaiee Forest 746.2 64.3 Wet Flatwoods 79.3 Hydric Hammock 28.3 Wet Prairie 40.9 41 Mesic Seen 3,005.6 207.2 Xeric Hammock 227.0 4.1 Wood wardia areolata,and Woodwardia virginica. aa common are Carex gigan- tea, Hypoxiscurtissii, [ris hexagona, Rhyncl i Saururuscernuus, Scleria triglomerata, and Spiranthes odorata. A non-native species found along the unimproved trails into the floodplain forest is Urena lobata. Basin Swamp.—A basin swamp is a large irregularly-shaped depression that is not part of a river, although during the rainy season it may be connected via surface flow to the river. This region is vegetated by species similar to the floodplain forest. The soils in the study site are Chobee and Sellers mucky loamy fine sand (Stankey 1982); both of these are frequently inundated with water. The study site contains 61.5 hectares of basin swamp. The largest of this com- munity type in the study site is located in the middle of the lower half of the site; it was probably an oxbow originating from the Anclote River. The dominant overstory species include Acer rubrum, Carpinus caroliniana, Diospyros virginiana, Fraxinus caroliniana, Gordonia lasianthus, Ilex cassine, Nyssa sylvatica var. biflora, Persea palustris, laxodium distichum, Ulmus alata, and Ulmus americana. The dominant woody understory species are Cephalanthus occidentalis, Lyonia lucida, and Myrica cerifera. The herba- ceous ground layer consists primarily of Amphicarpum muhlenbergianum, FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PRESERVE, FLORIDA 643 Carex spp., Cladium jamaicense, Cyperus spp., Eleocharis spp., Eriocaulon decangulare, Juncus spp., Leersia hexandra, Panicum hemitomon, Peltandra virginica, Pontederia cordata, Rhexia spp. Sagittaria lancifolia, Saururus cernuus,and Scleria spp. The basin swamp also contains a number of epiphytic species such as Pleopeltis polypodioides, Tillandsia recurvata, Tillandsia simulata, Tillandsia usneoides, and Tillandsia xfloridana. Dome Swamp.—Dome Swamps are found throughout the study site. The study site contains 10.5 hectares of dome swamp. These circular to oblong de- pressions contain Sellers mucky loamy fine sand, a very poorly-drained soil type (Stankey 1982). An impermeable clay layer beneath the soil helps main- tain water levels in these depressions. The depression is due to the soils slump- ing into sinkholes formed in the limestone rock. Most of the water found in the swamp is surface runoff from surrounding upland communities. The dome swamp may become completely desiccated at the end of the dry season, expos- ing the peat layer for a couple of weeks. These areas have a domed profile with smaller trees in the shallow water near the edge and larger trees in the deeper water in the middle of the swamp. The dominant tree in the dome swamp is Taxodium ascendens. Other prominent overstory species include Acer rubrum, Diospyros virginiana, Ilex cassine, Nyssa sylvatica var. biflora,and Persea palustris. Typical woody under- story species are Lyonia lucida, Myrica cerifera, Smilax laurifolia,and Toxico- dendron radicans. The herbaceous ground layer includes the pteridophytes Thelypteris interrupta, Woodwardia areolata,and Wood wardia virginica. Other ground layer species are Drosera capillaris, Helenium pinnatifidum, Polygala cymosa, Polygonum hydropiperoides, Rhynchospora latifolia, Spiranthes laciniata,and Xyrisspp. The epiphytic flora is similar to that of the floodplain forest and basin swamp and includes Pleopeltis polypodioides, Tillandsia recurvata, lillandsia simulata, Tillandsia usneoides, and Tillandsia xfloridana. A rare find in one of the swamps was Vittaria lineata. Depression Marsh.—Depression marshes are characterized by low depres- sions in the flatwoods ity due to slumping around the edges of a sink- hole or water collecting on top of a hardpan subsurface layer. Two hectares of depression marsh occur in the study site. The soils are Adamsville fine sand and Basinger fine sand (Stankey 1982). These regions typically do not contain any overstory species. Woody spe- cies include Hypericum fasciculatum, Ilex gla bra, Myrica cerifera, and Stillingia aquatica. The herbaceous ground layer contain hicarpum muhlenbergianum, Carex spp., Cyperus spp., Eleocharis spp., Eriocaulon decangulare, Juncus spp., Lachnanthes caroliana, Panicum hemitomon, Rhexia spp., Sabatia grandiflora, Scleria spp.,and Xyris spp. Wet Prairie.—About 4.1 hectares of the study site is covered by this open wetland herbaceous community. It is often found within the mesic flatwoods 644 BRIT.ORG/SIDA 22(1) or surrounding the fringe of a dome or basin swamp. The soils are Basinger fine sand and Myakka fine sand (Stankey 1982). The area is poorly drained due to the subsoil layer of hardpan clay. The wet prairie includes some native invading woody species: Hypericum fasciculatum, Hex glabra, Myrica cerifera,and Stillingia aquatica. The herbaceous ground layer primarily consists of Aletris lutea, Amphicarpum muhlenbergianum, Aristida stricta, Carex spp., Cyperus spp., Drosera capillaris, Eleocharis spp., Eriocaulon decangulare, Eupatorium mohrii, Fimbrystylis spp., Juncus spp., Lachnanthes caroliana, Panicum hemitomon, Polygala spp., Rhexia mariana, Sabatia spp., Scleria spp., Syngonanthus flavidulus, and Xyris spp Mesic Flatwoods.—T he most dominant fire adapted community found in the study site (207.2 hectares) is mesic flatwoods. This community has a rela- tively flat topography. The dominant soils are Immokalee fine sand, Myakka fine sand, and Smyrna fine sand (Stankey 1982). These soils normally have low levels of nutrients and organic matter. A hardpan clay layer under the soil causes poor water drainage in the rainy season and xeric conditions during the dry season. The overstory layer is composed of Pinus elliotti and Pinus palustris. The pines are widely spaced across the community allowing for a dense woody un- derstory layer. The understory layer includes Ilex glabra, Lyonia lucida, and Ser- enoa repens. Smaller understory layer species are Gaylussacia frondosa, Quercus minima, Vaccinium corymbosum, and Vaccinium myrsinites. The rich herba- ceous ground layer includes Amphicarpum muhlenbergianum, Aristida stricta, Dichanthelium ensifolium, Dichanthelium portoricense, Elephantopus elatus, Eupatorium mohrii, Euthamia caroliniana, Pityopsis graminifolia, Polygala lutea, Polygala setacea, Pteridium aquilinum, Pterocaulon pycnostachyum, Seymerid cassioides, and Sorghastrum secundum. Sandhill.—The sandhill community makes up 34.4 hectares of the study ite. There are two areas; both are relatively high in elevation at 12.2-15.2 m and have topography consisting of rolling hills and gentle slopes. The soil of both areas is Adamsville fine sand (Stankey 1982), which is also found under the scrubby flatwoods community within the study site. The deep sandy soils that are easily leached and well drained create xeric characteristics in this commu- nity. The overstory layer is sparse, allowing sunlight to reach the ground layer, adding to the xeric condition. The dominant overstory species include Pinus palustris, Quercus incana, Quercus laevis, and Quercus margaretta. Understory species include Asimina obovata, Asimina reticulata, Diospyros virginiana, Gaylussacia dumosa, Licania michduxii, Myrica cerifera, oes minima, Rhus Se and Serenod repens. The herbaceous ground cover layet consists pl imat ily of Aristida stricta, Asclepias humistrata, Balduina angustyold. Dichanthelium portoricense, Phoebanthus grandiflorus, Pityopsis graminifolia, Pterocaulon pycnostachyum, Rhynchospora megalocarpa, and Sorghastrum secundum. — WY FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PRESERVE, FLORIDA 645 Scrubby Flatwoods.—This community consists of 40.5 hectares within the study area. The soil is Adamsville fine sand (Stankey 1982). The vegetation con- sists of a mixture of mesic flatwoods and sand pine scrub species. It is charac- terized by an open pine canopy with sparse scattered clumps of oak species and patches of open sand. The species found in the overstory include Pinus clausa, Pinus palustris, Quercus chapmanii, Quercus geminata, and Quercus myrtifolia. The dense un- derstory layer contains Garberia heterophylla, Lyonia ferruginea, Lyonia fruticosa, Serenoa repens,and Ximenia americana. The herbaceous ground layer contains Aristida spiciformis, Aristida stricta, Carphephorus corymbosus, Dichan- thelium portoricense, Lupinus diffusus, Pteridium aquilinum,and Rhynchospora megalocarpa. Several Pinus palustris have the epiphyte Tillandsia xfloridana. Sand Pine Scrub.—One area containing a scrub community totaling 20.2 hectares occurs within the study site along the western boundary. This com- munity develops along the ridgeline of ancient dune formations composed of very fine sand sometimes referred to as “sugar sand.” The soil is Pomello fine sand (Stankey 1982). These fine sands allow rain water to quickly percolate down to the aquifer creating a xeric growing condition. The scrub can have an open or closed canopy consisting of Pinus clausa with various oaks and shrubs dominating the understory. Normally this community type exhibits patches of exposed sand; however, in the study site these are very few. The scrub found within the study site has mature Pinus clausa, an indication of fire suppres- sion. Under the Pinus clausa, the dominant overstory also contains Quercus chapmanii, Quercus geminata, and Quercus myrtifolia. The understory shrub layer contains Asimina obovata, Ceratiola ericoides, Garberia heterophylla, Licania michauxii, Lyonia ferruginea, Lyonia fruticosa, Serenoa repens, and Ximenia americana. The herbaceous ground cover is sparse; however some com- mon herbs found there are Palafoxia intergrifolia, Pityopsis graminifolia, and Rhynchospora megalocarpa. Xeric Hammock.— Xeric hammock is generally considered a degraded fire- excluded scrub or sandhill community. Thi unity in the study site is an advanced sandhill community and consists of 4.1 hectares on the fringes of the northern sandhill community. The xeric hammock occurs on two soil types, Adamsville fine sand and Immokalee fine sand (Stankey 1982). The commu- nity lacks a closed overstory canopy, but a low canopy is present from the shrub layer dominated by oaks The overstory species consist of Pinus palustris (few), Quercus geminata, and Quercus laevis. The dense woody understory consists primarily of Licania michauxii, Lyonia fruticosa, Quercus chapmanii, Quercus myrtifolia,and Serenoa repens. The herbaceous ground es Sue ae stricta, Dichanthelium por- toricense, Galactia elliottii, Pteri mand Rhynchospora megalocarpa. L 646 BRIT.ORG/SIDA 22(1) Ruderal.—Ruderal areas within the study site include regions of distur- bance due to anthropogenic changes. This encompasses 0.4 hectare of road- sides, hiking trails, horse riding trails, well pump houses, improved pasture, and the fringe of the power line artery. These areas contain a high number of non- native species because of the recent disturbances caused by their construction and subsequent use. The most prominent non-native is Paspalum notatum followed by Crotalaria spp., Indigofera hirsuta, and Ludwigia peruviana. The Southwest Florida Water Management District (SWFWMD) has worked to manage these ruderal areas to encourage recruitment of native species. ANNOTATED LIST OF THE VASCULAR FLORA The vascular flora of the study site within the Starkey Wilderness Preserve in the following list is documented by vouchered specimens in the University of South Florida herbarium (USF). The list is divided into four major sections: ptert- dophytes, gymnosperms, monocots, and dicots (basal angiosperms and eudicots). Within these sections the list is arranged alphabetically by family, genus, and species. The nomenclature follows Wunderlin and Hansen (2003, 2005). Each species and infraspecific taxon is followed by a common name and its frequency of oes The frequency of occurrence is given as: R, rare less than four individuals found; O, occasional, between four and 15 plants aoe and C, common, more than 15 plants in the immediate area. Following the fre- quency of occurrence, the primary plant community from which the . was collected is noted. The plant communities include: basin swamp (BS); de- pression marsh (DM); dome swamp (DW); floodplain forest (FF); mesic flatwoods (MF); ruderal (RD); sandhill (SH); scrubby nee (SF); sand pine scrub (SP); wet prairie (WP); and xeric hammock (XH). Following this, taxa listed as en- dangered, threatened, or commercially exploited in Florida (Coile & Garland 2003) are noted. The number(s) in brackets at the end of each species are the collection number(s) of the first author. Species endemic to Florida are listed in bold font. Non-native species are marked with an asterisk (*). New records for Pasco County are indicated by an underline. Exotic species listed by the Florida Exotic Pest Plant Council (FLEEPC 2005) are listed with their ranking: Category I—species altering Florida’s natu- ral plant community by displacing native species and changing the structure of the community; Category I[—species having the ability in the future to alter Florida’s natural plant communities by displacing natives and changing the community structure. FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PRESERVE, FLORIDA 647 PTERIDOPHYTES Blechnaceae Blechnum serrulatum Rich.—toothed midsorus fern; C; DS (116, 498, 538) Woodwardia areolata (L.) T. Moore—netted chain fern; C; FF (552, 822) Woodwardia virginica (L.) Sm.—Virginia chain fern; C; FF (523, 499, 418) SL haa a Pteri aquilinum (L.) Kuhn © var. ae eee (Clute) Clute ex A. Heller— tailed bracken fern; C; MF (249, 823 Lycopodiaceae Lycopodiella alopecuroides (L.) Cranfill—foxtail club-moss; R; BS (266) Osmundaceae Osmunda cinnamomea L.—cinnamon fern; C;FF; commercially exploited (347) Osmunda regalis L. var. spectabilis (Willd.) A ray—royal fern; O; FF; commercially ex- ploited (553,655) — Polypodiaceae Poe aureum (L.) J. Sm.—golden poly- pody; O; DS (579) Pleopeltis pobpodotes (L.) E.G. Andrews & Windham var. michauxiana (Weath.) E.G. Andrews & on eee fern; C; FF (398) Thelypteridaceae *Thelypteris dentata (Forssk.) E.P. St. Jonn— downy maiden fern; O; FF (803) lyoteris interrupta (Willd.) K.lwats.—hottentot fern; C; FF (354) Thelypteris kunthii (Desv.) C.V. Morton—wide- spread maiden fern; C; FF (554) The — Vittariaceae Vittaria lineata (L.) Sm.—shoestring fern; R; DS (765) GYMNOSPERMS Cupressaceae Juniperus virginiana —red cedar; R; RD (782) Taxodium lens Brongn.—pond-cypress;C; S (170) Taxodium distichum (L.) Rich.—bald-cypress; C; Pinaceae Pinus clausa (Chapm. ex Engelm.) Vasey ex rg.—sand pine; C; SF (290) Pinus elliottii Engelm.—slash pine; C; MF (595) Pinus palustris Mill—longleaf pine; C; MF (581) Zamiaceae Zamia pumila L.—Florida arrowroot;R; MF;com- mercially exploited — Ww — oO = MONOCOTS Agavaceae Yucca filamentosa L.—Adam’s needle; O; SH (91) Alismataceae Sagittaria graminea Michx. var. graminea— 15, 380) ee a a inea Michx. var. chapmanil J.G. d; O; FF (480) grassy arro\ shead:O; DS (1 rrowhea —Chapman’'s a a ittaria lameifal ia L—bulltongue arrowhead; Amaryllidaceae Zephyranthes atamasca (L.) Herb. var. treatiae (S. Watson) Meerow—Treat’s rainlily; R; FF; renee (665) Araceae Arisaema Hak (L.) Schott—Jack-in-the- pulpit; R; FF (740 Lemnava liane, Phil—Valdivia duckweed; C;BS 2\5 Peltandra virginica (L.) Schott—green arrow arum; R; BS (776) ecaceae Sabal minor (Jacq.) Pers.—dwarf palmetto; C; FF 316 Sabal palmetto (Walter) Lodd. ex Schult. & Schult.£.—cabbage palm; O; FF (794) Serenoa repens (W. Bartram) Small—saw pal- metto; C; MF (80) Bromeliaceae Ts if y (ret L 494) Tillandsia recurvata (L.) L—ballmoss; C; SH (270) Tillandsia setacea Sw.—southern neecleleaf; C; FF (751) Tillandsia simulata Small—airplant; C; FF (305, = — Tillandsia usneoides (L.) L—Spanish moss; CG; SH (452) 648 Tillandsia utriculata L.—giant airplant;O; SH; en- dangered (837) Tillandsia Xfloridana (L.B. Sm.) H. Luther; C; SF Burmanniaceae Burmannia capitata (J.F.Gmel.) Mart.—southern bluethread; O; DS (509) Commelinaceae Callisia ornata cD G, : eo scrub seling; O; MF (75 oe erecta L. ane dayflower;O; RD (77, 781) Cyperaceae “Bulbostylis barbata (Rottb.) C.B. Clarke— watergrass; O; RD (838) Carex gigantea Rudge—giant sedge; C; FF (556, 661) Carex longi Mack.—Lonq's sedge; C; FF (546, 558, 602, 830) Carex verrucosa Muhl.—warty sedge; C; DS (383 06) Cladium jamaicense Crantz—Jamaica swamp sawgrass; C; DS coc Cyperus croceus Vahl | lge;C;D (224) Cyperus haspan L.—haspan flatsedge; C; DM (571, 539) Cyperus odoratus L.—fragrant flatsedge; C; FF (545, 774) Cyperus polystachyos Rottb.—manyspike flatsedge; C; DM (160, 243, 295) Cyperus retrorsus Chapm.—pinebarren flatsedge; C; DM (136, 247, 293,410 Ta) | 4 C; DM (104, 232, 294, 437) Eleocharis baldwinii (Torr.) Chapm.—Baldwin's spikerush; C; DS (423, 430, 876) Eleocharis flavescens (Poir.) Urb.—yellow spikerush; C; WP (231, 547) Eleocharis vivipara Link—viviparous spikerush; C; (836) Fimbristylis autumnalis (L.) Roem. & Schult.— slender fimbry; C; RD (585,878) Fimbristylis cymosa R.Br—hurricanegrass;O;DS (515) Fimbristylis dichotoma (L.) Vahl—forked fimbry; *Fimbristylis schoenoides (Retz.) Vahl—ditch fimbry; O; RD (851,879, 880) BRIT.ORG/SIDA 22(1) Fimbristylis puberula (Michx.) Vahl—hairy fimbry; O; WP (221, 864) Fuirena breviseta (Coville) Coville—saltmarsh mbrella-sedge; C; DS (169, 513) Fulrena pumila (Torr.) Soreng.—dwarf umbrella- sedge; C; DS (233) Fulrena scirpoidea Michx.—southern umbrella- sedge; C; DS (429) Lipocarpha maculata (Michx.) Torr—American halfchaff sedge; O; DS (847) Rhynchospora cephalantha A. Gray—bunched eaksedge; C; FF (521) Rhynchospora colorata (L.) H. Pfeiff—starrush whitetop; O; WP (111) Rhynchospora corniculata (Lam.) A. Gray— shortbristle horned beaksedge; O; RD (833) ee fascicularis (Michx.) Vah|—fas- CDM (122, 135, 182, 255, aye 7,485) Rhynchospora _fernaldii Gale—Ffernald's MF (329 Rhynchospora inundata (Oakes) Fernald— narrowfruit horned beaksedge; C; DS (223, 5,888 ote aoe latifolia (Baldwin) W.W.Thomas— giant whitetop; O; DS (758) Rhynchospora Gals A.Gray—sandyfield beaksedge; C; SP (432, 786) arpa nee ex A. Gray— southern beaksedge; C; DM (831) Rhynchospora microcephala (Britton) Britton ex Small—bunched beaksedge; C; DS (529, 573) Rhyncospora miliacea (Lam.) A. Gray—millet beaksedge; C; FF (352, 687) Rhynchospora plumosa_ Elliott—plumed beaksedge; C; MF (189, 416, 678, 681) Scirpus cyperinus (L.) Kunth—woolgrass; O; DM Rhynchosnorg micr re 80) Scleria baldwinii eh Steud.—Baldwin’s nutrush; C; WP (742) Scleria reticularis Michx.—netted nutrush; C;WP 428,859 MF (119) Hyptis alata (Raf.) Shinners—musky mint; C; DM (274, 331, 345) *Hyptis are is (Rich.) bushmint; O; FF (173, 276) Lycopus rubellus Moench—taperleaf water- rehound; F;DM (511) Piloblephis rigida (W. Bartram ex Benth.) Raf— wild pennyroyal; O; MF (63 a alvia eae O; BS (606) mall—Florida scrub skull- Brig.—tropical larin nreni cap; C; SH (132) Teucrium canadense L.—woodsage;C; DM (113) Trichostema dichotomum L.—forked bluecurls;C; MF (377) Lauraceae *Cinnamomum camphora (L.) J. C hortree; R; BS; FLEPPC-I (734) Persea palustris (Raf.) Sarg—swamp bay; C; FF (289, 353) Pres|— Lentibulariaceae Pinguicula pumila Michx.—small butterwort; O; MF (588) Utricularia foliosa L_—leafy bladderwort; O; DS O; BS (626) Utricularia juncea Vah|—s ba | wthern bladderwort: Utricularia subulata L.—zigzag bladderwort; O; Linaceae Linum medium (Planch.) Britton var. texanum (Planch.) Fernald—stiff yellow flax; O; WP (797) Logan oor eee (.F. Gmel.) Torr. & A. Gray—lax or 97) Mitreola sessilifolia (J.F. Gmel.) G. Don—swamp ornpod; C; WP (148, 370, 379, 852) Lythra ae nee (Jacq.) J.F. Macbr.—Co- lombian waxweed; C; FF (219, 508, 799) 656 Lythrum alatum Pursh var. lanceolatum (Elliott) rr.& A. Gray ex Rothr.—winged loosestrife; O; MF (277) Magnoliaceae Magnolia virginiana L.—sweetbay; R; DS (658) Malvaceae *Urena lobata L.—caesarweed; C; FF; FLEPPC-I| (500) Melastomataceae Rhexia cubensis Griseb.—West Indian meadow- beauty; O; DS (183 Rhexia mariana L._—pale meadowbeauty; C; DS (84, 697 Rhexia nuttallii CW. James—Nuttall’s meadow- beauty; O; MF (185, 506) Myricaceae Myrica cerifera L—wax myrtle; C; MF (332, 693) Nymphaeaceae Nymphaea odorata Aiton—American white WatenINys O; ae 2) 4 Aj Nuphar UGVETIC VANILOTT) F | BS (754) Olacaceae Ximenia americana L.—tallow wood; O; SP (460) eaceae Fraxinus caroliniana Mill.—Carolina ash; C; FF (107, 337, 778) Onagraceae Gaura angustifolia Michx.—southern beeblos- m; O; MF (125, 128) pce linearis Walter—narrowleaf prim- willow; C; DS (280, 385,451) eee linifolia Poir.—southeastern prim- rosewillow; C; DM (265, 699) Ludwigia maritima R.M. Harper—seaside prim- rosewillow; O; MF (147, 201, 263) Ludwigia dial Michx.—smallfruit prim- rosewillow; O; DS (297) Ludwigia ets (Jacq.) Raven—Mexican primrosewillow; O; DM (384, 541) “Ludwigia peruviana (L.) H. Hara—Peruvian primrosewillow; O; RD (825) Ludwigia repens J.R. Forst.—creeping prim- rosewillow; C; FF (772) Ludwigia suffruticosa Walter—shrubby prim- rosewillow; O; DS (240) BRIT.ORG/SIDA 22(1) Oenothera laciniata Hill—cutleaf eveningprim- se; O; RD (204, 638) Orobanchaceae Agalinis fasciculata (Elliott) Raf —beach false fox- glove; C; MF (359, 381) Agalinis linifolia (Nutt.) Britton—flaxleaf false floxglove; O; DM (846) Agalinis tenuifolia (Vahl) Raf—slenderleaf false -O; MF (502) Buchnera americana L.—American bluehearts;O; MF (123, 755) ee cassioides (J.F. Gmel.) S.F. Blake yaupon blacksenna; O; MF (237,376 422) ie pectinata Pursh—piedmont black- senna; O; SH (368) — x< ‘Q Oxalidaceae Oxalis corniculata L—common yellow wood- sorrel; O; RD (617) Passifloraceae Passiflora incarnata L_—purple passionflower;O; D (155) Plantaginaceae Plantago virginica L.—Virginia plantain; O; RD (648) Polygalaceae Polygala cymosa Walter—tall pinebarren milk- wort; O; DS (724) Polygala lutea L—orange milkwort; CG; MF (83, 188) Polygala nana (Michx.) DC._—candyroot;C; MF Polygala rugelii Shuttlew. ex Chapm.—yellow milkwort;O;WP (175 / i) NAA ta lAlai rie pe) C; MF (98, 211, 748) Polygala violacea Aubl.—shov y milkwot t;C; MF (93) Polygonaceae Polygonella gracilis Meisn.—tall jointweed; O; SH (367, 404, 567) Polygonella polygama (Vent.) Engelm. & A. Gray—October flower; C; SH (395, 461) Polygonum hydropiperoides Michx.—swamp smartweed; C; DS (114, 399, 482) Polygonum punctatum Elliott—dotted smart- weed; C; RD (400) Rumex hastatulus Baldwin—heartwing dock; O; RD (719) FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PRESERVE, FLORIDA 657 Portulacaceae *Portulaca amilis Soeg.—Paraguayan purslane; O; RD (8 08) Primulaceae Samolus valerandi L. subsp. parviflorus (Raf. Hulten—pineland pimpernel; O; FF (349) a Rhamnaceae Berchemia scandens (Hill) K. Koch.—rattan vine; O; FF (336) Rosaceae Photinia pyrifolia (Lam.) K.R. Robertson & J.B. Phipps—red chokecherry; O; DS (628) Prunus caroliniana (Mill.) Aiton—Carolina laurelcherry; O; RD (631) Prunus serotina Ehrh.—black cherry; O; RD (632) Rubus argutus Link—sawtooth blackberry;C; DS (645, 663) Sapindaceae Acer rubrum L.—red maple; C; FF (590) Saururaceae Saururus cernuus L.—lizard’s tail; C; FF (585) Solanaceae Solanum americanum Mill.—American black nightshade; O; FF (593) Tetrachondraceae Polypremum procumbens L.—rustweed; C; RD (130) Theaceae Gordonia lasianthus (L.) J. Ellis—loblolly bay; O; BS (288, 559) Ulmaceae Ulmus alata Michx.—winged elm; O; FF (682) Ulmus americana L.—American elm; O; FF (619 Rubus cuneifolius Pursh—sand blackberry:C:MF 35) (730) ah Ulelcaceas uthern dewberry;O; FF ylindrica (L.) Sw.—false nettle; C; DS (792) (342, 584) biaceae Cephalanthus occidentalis L—common button- bush; C; DS (524, 770) Diodia teres Walter—rough buttonweed; C; SH (304) Diodia virginiana L—Virginia buttonweed;C;BS (106, 199, 388) Galium tinctorium L.—stiff marsh bedstraw;C; RD (598) Houstonia ee tie (J.F. Gmel.) Standl.—in- e; C; MF (589, 604) Mee ay repens L.—partridgeberry; O; FF 688) — 340, Oldenlandia uniflora L.—clustered mille graine; C; DS (413, 882 — ) Psychotria nervosa wild coffee; C; FF (339) a sulzneri ie coffee; FF (338 ey Sie +. coal AA clover; CG; RD (202) Spermacoce assurgens Ruiz & Pav.—woodland false buttonweed; O;DM (241) ean prostrata Aubl.—prostrate false buttonweed; O; FF (299) Salicaceae Salix caroliniana Michx.—Carolina willow; C; RD (633) Verbena Phyla ae nee (L.) Greene—turkey tangle fogfruit; C; RD (89) Bae brag iensis Vell—Brazilian vervain; O; D (775) ie scabra Vah|—sandpaper vervain; O; MF (849) Veronicaceae Bacopa monnieri (L.) Pennell—herb-of-qrace; O; DM (625 Gratiola hispida (Benth. ex Lindl.) Pollard-——rough gehyssop; O; MF (140, 141) Gratiola pilosa Michx.—shaggy hedgehyssop;O; B , 167, 766) Gratiola ramosa Walter—branched hedgehys- sop; O; DS (698) Linaria canadensis (L.) Chaz —Canadian toadflax; C; DS (622, 643) *Lindernia crustacea (L.) F. Muell—M false pimpernel; O; RD (440) Mecardonia acuminata (Walter) Small subsp. peninsularis (Pennell) Rossow—axilflower;C; WP (230, 743) Micranthemum glomeratum = (Chapm.) ners—manatee mudflower;O;WP (623) ee dulcis L—sweetbroom; C; MF (151) alaysian Violaceae Viola lanceolata L—bog white violet; O;DS (624) Viola palmata L.—early blue violet; O; MF (647) Viola primulifolia L.—primroseleaf violet; C; DS BRIT.ORG/SIDA 22(1) Vitaceae Ampelopsis arborea (L.) Koehne—peppervine;O; 50) Parthenocissus quinquefolia (L.) Planch.—Virginia ) Viola sororia Willd—common blue violet;O;FF Vitis rotundifolia Michx. == CeIn) C;MF (360) 21) Vitis shuttleworthi House -O;MF (736) Viscaceae Phoradendron leucarpum (Raf.) Reveal & M.C. Johnst.—oak mistletoe; O; BS (465) ACKNOWLEDGMENTS We are grateful to Darrell Freeman and Eugene Kelly of the Southwest Florida Management District for providing collection permits, access to the property, and other kinds of assistance. We thank Doug Coleman and John Kunzer for assistance in the field. We also thank Bruce F Hansen for assistance in plant identification and for his helpful comments on the manuscript and Kathleen Hotchkiss for assistance with graphics. REFERENCES CHEN, E.and J.F.Gereer. 1990. Climate. In: R.L. Myers and J.J.Ewel,eds. Ecosystems of Florida. University Presses of Florida, Gainesville. Pp. 11-34. Coie, N.C.and M.A. Gartano. 2003. Notes on Florida’s endangered and threatened plants. Florida Department of Agriculture and Consumer Services, Gainesville. FLorIDA Exotic Pest PLANT Council (FLEPPC). 2005. List of Florida's invasive species. Florida Exotic Pest Plant Council. (http: FLoriIDA NaturaAt AREAS INveNToRY (FNAI). Tallahassee. FLORIDA NATURAL AREAS INVENTORY (FNAI). 2004. Natural community mapping project of Starkey Wilderness Preserve, Southwest Florida Water Management District. Florida Natural Areas Inventory, Tallahassee. Lawson, S.F., R.P. INcatts, and C. Bayess. 1981. Preserving for the future: A history of the J.B. Starkey Wilderness Park. Department of History, University of South Florida, Tampa. Reprinted by Southwest Florida Water Management District, Brooksville. Mitanicu, J.T. 1994, Archaeology of Precolumbian Florida. University Press of Florida, Gainesville. Scott, 1.M, K.M. CAMPBELL, F.R. Rupert, J.D. ArtHur, T.M. Missimer, JM. Ltoyp, JW. Yon, and J.G. Dun- cAN. 2001. Geologic map of the state of Florida. Florida Geological Survey in coopera- tion with the Florida Department of Environmental Protection.U.S. Department of the Interior, U.S. Geological Survey, Center for Coastal Geology, Washington. SouTHwest Florina Water MANAGEMENT District (SWFWMD).1988.Ground-water resource avail- ability inventory: Pasco County, Florida. Brooksville VV ww.fleppc.org). 1990. Guide to the natural communities of Florida, FERGUSON AND WUNDERLIN, INVENTORY OF STARKEY WILDERNESS PRESERVE, FLORIDA 659 SoutHwest Floriba WaTER MANAGEMENT District (SWFWMD). 1990.A plan for the use and man- agement of the Starkey tract. Brooksville Sankey, D.L. 1982. Soil survey of Pasco County, Florida. U.S. Department of Agriculture, Soil Conservation Service in cooperation with University of Florida, Institute of Food and Agriculture Sciences Department and Florida Department of Agriculture and Con- sumer Services. U.S. Government Printing Office, Washington. United States Geotocicat Survey (USGS). 1974.Odessa Quadrangle, Florida 7.5 Minute Series (topographic). United States Geological Survey, Department of the Interior, Washington Weis, M.2004. Pasco county property appraiser. Dade City. (http://appraiser.pascogov.com/). WETTERHALL. W.S. 1964. Geohydrologic reconnaissance of Pasco and southern Hernando Counties, Florida. Florida Geological Survey Report. Florida Geological Survey, Talla- hassee. Witey,G.R.1949. Archeology of the Florida Gulf Coast. Smithsonian Miscellaneous Collec- tions, 113:1-559. Smithsonian Institute, Washington. Wunoerun, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida, Gainesville. WunNDERLIN, R.P.and B.F. HANseN. 2003. Guide to the vascular plants of Florida, 2nd ed. Univer- sity Press of Florida, Gainesville. Wunpbertin, R.P. and B.F. Hansen. 2005. Atlas of Florida vascular plants [S.M. Landry and K.N. Campbell (application development), Florida Center for Community Design and Re- search] Institute for Systematic Botany, University of South Florida, Tampa. (http:// www.plantatlas.usfedu). BRIT.ORG/SIDA 22(1) Book NOTICE JULIAN A. STEYERMARK, PAUL E. Berry, Kay YatskiEvycH, and Bruce K. Horst (Eds.). Flora of the Venezuelan Guayana. Vol. 9, Rutaceae-Zygophyllaceae. (ISBN 1-930723-47-4, hbk.). Missouri Botanical Garden Press, PO. Box.299, Saint Louis, MO 63166-0299, U.S.A. (Orders: http://www.mbgpress. EOE Oe 608 pp., 502 b/w line drawings, 71/4" x 10 1/2". The final volume of the series, concluding the taxonomic treatment of all native and naturalized vascular plant families known to occur in the Venezuelan Guayana. Forty-six contributors in Vol- ume a treat 48 uli oe eeners and 971 species lystrated es he figures. C eratophyllaceae is (from Theagens: Muntingiaceae (from Tiliaceae), oF Semaine (from Simaroubacene)~ —Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, Editor’s note—Two new names are published: Sterculia kayae PE. Berry, nom. nov. a Xyris sulcatifolia Kral, nom. nov. SIDA 22(1): 660. 2006 VASCULAR FLORAS OF SONOITA CREEK STATE NATURAL AREA AND SAN RAFAEL STATE PARK: ARIZONA'S FIRST NATURAL-AREA PARKS Steven P. McLaughlin Offi FA id | q eoales ay ed at THE GEIIVE Tucson, Aaa 85721 U S A. spmcl@ag.arizona.edu ABSTRACT J CRC - ft Sonoita Creek State N 1 Area (SCSNA) and San Rafael State P two natural- area parks in Arizona established with funding provided by the Heritage Initiative. Both sites are in Santa Cruz County, and both have riparian communities that are rare in the arid Southwest. The upland vegetation of SCSNA is primarily semidesert grassland, while that of SRSP is Plains grass- land. The flora of SCSNA includes 525 native species and 36 exotic species; that of SRSP includes 407 native species and 50 Significant taxa include one new species, Phacelia sonoitensis, found at SCSNA. Taxa new to fie flora of Arizona include Pennellia robinsonii Pa Aes minima (SCSNA), and Sibara virginica (SRSP). Lilaeopsis schaffneriana ssp. recury species, was found at SRSP. The ies of both SCSNA and SRSP are strongly eee in their ee ties. Species found at SCSNA but not at SRSP have relatively greater affinities with the flora of the Sonoran Floristic Province, while those ae at SRSP but not SCSNA have a greater affinity with the Great Plains. RESUMEN Sonoita Creek state Nancie ance SONA) y San Rafael State Park (SRSP) son los primeros parques aaa ge Herencia. Ambos lugares estan en el ondado de Santa Cruz, i id raras en el Suroeste arido. La vegeta del aaeao de SCSNA es un pastizal semidesértico y la de SRSP es un pastizal de las andes Llanuras. La flora de SCSNA ee 525 pees taclesnes y 36 a sce exoticas; i de incluye 407 especies indigenas y 50 una especies nueva, pagers noite nsis, s encontrada € en eae Entre De especies nuevas para la flora de Arizona se incluye ), y Sibara virginica (de SRSP). ent schaffneriana ssp. recurva, una especie en peligro fue aren en SRSP. Las floras de SCSNA y SRSP son fuertemente Apachianas en sus afinidades floristicas. Las especies encontradas en SCSNA pero no en SRSP tienen una afinidad mas grande con la flora de la provincia floristica de Sonora y las especies encontradas en SRSP pero no en SCSNA tienen una uras. afinidad mas grande con las Grandes Llanur INTRODUCTION Arizona voters passed the Heritage Initiative by a nearly two-thirds margin in the November 1990 general election. The Heritage Initiative allocates funds from the state lottery to Arizona State Parks and Arizona Game and Fish Depart- ment. Part of the funds for Arizona State Parks are designated for the acquisi- SIDA 22(1): 661 ~ 704, 2006 662 BRIT.ORG/SIDA 22(1) tion of “natural areas,” defined in the initiative as “parcels of land or water that contain examples of unique natural terrestrial or aquatic ecosystems, rare spe- cies of plants and animals, or outstanding geologic or hydrologic features.” The first two acquisitions of large acreage for natural areas were both in Santa Cruz County, in southern Arizona, adjacent to the Mexican state of Sonora. The Sonoita Creek State Natural Area (SCSNA) in central Santa Cruz County was purchased in January 1994 for $2.8 million. The natural area feature justi- fying this purchase was Sonoita Creek and its associated gallery forest. Sonoita Creek is one of the few perennial or near perennial streams in southern Ari- zona. The SCSNA purchase included 4.5 km of perennial stream, along with 1,990 ha of watershed north of the creek, including intermittent wetlands in the streams of Fresno Canyon and Coal Mine Canyon. Sonoita Creek, Fresno Canyon, and Coal Mine Canyon provide habitat for the Gila topminnow (Poeciliopsis occidentalis occidentalis), a federally listed endangered species of fish. The second acquisition was San Rafael State Park (SRSP) in extreme south- eastern Santa Cruz County. The 8,900 ha San Rafael Land and Cattle Company, which included the San Rafael de la Zanja land grant, was purchased by the Nature Conservancy in December 1998 for $1] million. Arizona State Parks pur- chased the 1,440 ha of patented land along the Arizona-Sonora border along with a conservation easement on the entire ranch from the Nature Conservancy for $8.6 million in January 1999. Natural-area values cited for this purchase in- cluded the unique grassland communities of the San Rafael Valley and signifi- cant riparian and wetland habitat along the upper Santa Cruz River and at Sharp Spring and Heron Spring. The wetlands provide critical habitat for at least two endangered species, the Huachuca water umbel (Lilaeopsis schaffneriana ssp. recurva) and Gila topminnow. Most of the watershed adjacent to SCSNA is mapped as semidesert grass- land (Brown 1994), which is an extensive area in Arizona and adjacent New Mexico, Sonora, and Chihuahua varying in physiognomy and species compo- sition. Grasses generally are not the dominant lifeform; instead there is a com- plex mixture of shrubs, small trees, succulents, rosette plants, and subshrubs, with the exact physiognomy and species composition determined by drought, the seasonal distribution of precipitation, periodic freezes, soils, grazing his- tory, and fire (Burgess 1995). Burgess suggested the term “Apacherian mixed shrub savanna” for this complex mosaic of subtropical, semiarid vegetation found above desert and below oak savannas. In contrast, the San Rafael Valley is mapped as “Plains grassland” (Brown 1994) and is dominated by perennial bunchgrasses and oak savannas with very few leguminous trees and shrubs, subshrubs, cacti, or rosette succulents. The primary objective of this study was to inventory the floras of these two natural area parks. The study also provided an opportunity to address a MCLAUGHLIN, FLORA OF SONOITA CREEK NATURAL AREA AND SAN RAFAEL STATE PARK 663 second objective, to contrast the floras of semidesert grassland and Plains grass- lands in southeastern Arizona. STUDY AREAS Table 1 lists the latitude, longitude, area, elevational relief, mean annual pre- cipitation, and mean January and July temperatures for both SCSNA and SRSP. Below are brief descriptions of the major physical features, soils, vegetation, and recent history of each park. Sonoita Creek State Natural Area The SCSNA is located along and north of Sonoita Creek downstream from Patagonia Lake State Park. It isan area of rugged terrain with moderate to steep, rocky slopes. Fresno Canyon hasan intermittent stream and extends from north to south across the center of the park. Coal Mine Canyon is an east-west trend- ing tributary to Fresno Canyon; the north-facing slopes of Coal Mine Canyon provide the most mesic terrestrial habitats within the park. A spring in the northeast corner of the park appears to be permanently wet. Elevations range from 1,080 m near the junction of Sonoita Creek and Fresno Canyon in the south- west corner of the park, to 1310 m at a point near the northern boundary. Fig- ure 1 includes a map of SCSNA showing the major drainages, representative upland and riparian habitats are shown in Figure 2. The exposed surface of the uplands adjacent to Sonoita Creek is that of the relatively impervious Nogales Formation, formed by the deposition of fine to coarse-grained alluvial fan deposits derived from pre-existing volcanic rocks. These deposits have been subject to subsequent folding and erosion, creating a complex topography of incised drainages and slopes of all aspects. Upland soils at SCSNA include: (1) Graham soils, mostly gravelly or cobbly loams on more gentle slopes at lower elevations in canyons, usually at the base of tuffaceous mountains; (2) Lampshire-Graham-Rock Outcrop associations, shallow, cobbly loams on steeper slopes, and (3) Rock Outcrop-Lithic Haplustolls, cobbly loams found in the dissected terrain in the southeast corner of the park (Richardson et al. 1979). The soils on the flood plain of Sonoita Creek are deep, well-drained sandy loams of the Comoro Association. The gallery forest along Sonoita Creek is dominated by cottonwood (Populus fremontii), ash (Fraxinus velutina), willows (Salix gooddingii and S. bonplandiana), and walnut (JJuglans major). On terraces adjacent to the gallery forest there is a dense bosque of mesquite (Prosopis velutina), hackberry (Celtis reticulata), elderberry (Sambucus mexicana), and one large colony of tree of eaven (Ailanthus altissima) The vegetation of the upland portions of SCSNA is mostly semidesert grass- land, with small sections of oak (Quercus emoryi and Q. oblongifolia) savanna, mostly confined to the north-facing slope of Coal Mine Canyon. Species com- position of the semidesert grassland is highly variable. The most common plants 664 BRIT.ORG/SIDA 22(1) Taste 1. Physical features of Sonoita Creek State Natural Area (SCSNA) and San Rafael State Park SRSP). Climate data are from Sellers et al. (1985); representative stations used are Nogales 6 N (31° 26'N,110° 55'W, 1160 m elev.) for SCSNA and San Rafael Ranch (31° 21' N, 110° 37' W, 1446 m elev) for SRSP. Feature SCSNA SRSP Latitude 31° 30'N 31° 21'N Longitude 110° 54'W 110° 36'W Area (ha) 1990 1440 Minimum elevation (m) 1080 1395 Maximum elevation (m) 1310 1475 Elevational relief (m) 230 80 Mean annual precipitation (mm) 430 440 Mean January temperate (°C) 74 5.6 Mean July temperature (°C) 25.9 23.5 include mesquite, hopbush (Dodonaea viscosa), coral-bean (Erythrina flabelliformis), beargrass (Nolina microcarpa), sotol (Dasylirion wheeleri), yucca (Yucca thornberi), shindagger (Agave schottii), acacia (Acacia greggii), mimosa (Mimosa aculeaticarpa var. biuncifera), and prickly pears (Opuntia spp.). Prior to acquisition the site was primarily used for cattle grazing: trespass cattle were frequently observed during the floristic inventory. There is also evi- dence of historic small-scale mining activity on the site. The New Mexico and Arizona Railroad was constructed along Sonoita Creek during 1881-1882. Its bridges washed out on many occasions, and after a particularly devastating flood in 1929 the section of the line along Sonoita Creek was abandoned (Myrick 1975). Sections of the railroad bed are evident along most of the reach of the stream, usually at the edge of the floodplain. San Rafael State Park The SRSP is located in the San Rafael Valley, an isolated valley bounded on the west by the Patagonia Mountains, on the north by the Canelo Hills, and on the east by the Huachuca Mountains. SRSP occurs at the southern end of the valley adjacent to the US-Mexico international border. The San Rafael Valley is the headwaters of the Santa Cruz River, which runs north to south down the cen- ter of the valley. During the period of field work for this study the Santa Cruz River ran continuously where it crosses the SRSP, supporting a variety of wet- land plant communities. Sharp Spring and Heron Spring are water courses flow- ing northeast to southwest into the Santa Cruz River: both have intermittent pools and support communities of wetland plants. Parker Canyon wash is a major drainage just entering SRSP in its southeastern corner. Figure lincludesa map of SRSP showing the major drainages; representative upland and riparian habitats are shown in Figure 3. RAI APFEA AND SAN RAFAFI STATE PARK 665 DCCVY NATII MCLAUGHLIN, FLORA OF SONOITA Sonoita Creek State Natural Area Morning Star Ranch i ov 7 spring ? c . c § Hl A : ‘\ Chivas Sige Sf ey *y, Tank r Cons 4 i : Mj ? \ ne Co §/ “om, on Rio Rico X . oS | Subdivision — | ) ss ; - ele : QA 7 * we Ss 2 1000 0 1000 2000 Meters — _ Park Headquarters (Historic Greene Ra ol k, Santa Cruz C ty, Arizona RA Fic. 1. Map Topography on the park consists mostly of low, gently sloping hills and shallow drainages. There are no bedrock outcrops. The Santa Cruz River is cur- rently entrenched about 2 to 4 m within its floodplain, which averages about 0.5 km in width. Soils along the floodplains of the Santa Cruz River, Sharp Spring, and Heron Spring are all mapped as Guest Soils, deep loams to clay loams (Richardson et al. 1979). The peepee soils of Parker Canyon are gravelly sandy loams of the Grabe-Com plex. Most of the upland areas have gentle slopes with grav- elly clay loans of the Bernardino-Hathaway Association, clay loams of the White House Association, or deep clay loams of the Pima Association. In the extreme southeast corner of SRSP on the south side of Parker Canyon, there are calcareous gravelly sandy loams of the Hathaway Association on slopes of 20 to 50 percent steepness. The high clay content of most of the soils of the San Rafael Valley is probably associated with their resilience to high grazing pres- sure and resistance to invasion by shrubs and cacti. The vegetation of the San Rafael Valley is primarily “Plains grassland”, a true grassland dominated by grama grasses (Bouteloua spp.), wolftail (Lycurus setosus), cane bluestem (Bothriochloa barbinodis), Plains lovegrass (Eragrostis ina), both on flood- bo an RY W TIQHIt], Yanler y £ (Cl ) Pre? (lower right). Fic 7 epee C¢ * tf a ge ee : Int Tene parse pea? soph a I I PI (Salix spp.) al S ita Creek (I Pi g 999 L)@Z VaIS/OYO'LIYS 10 VdO1d ‘NIHONV TDW MUVd JLVLS 1UVIVd NVS ONY Vad Fie 2 Tha be seit, +¢ l C4#a4 I 1 P| £C p6f TV. +h a | £D. L Lin Hy { * re - +) i fc Di a Se +4] IT ba \ kL bo x. j* . eae } ony . } y ) p ( pp.) adj | ghtii), cottonwood, and willow on fi Pe f1 1 £cClL L *1 I Dia I Md fl * L \ Pv \ t \ TIQHU). <) 668 BRIT.ORG/SIDA 22(1) intermedia), Texas bluestem (Schizachyrium cirratum), spider grass (Aristida ternipes), and other perennial bunchgrasses. Broadleaf annual and perennial herbs are an important component of the vegetation, particularly species in the families Euphorbiaceae, Asteraceae, and Fabaceae. The abundance of non- grass components in the vegetation is likely due to intensive grazing in the last 150 years. Oak savannas are present in the northwestern portion of the park and along Parker Canyon wash. There is an intermittent gallery forest along the Santa Cruz River, Sharp Spring, and Heron Spring composed mostly of cottonwood and willows (Salix gooddingii, S. lasiolepis, and S. exilifolia). Herbaceous vegetation along the Santa Cruz River is dominated by spikerushes (Eleocharis spp.) and true rushes (Juncus spp.). Grazing pressure in the wetlands along the Santa Cruz River, Sharp Spring, and Heron Spring appears to have been particularly heavy, and exotic species are locally abundant in these areas, particularly sweet clover (Melilotus officinalis), meadow fescue (Festuca arundinacea),and Kentucky blue grass (Poa pratensis). Most of the floodplain of the Santa Cruz River within SRSP was not cultivated, and this area contains an excellent example of sacaton (Sporobolus wrightii) grassland, a community that has disappeared from much of the South- west due to lowering of water tables. Hadley and Sheridan (1995) provided a detailed history of land use in the San Rafael Valley. The last 150 years have been a period of relatively intense cattle grazing. There have been numerous homesteads in the valley, and agri- culture has been practiced on a limited scale on the Santa Cruz River flood- plain. The San Rafael Ranch was the location for the filming of many movie westerns, including the musical Oklahoma! METHODS Field work was conducted from August, 2000, through November 2001. Dur- ing the 17 months of the project a total of 62 trips were made, 34 to SCSNA and 28 to SRSP. The field work thus covered three growing seasons: the summer/fall of 2000 and 2001 and the spring of 2001. The winter of 2000-2001 was unusu- ally wet, resulting in very productive collecting during spring of 2001. Since there was no direct vehicle access to the SCSNA, all of the collecting had to be done during long day-hikes. Plants were collected, mostly in duplicate, in plastic bags during the day and pressed either that same evening, or the following morn- ing after allowing collections to rehydrate somewhat. At SRSP most collections could be made on shorter hikes and pressed while still in good condition. A total of 1,286 specimens were collected, 689 from SCSNA and 597 from SRSP. Many trees, shrubs, and rosette succulents were not collected; rather their presence and distributions were recorded in field notes. Most cacti were identi- fied in the field using Benson (1969). A few species were included in the flora based on earlier collections from the study areas in the herbarium at the MCLAUGHLIN FIORA AREA AND SAN RAEAFI STATE PARK 669 University of Arizona. Rick Gagnon, the ranger at SCSNA during the period of the field work, took hundreds of photographs of plants he encountered on that study site; a few species not encountered during the collection trips but photo- graphed by Gagnon are also included in the flora. In 2004, 1,064 ha adjacent to SCSNA, located to the northeast and upstream along the Coal Mine Creek watershed, was added to the park. This area, which is even more remote that the original acquisition, is likely to have additional taxa not found during the field work for this study. Geographic affinities of the floras of SCSNA and SRSP were examined by mapping the percentage of native taxa found in 245 local floras from Mexico, the United States, and Canada, using a database compiled by the author. Such maps use contours (isolines of percentage overlap) to show in a concise way what proportion of the flora extends into neighboring geographic regions. Three maps were prepared: one for native species occurring in the floras of both SCSNA and SRSP, one for those occurring only at SCSNA, and one for those occurring only at SRSP. RESULTS AND DISCUSSION Floristic Summary The total known flora for SCSNA is 561 species, 525 native and 36 exotic. For SRSP the total flora is 457 species, 407 native and 50 exotic. The percentage of exotic species in the flora of SCSNA is 6.4%, a typical value for floras in south- eastern Arizona, but low for local floras in most other regions of the United States. Exotics at SRSP constitute 10.9% of the total flora. There are 277 species in common to the two sites, 252 native and 25 exotic. The floristic similarily (Otsuka’s Index, based on native species only) between SCSNA and SRSP is 54.5%, a somewhat low value for two areas in such close proximity (Fig. 1), indi- cating significant differences in the floras of semidesert and Plains grassland communities. There are 97 families and 354 genera in the flora of SCSNA, and 84 fami- lies and 274 genera at SRSP. The largest families in the two floras are listed in Table 2. The Asteraceae (composites), Poaceae (grasses), and Fabaceae (legumes) dominate in both floras, as they do in floras from most areas of the western United States. Even though the flora of SCSNA is larger than that of SRSP by more than 100 species, the number of Asteraceae are about the same and the number of Poaceae is actually larger at SRSP. The Poaceae also has the largest number of exotic species. The Euphorbiaceae is the fourth largest family at each site. One notable difference is the large number of Pteridaceae (ferns) at SCSNA, a family that is completely absent from the flora of SRSP Ferns in the Pteridaceae generally grow on rock outcrops or rocky slopes, a habitat not found at SRSP. The largest genera in the flora at SCSNA are Chamaesyce (Euphorbiaceae, 11 species), Bouteloua (Poaceae, 8 species), Cyperus (Cyperaceae, 8 species), Dalea 670 BRIT.ORG/SIDA 22(1) Taste 2. Largest families in the floras of Sonoita Creek State Natural Area and San Rafael State Park: numbers of genera (G), native species (NS), exotic species (ES), and total species (TS) Sonoita Creek State Natural Area San Rafael State Park Family G NS ES TS Family G NS ES TS Asteraceae 59 84 3 87 Asteraceae 46 79 3 82 Poac 35 50 16 66 Poaceae 38 Di 18 75 Fabaceae 31 51 2 53 Fabaceae 25 35 8 43 Euphorbiaceae 11 230 (0 23 Euphorbiaceae 6 21 0 21 Scrophulariaceae 11 15. 2 17 Brassicaceae 10 9 4 13 Brassicaceae 13. 13—~«3 16 Solanaceae 6 1] 0 11 Pteridaceae 7 13. (OO 13 Scrophulariaceae = 7 9 1 10 Cactaceae 5 12 0 12 Cyperaceae 4 10 0 10 Cyperaceae 4 12 O 12 Chenopodiaceae 4 7 1 8 Convolvulaceae 3 11 0 11 Convolvulaceae 3 8 0 8 Malvaceae 8 11 0 11 Malvaceae 5 / | 8 Solanaceae 7 TO + 4 1 Onagraceae 3 8 0 3 (Fabaceae, 8 species), Boerhavia (Nyctaginaceae, 7 species), Ipomoea (Convolvulaceae, 7 species), Brickellia (Asteraceae, 6 species), and Phacelia (Hydrophyllaceae, 6 species). At SRSP, the largest genera are Chamaesyce (9 species), Bouteloua (9 species), Dalea (7 species), Ascle 7 (Asclepiadaceae, 5 species), Euphorbia s.s. (Euphorbiaceae, 5 species), Ipomoea (5 species), Machaeranthera (Asteraceae, 5 species), and Oenothera ee 5 species). The percentage of the total flora (natives + exotics) that are epee: found only in aquatic and wetland habitats is 13.0% for SCSNA and 21.8% for SRSP. For those species found at SRSP but not at SCSNA the percentage of wetland plants is 34.8%, indicating the relatively greater importance of wetland habi- tats at SRSP. — Significant Taxa Ananthus squamulosa (A. Gray) King & H.E. Robinson (synonym Brickellia squamulosa A. Gray) (Asteraceae).—There is a relatively large population of this plant in Parker Canyon at SRSP. It is known in Arizona from only a few sites in the southeastern part of the state. Argyrochosma incana (C. Presl) Windham (Pteridaceae).—This is a rare fern in the United States, known only from a limited area in southern Arizona. It is uncommon on rock outcrops at SCSNA Cirsium grahamii A. Gray (Asteraceae).—This large, attractive, uncommon thistle is restricted to ciénega habitats. It occurs in the Heron Spring drainage on SRSP. Heterotheca rutteri (Rothrock) Shinners (Asteraceae).—This is a rare grass- land plant with a narrow range in southeastern Arizona. It is locally abundant at many localities in grasslands at SRSP. MCLAUGHLIN, FLORA OF REEK NATURAL AREA AND SAN RAFAEL STATE PARK 671 Lilaeopsis schaffneriana (Schltdl.) Coulter & Rose ssp. recurva (A.W. Hill) Affolter (Apiaceae ).—This is a listed endangered species under the Endangered Species Act, the only such species so far documented from the floras of the two sites. It was previously recorded from Sharp Spring at SRSP. This species was not found at either Sharp or Heron springs, but a few populations along the Santa Cruz River were found at SRSP. Pennellia robinsonii Rollins (Brassicaceae ).—This species is new to the flora of Arizona and the United States. It differs from P. micrantha (A. Gray) Nieuwl., a woodland and forest species, in having shorter, wider siliques, dark purple sepals, and purplish-tinged petals (Rollins 1995). Rollins gives its range as re- stricted to the Mexican state of Chihuahua. The species is locally common in grasslands at SRSP. Phacelia sonoitensis S.P. McLaughlin ined. (Hydrophyllaceae).—A previously unknown Phacelia was collected several times at SCSNA. The plant occurs on rock outcrops or on talus below rock outcrops; it was found at several widely scattered sites. It hasa combination of characters that do not match any known Phacelia from the Southwest: pinnately compound leaves, narrowly spathu- late sepals, and a capsule of several shallowly cymbiform seeds. A description of this proposed new taxon has been submitted to Novon. Phyllanthus polygonoides Nutt. ex Spreng. (Euphorbiaceae).—This perennial herb is rare in Arizona, but is more common in Texas and Mexico. It is a rare plant at SCSNA in Fresno Canyon along the stream. Potentilla wheeleri S. Watson (Rosaceae).—This species occurs at SRSP; it is known from relatively few locations in southeastern Arizona, all at much higher elevations in adjacent mountains or from rock walls in steep-sided mountain canyons. Its occurrence at such a low elevation is remarkable. It was common on a north-facing slope adjacent to Sharp Spring. Several other high-elevation taxa occurred at the same site, including Thlaspi montana var. fendleri and Houstonia wrightii. Rhynchosia minima (L.) DC. (Fabaceae).—This is a new state record for Ari- zona. The species is common in northern Mexico and also occurs in the south- eastern United States. It is rare at SCSNA, found once on a rocky slope above Coal Mine Canyon and once in Fresno Canyon. Sibara virginica (Brassicaceae).—This is a new record for the state of Ari- zona; it was found on two occasions in dense sacaton stands adjacent to the Santa Cruz River in SRSP. Sibara virginica is a rather widespread species, and its status here as a native or exotic is not clear. Tephrosia vicioides Schlect.(Fabaceae).—This perennial herb was previously overlooked in the flora of Arizona, and is only known from the state from just a few collections. It was found in SCSNA ata single location on loose, rocky soil in the center of the par Tillandsia recurvata a) L. (Bromeliaceae).—The epiphytic species is com- 672 BRIT.ORG/SIDA 22(1) mon in the tropics but is known in Arizona from just a few sites. It is locally common on junipers in narrow tributary canyons to Sonoita Creek at SCSNA. Floristic Geography Patterns of floristic affinity are illustrated by three maps (Fig. 4-6). These maps show the percentage of native species found in both parks (Fig. 4), found only at SCSNA (Fig. 5), and found only at SRSP (Fig. 6), which are present in each of 245 other local floras from North America. The average range size (number of local floras out of the sample of 245 local floras) for native species occurring in both SCSNA and SRSP is 26.2 floras. These taxa extend throughout the Apachian floristic area (McLaughlin 1989, 1992) of southeastern Arizona, northwestern Sonora, southeastern New Mexico, and northwestern Chihuahua, with clear af- finities extending southeast into west Texas (Fig. 4). Ruderal species, wetland plants, and perennial grasses constitute a large proportion of the most wide- spread native species common to both parks. Those native species recorded only for SCSNA are more narrowly distrib- uted (mean range size = 15.6 floras) than those occurring in both parks. The average range size for the native species recorded only from SRSP is 25.2 local floras, similar to that for those species recorded for both floras. The higher proportion of wetland taxa among the native species found only at SRSP ac- counts in part for the greater average range size of these taxa. While the species found only at SCSNA are mostly Apachian in distribution, a relatively larger proportion extend into the Sonoran Floristic Province (McLaughlin 1992) of western Arizona, southeastern California, and throughout Sonora (Fig. 5). A much smaller percentage of those species recorded only from SRSP extend into the Sonoran Floristic Province, but a higher percentage extend southeast into the Madrean mixed-grass prairies (Bock & Bock 2000) of Chihuahua and Durango and east and northeast into the short-grass plains of eastern New Mexico and Colorado (Fig. 6). The climates of SCSNA and SRSP are not that different (Table 1); in both areas precipitation occurring during the summer monsoon (July to Septem- ber) accounts for 61% of the mean annual precipitation. The small differences in temperature and precipitation, however, are consistent with the relatively higher affinity of the warmer, drier SCSNA with the Sonoran Floristic Prov- ince, and for the higher affinity of the cooler, somewhat moister SRSP with the short-grass plains. The coarser textured, shallower soils of SCSNA may favor desert species while the deep clay soils of SRSP may favor species from the Great Plains. The floras of both parks, however, are overwhelmingly Apachian in their species com position I : MCLAUGHLIN, FLORA OF ONQOITA CREEK NATIIRA! AREA AND CAN RALCAEI STATE PARK 673 55 \ 2 SO he, ean KO ees ‘a6 A5- LP et id (i Gor i ‘ 4 Lope 40 aoe: Ce 35- — a 30 SAN i 254 i . Catal Affinities of Speci ost 4 inities of Species o0- we ip. Occurringin both = L ay S, Sonoita Creek SNA << and San Rafael SP 15 T T T aoe ies -120 -110 -100 -90 -30 -70 in both Sonoita Creek State Natural A 1 San Rafael S Park. Isolines Fic. 4. Affinities of nat based on a sample of 245 local floras. ANNOTATED CHECKLIST Nomenclature in the following checklist, with few exceptions, is based on Kartesz (1999). Nomenclature in Cactaceae follows Parfitt and Gibson (2003). Synonyms, when given, are for taxa listed under a different name in Arizona Flora (Kearney & Peebles 1960). Exotic species are preceded by an asterisk (*). Annotations for each species include: common name(s) when available, habit, collection numbers, and abundance and distribution for each study area (SC = Sonoita Creek State Natural Area; SR = San Rafael State Park). Spanish common names are those enco din northeastern Sonora, as listed by White (1948). Common names and habits are not repeated for varieties with the same data. Voucher specimens were deposited at the University of Arizona Herbarium (ARIZ); duplicates were used to establish field herbaria for the two state parks. Several species, including most cacti and succulents, many trees and shrubs, and species not found in flower, were identified in the field and not vouchered. Unless otherwise indicated, collection numbers are those of the author. BRIT.ORG/SIDA 22(1) Affinities of Species ccurring in _ Sonoita Creek SNA Only 465 I T T T T T -120 -110 -100 -30 -80 -70 Fic. 5. Affinities of | 4 ry ly fa ra ft Lb C#eat iat LA ( z :< Cc Daf eae ees Park). Isolines represent the percentage of speci ly from SCSNA found in other local floras from North America, based on a sample of 245 local floras. LYCOPODIOPHYTA POLYPODIOPHYTA Selaginellaceae Azollaceae Selaginella rupincola Underwood. Spike moss; — Azolla mexicana C. Presl. Water fern; floating yerbaceous perennial.SC:8357; abundant on quatic. SC: 9330; rare in pools along Sonoita canyon slopes. EQUISETOPHTYA Equisetaceae Equisetum x ferrissii Clute (pro sp.).Scouring rush; herbaceous perennial. SR: 8268; asional, Santa Cruz River bottom Equisetum laevigatum A. Braun. Scouring rush; herb 8 TA c eous perennial. SC: 8745; uncommon along Sonoita Creek. SR: 9432; common, Santa Cruz River bottom. Creek. SR: 8784, uncommon, Sharp Spring. Marsileaceae Marsilea mollis B.L. Robinson & Fernald. Pepper- wort; herbaceous perennial. SR: Niles 698 (ARIZ); collected last in 1965 from a pool adjacent to the Santa Cruz River. Marsilea vestita Hook. & Grev. [M. mucronata A. Br]. Clover fern; herbaceous perennial. SC: 8498, 8532, uncommon along streams In Fresno and Coal Mine Canyons. SR: Van Devender 84-443 (ARIZ);collected last in 1984 from a pool adjacent to the Santa Cruz River. MCLAUGHLIN, AREA AND SAN RAFAEL STATE PARK 675 T 45-4 40- 35-4 Affinities of Species Occurring in San Rafael SP Only 18 T T 1 -120 -110 -100 -90 -B0 -70 Fic 6 ALE an 4 i + I a ul 1 L fa rs ic dR { £ j fa cr Le. Alas 1 Area) | 1 al | ‘| £ f ly £ SRSP fi P| L 1 ims] L. L America kK ld | {7 cl im si fod ‘he Te 5 Pteridaceae Adiantum capillus-veneris L. Maidenhair fern; her- ceous perennial. SC: 9419; below rock overhangs, west park boundary on south side of Sonoita Creek. Argyrochosma incana (K. Presl) Windham. Cloak ern; herbaceous perennial. SC:8407; uncom- mon on (i walls. n) Windham SSP. limitanea [Pellaea viva (Maxon) Mor- ton]. Cloak fern; herbaceous perennial. SC: 8806; uncommon on rock walls. Astrolepis integerrima (Hook.) D.M. Benham & Windham [Notholaena sinuata (Lagasca Kaulf.var.integerrima Hook.]. Canahuala; her- baceous perennial. SC: 8360, 8454; uncom- mon on rock walls. a Astrolepis sinuata (Lagasca ex Swartz) D.M. R Kk PO \AL 1} M TAT L | ra sinuata (Lagasca) Kaulf. var. sinuata]. sae herbaceous perennial. SC: 8406; mmon on rocky slopes and rock walls. Sees ia hispida (Mett. ex Kuhn) Underwood. A ommeria; herbaceous perennial. SC: 8400; uncommon, mostly below oaks. Cheil sna eatonii Baker. Lip fern; ieee perennial. SC: 9677; rare, canyon walls. Cheilanthes lindh k. Fairy swords; her- baceous perennial. SC: 8380; common on slopes and rock walls. Chei ne wrightii Hook. Wright's lip fern; her- baceous perennial. SC:8416,8820,8871;com- mon on rocky slopes Notholaena grayi Davenport. Cloak fern; herba- elmer) Hoo ceous perennial. SC: 8357; uncommon on rock walls Notholaena standleyi Maxon. Cloak fern; herba- eous perennial. SC: 8410, 8822; uncommon on rock walls. Pellaea intermedia Mett.ex Kuhn. Cliff brake; her- baceous perennial. SC: 6846, 9217; rare on ny is Pellaea wrightiana Hook. Cliff brake; herbaceous perennial. SC: 8878, 8933, 9217, rare on rock walls. PINOPHYTA Cupressaceae | , L ‘ 7 NA 7 pp Adams (J. monosperma (Torr.) Little sensu Kearney and Peebles (1960), i huata, tascate, tdscale; small tree. SC: 9408; nin canyons, uplands Juniperus deppeana Steud. Alligator juniper, huata, tascale, tascate; small tree. SR: 9392; uncommon,grasslands east of Santa Cruz River. n part]. Juniper, MAGNOLIOPHYTA: MAGNOLIOPSIDA Acanthaceae Anisacanthus thurberi (Torr.) A.Gray. Desert hon- re large shrub. SC: 9280; commen in anyon cago arizonica A. Gray. Herbaceous pe- nial. SC: 9291, 9297; uncommon in mes- quite gra ae ds. Sia resupinata (Vahl) Juss. Herbaceous f nial. SC: 8734; common in shade o gallery forests along Sonoita Creek. ees decumbens (A. Gray) Kuntze. Herba- ceous perennial. SR: 8287; common, grass- lands. Elytraria imbricata (Vahl) Pers. P Purple scaly stem, cordoncillo; herbaceous perennial. SC: 8369; anyons, rocky slopes. aenetate ium nervosum Nees [7. hispidum Nees]. Herbaceous perennial. SC: 8348, 9232; un- common, mostly in canyon bottoms. Aizoaceae Trianthema portulacastrum L. Horse pveRne summer annual. SC: 841 9;0c over- grazed sites. SR: 8791; uncommon, sacaton stands BRIT.ORG/SIDA 22(1) Amaranthaceae Alternanthera pungens Kunth [A. repens (L.) Kuntze]. Khakiweed; mat-forming herba- ceous perennial. SR: 87/2; occasional, grass- lands. Amaranthus palmeri S.Watson. te aay bledo, quelite; summer annual. 3c: 8352 common, canyons at 8270, common, mostly along drainages. Amaranthus torreyi (A.Gray) Benth. ex S.Watson Torr.’s amaranth; summer annual. SC: 8392, 8422, uncommon, mostly in canyons. Froelichia arizonica Tho cotton; herbaceous ete SC: 8809; rare, found once in Fresno COMP aiena caespitosa i Globe almakenisiy * a 1] Cc | rnbe rex standley. snake CD. O072 grasslands:S E Gomphrena nitida Rothrock. Globe amaranth; herbaceous perennial. SC: 8487; common, 10; occasional, grass- grasslands grasslands. SR: 8283, 83 lands. Gomphrena sonorae Torr. Globe amaranth; her- aceous perennial. SC:8370; common, grass- ei OF nds. Guilleminea densa (Humb. & Bonpl. ex Willd.) Mog. [Brayulinea densa (Humb. & Bonpl.) Small]. H ianal baceous perennial SC: 8415; 0cca- J Cr. Sap of | pee Iresine heterophylla Standley. Bloodleaf; herba- C ; eous perennial. SC: 8751; rare, canyons Anacardiaceae Rhus trilobata Nutt. var. anisophylla (Greene) Jepson. Sumac 9630; rare, oak woodland adjacent to Parker quawbush, lima; shrub. S anyon. Rhus trilobata Nutt. var. pilosissima Engelm. Shrub. SC: 9209; uncommon, canyons Rhus virens Lindheimer ex A. Gray var. choriophylla (Wooton & Standley) L.Benson [R. choriophylla Wooton & Standley]. Lima; rare, canyons. Toxicodendron rydbergii (Small ex Rydberg) Greene [Rhus radicans L.var.rydbergii (Small) Rehder]. Poison ivy, hiedra; woody vine. SC: 9420; uncommon, canyons. Apiaceae Berula erecta (Huds.) Coville. Water parsnip; her- baceous perennial. SR:8276;commMon, springs. MCLAUGHLIN, FLORA OF AREA ANN CAN RAEAFI STATE PARK Bowlesia incana Ruiz & Pavoén. Hairy bowlesia; winter annual. SC:8875;abundant, especially below mesquites. *Conium maculatum L.Poison hemlock; biennial C: 9474; rare, along Sonoita Creek. Daucus pusillus Michx. American carrot; winter annual. SC:8926;common on rocky uplands. ro herbaceous perennial. SR: 8375; occasional, grasslands. Hydrocotyle ranunculoides L.f.Water pennywort; atic herbaceous perennial. SC:8496; rare, pools in Fresno and Coal Mine Canyons. SR: 8787, occasional; Santa Cruz River bottom. Lilaeopsis schaffneriana (Schitdl.) J.M. Coulter & Rose var. recurva (A.W. Hill) Affolter. Huachuca water ae inconspicuous parce peren SR: 9434 allo ter fe sont Cruz River. Lomatium nevadense (S.Watson) J. M.Coulter & Rose var.parishii J.M.Coulter & Rose) Jepson. Indian root; herbaceous perennial. SC: 8843, 9084; rare, mesquite grasslands Spermolepis echinata (Nutt. ex DC.) Heller. Scale seed; winter annual. SC:8903; occasional, can- yons and uplands. SR: 91704; uncommon, grasslands. Yabea microcarpa (Hook. & Arn.) K.-Pol [Caucalis microcarpa Hook. & Arn.].Winter annual. SC: 9024, 9025; locally abundant, north-facing slopes in oak savanna. Apocynaceae Apocynum cannabinum L. Dogbane, Indian hemp; herbaceous perennial. SC: 9360; rare, one population in Coal Mine Canyon Haplophyton crooksii (L.Benson) L.Benson.Cock- roach plant, hierba de la cucuracha; herba- ceous perennial. SC: 9500, E.A. Lewis 012; rare, slopes, canyons Telosiphonia brachysiphon (Torr.) Henrickson ee brachysiphon (Torr.) A.Gray]. Rock trumpet; herbaceous perennial. SC: 8523, a Lewis 013, locally abundant on rocky slopes. SR: 9579; rare, grasslands Araliaceae Aralia humilis Cav. Tepetate; shrub. SC: 9703; rare, cky north-facing slope Aristolochiaceae — WW, & Standley. Indian Ar istoloct HCA VV CACIUTETT root; herbaceous perennial. SC: 8907, 9550; uncommon and infrequent, mostly on rock walls. Asclepiadaceae Asclepias asperula (Dcne.) Woodson ssp.asperula (A. capricornu Woodson sensu Kearny and Peebles (1960)].Antelope horns; herbaceous perennial. SC: 9220; rare, grasslands. SR: 9397; uncommon, grasslands lepias involucrata oe Milkweed; herba- ceous perennial.SR:9154 gras lands. apace inaria Cav. Milkweed; shrub. SC: 6437; rare, r Asclepias ae a Torr. Milkweed, yerba de cuerve; herbaceous perennial. SC: 9279; rare, grasslands, rocky slopes. SR: 9394; occasional, grasslands. Asclepias nyctaginifolia A.Gray.Four o'clock milk- weed; herbaceous perennial. SC: 9551; rare, canyons, rocky slopes. SR: uncommen, grass- lands. Asclepias subverticillata (A.Gray) Vail. Poison milk- weed, orled milkweed; herba- eOus sect SR: 8239; common, grass- onde Sharp Spring drainage. Asclepias tuberosa L. ssp. interior Woodson. But- terfly weed, pleurisy root, orange m |kweed; grasslands. Funastrum cynanchoides Decne. ssp. hetero- payin (Engelm. ex Torr.) Kartesz [F het wllum (Engelm.) Standley].Climbing siloeee herbaceous Peleg, vine. SC: 929): rare, cCanyo shrubs in sec ,on Asteraceae A.Gray) Reveal & King [Perezia compe A. Gia Large herbaceous peren- Se 8771; rare, canyons C Gray) fave & King [Perezia wrightii A.Gray]. Brownfoot; herbaceous pe- rennial. SC: 8922; rare, uplands Ambrosia confertiflora DC. [Franseria confertiflora C.) Rydberg] SU ea ues, herbaceous perennial. SR: 8 grasslan Ambrosia pallostachy DC. Western cel h us perennial.SR:8599; locally abun- dant ae Parker Canyon. ACOuUI CU tl er beri = OWI wrig! PUT i (A. 678 Ambrosia trifida L. var. texana Scheele [A. oe DC.].Ragweed, chuchuni; herbaceous peren nial. SC:8656;abundant along Sonoita Creek. SR:8302;common, roadsides, low areas, Santa Cruz River bottom Ananthus squamulosa (A. Gray) King & H.E. Rob- inson [Brickellia squamulosa A. Gray]. Herba- ceous perennial. SR: 9728, 9736; locally com- mon, Parker Canyon. Antheropeas lanosum (A. Gray) Rydberg. [Eriophyllum lanosum A. Gray].Woolly-daisy; winter annual. SC: 8920A, 9076; common in mesquite grasslands. Artemisia campestris L. ssp. borealis (Pallas) Hall lements var. scouleriana (Hook.) Cronq.[A. pacifica Nutt.]. Wormwood; herbaceous pe- rennial. SR: 9623; rare, Parker Canyon Artemisia dracunculus L. [A. dracunculoides Pursh]. False tarragon; herbaceous perennial. SC:970/, rare, north-facing slope in Coal Mine Canyon. Artemisia ludoviciana Nutt.ssp.a/bula (Wooton) Keck \A/ 1.1 | pee SC: 8673; common, north-facing slopes. SR: oc- casional, oak woodlands and savannas. Artemisia ee Nutt. “SSP. eae (Rydberg) Keck _oak woodlands and savannas. ii A. Gray. Shrub. SR: 9694, 9726; rare,Oa siAwoudiand adjacent to Parker Canyon. ee DC. Yerba de pasmo; small shrub ommon in grasslands. SR: 9376; asslands Baccharis eae ia ete & Pavon) Pers. [B. glutinosa Pers.].Seep willow, batamote jarilla; shrub. SC: 9651; occasional along Sonoita Creek, Fresno Canyon. SR: 9660; occasional, Parker C n. Baccharis ere A, ey Desert broom, romerillo; shrub. SC:881 1 common, mostly roadsides. ee eines Kunth. Shrub. SC: 8759; rare, canyon one! north-facing slopes. ANIA > common, canyons absinthifolia Benth. var. dealbata (A. Gray) A. Gray. Perennial herb. SC: 9277; rare, rocky slopes. Baileya multiradiata Harvey & A. Gray ex A. Gray. Desert marigold; perennial herb. SC: 9742; rare, grasslands. Barkleyanthus salicifolius (Kunth) H.E. Robinson BRIT.ORG/SIDA 22(1) & Brett. [Senecio salignus DC] Shrub.$C:8974; common in gallery forest along Sonoita reek, Bebbia juncea (Benth.) Greene sss cnuseeds Sweet bush;shrub.SC:9414:ra Berlandiera lyrata Benth. Cheese flower ae de la vibora; herbaceous ie SR: 9160; occasional, grasslands, roadside Bidens aurea (Ait.) Sherff. Bur eee peren- nial herb. SC: ‘as 8773, 9719; uncommon, along stream Bidens bigelovil a Gray. Beggar ticks, Spanish edles; summer annual. SR: 9619, 9692; un- n, Parker Canyon. nape ferulaetot Waed) DC sbegdal WeNs apa: Bidens laevis (L.) B.S.P. Bur marigold; herbaceous pere ae SR: 878 | 73; common, springs, z River bot Pe ena met on seal ticks, Spanish needles; summer annual. SC:8356;abundant, canyons, oak woodlands. SR; 8700, common below oaks. Brickellia amplexicaulis B. L. Robinson var. amplexicaulis. Perennial herb. SC: 8763; rare, upper Fresno Canyon. | lia baccharidea A. Gray. Shrub. SC: 9680; rare, o Can een. nena ia A.Gray. Herbaceous peren- nial. SR: 9699; rare, oak sav Brickellia californica (Torr. & < Ges) A. Gray vat. californica. Shrub. SC: 8807; uncommon, canyons. Brickellia coulteri A.Gray. Perennial herb. SC:8837; rare, canyon Brickellia eupat are (L.) Shinners. var. chlorolepis (Wooton & Standley) B.L. Turner [Kuhnia rosmarinifolia Vent. var. chlorolepis (Wooton & Standley) Blake]. Herbaceous perennial. SR: 8563; common, grasslands. Brickellia floribunda A. Gray. Robust perennial herb. SC: uncommon, along Sonoita Creek. SR: 9717; common, Parker Canyon Brickellia venosa (Wooton & Standle B. L. Rob- n.Herbaceous perennial.SC:8676B, 9720; Me eee ons Calycoseris wrightii A. Gray. Tackstem; winter an- nual. SC: 9074; rare, uplands. DC. Plume weed; summer annual. SC: 8657; rare, shaded canyon walls. MCLAUGHLIN, FLORA OF SONOITA CREEK NATURAL AREA AND SAN RAFAEL STATE PARK 679 Carphochaete bigelovii A. Gray. Bristlehead; low shrub. SC: 9283: rare, rocky uplands. SR: 9102; rare,oak woodland adjacent to Parker Canyon. Chaetopappa ericoides (Torr.) Nesom [Aster arenosus (Heller) Blake, A. hirtifolius Blake]. White aster, rose heath; herbaceous peren- nial. SC: 9029; occasional, mesquite grass- lands. SR: 9377; grasslands Cirsium gana A. Gray. Thistle; large biennial. uncommon, springs. Cirsium neomexicanum A. Gray. Thistle; biennial. : 9136; occasional, mostly in canyons. SR: oscasionial eee Vall ; (Bees | biennial. SR: 9368; common, aeacs| ands. Conyza canadensis (L.) Cronq. [Erigeron cress L.]. Horseweed; summer annual. SC: 8 common, canyons. SR: 8696; locally ae dant, grasslands. Cosmos Brees (Jacq.) Pers.Cosmos;summer annual. SR: 9714; rare, Parker Can Eclipta eal. .) L.[E. alba (L.) Tee Yerba tajo; summer annual. SC: 9669; rare, wet along Sonoita Creek. Ericameria taricifolia (A. Gray) Shinners [Aplopappus laricifolius A. Gray]. Turpentine bush; shrub. SC: 9730; common, north-facing slopes Caen Ericameria nauseosa 5 Pals ex Pursh) ow & aird ssp.nauseosa var.latisquamea A.Gray) Nesom & Baird en a nauseosus (Pallas ex Pursh) Britt. ssp. fatisquameus (A. ay): Hie & einen) Rabbitbrush; shrub. P n 4: abur Wd tlt, a8 Sieeron ariosolus Nesom [E. divergens Torr. & A. Gray, in part]. Fleabane; annual or herba- ceous perennial. SR: 8237, 8279, 8549B; com- mon, grasslands. Frigeron colomexicanus A. Nelson [E. divergens Torr.& A. Gray, in Ae Fleabane; herbaceous ncommon, along stream, perennial.SC: 9271 resno Can eerie ten Torr.& A.Gray. Spreading flea- bane; annual or herbaceous ees SE he oe 908?,9224;com yons and uplands.SR: 9700, roe concn toa Erigeron flagellaris A.Gray.Fleabane; herbaceous perennial. SR: 9246, 9252, common, grasslands and sacaton flat Erigeron oreophilus Greenm. Ftabere herba- ceous perennial. SC: 8477, yons, below Evax verna Raf. var. verna [E. multicaulis DC. var. multicaulis]. Rabbit tobacco, cotton rose; winter annual. SC: 9073; common, mesquite grasslands. SR: 9710, 9250; occasional, grass- ianiGS Fil ica Nutt. Yert i annual. SC: 9014B, 9017, 9087;common, canyons and _can- slopes. Fleischmannia pycnocephala (Less.) King & H.E. ROpInSen LEP acorn pycnccenagiom Less ].T SC: 8774; rare, north: facing slo opes. Gaillardia pinnatifida Torr. Blanket flower; herba- ceous perennial. SR: 8726; occasional, grass- lands and roadsides Gamochaeta purpurea (L.) Cabrera [Gnaphalium purpureum L.]. Cudweed; winter annual. SC: . ci 9026; abundant, mesquite grasslands. “common, grasslands. eneee eee A. Gray. Shrub. SC: 9552; uncommon, Fresno Canyon. ag alee eee (DC.) A. Gray. nakeweed; subshrub. SC: 8772; common in ee ; Helenium thurberi A. Gray. Sneezeweed; annual. SC: 8659, 8747; uncommon along streams. i th | (é f] ; | J>/ tall summer annual. SC: 9653; occasional, along Sonoita Creek. SR:8303; abundant, low areas, roadsides. Helianthus petiolaris Nutt. Sunflower, girasol; summer annual. 3; rare, canyons. SR: 8731;common, Parker Can ayaa longifol ia eo. & Greenm.) Cockerell var. annua (M.E. Jones) Yates [Viguiera annua (Jones) Blake]. ean gold- eneye; late summer Falpual ae eke com- mon, uplands Heliomeris multiflora Nutt. ee “multiflora (Nutt.) Blake]. Golden eye; herbaceous pe- ee Heterosperma pinnatum Cav.Summer annual. SC: common, north-facing slope, Coal Mine Canyon. SR; 8590, common, Parker anyon. Heterotheca rutteri (Rothrock) Shinners. Herba- erennial. SR: 8264; locally common, grasslands. 680 Heterotheca subaxillaris plant; herbaceous perennial. SC: ae common, along Sonoita Creek, can- yons. SR: 8553; common, grasslands and roadsides. EOSES mencgyd Torr. & A. Gray ex A. Gray. 8812; common along Sonoita Creek. SR: 9693; rare, Parker Canyon. Ba wislizenil A. Gray. Summer annual. 583; occasional, grasslan pienoninwe ightli A.Gray. cr ean nial. SC: 8768, 9732; uncommon, Fres a | | Bul robush; PIRI Bele SC: lsocoma tenulsecta Greene [Aplopappus tenuisectus potas Blake]. Burroweed; sub- shrub. 9707, common, ers aed nds. SR: 971 3; uncommon, gq etal lon solidaginifolium (A. se ca & Robin Sa eel iallal, daginifol ow wih s (Lam.) Britton & Rusby. BRIT.ORG/SIDA 22(1) Machaeranthera tanacetifolia (Kunth) Nees ter tanacetifolius (Kunth) Nees]. oe perennial. SC: ; uncommon, along ie Creek. SR: 8343, 8703, pe 9677. common, roadsides, Parker Canyon, Santa Cruz floodplain Malacothrix clevel andl A. Gray. eae casa saucers; winter annual. SC: 9079, 9121; un common, canyons and bees Jones SR: 9242; uncommon, mostly below Malacothrix fendleri A. fe or dandelion; winter annual. SC: 9015; common, me i arasslands. SR:9097 9157 clans MmMon Melampodium longicorne A. Gray. Summer an- nual.SC:8385; uncommon, canyons. SR:85917; tale Pane! Ca YOR. g.[M hispid } h]. Summer aes oe 8663; uncommon, can- yons. SR:8309, 8701: arker Canyon. Packera (A. ani [ A.Weber & A. Gr SC: 9506, 9731: ;rare, north-facing slopes. *“Lactuca serriola L. Prickly lettuce, wild lettuce; nnual. ee, eee Saas rocky slopes. SR nds Laennecia euler (A. Gray) Nesom [Conyza coulteri A. = SUING annual. SC: 9649; uncommo along Sonoita Creek. SR: 8338; oc fasta Laennecia sophiifolia (Kunth) Nesom sai ied sophiaefolia Kunth]. Annual. SC: 8765 common, canyons. SR: 9657; eal gr grasslands. Lasianthaea podocephala (A. Gray) K. Becker [Zexmenia podocephala A. Gray]. Herbaceous perennial. SC: 85 canyons. SR: 9629; rare, oak woodland adja- cent to Parker Canyon. Machaeranthera gracilis (Nutt.) Shinners [Aplopappus gracilis (Nutt.) A. Gray]. Annual. ep G009, common, ioe uplands. SR: 5 9583:rare, below trees in filachaerantherani aril eet ) Shinners ssp innatifida var. pinnatifida [Aplopappus spinulosus (Pursh) DC. var. turbinellus (Rydberg) Blake]. abnectars ems SR: paul gO common, grass [Act agetinus (Greene) Blake]. Hereaeeous. pee SC: 8395, 9586; common, below mesquite. SR: 9572: al grasslands Love Gace neomexicanus A. Gray]. Hew NA SR: 9387. rare, drainages. Parthenice mollis A.Gray.Robust summer annual. rare, canyons Pectis ie Harvey & A, Giay Fetid marigold; summer annual.SC:8477; uncommon, uplands. ae longipes A. Gray. Fetid marigold; herba- ceous perennial. SC: 9228; uncommon, mes- quite grassland. SR: 8777, 9375; uncommon, grasslands Pectis prostrata Cav. Fetid marigold;summer an- ual. SC: 8537; uncommon, grasslands. SR: 9618; rare, Parker Canyon ec gracile Benth. Galen: herbaceous perennial. SC: 9364; rare, uplan ee um ruderale (Jacq.) a SSp. macrocephalum i. R.R. Johnson [P san eee um DC]. Yerba del venado;sum mer annual. SC: 8423; uncommon, canyons. Se aie canescens (DC.) W. A. Weber ssp.canescens [Gnaphalium wrightii A. Gray]. Cudweed; eae perennial. SC: 8766; uncommon, canyons. SR: 8318, 8699: locally mon, grasslands eee di leucocephalum (A. Gray) A. Anderb. [Gnaphalium leucocephalum A. Gray]. White cudweed; herbaceous peren- nial. SC:8388, 8755; uncommon, canyon bot- tom. SR: 9697; uncommon, Parker Canyon. =a MCLAUGHLIN, FLORA OF RAL AREA AND SAN RAFAEL STATE PARK 681 Pseudognaphalium pringlei (A. Gray) A. Anderb. as ium pringlei A.Gray].Cudweed;her- baceous perennial. SR: 9519; occasional, a Pseudognaphalium stramineum (Kunth) W. A. r [Gnaphalium chilense es Cud- weed: annual.SC: 9 an SR:8267; occasional, Santa Cruz River es aa ate aes A. Gray. Summer annual. SR: t,low areas, roadsides, washes. Buinesanees pauciflorus (D. Don ee multicaulis DC.].False dandelion; hereacdeus perennial. SR: 8305 or 8 Q2 2} i UNICO TIOTT, rings and Santa Cruz River bottom ica: a californica Nutt. California saicony winter annual. SC: 9362; uncommon, rocky slopes. Rafinesquia neomexicana A.Gray. Desert chicory; winter annual. SC: 9012, 9016; common, can- yons, rocky slopes. Sanvitalia abertii A. Gray. Summer annual. SC: see rare, canyons. SR: 8778, 9625; uncom- on, Parker Canyon oe iO oe idus oe val. enaceaus [S.longilobus Ben f groundsel;shrub. SC: 9284; les canyons. SR: 8778; occasional, grasslands. olidago velutina DC.[S. sparsiflora A.Gray].Gold- enrod; herbaceous perennial. SC: 9/12; rare, one patch found in canyon west of Fresno Canyon. SR: 8578; uncommon, mostly oak woodlands *Sonchus asper (L.) Hill. Sow thistle; annual. SC: 9151; occasional, mostly in canyons. SR: 9156, 1 joccasional Santa Cruz River bottom. Nit lA latt 4 | ce 9471: rare, canyon sepharomea pcr ci A. Bd He erennial. S Oc- lettuce; herbac casional, canyons 7 ce ms os ae occasional, grasslands ee thurberi A. Gray. Wire lettuce; mer annual. SR: 9450; occasional, grass- an ee oan divaricatum . ee Nesom [As- ter exilis Ell.]. Annual. SC: 8 common in ere ene LM River bottom. Symphyotrichum falcatum (Lindley) Nesom var. commutatum (Torr. & A. Gray) Nesom [Aster commutatus (Torr. & A. Gray) A. Gray]. White prairie daisy; perennial herb. SR: 8577; com- mon, springs. *Taraxacum officinale G. H. Weber ex F.H.Wigg. Dandelion; annual. SC: 9475; rare in wet ground along streams. SR:97712;uncommon, prings. at enn megapotamicum (Spreng.) Kuntze. Her ous perennial. SR: 9369; common, aie a Thymophylla pentachaeta (DC.) Small var. hartwegii (A. Gray) Strother [Dyssodia pentachaeta (DC.) B. L. Robinson]. Herba- eOUS perennial SC: 9271,9?72.uncommon, uplands. tea ia POE A. Gray. Summer annual. SC: north-facing slopes. Towinceneis (Richardson) Porter. Herba- Cceous sefeitial SR: 9257; rare, grasslands Trixis californica Kellogg. shrub. SC: 8364; occa- sional, canyons and rocky slop Tee: lindleyi (DC.) ee Gea: linearifolia (DC.) Schultz-Bip.]. Silver puffs; winter annual. SC: 8867, 8924; occasional, uplands. SR: 9054 |, grasslands, be- low oaks. Koc hioid, v.) Benth. & Hook.f. ssp. exauriculata (Robinson & Greenm.) J.R.Cole- Crownbeard;summer annual. SC:86917; weedy terrace adjacent to Sonoita Creek. SR:uncommon, Parker Canyon. Viguiera cordifolia A. Gray. Golden eye; herba- eous perennial. SR: 8598; locally common, Viguiera dentata (Cav.) Spreng. var. dentata. Golden eye; Ded perennial. SC: 8672; abundant in mesquite bosques adjacent to Sonoita Creek. SR: one sbunaant oak odlands OQI4 . 00. Viguiera dentata (Cav.) Spreng. var. lancifolia i | ro Xanthium strumarium L. var. canadense (P. Mill.) Torr. & A. Gray [X. saccharatum Wallr]. Cock- lebur,abrojo;summer annual. SC:8758; abun- dant in moist ground along streams. 5R:8597; common, low areas, washes, Santa Cruz River ottom. Xanthocephalum gymnospermoides (A. Gray) th. & Hook. Summer annual. SR: 8262; abundant, grasslands pak Zinnia acerosa (DC,) A. Gray [Z pumila A. Gray]. ert zinnia; low shrub. SC: 9326; uncom- mon, uplands. Zinnia grandiflora Nutt. Prairie zinnia; herbaceous erennial. SR: 9400 ional | grasslands bas Zinnia peruviana (L.) L. [Z.multiflora LJ]. Summer annual. SR:8720; uncommon, Parker Canyon. Berberidaceae Berberis wilcoxiik Bart “shrub. SC: 9276; rare, north-facing slope in Fresno Canyon. Bignoniaceae Chilopsis linearis (Cav.) Sweet ssp. arcuata (Fosberg) Henrickson. Desert willow, mimbre; small tree. SC: 9468; uncommon, along Sonoita Creek. SR: 9621; abundant, Parker Canyon. Tecoma stans (L.) Juss.ex Kunth var.angustatum Rehder. Trumpet flower, l/uvia de oro, palo de arco, gloria; shrub. SC:882]:rare, rocky slopes. Bixaceae Amoreuxia palmatifida Moc. & Sessé ex DC. lic ue perennial herb.£.A. Lewis 071; mon, grasslands and uplands. SR: 9526; ae cee Boraginaceae Amsinckia menziesii (Lenm.) A. Nelson & J. F. Macbride var. intermedia (Fischer & C. A. Meyer) Ganders IA. intermedia Fischer & C.A. Meyer]. Fiddleneck; winter annual. SC: 9021; rare, Coal Mine Canyon. Cryptantha angustifolia (Torr) Greene. Narrow- annual. SC: i barbigera (A. Gray) Greene. Bearded popco ower; mainte anvil SC:8902, 8906 canyons and uplands. 8925, 9 SR: aie rare, oak savannas so ceerate micrantha (Torr) |. M. Johnston var. micrantha. Purple-rooted popcorn flower: ence SC: 9133; ndy floodplain of Sonoita Creek. Cryptantha pterocarya (Torr.) Greene var. cycloptera (Greene) J. F. Macbride. Winter annual. SC: 8866, 9179: abundant, mostly on ocky slopes. Cryptantha pusilla (Torr. & A. Gray) alg Pop- corn flower; winter annual. SC: 8865, 9069, 9139B; uncommon, canyons and sl up- lands. SR: 9755; uncommon, grasslands. BRIT.ORG/SIDA 22(1) Lithospermum cabrense Greene. Puccoon; herba- ceous perennial. SR: 9482, 9486; common, grasslands Lithospermum incisum Lehm. Puccoon; herba- eous perennial. SR: 9302; occasional, grass- lands. Pectocarya recurvata |.M. Johnston. Arch-nutted bbur; winter annual. oe aed wel 9132; oC. casional, canyons, slopes. S grasslands, disturbed areas. Plagiobothrys arizonicus (A. Gray) Greene ex A. Gray. Blood weed, Arizona popcorn a winter annual.SC:8855,9116;common,gra lands and canyons. SR: 894 ton flats, grasslands. /$aca- Brassicaceae Arabis perennans S. Watson. Rock cress; herba- ceous perennial. SC:8831,8858; uncommon, walls. eo bursa- patel (L purse, rare, gallery forest along Sonoita sae . 9047. disturbed areas. Descurainia pin (Walt.) Britton ssp. ‘an a Detling. Tansy mus- tard; winter annual. SC: ;common, can- yons and uplands. SR: 9058; abundant, road- sides, grasslands Descurainia pinnata (Walt.) Britton. ssp. ochroleuca (Wooton) Detling. Tansy mustard; winter annual. SR:8939; abundant, roadsides, grasslands. “Descurainia sophia (L.) Webb ex Prantl. Flixweed; winter annual. SC: 9137; uncommon, along 9048 oadsides .) Medik. oe S panique illo; winter annual. S asslands. Draba cuneifolia Nutt. ex Torr. & A. Gray var. a. S.Watson. Whitlow grass; winter nnual.SC: 8826, 8840; common, canyons. SR: ae uncommon, clara Gray. Winter annual. Ne 88/73, 9013; rare, oe walls, Fresno and Coal Mine canyons. *Lepidium campestre (L.) Ait. f. Peppergrass; win- ter annual. SR: 9300; rare, springs Lepidium lasiocarpum Nutt. var. lasiocarpum. Sand peppergrass; winter annual. SC: ioe commo land d drai SR:8 Dry opeta lo inathim abundant, ‘low areas, grasslan Lepidium ie Wooton. Spalciny spring- MCLAUGHLIN, FLORA OF SONOITA CREEK NATURAL AREA AND SAN RAFAEL STATE PARK 683 summer annual. SC: 9491; uncommon, can- yons. SR: 9303; locally abundant, Parker Can- yon, grasslan nds. Lepidium virginicum L.var.medium (Greene) C.L. Hitchcock [1 ae Greene]. Peppergrass; winter annual.SC:8842;common, rocky slopes. sae gordonii (A. Gray) S. Watson var. i, Bladder oe winter annual. SC Penne micrantha ne Gray) Nieuwl. [Thelypo- anthum (A. Gray) S. Watson]. An- nual or am SC: 9510, 9645; rare, north- facing slopes, Coal Mine Canyon Pennellia robinsonii Rollins. Annual or biennial. SR: 8256, 8307, 9567, 9597, occasional, grass- lands, springs ales nasturtium-aquaticum (L.) Hayek.Water- DEO, aqu Halle Dla perennial. n streams. SR: 9/60; oc- Srl springs, Santa Cruz River. Schoenocrambe linearifolia (A. Gray) Rollins [Sisymbrium linearifolium (A. Gray) Payson]. Herbaceous perennial. SC: 8379, 8476; com- mon, rocky slopes. SR: 87271; uncommon, grasslands bes pie (L.) Rollins. Winter annual. SR: 9; uncommon, sacaton flats. a, irio L.London rocket; winter annual. SC:9013;common, particularly in overgrazed mesquite grasslands. SR: 9046; occasional, disturbed areas. Streptanthus carinatus C.Wright ex A. Gray ssp. arizonicus Weise) Kruckeberg, Rodman on]. Twist arizonicus S.Wats flower; winter annual. SC: 9075, 9090; rare, desert grasslands Thlaspi montanum L. var. fendleri (A. Gray Holmgren [T7.fendleri A.Gray]. oe candytuft; herbaceous perennial.SR:8947, uncommon, = ae) north-facing slope adjacent to ae Spring. Thysanocarpus curvipes Hook. Lace pod; winter annual. SC:8 |, rocky slopes. Gactaceae \Rritton & Roce in N phantha vivipara (Nutt. i" Britton & A.Brown [C. vivipara var.bisbeeana (Orcutt) L.D. Benson]. Beehive cactus; small stem succulent. SC: uncommon, grasslands Oc casional, grasslands Cylindropuntia le is (DC.) FM. Knuth in C. Backeberg & F.M.Knuth [O; ial li DC]. Desert Christmas cactus, st aecilent SC: rare, included based on a photo by R. Cyli rerun ee ano Nite M. Kunth i in keber unt ae ree Cane cholla, Holle large succulent. SC: common, grasslands and rocky slopes. SR: occasional, grasslands, Parker Canyon Cylindropuntia versicolor (Engelm. ex J. M. Coulter) FM. Knuth in C. Backeberg & F. M. Kunth [Opuntia versicolor Engelm. ex J.M. Coulter]. Staghorn cholla, cholla, siviri; large succulent. SC: rare, a single, sterile plant ina drainage on the north end of the park ap- e this species. Echinocereus fendleri (Engelm.) Sencke ex J.N. Haage. Hedgehog cactus;small stem succu- lent. SC: occasional, grasslands and rocky slopes. Echinocereus — idissimus (Engelm.) Engelm. ex Ha ectinatus (Scheidw.) Engelm. var. rigidissimus is (enalan) Engelm. ex Rumpler]. ainbow cactus; small stem succulent. SC: occasional, rocky slopes. SR: uncommon, grasslands. Echinomastus intertextus (Engelm.) Britton & Rose]. Small stem succulent. SR: 8953; locally slands pall common gras Ferocactus wislizeni (Engelm.) Britton & Rose. Barrel cactus; large stem succulent. SC: oc- casional, grasslands Mammillaria grahamii Engelm. [M. microcarpa Engelm.]. Fishhook cactus; small stem suc- culent. SC: occasional, rocky slopes Mammillaria macdougalli Rose in L.H. Bailey]. Small, flat succulent. SC: occasional, rocky slopes, canyons Opuntia engelmannii Salm-Dyck ex Engelm. var engelmannii.Engelmann prickly pear, nopal; eae parca common, mostly in mes- grasslands Ba oedl ds. ae macrorhiza Engelm. [O. plumbea Rose]. ckly pear; small succulent. SR: occasional grasslands. Opuntia phaeacantha Engelm. Prickly pear, no- pal; sprawling succulent. SC: occasional, grasslands. SR: uncommon, grasslands. 684 Opuntia santa-rita (Griffiths & Hare) Rose. Santa Rita prickly pear, nopal; large succulent. SC: abundant on rocky slopes Campanulaceae Lobelia os nalis L. Cardinal flower; perennial erb. SC: 8490; uncommon, wet soil along streams Lobelia fenestralis Cav. Summer annual. SR: 8555; incommon, grasslands, sacaton flat Nemacladus lees liferus Jepson var. ori Sais Thread plant; winter annual. SC: 9070, 9135; or common, rocky slopes, canyons, Triodanis holzingeri McVaugh. Venus ane Ss; annual. SC: 9359; rare, canyon a perfoliata (L.) Nieuwl. rane looking ss; annual. SC: 9203, 9223; occasional, can- yons. oO | sav) Capp Cleome lutea Hook. var. jonesii J.F. Macbride [C. jonesil J.F. Macbride]. Bee plant; robust sum- mer annual. a soils adjacent to Sonoita Polanisia Te dra (L.) De ssp. trachysperma (Torr.& A. Gray) Iltis [PR trachysperma Torr. & A. Gray]. Clammy weed; summer annual. SC: 8464; occasional, along streams. SR: 8717 common, Parker Canyon. Caprifolia | en mexicana K. Presl ex DC. Mexican el- der, tdpiro; small tree. SC: 8915; common along Sonoita Creek. SR: 9440; occasional, Santa Cruz River floodplain. Kartesz (1999) includes this taxon as a synonym of the widespread S. nigra L. ssp. canadensis (L.) R. Bolli. Mexican elder is a low-elevation, early spring-flowering species, while the variety of S. nigra [S. caerulea Raf.] found in Arizona occurs at much higher elevations and flow- ers in the summer and fall. — coy Caryophyllaceae Cerastium texanum Britton. Mouse-ear chick- weed; annual. SC: 9020; rare, ee: slopes, Fresno and Coal Mine canyons. a molluginea (Lag.) Didr. [D. sperguloides Gray]. Drymary; summer annual. SC: 8472; mmon, rocky upland uNnCcOo Silene antirrhina L. Sleepy catchfly; winter annual. : 9027; common, rocky slopes. SR: 9307; BRIT.ORG/SIDA 22(1) uncommon, oak woodland adjacent to Parker Canyon. Chenopodiaceae Atriplex canescens (Pursh) Nutt. var. canescens. Saltbush, orache;shrub. SC overgrazed ridgetops. Atriplex elegans (Moq.) D. Dietr. var.elegans. Sum- annual. SR: 9513; occasional, grasslands roadsides. Atriplex wrightit S. sae Wright saltbush; sum- nnual. SR: 8277, 9560; ee Santa Cruz ne *Chenopodium Giro ONaE: : Goosefoo hipasote;annual.SC:8 ee roadsi Sonoita Creek. Chenopodium berlandieri Moq. var. sinuatum (Murr) Wahl. Pitseed goosefoot; summer an- nual. SR: 8704, 8732, 9615; occasional, Parker Canyon, grasslan ds Chenopodium fremontii S.Watson var. fremontii. Go ual. SC:8382; occasional, can yons. SR: 9690; uncommon, Parker in. lisa neomexicanum Standley var. n xicanum. Goosefoot;summer annual. SC: a 9548, 9670; Common, Canyons, up- lands. SR: 9667: uncommon, below oaks. Chenop ? watsoni A. Nelson. nen ee ae ar- eas, grasslands Monolepis nuttalliana (J.A. Schultes) Greene. a winter annual. SR: 8941; locally abun dant, grasslands. *Salsola tragus L.[S. kali L. var. tenuifolia Tausch]. Tumbleweed, Russian thistle; summer an- nual, SC:uncommon, disturbed sites. SR: 9564; abundant, roadsides Convolvulaceae Convolvulus equitans Benth. [C. incanus Vahl. Hoary disc dea ipeae perennial vine. C: 9585; uncommon, mostly in canyons. SR: 8710;0 paces grasslands Evolvulus a sinoides (L.) L. var. angustifolia Torr. H ous perennial. SC: 9127; rare, rocky Hes Evolvulus arizonicus ae Gray. MGI BRESOUS peren- nial.SC:8575, | canyons, rocky slopes. SR: 8308; occasional, grasslands. Evolvulus sericeus Sw. var. sericeus. Herbaceous ial. SC: 9509; rare, canyons I; ae ee gion oak woodenes MCLAUGHLIN, FLOR AREA AND CAN RAEFCAEI STATE PARK 685 Ipomoea barbatisepala A. Gray. Morning glory; summer annual vine. SC: 8394, 9582; com- mon, canyons Ipomoea capillacea (Kunth) G. Don [/. muricata Cav. sensu Kearney & Peebles (1960)]. Morn- ing glory; herbaceous perennial. SR: 9520; uncommon, grasslands Ipomoea costellata Torr.M Cucurbita foetidissima Kunth. Buffalo gourd, calabacilla; herbaceous perennial vine. SR: 456; occasional, grasslands Echinopepon wrightii (A. Gray) S. Watson. Wild balsam apple; herbaceous annual vine. SC: 8383; uncommon, canyons Cuscutaceae annual vine. oS eg 851 . common, can sand uy 07, occasional, grasslands, Parker Canyon. Ipomoea cristulata Hallier f. [ coccinea L. sensu Kearney and Peebles (1960)].Star glory, trom- pillo;sumn er annual vine. SC:8466;common, canyons. SR: 8593; common, Parker Canyon. Ipomoea hederacea oo ee ae suM- mer annual vine.SC:9 ns SR:8595, 9610,9727;common, arn Ipomoea longifolia Benth. Morning glory; long- stemmed ae SR: 9515;common, Oak sav ales purpurea .) Roth [/ hirsutula Jacq. f]. Tall morning 8376; occasional, canyons ? av. var. leptotoma (Torr) J.A. ~ McDonald [ [/. leptotoma Torr].Morning glory; summer perennial vine. SC: 8365; common, canyons Ipomoea thurberi A. Gray. Morning glory; sum- er perennial vine. SC: 8473; rare, canyons. fa) nnual vine. SC: CHMHIOHG Crassulaceae Crassula connata (Ruiz & Pavén) Berger var. ta [Tillaea erecta Hook. & Arn]. Pigmy weed; winter annual. SC: 8937; uncommon, grasslands Sedum cockerellii Britton. Stonecrop; succulent perennial herb. SC: 9704; rare, rock walls. Cucurbitaceae es undulata A.Gray.Meldn loco,melén de coyote; DelPadcous Peele) vine. oo 8561; occasional, grasslands. *Citrullus lanatus (Thunb.) Matsum. & Nakai Watermelon; annual. SR: 9737; rare, Parker Can Cucurbi : ee A. Gray. FINget: leaved gourd, calabacilla, chichicayota nial vine. SC: uncommon, washes aR Unedate mon, grasslands. Cuscuta erosa Yuncker. Dodder; parasitic vine. SC: 8504, 8526; uncommon, found on Tragia nepetifolia. Ericaceae i] AA : | | SR: 9055; rare,oak savanna.One collection from near the north (unfenced) park boundary. Euphorbiaceae Acalypha neomexicana Muell.-Arg. New Mexican ie summer annual. SC: 8389; un- on, canyons. SR: 8248, 9590; common, grass sland S. Acalypha phleoides Cav. [A. lindheimeri Muell.- Arg.]. Herbaceous perennial. SR: 8242; occa- sional, wash Acalypha esmole Riddell. Hornbeam three- seeded mercury; summer annual. SC ; uncommon, canyons and slopes. SR: 8245; occasional, washes. Argythamnia neomexicana Muell.-Arg. [Ditaxis neomexicana (Muell.-Arg.) Heller]. Herba- ceous perennial. SC:8458,8835; uncommon, rocky slope Chamaesyce abomerennata (Torr. & A. Gray) Small [Euphorbia albomarginata Torr.& Gray]. ee TS herbaceous eos sslands. SR:8243; oc- ne disturbed a Chamaesyce arizonica ae Arthur (Euphor- bia arizonica Engelm.]. Spurge, golondrina, herbaceous perennial. SC: 8377, 8412; occa- sional, rocky slope Chamaesyce capitellata (Engelm.) Millso. [Eu- phorbia capitellata Engelm.]. Spurge, golondrina; herbaceous Salle SR: 8340, 8565; occasional, grasslan Chamaesyce dioica (Kunth) ie [Euphorbia indivisa (Engelm.) Tidestrom]. se ae summer annual. SC: 8463; com- watercourses. SR: 8332; a ee pas Chamaesyce florida (Engelm.) Millsp. [Euphorbia florida Englem.].Golondrina;summer annual. common,grasslands and uplands. Chamaesyce hirta (L.) Millsp. [Euphorbia hirta LJ. Spurge, golondrina; summer annual. SC:8349, occasional, canyons. SR: 8587; occasional, washes. Chamaesyce hyssopifolia (L.) Small oe hyssopifolia L.].Golondrina, summer annual. SC: 8387; uncommon, canyons. SR: a ,OC- casional, washes, grasslands, roadsides Chamaesyce melanadenia (Torr) Millsp. [Euphor- bia melanadenia Torr]. Spurge, golondrina; herbaceous perennial. SC: 8832, 9415; occa- sional, rock walls. Chamaesyce micramera (Boiss. ex Engelm.) Wooton & Standley [Euphorbia micromera Boiss.]. pulse golondrina; summer annual. SC: 8661; uncommon, drainages. Chamaesyce pediculifera (Engelm.) Rose & Standley var. pediculifera [Euphorbia pediculifera Engelm.]. Spurge, golondrina; herbaceous perennial. SC: 8359; occasional, rocky slopes. Chamaesyce revoluta (Engelm.) Small [Euphor- bia revoluta Engelm.]. Golondrina; summer annual. SC: 8521; rare, rocky slopes. SR: 8716; rare, Parker Canyon Chamaesyce serpyllifolia (Pers.) Small [Euphorbia sea i Sy coal ee summer nnual.SR:8 8695 i | sslands Chamaesyce ne (Engelm. ex di cite eX Parish [Euphorbia setiloba Engelm.]. Spurge, golondrina; summer annual. SC: 86/0; rare, along Sonoita Creek. Chamaesyce stictospora (Engelm.) Small [Eu- horbia stictospora Engelm.]. Spurge, golondrina, summer annnal. SR: 8321, 8600; occasional, grasslands. Chamaesyce vermiculata (Raf.) House [Euphorbia vermiculata Raf]. Spurge, golondrina; sum- mer annual. SR: 8304; occasional, springs Cnidoscolus angustidens Torr. Mala mujer, sla a; Liles perennial. SC: 8456; common, S. Croton ae halen ce ee [C. L / | ] | | Jef | ceous perennial. SC: 9542 ional, rocky slopes. SR: 8601 - occasional, grasslands. Croton texensis (Klotzsch) Muell.-Arg. var. texensis. BRIT.ORG/SIDA 22(1) Dove weed, yerba de la tértola, summer an- nual. SC: 8668, 8692; uncommon, sce soils along Sonoita Creek. S i, Santa Cruz River patton. Euphorbia bilobata Engelm.Summer annual. SR: 9613, 9718; uncommon, Parker Canyon. Euphorbia chamaesula Boiss. Herbaceous peren- nial. SR: 8281, 8333, 9312; occasional, grass- lands, roadsides. ss cuphosperma (Engelm.) Boiss. [E. dentata Michx. var. cuphosperma (Engelm.) Fein Painted ae summer annual. SC: nds ee cae fae Spurge; summer annual. SC: ee uncommon, mesquite shes Euphorbia ae ee bainted ae sum- mer annual. SC: 8350; uncommon, canyons and uplands.SR:8377;occasional, grasslands springs. Euphorbia radians Benth. Painted spurge; herba- ceous perennial. SR: 9/09; common, grass- lands. Jatropha macrorhiza Benth. var. septemfida Engelm. Herbaceous perennial. SC: 9472; uncommon, canyons and uplan S Manihot angustiloba (Torr) Muell.-Arg. Herba- ous perennial. SC: 9547; rare, canyons. or ens polygonoides Nutt. ex Spreng. Her- baceous perennial. SC: 8488; rare, Fresno anyon. Tragia laciniata (Torr.) Muell.-Arg. Nose burn; h ous perennial. SR: 8379; occasional, a on north-facing slopes. Tragia het Cav. var. dissecta Muell.-Arg. se burn; herbaceous perennial. SC: 8358, 8500; eae rocky slopes. Fabaceae Acacia angustissima (P. Mill.) Kuntze. White ball acacia; oe a ial 8; occasional, ridge crests. SR: | Plants appearing to belong to different va- rieties may occur mixed in the same popu- lations. Acacia greggii A. Gray. Catclaw, una o_o ae os coe Fonlbae ca n.Large ae ee 8503, grasslands 9492; rare, aoe Amorpha fruticosa L. ao ae large shrub. SC: 9286; rare, one patch found in Fresno Canyon. MCLAUGHLIN, FLORA OF AREA AND SAN RAFAEL STATE PARK 687 Astragalus allochrous A. Gray var. allochrous. Halfmoon loco; winter annual. SC: 9730; un- common, along Sonoita Creek. Astragalus arizonicus A. Gray. Locoweed; herba- Cc s perennial. SC: 9729, 9219; common, uplands. ee humistratus A. Gray var. sonorae (A. rdy Astragalus nothoxys A. Gray. Locoweed; herba- ceous perennial. SC: 9237; uncommon, can- yons and uplands. SR: 9306; common, grass- lands. en nuttallianus DC. var. tri a. Torr. ocoweed; winter annual. S 04; sundae tae SR:9093; sbuneants grass- lan ae pinia gilliesii (Hook.) Wallich ex D. Dietr. Bird-of- Ghee ale 9473; rare, gallery he along Sonoita Cre Call eee eriophylla ey var. eriophylla. Fairy duster;low shrub.$C:932/,occasional, uplands. Calli aa humilis Benth. var. reticulata (A. Gray) n [C. reticulata A. Gray]. False mes- ones perennial a 8559: occa- ands Chamaecrista nictitans (L.) Moench ssp. nictitans var. leptadenia (Greenman) Gandhi & Hatch [Cassia leptadenia Greenman]. Yerba del piojo; summer annual. SC: 8403; occasional, grass- lands and uplands. SR: 8257; occasional, grasslands. Cologania angustifolia Kunth [C. longifolia A. Gray]. Herbaceous perennial. SC: 9682; rare, deel ae SR: E.A. Lewis 004; uncom- ker Cany ee ane ee Lavin var. caribaea a edwardsii A. Gray]. Herbaceous pe- Ae rae ie ne on north- facing slopes. Crotalaria js Ortega. Rattle box; summer |. SC: 7, occasional, canyons and Ge oie in Parker Canyon. Dalea albiflora A.Gray. Herbaceous perennial.SC: uncommon Sey aes SR: 8507, 8579; baa ae eae ex ie var. oligophylla rr.) Shinners [Petalostemon candidum ae Michx. var. oligophyllum (Torr.) Her- = mann].White prairie clover; herbaceous pe- rennial. SR: 9390; occasional, grasslands. Dalea formosa Torr. Feather plume, shrub. SC: 8674,9 9068; locally common uplands Dalea grayi (Vail) L.O. Williams. Herbaceous pe- rennial. SR: 8506; uncommon, grasslands. Dalea lachnostachys A.Gray. Herbaceous peren- nial. SC: 9322; uncommon, oi uplands. SR: 8722; uncommon, gras cs Dalea leporina (Ait.) niece ee dalea; sum- mer annual. SR:8620;uncommon, grasslands. Dalea nana Torr. ex A. Gray var. carnescens Kear- ney & Peebles. Herbaceous perennial. SC: 8524, 9287, occasional, rocky uplands. SR: 8549C; occasional, grasslands Dalea neomexicana (A. Gray) A. Gray var. neomexicana. Herbaceous perennial. SC: 8682, 9273; uncommon, rocky uplands. Dalea pogonathera A. Gray var. pogonathera. Yerba del coraz6n; herbaceous perennial. SC: 8512?:0c casional, uplands Dalea pringlei A.Gray. Herbaceous perennial. SC: 8752, 9222, 9288; occasional, canyon, rocky slo Dalea ee H.S. Gentry. Indigo bush, shrub. SC: 8839, 9126; occasional, canyons, rocky slopes. Dalea versicolor Zucc. ssp. versicolor var. sessilis A. Gray) BanSDy ingige bush; shrub. SR: 9305, rare, oal d adjacent to Parker Canyon. Desmanthus cooleyi (Eaton) Trel. Herbaceous perennial.SC: 957 T;rare, mesquite grasslands. SR: 9529, E.A. Lewis 002; common, grasslands and roadsides. maditim bor qulon A. Gray. Tick clover, her- baceous perennial. SC: 8489; rare, canyon Desmodium neomexicanum A. Gray. Tick an summer annual. SC: 8541; rare, en SR: Parker C Desnouilinn ee (P. ai, Hitche. var. exiguum (A. Gray) Schub. Tick clover, summer annual. SC: 8681, 8687; rare, canyons y. Tick clover, ro- bust hetbacsous eer SC: 8436, 8443, 9553, 9701; common, canyons and rocky Ooditym nsilocarnism A. Gra tf slopes. ee rosei Schub. Tick clover; summer annual. SR: 9677; uncommon, Parker Canyon. Erythrina flabelliformis Kearney. Southwestern 688 coralbean, chilicote, pionilla; shrub. SC: 9411; common, rocky slopes. Eysenhardtia orthocarpa (A. Gray) S. Watson [E. polystachya (Ortega) Sarg. sensu Kearney & Peebles (1960)]. Kidneywood, palo dulce; large shrub. SC: 84317, 9412; common, rocky slopes. Galactia wrightii A. Gray var. mollissima Kearney & Peebles. Herbaceous perennial vine. SC: 8373; occasional, canyons. SR: 9459; rare, grasslands. Indigofera sphaerocarpa A.Gray.|ndigo, shrub. SC: orth-facing slope, Coal Mine Canyon. *Lotus aoe sel Bicasieot trefoil; herba- sprin Note River botto me greenei Ottley ex ae & Peebles. Deer vetch. 9022. Herbaceous perennial; uncom- mon, oak savannas. SR:8949 grasslands. Lotus humistratus Greene. Winter annual. SC: 8 ommon, es coals grasslands. SR: | | grasslands Lupinus brevicaulis S. ia Short-stemmed lupine; winter annual. SC: 9080; occasional, mostly along drainages. SR: 9/05; common, grasslands. Lupinus concinnus J.G. Agardh. ssp. orcuttil . Watson) D. Dunn. Elegant lupine; winte nual.SC:8930, 9063; occasional, mostly fone drainages. SR: 9107; occasional, Parker Canyon. Lupinus sparsiflora Benth. ssp. mohavensis Dzie cen & D. Dunn. Lupine; winter an- 08; locally abundant, slopes. Macroptil paae ce (Ortega) Delgado [Phaseolus heterophyllus Willd.]. Herbaceous perennial vine. SC: 8479; uncommon, can yons. SR: 9517, 9569, 9659; common, grass- lands Marina calycosa (A. Gray) Barneby [Dalea calycosa A. Gray]. Annual. SC: 8679, 9274; uncommon, uplands. SR: 9382; uncommon, oak savannas *Medicago lupulina L. Black medic; annual. SR: 9313, 9386; common, springs, Santa Cruz River bottom. lea sativa L. Alfalfa; se perennial. SR: side *Meli fe indicus (L.) All. one ion sweet BRIT.ORG/SIDA 22(1) clover, alfalfilla; annual. SC: 9265; occasional along Sonoita Creek. SR: 9298; occasional, Santa Cruz River floodplain. *Melilotus officinalis (L.) Lam. Sweet clover; tall annual. SR 8790, 9426; both white and yel- low-flowered forms abundant, Santa Cruz River bottom Mimosa aculeaticarpa Ortega var. biuncifera Benth.) Barneby [M. biuncifera Benth.].Wait- a-minute, cat's claw; shru > : 9470; com- mon, rocky slopes, canyons. SR:| llyab dant, lastee Ai Benth. Shrub. SC: 9353: com- mon, rocky slopes. SR:8337,occasional, grass- ands. — | Nissolia schottii (Torr.) A. Gray. Herbaceous pe- nnial vine. SC:8409; uncommon, north- fac- ing slop Phaseolus Stee A. Gray var. acut uae ep >summer annual. SC: 9639; un- ommon, oe SR: 9616; uncommon, Parker Canyon Prosopis velutina W [Pjuliflora(S DC. var. velutina (Wooton) Sarg.].Velvet mesquite mezquite; small tree. SC: 9587; common, washes, ridgetops, gentle slopes. SR: uncom- mon, grasslands. eAIGHGIE ium a aOR a (Pursh) Rydberg [Psoralea tenuiflora Pursh]. Scurf pea; herba- ceous perennial. SC: 9287; rare, canyons. SR: 939 | {land adj to Parker Canyon. Rhynchosia minima (L.) DC. Rosary bean; herba- eous perennial vine. SC: 9706, 9733; rare, rocky slopes, canyons Rhynchosia senna Gillies ex Hook. var. texana (Torr.& A.Gray) M.C. Johnston [R. texana Torr. & A. Gray]. Rosary bean; herbaceous peren- nial vine. SC: uncommon, north-facing slopes, canyons. SR: 8558, 9580; occasional, grasslands. Senna bauhinioides (A. Gray) Irwin & Barneby [Cassia bauhinioides A. Gray]. Herbaceous perennial. SC: 9325; uncommon, uplands Senna hirsuta (L.) Irwin & Barneby var.glaberrima M.E. Jones) Irwin & Barneby [Cassia leptocarpa flees Robust herbaceous pe- rennial. SC: 8381, 8465; uncommon, canyon bottom. SR: uncommon, grasslands. Sphinctospermum constrictum (S.Watson) Rose. MCLAUGHLIN, FLORA OF SONOITA CREEK NATURAL AREA AND SAN RAFAFL STATE PARK 689 Summer annual. SC: 8460; rare, mesquite grasslands. Tephrosia tenella A. Gray. Summer annual. SC: 8542; uncommon, a dal SR: 8584; unco Parker Can Tephrosia ae Schlecht. Herbaeou peren- nial.SC:8483; rare, found trop, where it was locally abundant. *Trifolium fragiferum L. Clover; herbaceous pe- rennial. SR: 9437; uncommon, Santa Cruz River bottom Trifolium mucronatum Willd. ex Spreng. ssp lacerum (Greene) J. Gillett [7. lacerum Greene]. Clover; herbaceous perennial. SR: 9523, rare, wet groun *Trifolium pratense L. Red clover, eae perennial. SR: 9489; a Cruz Vicia ludoviciana Nutt. ssp. /[udoviciana. Vetch, winter annual vine. SC: 8874; common, ashes. ee gemella Vogel [Z. diphylla (L.) Pers. sensu Kearney & Peebles (1960)]. Herbaceous pe- Cp. nez. rennial. SR: ip) 7; unc ommon, arasslands I ,encino Quercus arizonica Sarg. Arizona blanco; tree. SR: 9452, 9457, 9663, 9738 com- mon in woodland adjacent to Parker Canyon. Quercus emoryi Torr.Emory oak, bellota; tree. SC: 8533; uncommon, canyons, north-facing slopes. SR: 9516; common, oak savannas and oak woodlands ales grisea Liebm. Gray oak; tree. SR: 8344, afew trees just east of the Santa Cruz River appear to be this species. oe ob! ee, Torr. Mexican blue oak, 8775, occasional, north- Hee slopes, canyons. oazul tre Fouquieriaceae Fouquieria splendens Engelm. Coach whip, oco- tillo; shrub. SC: uncommon, rocky slopes Fumariaceae Corydalis curvisiliqua Engelm. ssp. occidentalis Engelm. ex A. Gray) W.A. Weber [C. aurea Willd. ssp. occidentalis (Englem. ex A. Gray) G.B. Ownbey]. Golden corydalis; winter an- nual. SC: 8869; common, rocky slopes, can- yons. SR: 895 1; occasional, Parker Canyon. Garryaceae Garrya wrightii Torr. Silk tassel; al shrub, SC: ;uncommon, ca Gentianaceae Centaurium calycosum (Buckley) Fernald. Buckley's centaury, annual. SC: 9272; uncom- alon ; Centaurium nudicaule (Engelm.) B. L. Robinson. Annual. SC: 9230, 9295; uncommon along streams. Gereningese Erodium cicutarium (L.) L’Her. ex Ait. ssp. cicutarium. Filaree, alfilaria, afilerillo; winter annual. SC: 9134 ional, mostly in mes- quite grasslands. SR: 9050; uncommon, dis- rbed areas. Erodium texanum A. Gray. Heron bill; winter an- nual. SC: 9072; uncommon, canyons and mesquite grasslands. Grossulariaceae Ribes aqureum Pursh var.aureum.Golden current, shrub. SR: 9759; rare, Sharp Spring, Santa Cruz River bottom. Haloragaceae Myriophyllum cf. sibiricum Komarov [M. exalbescens Fern.].Water milfoil; subrnerged aquatic perennial. SR: 9247; rare, Santa Cruz River bottom; not found in flower. Hydrangeaceae Fendlera wrightii (A. Gray) Heller [Ff rupicola A. Gray var. wrightii A. Gray]. Large shrub. SC 8407, 8836, uncommon, north-facing slobes anyons. Been: microphyllus A.Gray. Mock orange, shrub. SC: 9218, 9357; rare, north-facing slope in Fresno Canyon. An atypically low-eleva- tion record for this species. Hydrophyllaceae ees buts oe Heller. Winter annual. 85 canyons, rocky fae Nama upigdrn A. Gray. eae ae fare ae C: 8913 s adjacen stre ee Phacelia al 5 A. Gray. purole < phacelia;win- - 9062, 9207, 9268; Common, mostly ae mesquites. [ea prewar . 4: it Phacelia arizonica A. Gray.Winter annual. SC: 8856; common, ec nes quite eine! pune aiercd areas.$ sslands Phacelia bycina Wooton & atta Winter annual. SC: 8905, 9206; occasional, rocky slopes. Phacelia caerulea Greene.Winter annual. SC:897 7 nal, grasslands, c ha ee eve ae winter annual. SC: 9071; locally abundant, rocky slopes. Phacelia sonoitensis S.P. McLaughlin ined. Win- ter annual. SC: 9088, 9123,9206, 9226; uncom- ene slopes. Hypericaceae Hypericum scouleri Hook. ssp. scouleri [H. formosum Kunth]. Herbaceous perennial. SR: 9577, rare, SDrings. Juglandaceae Juglans major (Torr.) lee Arizona walnut, nogal; large tree. SC: 9 |,along eG Creek and in the larger canyons. SR: ‘ , Parker Canyon. Krameriaceae Krameria erecta Willd.ex JA on ites [K. parvifolia Benth.]. Ratany, tamichi; low hemiparasitic shrub, SC: 9227, pe rocky slopes. SR: 9380; uncommon, oak savannas. Krameria lanceolata Torr. Ratany; hemiparasitic herbaceou | ennial SR:9393 grasslands, Lamiaceae Hedeoma dentata Torr.Mock pennyroyal;herba- ceous Hees SC: 851]; uncommon, can- yons. SR: 8727; uncommon, oak woodland adjacent to Parker Can Hedeoma oblongifolia (A.Gray) Heller Mock pen- yroyal; herbaceous perennial. SC: 9584; uncommon, canyons *Lamium amplexicaule L. Dead nettle; winter annual. SC: 8970; uncommon, mostly along on Sonoita Creek. *Marrubium vulgare L. Common horehound, marrubio; herbaceous persis) alt 8326, occasional, grasslands sist around abandoned homesteads. Monarda citriodora Cerv.ex Lag. ssp. austromon- a (Epling) Scora [M. austromontana Epling]. Bee balm, horsemint, orégano; her- BRIT.ORG/SIDA 22(1) baceous perennial. SR: 8238; occasional, grasslands. Prunella vulgaris L. ssp. lanceolatus (W. Bart.) Hultén. Self-heal; herbaceous perennial. SR: 9522, rare, wet ground. Salvi peal i A. Gray. Sage; low shrub. SC: 8683, ,Canyons Salvia ree Benth.Summer annual. SC: 8398, 1; occasional, canyons and uplands. SR: 82 a occasional, grassland. Stachys coccinea Ortega. Texas betony; herba- ceous perennial. SC: 8872; locally common in canyons Trichostema arizonicum A. Gray. Blue curls; her- baceous perennial. SC: 8680, 9546; rare, canyons Linaceae Linum deseo (Engelm.) Heller. Plains flax; a al.SC: 9140, 9278; uncommon, uplands -§317-uncommon. arasslands Loasaceae Menitzelia albicaulis (Douglas ex Hook.) Douglas ex Torr.& A.Gray. Small flowered blazing star; winter annual. SC: 8847, 885 7;occasional, rocky slopes. SR: 9098; occasional, Parker Canyon. alee isolata H.S.Gentry [M. asperula Wooton tandley sensu Kearney and Peebles ae ure ee 8404; occasional, rocky slopes Mentzelia multiflora (Nutt.) A. Gray. Adonis blaz- ing star, yerba pegajosa; biennial or herba- ceous perennial. SC: 8690; occasional, sandy soils adjacent to Sonoita Creek. SR: 9373: ac- casional, grasslands, roadsides Lythraceae Lythrum californicum Torr. & A. Gray. Loosestrife, yerba del cancer, herbaceous perennial. SR: springs Rotala ramosior (L.) Koehne. Toothe cup; sum- mer annual. SC: 9674; rare, wet soil of canyon bottom Malpighiac PaPICa DS en a Rich. Herbaceous perennial. SR: oak savannas, grasslands Janusia gracilis A. Gray. Twining shrub. SC: 8516 rare, ridgetops Malvaceae Abutilon incanum (Link) Sweet. Indian mallow, MCLAUGHLIN, FLORA OF URAL AREA AND CAN RALFAFI STATE PARK 691 pelotazo; small shrub. SC: 8844; un rocky slopes. Abutilon mollicomum (Willd.) Sweet [A. sonorae A. Gray]. Tall herbaceous perennial. SC: 8447; = = unco /canyons. Abutilon parvulum A.Gray.Small-leaved abutilon; herbaceous perennial. SC: 9556; rare, can- yons. Anoda abutiloides A. Gray. Tall herbaceous pe- rennial.SC:8386, 8442, 8753;occasional, rocky slopes and canyons. Anoda cristata (L.) Schlecht. Spurred anoda, uesito; summer annual. SC: 8462, 8662; oc- casional, mostly in canyons. SR: 8566; occa- sional, grasslands. Gossypium thurberi Todaro. Desert cotton, shrub. SC: 8414; locally common, canyons, slopes, ridgetops. Herissantia crispa (L.) Brizicky [Gayoides crispum (L.) Small]. Herbaceous perennial. SC: 8477 8517; uncommon, rocky slopes and ridgetops Hit [ S.Watson.R llow;herb ceous perennial. SC:8424 rocky slopes. *Malva ees L.Cheese weed; winter annual. SR: common, disturbed areas eee tan (A. Gray) Fryxell [Sida physocalyx A. Gray]. Herbaceous ie tie ncommon, canyons. SR: 8589; un- ker Canyon. sa cut P. Mill. [S. procumbens Sw.]. Pros- herbaceous perennial. SC: 8546; occa- Bee ede 8 oe eee lands. ae neomexicana A.Gray. Herbaceous perennial. 580; occasional, Tee | a pigs L. Prickly mallow; summer annual. R: 8336, 8568; aes grasslan eres angustifolia (Cav.) G. Don var. cuspidata (A. Gray) Kearney. Narrow-leaved globe mallow; herbaceous perennial. SR: 8552; occasional, mostly below oaks. Sphaeralcea fendleri A. Gray var. fendleri. Globe mallow; robust herbaceous perennial. SR: 9687; uncommon, Santa Cruz River flood- plain. Sphaeralcea laxa Wooton & Standley. Caliche globe mallow; herbaceous perennial. SC: 8811; occasional, rocky slopes Menispermaceae Cl Hf ait ital) is DC. Snail seed, correhuela; woody vine. SC: 9478; uncommon, canyons, rocky slopes. uginaceae *Mollugo verticillata L.lndian chickweed; annual. SC: 8413; uncommon, rocky slopes. SR: 8288; common grasslands Moraceae *Morus alba L. Mulberry, mora. Tree. SF: rare, a single plant persisting near ranch buildings. Morus ee EUKICY. Texas ee mora; small tre / anyons and rocky se Nyctaginaceae Allionia incarnata L. Trailing four o' clock, wind- ne sone herbaceous ee SC:8455, ] (0) ee coccinea i Mi Red ering ag srssonds. 5 SR: a 1; occasional, gras ae disturbed a Boerhavia erecta fe Summer annual. SC: 8397, 10; uncommon, canyons and uplands. SR: 592; occasional, Parker Canyon Boerhavia intermedia M.E. Jones. ace -winged ringstern; summer annual. SC: 8514; occa- sional, mesquite grasslan Boerhavia megaptera Standley. ae annual. 5- rare, grasslands Boerhavia purpurascens A.Gray. Purple spiderling; summer annual. SC: 8522; occasional, rocky slopes. SR:8775; uncommon, n, Pa rker Canyon. Boerhavia scandens L. [Commicarpus scandens (L.) Standley]. Herbaceous eens mon,below trees and shrubs in canyons. eee aG spicata ule Summer annual. SC: |, sandy soil of canyon bot- tom. SR: 8714, 9715; uncommon, Parker Can- Mirabilis coccinea (Torr.) Benth. & Hook. f. [Oxybaphus coccineus Torr].Four o'clock, her- baceous ae SC: 9354: rare, Fresno Can- yon. SR: 9379, sslands Mirabilis eee (Pursh) Heimer| ae linearis (Pursh) B. L. Robinson]. Herbaceous perennial. SR: 8587, 8709; uncommon, grass- lands. Mirabilis longiflora L. var. wrightiana (A. Gray ex Britton & Kearney) Kearney & Peebles. Sweet four o'clock, maravilla; herbaceous perennial. SC: 9505; rare, north-facing slopes. SR: 8730, 9620; uncommon, oak savannas. Oleaceae Fraxinus velutina Torr. Velvet ash, fresno; tree. SC: 8399; common along Sonoita Creek and in larger canyons. SR: 8735; uncommon, Parker Canyon baie sabia A. Gray. Small shrub. SC: 8675, 8,932 4; locally common cany Onagraceae Camissonia californica (Nutt. ex Torr. & A. Gray en [Oenothera leptocarpa Greene}. Win- ter en SC: 9009, 9122; occasional, can- yons and rocky slopes Camissonia chamaenerioides (A. Gray) Raven [Oenothera chamaenerioides A.Gray].Winter annual. SC: 9 ncommon, canyons Camissonia claviformis (Torr. & Frémont) Raven ssp. peeblesii (Munz) Raven [Oenothera clavaeformis Torr. & Frémont var. peeblesii Munz]. Winter annual. SC: 9077; rare, one population seen in — of Sonoita Creek. Epilobium canum (Greene) Raven ssp. latifolium (Hook.) Raven ee ia (Hook.) Greene var. arizonica (Davidson) Hilend]. Hummingbird trumpet; herbaceous peren- nial. SC: 8750; i Ica Cnilnh; MNiar Rifle Pi |, W | | = ae Us 3K Willow weed; herbaceous perennial. SR:8274; common prings Gaura hexandra Ortega ssp. gracilis (Wooton & Standley) Raven & Gregory [G. gracilis Wenn & seat oo one, SC:8 ons. SR: 8260, 9481: common, cow, areas Gaura mollis James [G. parviflora Douglas ex Lehm.]. Lizard tail, velvet leaf gaura; tall her- baceous perennial. SC: 8654; occasional, mostly along Sonoita Creek. SR: 8263; com- mon, Sharp Spring, low areas, roadsides. Oenothera albicaulis Pursh. Evening primrose; annual. SR: 9239, 9477, 9514: locally sslands oer est Nutt. ssp. marginata (Nutt. x Hook. & Arn.) Munz. Evening primrose; reacts eine SC:9117; Nae a SR: assla BRIT.ORG/SIDA 22(1 = Oenothera primiveris A. Gray ssp. primiveris. Yel- low desert primrose; herbaceous perennial. SC: 8834,8918 asslands. SR:8940; common, south-facing slopes. Oenothera pubescens Willd. ex Spreng [O. laciniata Hill ssp. pubescens]. Evening prim- rose;annual. SR: 9378, 9480, 9578:uncommon, grasslands. aiid rosea L'Her. ex Ait. Sun drops; herba- eous perennial. SC: 8655; uncommon, wet a along edge of eae Creek. SR: 8278; common, springs Orobanchaceae Orobanche cooper! (A. Gray) Heller ssp. cooperi [O. ludoviciana Nut var, cooper! ( (A. Gra B herbaceous perennial. SC: 9269, 9328; occasional, grass- lands, canyons. Oxalidaceae Oxalis albicans Kunth ssp. albicans. Wood sorrel, ee herbaceous perennial. SC: 8746; on,mostly along Sonoita Creek. SR: ee occasional, grasslands. = ions A. Gray [O. amplifolia (Trel.) nsu Kearney and Peebles (1960)]. ie ne herbaceous perennial.SC: 9540, uncommon, delet) ery Ete along Sonoita Creek, canyons.S Parker Canyon ?4-UNn HTMTOT, Papaveraceae Argemone mexicana L. Prickly poppy; annual. SC: 9290; rare, along old railroad right-of-way ong Sonoita Creek, possibly introduced. pent pleiacantha Greene . saree Prickly poppy; annual. SC: 9 common along nae Creek and in canyons SR:9425; co oadsides, grasslan Eschscholzia californica Cham. ssp. mexicana crate Clark [E. Greene]. Mexi- an poppy, amapola del campo; winter an- ee nds Passifloraceae Passiflora mexicana Juss. Passion flower, Herba- perennial vine. SC:un forest, canyons, not found in flower. common gallery Pedaliaceae Proboscidea parviflora (Wooton) Wooton & Standley ssp. parviflora. Devil's claw, gatitio; MCLAUGHLIN, FLORA OF AREA AND CAN RAFAFI STATE PARK 693 summer annual. SC: 8374 soils adjacent to cone Creek. a 8335; common, grasslands Phytolaccaceae Rivina humilis L. Pigeon berry, rouge plant, coralito; herbaceous perennial. SC: 8362 commoan, canyons Plantaginaceae *Plantago major L.common plantain;annual.SC: 8742, 9314; rare, along Sonoita Cree Plantago patagonica Jacq. [P. purshii Roemer & Schultes]. Plantain; winter annual. SC: 8927; abundant, uplands. SR: 9092;abundant, grass- ds Plantago virginica L. Pale-seeded plantain; win- ter annual. SC: 9225, 9231; uncommon, mostly along pce in wet soil. SR: 9244; locally PVA RA comm ine oe and probably native in Arizona Platanaceae Platanus wrightii S. Watson. Arizona sycamore, aliso; tree. SC: rare, present as a few saplings along Sonoita Creek. Plumbaginaceae Plumbago scandens L. Leadwort, hierba de alacrdn, pitillo; herbaceous perennial. SC: 8408, uncommon, canyons, rocky slopes. Polemoniaceae pues diffusum (A. Gray) Mason. Winter an- nual. SC: 9725, 9143; uncommon, mesquite grasslands. SR: 9240; occasional grasslands Gilia flavocincta A. Nelson ssp. australis (A. & V. Grant) A. Day & V. Grant. Winter annual. SC: 8860, 9061;common, gallery forests, canyons, and uplands. SR: 8942 grasslands Gilia mexicana A. & V. Gane Winter annual. SE 9085;uncommon, mostly ridgetops. SR: 9108; common, grasslands. Ipomopsis macombii (Torr. ex A. Gray) V. Grant [Gilia macoumbii Torr]. Herbaceous peren- Ue eee ac ea c yons. lesen rasslands Nel Linanthus e ssp.aureus.Win- ae SE: 9089:1 rare, a Saab sa bigelovii (A. Gray) Greene. Winter SC: 8861; uncommon, canyons. > Phlox gracilis (Hook.) Greene [Microsteris gracilis (Hook.) Greene].Winter annual. SC: 9.214; rare Fresno Canyon Polygalac Polygal Hae te Milkwort; herbaceous peren- nial. SR: 8560, 8737; uncommon, grasslands. ies barbeyana Chod. [P. longa Blake]. Milk- t; herbaceous perennial. SC: 9405; un- common, canyons, rocky slopes. SR: 95217; nd uncommon, grassla Polygala obscura Benth. Milkwort; herbaceous perennial. SC: 8749, 9323; uncommon, can- yons, rocky slopes. SR: 9572; uncommon, grasslands. Polygona Eriogonum ae Torr.var.abertianum.Wild buckwheat; annual. SC:8384;common,all ter- restrial habitats. SR: 8330, 8331; occasional, rasslands. suena polycladon Benth. Sorrel buckwheat; mer annual. SC: 8517 oe sas po to eile Creek, other dis- turbed are 7,occasional, nen aerard ee i Torr. ex Benth. var. wrightii. ht buckwheat; low shrub. SC: 9729; oc- ae ae rocky slopes. SR: 8779; common, grasslands. *Polygonum aviculare L. Prostrate knotweed, chilillo; herbaceous perennial.SR:8342; locally abundant, tanks pay lapel ffolium L.Willow smarty |.SR:8266; pally abundant, cattle Anke Santa Cruz River bot- tom. Polygonum pensylvanica L. Pinkweed, chilillo; herbaceous perennial. cea ie aa dant, low areas, cattle ta Polygonum punctatum ale Var. - confertiflorum (Meisn.) Fassett. Water smartweed, chilillo; herbaceous perennial. SC: 8497, 8653; occa- sional, stream edges. *Rumex crispus L. Curley dock; herbaceous pe- rennial. SR: 8780, 9427; occasional, springs. Rumex hymenosepalus Torr. Wild rhubarb, cahaigre; herbaceous perennial. SC: uncom- mon, gallery forest along Sonoita Creek. Rumex violescens Rech. f. Herbaceous perennial. SC: 9318; uncommon, edge of Sonoita Creek. Portulacaceae Calandrinia ciliata (Ruiz & Pavén) DC. Red maids; winter annual. SC: 8854; common, canyons, grasslands. SR: 8938; common, low areas, roadsides Cistanthe ae (A. Gray) Hershkovitz var. arizonica (J. T. Howell) Kartesz & Gandhi [Cal ean parryi A. Gray]. Winter annual. SC: 8928; uncommon, grasslands Portulaca suffrutescens Engelm.Purslane; herba- ceous perennial. SC: 8420; common, mes- quite grasslands, ridgetops. SR:8567; uncom- mon, grasslands. Portulaca umbraticola Kunth ssp. coronata Small) Matthews & Ketron [2 coronata Small]. Purslane; summer annual. SC: 83717; common, uplands. SR: 8286, 8570; uncom- — n,u ands. inum aurantiacum Engelm. Flame flower; her Tal ‘ {| | baceous enn SC: occasional, grass- lands. | arasslands Talinum paniculatum (Jacq.) Gaertn. Pink baby reath SC: 8353; common, canyons. Primulaceae *“Anagallis arvensis De Pimpernel, yeiog ae Pale annual. Androsace occidentalis Pursh. Rock jasmine; win- ter annual. SC: 8827 canyons, up- lands. SR:8936, 8950: common, low areas, Sa- caton flats. Samolus vagans Greene. Water pimpernel; her- baceous perennial. SC: 8492; rare, wet soil along stream, Fresno Canyon. Ranunculaceae Anemone tuberosa Rydberg var. tuberosa. Desert windflower; herbaceous perennial. SC:8825; occasional, rocky slopes. Clematis drummondii Torr. & A. Gray. Texas virgin bower, barbas de chivito; woody vine. SC: 403; uncommon, Sonoita Creek. Delphinium scaposum Greene. Barestem lark- spur; herbaceous pais: SC: 9066; locally common, uplan Delphinium wootonii a berg [D. virescens Nutt. JE gallery forest along a WOON act ]. Larkspur; her- | I.SR:9357-rare ae Myosurus cupulat S Walco Mouseeilwiaer BRIT.ORG/SIDA 22(1) annual. SC: 8862, 9010, 9023; abundant, wet soil of canyon bottom, north-facing slopes. SR: 8934, 9056; abundant, low areas, sacaton flats. Myosurus minimus L. Mousetail; winter annual. SR: 9254, 9350; uncommon, springs. Ranunculus hydrocharoides A. Gray. var. taeg dasee Buttercup, herbaceous pe- ial. SR: 9259; locally co aoe macranthus Scheele. Large butter- cup;herbaceous perennial. SR: 9485; uncom- mmon, springs mon Thalictrum Pai Engelm.ex A. Gray var.fendleri. Meadow rue; herbaceous perennial.SC:8543; rare, canyons, north-facing slopes. Rhamnaceae Ceanothus greggii A. Gray. Buckbrush; shrub. SC: 9544; rare, Canyons Condalia warnockii M. C. Johnston. var. kearneyana M. C. Johnston [Condalia mexicana Schlecht. sensu Kearney & Peebles (1960)]. Mexican bluewood; shrub. SC: 8367; occasional, rocky slopes Sageretia wrightii S. Watson. Sarl SC: 8671; lo- cally common, canyons. — obtusifolia (Hook. ex Torr. & A. Gray) A Gray var. canescens (A. Gray) M.C. Johnston ae lycioides (A. Gray) Weberb.]. Gray thorn, abrojo, bachata; shrub. SC: 9494; occa- sional, mostly along edge of gallery forest along Sonoita Creek. osaceae *Malus pumilla P. Mill. Apple; tree. SR: 9484, a few trees persisting at Sharp Spring near junc- tion with Santa Cruz River floodplain Potentilla wheeleri S. Watson [P viscidula ayabi Cinquefoil; herbaceous perennial. SR: 9059; uncommon, north-facing slopes in grass- lands, usually found at much higher eleva- tions in southeastern Arizona Purshia stansburiana (Torr.) Henrickson [Cowania mexicana D.Don var. stansburiana (Torr) Jespon]. Cliffrose, pofio; large shrub. SC: 44; occasional, uplands. SR: 8724; uncom- mon, grasslands. Rubiaceae Bouvardia ternifolia (Cav.) Schlecht.[B. glaberrima ngelm.]. Shrub. SC: 8435, 9361; rare, north- facing slopes, ridgetops. MCLAUGHLIN, FLORA OF SONOITA CREEK NATURAL AREA AND SAN RAFAEL STATE PARK 695 Diodia teres Walt. var. angustata A. Gray. Rough buttonweed, poor Joe; summer annual. SC: 8767. rare, canyons. SR: 8719; uncommon, grasslands Galium aparine L. Goosegrass bedstraw, cleav- ers; annual vine. SC: 9083; common in can- ons, below oaks. Galium microphyllum A.Gray. Herbaceous peren- nial. SC: 9355; uncommon, rock walls. Galium ee A.Gray.Great Basin bedstraw; annual. SC: 9114, 9275; uncommon, canyons. Galium Ae aloe perennial. SC: 02; uncommon, walls. Houstonia fled A. cy nce perennial. SR: 8 common, north-facing slo ssla ae bcc breviflorus A. Gray. Summer annual. SC: 8 9721; uncommon, ew canyon eee asslands Rutaceae Thamnosma texana (A. Gray) Torr. Turpentine broom; herbaceous perennial. SC: 9067; un- common, uplands Salicaceae Populus fremontii S. Watson. Fremont cotton- wood, dlamo; large tree. SC: commen, gal- lery forest al S eek.SR:common, springs, Santa Cruz River ek Salix bonplandiana Kunth. Bonpland willow, uce, sauz; tree. SC: 8651, 8665, 9264; occa- sional, gallery forest along Sonoita Creek. Salix exilifolia Dorn [Salix taxifolia Kunth]. Yew- ; small tree. SC: 8669, 9150; rare, Sonoita Creek. SR: 9267; uncommon, springs, Santa Cruz River bott Salix gooddingii Ball. Goodding ee sauce, sauz; tree. SC: 8876, 9148; abundant, gallery forest along Sonoita Creek. SR: 9429; com- mon, springs, Santa Cruz River bottom. Salix lasiolepis Benth. var. lasiolepis. Arroyo wil- low, sauce; large shrub. SC: 9064; common, Sonoita Creek.SR:8949, 91 11,9260, 9262;com- mon, springs, Santa Cruz River bottom. leaf willow, tardi. Sapindaceae Dodonaea viscosa (L deed Hopbush, jarilla, tara- C:9 canyons chiqui;shru ged saponaria L. var.drummondii (Ho rn.) L.Benson.Western soapberry, amoli ll, cirioni, jaboncillo; small tree. SC: 9498; occa- er canyons, rocky slopes. SR: 8596; un- common, Parker Canyon. Scrophulariaceae Castilleja exserta (Heller) Chuang & Heckard ssp. exserta [Orthocarpus purpurascens Benth,]. Owl's clover;hemiparasitic winter annual. SC: 208; rare, Mesquite grasslands. Castilleja integra A. Gray. Indian paintbrush; hemiparasitic herbaceous perennial. SR: 9103, 9251; occasional, grasslands Castilleja lanata A sp./anata.\indian paint- brush; hemiparasitic herbaceous perennial. SC: 8917, 9137, uncommon, canyons aoe minor (A. Gray) A. Gray ssp. minor. In- aintbrush; hemiparasitic annual. SC: ee e, canyon bot : Maurandella antirrhiniflora (Humb. & Bonpl. ex Willd.) Rothm. [Maurandya antirrhiflora Humb.& Bonpl.]. Herbaceous perennial vine. SC: 8677: ional, canyons, rock walls. Mecardonia procumbens (P. Mill.) Small [M. vandellioides (Kunth) Pennell]. Herbaceous perennial. SC: 8445, 8468; occasional, canyon bottom. Mimulus floribundus Lindley. Monkey flower, baisémari, herbaceous perennial. SC: 9234; rare, Coal Mine Canyon Mimulus guttatus DC. Monkey flower, baisomari; baceous perennial. SC: 8838; common, ri- parian areas. SR: 9096; common, Santa Cruz River bottom. Mimulus rubellus A.Gray.Red-stemmed mimulus; winter annual. SC: 8828; locally common, mostly in canyons. SR:9053;occasional, grass- lands. Nuttallanthus texanus (Scheele) D.A. Sutton [Linaria texana Scheele]. Texas toad flax; win- ter annual. SC: 8864, 8912; common, rocky slopes. SR: 9094; grasslands Penstemon barbatus (Cav.) Roth ssp. barbatus. Scarlet bugler; herbaceous perennial. SC: 9407, rare, Fresno Canyon. SR: 9453; rare, oak woodlands adjacent to Parker Canyon. Penstemon linarioides A. Gray ssp. linarioides. ous perennial. SR: 9383; occasional, grass a ae Penstemon parryi (A. Gray) A. Gray. Herbaceous perennial. SC: 9720; occasional, rocky slopes, rock walls. Penstemon ei (A. Gray) T. J. Howell. ngue; herbaceous perennial. SR: 9525,9568; uncommon, north-facing slopes in oak woodlands and grasslands. Sairocarpus nuttallianus (Benth. ex A. DC.) D. A. Sutton [Antirrhinum nuttallianum Benth,]. Winter annual. SC: 9127; rare, rocky slopes. Schi alae ee gsi (A. Gray) Pennell. annual. SC: 8480; uncommon, eae a ee (L.) Sw. Herbaceous peren- nial, SC: 9404; rare, rock walls, Fresno Canyon. SR: 8734; rare, Parker Canyon. Verbascum vi Stokes. Mull bust an- nual. SC: 9476; uncommon, along Sonoita Creek. SR: 8306; uncommon, springs *Veronica anagallis-aquatica L.Water speedwell, herbaceous perennial. SC: 9145; occasional, riparian areas Veronica Sa eate 2 i880. xalapensis (Kunth) Pen lly winter annual. SC: 8859 on 5. sbundant canyons and nid drainages. SR:8 es Simaroubaceae *“Ailanthus altissimus (P. Mill.) Swingle. Tree of ven, Copal tree; tree. SC:one large colony in the gallery forest of Sonoita Creek. SR:8272; Il col | road east of ¥ dlONnG uncommon the Santa Cruz River. Solanaceae Calibrachoa parviflora (Juss.) D'Arcy aad parviflora Juss.]. Wild petunia; annual. $ 9149; uncommon, wet soil along Sono Creek. SR:9 tanks. ee ede coniodes ee ex Dunal) Brit- ton Small ground cl nial. SC: 9296; uncommon, grasslands, SR: 944]: roadsides Chamaesaracha coronopus (Dunal) A.Gray. False nightshade; herbaceous perennial. SR: 9367, 9402; uncommon, roa Datura quercifolia Kunth. eon weed toloache; herbaceous perennial.SR:8577; uncommon, roadsides Datura wight Regel [D. meteloides DC]. Sacred datura, toloache; herbaceous perennial. SC: ccasional, sandy soils adjacent to Sonoita Creek. SR:8564; occasional, roadsides, BRIT.ORG/SIDA 22(1) Lycium berlandieri Dunal var. parviflorum (A.Gray) Terracc. Wolfberry; shrub. SC: 8874; uncom- on, uplands. lycium exsertum A. Gray. Wolfberry; shrub. SC: 8817, uncommon, uplands. Margaranthus solanaceus Schlecht. Netted globe berry; summer annual. SR: 8725; un- common, oak woodland adjacent to Parker Canyon *Nicotiana glauca Graham. Tree tobacco, corneton, juan loco, palo loco; robust herba- ceous perennial. SC: 8693; occasional along onoita Creek. Nicotiana obtusifolia Martens & Galleotti var. obtusifolia [N. trigonophylla Dunal var. trigonophylla]. Desert tobacco, tabaquillo; herbaceous perennial. SC: 8684; occasional, nyon walls Physalis acutifolia (Miers) Sandw. [P wrightii A. Gray].Wright sual cherry, tomatito; sum- mer annual.SR: 8265, 8341: |, grass- lands, springs Physalis hederifolia A. Gray var. hederifolia. lvy- leaved ground cherry, tomatito; herbaceous perennial.SR:8588; uncommon, Parker Canyon. Physalis longifolia Nutt. Tomatito; herbaceous perennial. SR: 8299, 8300, 8706; common, springs. Physalis pubescens L. var. integrifolia (Dunal) Wa- terfall. Hairy ground cherry, tomate, fresadilla; DL rn dn Solanum douglasii Dunal. Nightshade; herba- ceous perennial. SC: 8491; rare, Fresno Canyon. Solanum elaeagnifolium Cav. Silverleaf night- shade, trompillo; herbaceous perennial. SC: 9363; occasional, mesquite grasslands. SR: 9398; occasional, disturbed areas. Solanum rostratum Dunal. Buffalo bur; herba- ceous perennial. SC: 9633; uncommon mostly along Sonoita Creek. SR: uncommon, roadsides Sterculiaceae Ayenia filiformis S.Watson [A. pusilla L.sensu Kear- ney and Peebles (1960)]. Herbaceous peren- nial. SC: 8372; occasional, rocky slopes. Tamaricaceae aaa Moss iin Ledeb. [7. pentandra Pall. SETISU KEAINey dla | Peet sles (1 960)] Tamarisk, MCLAUGHLIN, FLORA OF SONOITA CREEK NATURAL AREA AND SAN RAFAEL STATE PARK 697 pino salado; small tree. SR: 9348; rare, Santa Cruz River bottom Ulmaceae Celtis reticulata Torr. Hackberry, palo blanco, cumaro; tree. SC common, gallery for- ests and canyons. $R:8345; f Santa Cruz River floodplain. Celtis pallida Torr. Desert hackberry, garambullo; — edge ~~ rub. SC: uncommon, rocky slopes. roe rticaceae Parietaria bes Hinton var.hespera [P floridana Nutt nd Peebles (1960)].Pel- litory, Saeuines winter annual. SC: 8830; oc- casional, rocky slopes Q Verbenaceae Aloysia wrightii Heller ex Abrams. Wright lippia, oreganillo; shrub. SC: 8525; common, rock slopes. Bouchea prismatica (L.) Kuntze. Summer annual. 59; uncommon, low areas. Glandularia bipinnatifida (Nutt.) Nutt. var. bipinnatifida. [Verbena bipinnatifida Nutt]. Verbena; herbaceous oe SR: 8261], 9101;common, grasslan Phyla nodiflora (L.) Greene. ee peren- nial. SR: 9443; one a near ranch build- ings, probably introduc Verbena carolina L. Fese-oul perennial. SC: uncommon, canyons Verbena ae is Desf. orbacealls perennial. SR: Small var, xylopoda Perry. Hillside vervain; herbaceous perennial. SC: 8478, 8739; occasional, mostly on shaded slopes. Necheeae alifornicum Nutt. Desert mistle- toe; enaicl herbaceous epiphyte. SC: 8849, occasional on Prosopis, Condalia. Phoradendron capitellatum Torr. ex Trel. [P. bolleanum (Seem.) Eichler var. capitellatum (Torr. ex Trel.) Kearney & Peebles]. Mistletoe; | us ee SC:8685, hemiparastic I er coryae Trel, ene hemipara- sitic herbaceous epiphyte. SC: 8868; occa- sional on Quer blongifolia. SR: : casional, on Ouen rCUS SPP. Phoradendron tomentosum (DC.) Engelm. ex A. Gray [Pflavescens (Pursh) Nutt. var. pubescens Engelm.]. Mistletoe, inierto; hemiparasitic herbaceous epiphyte. SC: 9018; uncommon on Fraxinus Vitaceae Vitis arizonica Bnet) sn grape, parra del monte; woody vine. 417;occasional,can- yons. SR: 9395; nae common, Parker Canyon. Zygophyllaceae Kallstroemia grandiflora Torr.ex A.Gray. Mexican poppy, summer poppy, baiburin, mal de ojo; summer annual. SC: 8361; occasional, can- yons and slopes. Kallstroemia parviflora Norton. Caltrop; summer annual.SR:8339 Parker Canyon. *Tribulus terrestris L. Caltrop; summer annual. SR: 9591; uncommon, disturbed sites. 8713;:occasional, grasslands MAGNOLIOPHYTA: LILIOPSIDA Agavaceae Agave palmeri Engelm.Palmer agave, lechuguilla; large leaf succulent. SC: occasional, canyons and rocky slopes. SR:uncommon, grasslands. Agave parryi Engelm. var. parryi. Century plant; large leaf succulent. SR: uncommon, grass- lands. Agave schottii Engelm. var. schottii. Shindagger. amolillo ll leaf succulent. SC: 9413; bundant on uplands Yucca elata Engelm. Soap tree yucca, palmilla; caulescent leaf succulent. SC: 9408; uncom- mon, canyons. SR: rare, a single individual observed in grassland west of ranch house. Yucca schottii Engelm. Mountain yucca; caulescent leaf succulent. SC: uncommon, north- facing slope, Coal Mine Canyon. SR: rare, a single individual observed with oaks east of Santa Cruz Yucca oe ee Banana yucca, datil: acaulescent or short-caulescent leaf succu- lent. SC: abundant, uplands. Kartesz (1999) treats this as a variety of Yucca baccata; the latter taxon has strongly glaucous foliage, thick leaf fibers, and tends to form dense colonies of acaulescent rosettes. The plants of southern Arizona have green foliage, thin- ner fibers, and are often caulescent amol:sma 698 Alismataceae Sagittaria se Engelm. ex J. G. Sm. Arrow- head; aquatic perennial. SC: 6488, J.Kaiser 1 ber ae collected four times from a Cattle tank adjacent to Santa Cruz River, formerly plentiful’; last collected in 1991, Bromeliaceae Tillandsia recurvata (L.) L. Ball moss, gallitos; epi- La leaf Succulent, SC: 8678; locally abun- niperus coahuilensis in Narrow Canyons. Commelinace Commelina di itl ifolia Delile. Day flower, tale de Maria; herbaceous perennial. SC: ons and rocky slopes. n, grassland Coramnelien erecta L. var. angustifolia (Michx.) Fern. Day flower, yerba del pollo; herbaceous perennial.SC:8355;uncommon,canyons and rocky slopes. SR: 9622; uncommon, Parker SR 8314 Canyon. Tradescantia pinetorum Greene. Spiderwort; her- aceous perennial. SR: 9565; rare, grasslands. Cyperaceae Carex praegracilis W. Boott. Sedge, herbaceous i re ae SR: 9248, 253, ‘ona 9463; abun- |\ y collections are atc to eran om C.chihuahuensis = ackenzie. oped Liebamann [C. wrightii aaa edge; herbaceous perennial. SC: 8 common,north-facing slopes below oa ee esculentus L. eed hutseelge) herba- ceous perennial. SC: 9 an- yons. SR: 9593, 9598, 9611 7 occasional, grass- lands, roadsides. Cyperus none Michx. [C. albomarginatus Mart. & Schrad.]. Flat sedge; herbaceous pe- rennial. SC: 8467, 8539, 9647; common along ream in Fresno Canyon. Cyperus mutisii (Kunth) Griseb. Flat sedge; her- baceous perennial. SC: 9708; rare, Fresno Canyon, ne niger Ruiz & Pavon. Flag sedge; herba- ous perennial. SC: 8776; uncommon, a Canyon. SR:8324,8789, 9599;common, nta Cruz River bottom ope odoratus L. Flat sedge; herbaceous pe- rennial.SC:8495, 9668, 9684; common in wet BRIT.ORG/SIDA 22(1) soil ae Sonoita Creek. SR: 9668; common, z River bottom. Cyperus pal iin or (Kukenthal) G. Tucker [C. flavus (Vahl) Nees, in part]. Flat na her- baceous perennial. SC:8438; rare, below cane sphaerolepis Boeckl.[C. rusbyi en i ge; herbaceous perennial. SR:8508; occa- he oak sav Cyperus SqUBHOSUS ai [C. aristatus Rottb.]. Flat sedge; annual. SC: 9497, 9640; uncommon, moist af of canyon bottom Eleocharis bella (Piper) Svenson. Spike rush;sum- ual. SR: 9575 |, Santa Cruz River bottom. Eleocharis montevidensis Kunth. Spike rush; her- ; -41 SC-939()-a} along streams. SR: 9437; abundant, springs, Santa Cruz River bottom Eleocharis palustris (L.) Roemer & J.A.Schultes [. macrostachya Britton sensu Kearney & Peebles (1960), in part]. Spike rush, herba- ceous perennial. SC: 9293: occasional, mostly in canyons. nr aca all Cruz River bot Fimbristylis annua a Roemer & J.A. Schultes [F. baldwiniana (Schultes) Torr]. ara an- nual. SC:8539; uncommon, canyon bot Lipocarpa micrantha (Vahl) G. Tucker heee carpha micrantha (Vahl) ie Summer an- nual. SC: 9675; n botto sae ore acutus (Muhl. ex oa) A. ‘é ssp. occidentalis (S.Watson) $.G.Sm. ae acutus Muhl.]. Bullrush; herbaceous perennial. SR: 9461; uncommon, springs. Schoenoplectus americanus (Pers.) Volk.ex Schinz &R. a istaabe americanus assets Bullrush; CPOUS p — Ut] MT IOrd, Santa Cruz River bottom. lridaceae Sisyrinchium demissum Greene. Blue eye grass; herbaceous perennial. SR: 9376; common, springs JUNCACEAE Juncus bufonius L. Toad rush; annual. SC: 97.73, mmon, moist soils of canyon bottom. Juncus interior i eeaealereaciae a perennial. SC: 9236, 9294 ms. Juncus mexicanus Willd. ex J A & J.H. Schultes. Rush; herbaceous perennial. SR: 9325, 9388, gq =) MCLAUGHLIN, FLORA OF SONOITA CREEK NATU AREA AND SAN RAFAEL STATE PARK 699 9430, 9433; abundant, springs, Santa Cruz River bot Juncus saximontanus A. Nelson. Rush; herba- ceous perennial.SC:8493, 9210;uncommon, along streams. SR: 9315, 9438, 9464, 9487. _ springs, Santa Cruz River bottom. Juncus torreyi ae Rush; herbaceous peren- nial. SR: 8284, 9435; common, springs, Santa Cruz River bottom. commo { {| €] eae Lemna gibba L.Windbags, duck ing aquatic. SC: rare, Fresno Canyon. SR:8373; occasional, springs. Usually found as a minor component in mixed colonies with L. minor. Lemna minor L. Water lentil, duckweed: small floating aquatic. SC: 8499; locally abundant in pools of canyons. SR: 8372; common, springs Liliaceae Allium kunthii G. Don. Wild onion; herbaceous perennial. SC: 9644; rare, Coal Mine Canyon. Calochortus ambiguus (M.E. Jones) Ownbey. Mari lily;hert ial. SC: 9282. locally abundant, Fresno Canyon. Calochortus kennedyi Porter var. kennedy. so ariposa; herbaceous perennial. SC: 9724; occasional, uplands. SR: 9241; locally com- mon, grasslands. Dichelostemma capitatum (Benth.) Wood ssp. pauciflorum (Torr.) G. Keator [D. pulchellum (Salisb.) Heller].Bluedicks; herbaceous peren- nial. SC: 8929; common, uplands. SR: 9106; grasslan Pee as pee U. 7 & H.H.Schultes) Cruden [Anthericum torreyi Baker]. Crag lily; herba- eous perennial SR: 95278 grasslands. Milla biflora Cav. Mexican Star, herbaceous pe- rennial. SC: rare, included based on a photo by R. Gagnon. SR: 9631; rare, grasslands Nothoscordum texanum M.E. Jones. Herbaceous perennial.SC:8440, 9235; uncommon,mostly in seeps in Coal Mine Canyon. Nolinaceae Dasylirion wheeleri S. Watson. Desert spoon, so- tol; leaf rosette shrub. SC: common, rocky slopes. Nolina microcarpa S. Watson. Beargrass, sacahuista, palmilla; leaf rosette shrub. SC: common, rocky slopes, grasslands. SR: locally abundant, grasslands and oak wooclands. Poaceae Agrostis exarata Trinius. Spike bent; herbaceous perennial. SR: 9439; Santa Cruz Alopecurus carolinianus Walt. Fox tail; summer annual. SR: 9255; uncommon, springs. Aristida adscencionis L. Six weeks three-awn; summer annual. SC: 8426, 9229: common, uplands. SR: 9588; occasional, grasslands, dsides. Aristida divaricata Humb,& Bonpl.ex Willd. Three- awn; herbaceous perennial. SR: 9698; occa- sional, grasslands Aristida purpurea Nutt. var. fendleriana (Steud.) Vasey [A. fendleriana Steud.]. Purple three- awn; herbaceous perennial. SR: 9374; occa- sional, grasslands Aristida purpurea Nutt. var. neallyi (Vasey) Allred a glauca ise ia SC: 9321, 9543; u n, uplan Pee purpurea ea var. wrightii (Nash) Allred A. wrightii Nash]. Purple three-awn; herba- ceous perennial. SC: 8573; occasional, grass- lands. SR: 9605; common, gra asslands ristida ternipes Cav. var.gentilis (Henr.) A lred [A. mulosa Henr]J. Spider grass; herbaceous perennial SC: 8428; common, canyons. SR: grasslands = co Asti as Cav. var. ternipes. SC: 8505, 8520 ceous perennial; Common, canyons. mt fatua L.Wild oats; winter annual. SC: 9238; rare, canyons. SR: 9264; uncommon, dis- turbed areas. *Avena sativa L. Oats; winter annual. SR: 9752; incommon, disturbed a birch barbinodis a Herter [Andro- rbinodis Lag.J. Cane beardstem; feat perennial. SC: 8430; abundant, uplands. SR: 8329; abundant, grasslands. Bothriochloa laguroides (DC.) Herter ssp. torreyana (Steud.) Allred & Gould [Andro- pogon ISO Swartz]. alee rp Spring perennial.S drainage. eee aristidoides (Kunth) Griseb. var. aristidoides. Six-weeks needle grama; sum- mer annual. SC: 8434; occasional, canyons and uplands. SR: 8572; common, grasslands 700 Bouteloua chondrosioides (Kunth) Benth. ex S. Watson. Spruce-top grama; herbaceous ES ee SC:8433,8502;common,uplands.SR 8708 grasslands — curti inenctale (Michx.) Torr. Side oats erbaceous perennial. SC:8402; abun- ee canyons and uplands. SR: 8550; abun- dant grasslands woodlands sasaueson a ae selects peren- nial.S and adjacent to BRIT.ORG/SIDA 22(1) areas. SR: 8549; | grasslands, road- sides. Cottea pappophoroides Kunth. Cotta grass; her- baceous perennial. SC: 9679; rare, Fresno Canyon. *Cynodon dactylon (L.) Pers.Bermuda grass; her- aceous perennial. SC: 9284; occasional, mostly along streams. SR: 9563; ss aahdiad fale Santa Cruz River botto pulchella (Kunth) Willd. ex caviar Parker ee Bouteloua ene ou (Torr.) Torr. Black grama;her- s perennial. SC: 9637; uncommon, tee aa SR: 9604; sslands Bouteloua gracilis (Willd. ex it Lag. ex Griffiths. Blue ist eae perennial. grasslands ~~ SR: 9 Bouteloua hirsuta Lag. var. aa Hairy grama; NELDacSOUs Perennial SC: 8429; common, uplands grasslands Bouteloua railcar (E. Fourn.) Griffiths. Herba- eous perennial. SC: 8761; occasional, up- lands. SR: 9626; uncommon, oak woodland adjacent to Parker Can Bouteloua repens (Kunth) cation: & Merr. [B. filiformis (Fourn.) Griffiths]. Slender grama; erbaceous perennial. SC: 8427; common, uplands. Bouteloua rothrockii Vasey. Rothrock grama;her- ceous perennial. SC: 8528A; occasional, mesquite grasslands. SR: 9601; occasional, low areas, sacaton flats. Bromus anomalus Rupr. ex E.Fourn. Brome; her- eous perennial. SR: 9696; rare, Parker Canyon. Bromus carinatus Hook. & Arn. California brome; herbaceous perennial. SC: 9215, 9267; com- mon, gallery ies along Sonoita Creek. *Bromus catharticus Vahl.Brome; annual. SR: 9052 9158, 7 commen Sn Cruz River flood- plain, disturbed are *Bromus japonicus =n ex Murr. Japanese bro rama: annual. SR: 9389; common, Sharp Spring drain pence, spinifex a c nay orus Benth. Sandbur;summer annual.SC:8667;common, mostly in su ae ieamaa areas. SR:85 76; mmon, grassla Chloris virgata her fingergrass; sum- mer eae SC: 8398B; its, disturbed Swal L pee ‘pulchellus (Kunth) Hitchc.]. Fluff grass; herbaceous perennial. SC: 9547, 9648; locally common, uplands Digitaria californica (Benth.) Henr. [Trichachne californica (Benth.) Chase]. Arizona cottontop; herbaceous perennial. SC: 8432; uncommon, uplands. SR: 9609; uncommon, grasslands. Digitaria cognata (J.A. Schultes) Pilger var. pubiflora Vasey ex L.H. Dewey [Leptoloma cognatum (Schultes) Chase]. Fall witch grass; teease| “41 SC-9028 ae Coal Mine Canyon. “Digitaria sanguinalis (L.) Scop. Common c grass; summer annual. SC: 8660; ator wet areas. SR: 9589; occasional, disturbed areas. “Echinochloa colana (L.) Link. Jungle rice; sum- mer annual. SC: 8537; ional, canyons *Echinochloa crus-galli (L.) P. Beauv. Barnyard rass; summer annual. SC: 8452, 8487; occa- sional, along streams. SR: 8269; common, Santa a River bottom, abundant around cattle Elionurus ae tibial s Hack. Herbaceous peren- nial. SC: 97108; rare, shaded slopes Elymus canadensis L.Wild rye; herbaeous peren- nial. SR: 94 Santa Cruz River bot- tom, springs. Elymus elymoides (Raf.) Swezey [Sitanion hystrix (Nutt.) J.G. Smith]. Bottlebrush squirrel tail; herbaceous Lear SC: 91] 8; Secsleiiel R: 9263; |, Sharp canyons, slopes Spring drainage. Enneapogon desvauxii Desv. ex Beauv. Spike pappusarass;summer annual. SC:8474, 8482; uncommon, rocky slopes. SR: 8777; uncom- n, grasslands. eet ae (All.) Vign.ex Janchen. Stink grass; annual. SC: 8450; mostly in heavily MCLAUGHLIN, FLORA OF SONOITA CREEK NATURAL AREA AND SAN RAFAEI STATE PARK 701 Cp.ea74 grazed areas. SR: 82 |, disturbed areas. *Eragrostis curvula (Schrad.) Nees var. conferta nial. SC: 9277; eee intermedia Hitchc. Plains lovegrass; er ePSTeniich SC: 8439; common, me a 8548 dant, grasslands ks lehmanniana Nees. Lehmann vegrass; herbaceous perennial. SC: 9499; Bi mostly oe saddles. SR: 9483: locally co Eragrostis pectinacea (Michx.) Nees ex Steud. var. pectinacea [E. diffusa Buckley]. Summer an- ual. SC: 8530, 9495: occasional, canyons. SR: 9612; occasional, grasslands, ene Eriochloa acuminata (Presl) Kunth var.acuminata (Vasey) R.B. Shaw [E. gracilis (Fourn.) Hitche. var. gracilis (Vasey) Hitchc.]. Cupgrass; sum- mer annual. SC: 8375; common, gallery for- ests, Canyons. Eriochloa acuminata (Presl) Kunth var. minor (Vasey) R.B. Shaw [E. gracilis (Fourn.) Hitch. var.minor (Vasey) pcue Cupgrass;summer canyons mmon, grasslan LO) lox oO 2 se ual. SR: QI Hl + | OL ann ne arundinacea Schreber. [F. elatior L.]. Meadow fescue; herbaceous perennial. SR: 9299, 9314, 9352; abundant, springs Heteropogon contortus (L.) Beauv. ex Roemer & J.A.Schultes. Tangle head; herbaceous peren- nial. SC: 8770; rocky slopes Heteropogon melanocarpus (EIl.) Ell. ex Benth. Sweet tangle head;summer annual. SC:8396; common, rocky slopes. SR: 8702; occasional, oak savannas. Hilaria belangeri (Steud.) Nash var. belangeri. Curly mesquite grass; herbaceous perennial. SC: 8816; locally abundant, grasslands. SR: 9581; occasional, grasslands. Hordeum jubatum L.ssp.jubatum. Foxtail;herba- ceous Beicuael SR: 9308B, 9349; common, springs, low areas *Hordeum murinum L. ssp. leporinum (Link) Arcang. Wild barley; winter annual. SR: 9049; asional, margins of cattle tanks Hordeum eid Nutt. Little barley; annual. SR: 8A; uncommon, margins of cattle tanks. Leersia oryzoides (L.) Sw. Rice cutgrass; herba- ceous perennial. SR:8783, 8786; uncommon, Sharp Spring. Leptochloa dubia (Kunth) Nees.Green sprangle- top; herbaceous perennial. SC:8421;uncom- mon, rocky slopes. SR:8707; occasional, grass- lands. ec oa fusca (L.) Kunth ssp. fascicularis (Lam.) w [L. fascicularis (Lam.) A. Gray var. aaa Beaded sprangletop; summer i aoe disturbed are Leptochloa ie (Retz) Ohwi ssp. lies (Steudl.) N. Snow [L. filiformis Lam.].Sum annual. SC: 8529; common, canyons. Leptochloa viscida (Scribn.) Beal.Summer annual. SC: 9496; occasional, mostly in canyons. SR: F mmon, washes. Lycurus setosus (Nutt.) C. G. Reeder [L. phleoides Kunth sensu Kearney and Peebles (1960), in part]. Wolf-tail; herbaceous perennial. SC: 8425;common, uplands. SR: 9570; abundant, — grasslands. *Melinis repens (Willd.) Zizka [Rhynchelytrum roseum (Nees) Stapf & C. E. Hubbards ex Bews]. Natal grass; herbaceous perennial. SC: 813:rare, canyons. SR: 9736; rare, Parker Can- yon. NArihl ica Scribn. Herbaceous pe- enna SC: 8760, 9078; locally abundant, up- lands. SR: 9658; uncommon, grassland Muhlenbergia oe (Nees & ee ex Trinius) Parodi. Scratchgrass; herbaceous perennial. SR: 9656; locally abundant, Sharp Spring drainage. Muhlenbergia emersleyi Vasey. Bullgrass; herba- ceous perennial. SC: 8676, 8762; occasional, mostly north-facing slopes. a fragilis Swallen. Summer annual. ; asional, canyons and rocky oe SR:8730, 9725; uncommon, oak wood- land adjacent to Parker Can a ate rigens (Benth. oe Deer grass; herbaceous ane SC: 8764; abundant, m. SR: 9686; occasional, drain- canyon b ages. Muhlenbergia tenuifolia (Kunth) Trinius [M. monticola Buckley]. Herbaceous perennial. SC: 9642A, 9676, 9705; occasional, rock walls. Panicum bulbosum Kunth. Bulb panicum;herba- ceous perennial. SC:8378,8507; uncommon, canyons. SR: 9628; uncommon, oak wood- land adjacent to Parker Canyon. Panicum hallii Vasey var. hallii. Herbaceous pe- ial SR: rare, grasslands Sacon hirticaule J. Presl. Summer annual. SC: 8391, 8395, 8449; common, canyons and slopes. SR: 8285; common, grasslands. Panicum obtusum Kunth. Vine mesquite; herba- ceous perennial. SC: 9549; uncommon, washes. SR: 9578; occasional, grasslands, springs. “Paspalum dilatatum Poir. Dallis grass; herba- ceous Sal SC:8664; streams SR: 832 |, Sharp Spring. Paspalum - stichum L. Knotgrass; herbaceous perennial. SR: 8788; locally common, Santa Cruz River bottom Paspalum setaceum Michx ceous perennial. SR: 8 .[Pstramineum Nash]. 05, 9595: un- common, Oak 5 S. Phalaris caroliniana Walt. Canary grass; annual. SR: 9309, 9428: occasional, low areas. *Phalaris minor Retzius. Littleseed cama grass; ual. SC: 9204; /canyon Poa bigeloviiVasey & Seti: Bigelow's pie ae winter annual. SC canyons rocky slopes. *Poa pratensis L. ssp. pratensis. Kentucky blue- grass; herbaceous perennial. SR: 9256; abun- ant, Sharp and Heron springs drainages. *Polypogon monspeliensis (L.) Desf. Rabbitfoot grass; annual. SC: 9147; common along streams. SR: 9099, 9372; anta Cruz River bottom. *Polypogon viridis (Gouan) Breistr. [Agrostis semiverticillata (Forsk.) C. Christens.]. Water nt; herbaceous oe aia 9331, 9356; Schizachyrium cirratum (Hack.) Wooton & aes reo cirratus Hack.]. Texas bluestem; herbaceous oo SR: 9668; decal aca peer ae Altson var. ae Mees) | Hatha [Andropogon hirtiflorum (Nees) Kunth]. Herbaceous peren- nial. SC: 8545, 9681, 9710A; occasional, can- yons, shaded slopes. SR: 8785; occasional, nor Setaria grisebachii E. Fourn. Grisebach bristlegrass; summer annual. SC: 8538; occa- BRIT.ORG/SIDA 22(1) sional, north- — SR: 8792, 9620A; oc nal, Parker C Setaria irucouie ccibn He og K. Schum. [S. macrostachya Kunth sensu Kearney and Peebles (1960), in part]. sop bristle grass; herbaceous a ape SC:8441,8528B; occa- e grasslan ee ee (L.) Pers. eee grass;her- baceous perennial. SC: 9652; occasional, mostly along Sonoita Creek. SR: 9561; locally gene roadsides, springs, Santa Cruz River floodplain Pre abuse (Michx.) Scribn. Prairie wedge grass; herbaceous perennial. SR: 9460; uncommon spr in Sporobolus contractus aeiche Spike dropseed; herbaceous perennial. SC: 9635; occasional, along Sonoita Creek. SR: 9624; rare, Parker Canyon. Sporobolus cryptandrus (Torr.) A. Gray. Sand dropseed; herbaceous perennial. SR: 9592, 9611B,9695;uncommon, grasslands, washes. Sporobolus wrightii Munro ex Scribn. Sacaton. Herbaceous perennial. SC: rare, canyons. SR: 2,abundant on Santa Cruz River flood- plain. Trachypogon spicatus (L.) Kuntze [T. secundus (J. Presl) Scribn.]. Srinkis: awn; dali S pe- rennial.SC:8536,87 arte fac- ing slopes. SR: 9608, 9664; common, grass- lands. Urochloa arizonica (Scribn. & Merr.) O. Morrone & F. Zuloaga [Panicum arizonicum Scribn. & Merr]. Summer annual. SC: oe can- S. 4, occasional, low Vulpia aibiosaeiue (Nutt.) Gai var. ee (Scribn. ex Beal) Lonard & Gould [Festuca eae Piper]. Winter annual. SC: 8870; oc- asional, rocky slope Vulpia pene a (Walt.) a var. hirtella (Piper) He Aa octoflora Walt. var. hirtella pict Six-w ue; winter annual. SC: 8923: ene rocky slopes. SR: 9095; oc- casional, grasslands. Vulpia octoflora (Walt.) Rydb. var. octoflora. Six- weeks fescue; winter annual. SC: 908]: com- mon, grasslands, sandy soils along Sonoita Creek. Pontederiaceae Heteranthera limosa (Sw.) Willd. Mud plantain; MCLAUGHLIN, FLORA OF emergent aquatic perennial. SR: 6489: last collected from pond adjacent to Santa Cruz River in 1991. Potamogetonaceae Potamogeton foliosus Raf. Pondweed; sub- merged aquatic perennial. SR: 9462, 9465; AREA AND SAN RAFAEL STATE PARK 703 tion seen along Sonoita Creek. SR: 9574: uncommon, Santa Cruz River bottom. Zannichelliaceae Zannichellia palustris L. Horned pondweed,com- mon poolmat; submerged herbaceous aquatic. SC: 9332; rare, found once in pool uncommon, Sharp and Heron springs. along Sonoita Creek. SR: 9245: common, Typhaceae springs, Santa Cruz River. Typha domingensis Pers. Cattail; emergent peren- nial aquatic. SC: rare, a single small popula- ACKNOWLEDGMENTS I want to extend particular thanks to Betsy Lewis for her capable assistance with the field work, often under very difficult conditions. Matt Chew, formerly with Arizona State Parks, helped in initiating this study. Rick Gagnon, Lee Eseman, Carol Bercich, Dave Pawlik, and Steven Haas of State Parks provided extensive logistic support. Access to Sonoita Creek State Natural Area through private property was granted by Dave Blouin, Morningstar Ranch, and by Javier Badillo. Several individuals helped with the field work at various times; | par- ticularly want to acknowledge the assistance of George Montgomery, Rick Gagnon, Janice Bowers, Anne Thorne, Jim Jordan, Mills Tandy, Bob Zahner, and Glenda Zahner. Erika Geiger drafted the maps of the study areas. Richard S. Felger and Dale S. Turner provided very thorough and thoughtful reviews. REFERENCES Benson, L. 1969. The cacti of Arizona. 3rd ed. University of Arizona Press, Tucson. Bock, J.H.and C.E. Bock. 2000. The view from Bald Hill: Thirty years in an Arizona grassland. University of California Press, Berkeley. Brown, D.E. 1994. Biotic communities: southwestern United States and northwestern Mexico. University of Utah Press, Salt Lake City. Burcess, T.L. 1995. Desert grassland, mixed shrub savanna, shrub steppe, or semidesert scrub? The dilemma of coexisting growth forms. |n:M.P. McClaran and T.R.Van Devender, eds. The desert grassland. The University of Arizona Press, Tucson, Arizona. Pp. 31-67. Haotey, D.and T.E. SHerioan. 1995. Land use history of the San Rafael Valley, Arizona (1540- 1960). USDA Forest Service General Technical Report RM-GTR-269. Fort Collins, Colorado. Kartesz, J.T.1999.A synonymized checklist and atlas with biological attributes for the vas- cular flora of the United States, Canada, and Greenland. 1st ed. In:J.J. Kartesz and C.A. Meacham. Synthesis of the North American Flora, Version 1.0.North Carolina Botanical Garden, Chapel Hill. Kearney, TH. and R.H. Peestes. 1960. Arizona flora. 2nd ed. The University of California Press, Berkeley. 704 BRIT.ORG/SIDA 22(1) McLAUGHLIN, S.P. 1989. Natural floristic areas of the western United States. J. Biogeogr. 16: McLAauGHUuwN, S.P. 1992. Are floristic areas hierarchically arranged? J. Biogeogr. 19:21-32. Myrick, D.F. 1975. Railroads of Arizona. Vol. 1. The southern roads. Howell-North Books, Berkeley. Parritt, B.D. and A.C. Gisson. 2003. Cactaceae Jussieu. In: Flora of North America Editorial Committee, eds., Flora of North America. Volume 4. Magnoliophyta: Caryophyllidae, part 1.Oxford University Press, New York. Pp. 92-257. RicHarDsON, M.L., S.D. Clemmons, and J.C. Wacker. 1979. Soil survey of Santa Cruz and parts of Cochise and Pima counties, Arizona. USDA Soil Conservation Service and Forest Ser- vice, Washington, DC. Rotuins, R.C. 1993. The Cruciferae of Continental North America: systematics of the mus- tard family from the arctic to Panama. Stanford University Press, Stanford, California. Setters, W.D., RH. Hitt, and M. Sanperson-Rae. 1985. Arizona climate: the first hundred years. University of Arizona, Tucson. Wuite, S.S. 1948. The vegetation and flora of the Rio de Bavispe in northeastern Sonora, Mexico. Lloydia 11:229-302. ANNOTATED CHECKLIST OF THE VASCULAR FLORA OF THE BIG SANDY CREEK UNIT, BIG THICKET NATIONAL PRESERVE, TEXAS Barbara R. MacRoberts and Larry E.Brown Michael H. MacRoberts Houston Community College Bog Research 1300 Holman mbia Houston, Texas 77004, U.S.A. Shreveport, Louisiana 71104, U.S.A. and an Herbarium, Spring Branch Science Center Herbarium, Museum of Life Sciences 8856 Westview Drive Louisiana State University in Shreveport Houston, Texas 77055, U.S.A. Shreveport, Louisiana 71115, U.S.A. Paul A.Harcombe, Warren W. oe |. Sandra Elsik Stanley D. Jones Department of Ecology volutionary Biology Herbarium, Botanical ae Center Rice Universi’ PO. Box 6 Houston, Texas 77005, U.S.A. Bryan, Texas sr U.S.A. ABSTRACT An annotated, vouchered checklist is provided of the vascular plant taxa of the 5,806 hectare Big Sandy Creek Unit, Big Thicket National Preserve, Polk County, southeastern Texas. Documented for the unit are 693 total taxa with 45 of them exotic RESUMEN AAs a poe | ] Bead Pane oe Oa Ven | Se ofrece un listado con anotaciones, y de Big Sandy Creek Unit, Big Thicket National oe Polk ore Sureste de Texas. Se documentan para la unidad un total de 693 de los que 45 INTRODUCTION This is the third installment of an inventory, based upon available voucher speci- mens, of the vascular flora of the Big Thicket National Preserve, southeastern Texas. In the two previous papers we provided an annotated checklist of the Hickory Creek and the Turkey Creek units (MacRoberts et al. 2002b; Brown et al. 2005.). In this paper we do the same for the Big Sandy Creek Unit. SITE AND METHODS The Big Thicket is located mostly within the longleaf pine region of the West Gulf Coastal Plain in southeastern Texas (Parks & Cory 1936; McLeod 1971; Harcombe & Marks 1979; Marks & Harcombe 1981; Harcombe et al. 1993). The Big Thicket National Preserve consists of 12 units scattered over seven Texas SIDA 22(1): 705 — 723. 2006 706 BRIT.ORG/SIDA 22(1) counties. The units range from 222 to 10,100 hectares and total about 34,000 hectares (Peacock 1994). The Big Sandy Creek Unit consists of 5,806 hectares and is located in eastern Polk County (Fig. 1). It is about 14.5 km from north to south and is about 9.7 km at its widest east-west point. In elevation it ranges from about 53 to 94 meters. The Big Sandy Creek Unit was chosen for inclusion in the Big Thicket National Preserve because it is diverse, has a wide range of plant communities, is a stream corridor unit, and has “hilly” topography ab- sent from most of the other units (Ajilvsgi 1979; Watson 1979; Harcombe & Marks 1981). It is dominated by upper slope pine oak forest, mid slope oak pine forest, and lower slope hardwood pine forest. Also present are upland pine for- est, floodplain hardwood forest, floodplain hardwood pine forest, baygall, cy- press-tupelo swamp, and sandhill pine forest (Harcombe & Marks 1979). It has one pitcher plant bog that is badly degraded , no wetland pine savanna, no prai- ries, and no xeric sandylands although some of the upper slope pine oak forest and sandhill pine forest approach xeric sandylands conditions. A map show- ing the distribution of plant community types is in Harcombe and Marks (1979). Fire suppression over the past 30 years has resulted in extensive midstory de- velopment and shading out of herbaceous vegetation. Details regarding soils, topography, and climate can be found in Deshotels (1978), Watson (1979), Harcombe and Marks (1979), and Marks and Harcombe (1981). Like all units of the Big Thicket National Preserve, the Big Sandy Creek Unit had not previously been the subject of a detailed floristic inventory al- though it is one of the best documented units (Watson 1982). The MacRoberts collected 446 specimens, representing 348 taxa, during Ll field days beginning 3 April 2003 and ending 2 May 2004. Larry Brown col- lected 644 specimens , representing 435 taxa, during 12 field days beginning 4 April 2003 and ending 27 September 2005. Stanley Jones collected 252 speci- mens in 2003. Geraldine Watson collected 255 specimens over several years in the late 1970s and early 1980s. Other collectors gathered about 84 specimens. We searched the Texas A & M University (TAMU) herbarium consortium data- base for additional specimens but no new taxa were found there. Since our purpose is to produce a list of documented taxa known to occur on the Big Sandy Creek Unit, a vouchered specimen was considered to be the only evidence acceptable for inclusion in the list. However, two species, Phlox nivalis subsp. texensis a federally listed species, and Utricularia inflata were photographically documented and not collected. In all, about 1,650 herbarium specimens form the data for this report. All specimens have been annotated by Larry Brown including most of Bill Carr’s 21 collections at TEX/LL. All 6 of his single taxa records have been examined and annotated. In most cases, nomenclature follows Jones et al. (1997) and Kartesz and Meacham (1999). Some recent literature sources were followed, especially some BROWN ET AL., FLORA OF THE BIG SANDY BIG THICKET NATIONAL PRESERVE ALABAMA - COUSHATTA .. wooovi? ~ cai INDIAN RESERVATION UPPER NECHES RIVER { BEECH CORRIDOR UNIT CREEK UN HILLISTER(D WARREN (p Big Sandy | TURKEY Creek Unité! MENARD CREEK (CORRIDOR UNIT NECHES BOTTOM & JACK GORE BAYGALL UNIT \ ‘N KOUNTZE aprove = ane! a i ry oO LOWER NECHES RIVER ORRIDOR UNIT 2 oO F < co z == a ROSIER UNIT “ I \Y LITTLE pee ISLAND RIOOR UNIT ORANGE CO ‘| BEAUMONT UNIT Tain LAKE D: BEAUMONT — —_ \ JEFFERSDN CC | ¢ cs Fic. 1. Location of Big Sandy Creek Unit, Big Thicket National Preserve, Texas. nomenclature in the Flora of North America volumes. In cases of multiple col- lections, no more than four are listed for each taxon. RESULTS Table 1 is an annotated list of the Big Sandy Creek Unit taxa. The following ab- breviations are used in Table 1 CL=C. Liu. His specimen is at Rice University. CP = Charles Peterson. His two 2 aden are at SBSC. DR= David Rose 1? snecimens ar Be Hers GW = Geraldine yeeon Her specimens are at Rice University. 708 BRIT.ORG/SIDA 22 = 1) LB = Larry Brown. His specimens are currently at Rice University except for those few that are indicated to be at SBSC. MM = Barbara and Michael MacRoberts. Their specimens are currently at Rice niversity except for one at SBSC. PR = Paul Rolling. His collections are a clear picture of Utricularia inflata from the Beaverslide Trail. There is no doubt as to its identification. One speci- men is at SBSC and a duplicate is currently at Rice University SEWP = Sandra Elsik and Warren Pruess. Their specimens are currently at Rice University. SJ = Stanley Jones. His specimens are oo at Rice University. WEE Bill Carr. His 21 specimens are at TEX/LI = exotic =see notes at end of list Those specimens listed as currently at Rice University are mounted and will be deposited at a yet to be selected major Texas herbarium. Except for the Stan- ley Jones specimens, they were collected under National Park Service Coopera- tive Agreement No. CA 14001004. ACANTHACEAE ANNONACEAE Hygrophila lacustris (Schlecht. & Cham.) Nees, SJ Asimina parviflora (Michx.) Dunal, SJ 16093; LB 90;MM6929; LB 28616; GW 2582 28146, 28188 Justicia ovata (Walter) Lindau var. lanceolata APIACEAE (Chapm,) R.W. Long, 5J 16240; LB 28170 Centella erecta (Lf) Fernald, S) 17408; LB 28629: Ruellia humilis Nutt., SJ 16340; LB 28760, 28219; MM 6256 ae = Chaerophyllum tainturieri Hook. var dasycarpum ACERACEAE S.Watson, MM 6324 Acer barbatum Michx., SJ 16242; LB 27804; GW — Chaerophyllum tainturieri Hook.var. tainturieri, LB 2446 27843 Acer rubrum L., SJ 16099; MM 6091; LB 27804, — Cicuta maculata L., GW 3502 28147 *Cyclospermum leptophyllum (Pers.) Sprague ex AGAVACEAE Britton & Wilson, L Manfreda virginica (L,) Salisb. ex Rose, LB 28995, “/70sciadium beach DC, 5) 16429;GW : 04] Eryngium prostratum Nutt.ex DC. LB 2822 28775 : Yucca louisianensis Trel., LB 27791;5J 16429 Hydrocotyle verticitata Thunb., SJ 16236 LB 28657, 28243; GW 2606 ALISMATACEAE Ptilimnium capillaceum (Michx.) Raf., SJ 16362; paecadie coat ifolius (L ey LB 28622 GW 2596 Sagittaria la Willd., GW 2802 Sanicula canadensis L., MM 6275,6571;LB 28144 AMARANTHACEAE GW 248. Froelichia floridana (Nutt.) Moq., MM 6709 Sanicula smallii Bickn., SJ 1618 Spermolepis divaricata ie Raf. ex Ser., SJ ANACARDIACEAE | 16381: MM 6616 us aromatica Aiton var. aromatica, LB 28669; lepis echinata (DC. Heller. MM 61444 730 Rhus copallinum L., SJ 16392; MM 6081 sive up ae L., LB 28683 tron radicans (L.) Kuntze, LB 27802 Toxicodendron vernix (L.) Kuntze SJ 16188 Thaspium trifoliatum (L.) A.Gray var.aureum Brit- ton, GW 2470 Trepocarpus aethusae Nutt. ex DC., SJ 16254; LB BROWN ET AL., FLORA OF APOCYNACEAE Trachelospermum difforme (Walter) A. Gray, LB 28790 AQUIFOLIACEAE llex ambigua (Michx.) Torr., SJ 16233; LB 30452 (SBSC) llex coriacea (Pursh) Chapm., LB 28955, 28148, 28195; MM 6257 llex decidua Walter, GW 2460, oe 2694, 2881 llex longipes Chapm. ex Trel., SJ 16173; GW 2466, 2882, LB 30460 (SBSC) flex opaca Soland., LB 28595, 28189, 28734; MM 6100 llex vomitoria Aiton, SJ 16095;:MM 6121 ARACEAE Arisaema dracontium (L.) Schott, MM 6346; LB 28346 ee um (L oe LB 28242, 28621; MM 6258, GW 253 16294; GW Peltandra Wane. (L ; Schott, SJ 1 2464a, 2462a, 2641 ARALIACEAE Aralia spinosa L.,SJ 16097; LB 28786; MM 6341 *Hedera helix L., LB 28143 ARECACEAE Sabal minor Jacq.) Pers., SJ 16079; MM 6073;LB ARISTOLOCHIACEAE Aristolochia reticulata Jacq., MM 6315;LB 28240 Aristolochia serpentaria L., LB 28972; MM 6276, 6549; GW 2017 Aristolochia COMME LOISG Sims, GW 251 ] ASCLEPIADACEAE Asclepias ee Michx., SJ 16369; LB 28374; MM 6699 Asclepias perennis Walter, LB 30535 (SBSC) Asclepias tuberosa Woodson subsp. interior mee! cea variegata L., MM 6280; LB 28364 pias verticillata L,LB 28779; GW 2016 he. viridis Walter, MM 6310 Matelea Ce ides (Engelm.) Woodson, MM 656 Moteea gaa! ipiens et are SJ 16182;LB MM Maree Roneenos (Walter) cone SJ 16415 LB 28332,GW 2019 PRESERVE 709 ASPLENIACEAE aos ium platyneuron (L.) Britton, a & Poggenb., LB 28619, 28382; MM 6 ASTERACEAE isiifolia L., LB 28672, 28719 Ambrosia psilostachya DC., MM 6886; LB 28996 Baccharis halimifolia L., LB 28950; GW 2735 Berlandiera pumila (Michx.) Nutt. var. pumila, MM 6285, LB 27801 Ber! eo pumila (Michx.) Nutt. var. scabrella esom & B.L. Turner, LB 28431a; MM 6605 oe bipinnata L., MM 6933:LB 28723 Pie discoidea (Torr.& A.Gray) Britton, LB i 5 ens frondosa L., MM 6934 8,27: ce tonia diffusa Elliott, MM 6907, 6727 Chrysopsis mariana (L.) Elliott, LB 28943 Chrysopsis pilosa Nutt., LB 28674, 28778; MM 6614, 6611 Cirsium horridulum Michx., MM 6152 Conoclinium coelestinum (L.) DC., MM 6891: LB 28937 Conyza canadensis (L.) Cronquist var. canadensis, LB 28761 Conyza canadensis (L.) Cronquist var. pusilla (Nutt.) Cronquist, LB 28415; MM 6708; GW 2770 Coreopsis lanceolata L., MM se Goleop sat Hnetona Nutt., MM 6 Cr (Nutt) sr 16 29001; MM 6704; WRC 10827 +Croptilon hookerianum (Torr. & A. Gray) House, GW 2688 +Diaperia candida (Torr.& A.Gray) Bentham, MM 7049 Echinacea pallida (Nutt.) Nutt. var. sanguinea (Nutt.) pe i R.D.Thomas, SJ 16402; LB 28223a; MM 6 Elephantopus oy, nianus Raeusch., MM 6946: WRC 10831 Elephantopus nudatus A. Gray, MM 6896 Flephantopus tomentosus L., LB 28598a, 28353; Erechtites hieraciifolia (L.) Raf.ex — LB 28960 Erigeron annuus (L.) Pers., GW 262 ee strigosus ee ex Willd., - 28153, MM 084, 6107; GW 2 at apillifoli om all) Lam., LB 28935 Epaeaim lancifolium (Torr.& A.Gray) Small, LB 28756 = Fupatorium leucolepis (DC.) Torr. & A. Gray, MM 6730, 6594 Eupatorium perfoliatum L., LB 28613; MM 6344 Fupatorium rotundifolium L.,LB 28427;MM 6729 Eupatorium serotinum Michx., LB 28742, 28228 ae ree (Torr. & A. ae Greene ex er & Britton, LB 28962, Facelis retusa (Lam.) Schultz-Bip., . er 2490 Gaillardia aestivalis (Walter) H.Rock var.aestivalis, LB 28680 Gaillardia pulchella Foug. var. pulchella, MM 6236 Gamochaeta coarctata (Willd.) Kerguelen, LB 3045 1a (SBSC) Gamochaeta pensylvanica (Willd.) Cabrera, GW 2585 a Gamochaeta purpurea (L : - oe MM 6102;LB 27786, 30537; WRC 1 Helenium amarum ie - a var. amarum, MM 6545 Helianthus angustif L.,LB 28947 Helianthus hirsutus Raf.,LB 28402;MM 6750,6524 Helianthus mollis Lam., LB 28676, 20400; MM Helianthus simulans E.E.Wats., LB 30877 ber ee eet (Lam.) Britton & Rusby, LB 28932, 2 ae 0828 — L., LB 28594; MM 6565 Mi cnonenas artemisiifolius DC. var. artemisiifolius, SJ 16260;LB 28122; MM 6085 *Hypochaeris microcephala (Sch. oe var, albiflora (Kuntze) Cabrera, MM 6 lva annua L., GW 2052 Krigia cespitosa (Raf.) Chambers, GW 2493 Krigia virginica (L.) Willd, MM 6111;LB 27782 Lactuca canadensis L.,LB 28409, 28378; MM 6518, 6559 Liatris acidota Engelm.& A. Gray, MM 6917 Liatris Se he (Walter) Michx. var. elegans, MM 747, LB 28984, 28762 Liatris squarrosa (L.) Michx., var. hirsuta (Rydb.) r, LB 28990, 28684 bon ise (L.) — var. squarrosa, MM 6746; GW 2092, Wien ia ee. (L.) ci . 27807 Oligoneuron nitidum yale : a en Small, LB 28985, 28396; MM 669 Packera glabella (Poir.) C. ee SJ 16237; MM 6342; GW 2526 BRIT.ORG/SIDA 22(1) Packera obovata (Muhl. ex Willd.) W. A.Weber & A.Love, LB 27805 Pityopsis graminifolia (Michx.) Nutt., MM 6898, 6080; LB 28772 Pluchea camphorata (L.) DC.,LB 28934; GW 2785 Pluchea foetida (L.) DC., LB 28710 Pseudognaphalium obtusifolium (L.) Hillard & Burtt, LB 28934, 28673; MM 6716 Pyrrhopappus carolinianus (Walter) DC., MM 6313; SJ 15348 Ratibida columnifera (Nutt.) Woot. & Standl., MM 6543 Rudbeckia hirta L., MM 6278, LB 28200; $/16353 Smallanthus uvedalius (L.) Mack. ex Small, LB 28/29a Solidago canadensis L., LB 28968; GW 2885 Solidago ludoviciana (A. Gray) Small, LB 28749, 28334, 27801a; MM 6294 Solidago odora Aiton var. eee - 28942, 28660 patie petiolaris Aiton., LB olidago rugosa Mill. nee asper (Aiton) Cronquist, LB 28946,22873 Solidago rugosa Mill. subsp. rugosa, LB 28752,GW 2424, 2423A;CL 901179 *Soliva sessilis Ruiz & Pavon., GW 2476 *Sonchus asper (L.) Hill, LB 29012 *Sonchus oleraceus L., LB 2811 Symphyotrichum dumosum (L.) Nesom, LB Roos, 6a Symphyotrichum patens (Aiton) Nesom, LB 28981, 27776 Symphyotrichum pratense (Raf. 6909 <-> Nesom, MM Verbesina virginica L., LB 28780 Vernonia missurica Raf., LB 30766 (SBSC) Vernonia texana (A. Gray) Small, MM 6725,6527; 28731; GW 2731 *Youngia japonica (L.) DC., LB 27839 BERBERIDACEAE Podophyllum peltatum L., MM 6151 BETULACEAE Alnus serrulata (Aiton) Willd., SJ 16144;LB 28172 Betula nigra L., SJ 16297; MM 6328; LB 28249a Carpinus caroliniana Walter, MM 6538, 6148 Ostrya virginiana (Mill.) K. Koch, SJ 16072, 16440; LB 28131;MM 6525 ae ag ecnuanaine aL. LB 27834a BROWN ET AL., FLORA OF TH PRESERVE 711 EC DIN SANUT meat Can ai ex Bureau, SJ 16422 rd t) Warder ex Engelm., SJ 16321; LB 27775 BLECHNACEAE Wanaivard lata (L.) T. Moore, MM 6338; LB 27846 BORAGINACEAE y! oglossum virginianum fin, G Lithosperma tuberosum Rugel ex a 2213 Myosotis macrosperma Engelm., GW 2491 BROMELIACEAE (L.) L, MM 6936, 6330, 6755 BUDDLEJACEAE Polypremum — L.,. MM 6566 BURMANNIACEA AO SUED Nut LB 28782;CP 1041:DR 2373 ta J.F.Gmel.) Mart., MM 6863 CABOMBA Brasenia ae J.F.Gmel.,SJ 16291 CACTACEAE Opuntia humifusa (Raf) Raf, SJ 16383; MM 6603 CALLITRICHACEAE Callitriche heterophylla Pursh, SJ 16147;GW 2527 CAMPANULACEAE Lobelia appendiculata A. - 6303;GW 2562 Lobelia cardinalis L.,LB 2 sere puberula Michx., ree 6878; GW 2822;LB 28958, 28606 Triodanis perfoliata (L.) Nieuwl., GW 2475 *Wahlenbergia marginata (Thunb.) A. DC., SJ bs tee Sa NAKA | Z CAPRIFOLIACEAE */ onicera japonica Thunb., MM 6263;LB 28156 Lonicera sempervirens L., LB 27815; GW 2469 Sambucus nigra L. subsp. canadensis (L.) R. Bolli, LB 28641 Viburnum acerifolium L., SJ 16322 Viburnum dentatum L., SJ 16396: MM 6074; LB28593; GW2465 Viburnum nudum L. var. nudum, SJ 16286; LB 28198 Viburnum rufidulum Raf., MM 6093: GW 2040, 3 CARYOPHYLLACEAE *+Cerastium pumilum Curtis, MM 6143 (SBSC) 072 Gee Loeflingia squarrosa Nutt., MM 7 +Minuartia muscorum (Fassett) Rabeler, SJ 16105 +Silene subciliata B.L. Robins., LB 28599, 28626 CELASTRACEAE Euonymus americanus L., MM 6071;LB 28163 CHENOPODIACEAE Dysphania ambrosioides (L.) Mosyakin & Clem- ents, LB 28646 CISTACEAE Helianthemum carolinianum (Walter) Michx, LB 28235a, 27797; MM 6110 Lechea mucronata Raf., LB 27783 Lechea tenuifolia Michx.,SJ 16360;LB 28229: MM 6609 COMMELINACEAE ey ina say L.. MM 6601,LB 28385, 28169; SJ] one ei ee Le LB 29035; MM 6940 Tradescantia reverchonii Bush, MM 6267 CONVOLVULACEAE ae os is Michx., MM 6125; LB oe ae ie Slee LB 28333 ipomoea lacunosa L., L 735 — pandurata (L.) G. FW. Mey., LB 28429; MM 2753, 6617,6570 Stylisma pickeringli A. Gray var. pattersonii (Fernald & B.G. Schub.) Myint, MM 6611 CORNACEAE Cornus florida L., LB 28154; MM 6105; SJ 16394 Nyssa aquatica L., SJ 16205, 16256; LB 28134, 30455 (SBSC Nyssa biflora Walter, SJ 16317;LB 28322 Nyssa sylvatica Marshall., MM 6429; LB 28759, 30540 (SBSC); SJ 16087 CUPRESSACEAE Juniperus virginiana L,MM 6114;LB 28598, 28356; SJ 16410 Taxodium distichum (L.) L. C. Rich., SJ 16209, 16339 CUSCUTACEAE Cuscuta compacta Juss. ex Choisy, LB 30765 (SBSC) CYPERACEAE Bulbostylis ee (Elliott) Fernald var ciliatifolia Bulbostylis Vee (Elliott) Fernald var coarctata (Elliott) Kral , SJ 16425 712 Carex abscondita Mack., SJ 16119; LB 30456 (SBSC) Carex albicans Willd. ex Spreng. subsp. australis eal) JH. Rettig, LB 27835 Carex chwein. 0566 Carex atlantica Bailey subsp: capillacea (Bailey) Reznicek, MM 6244: LB 7 i Carex basiantha Steud., SJ 16 Carex b/ ae ‘Dewey, SJ 161 . Carex Schwein, SJ 16115;LB 27820 Carex ee mep nota Muhl. ex ae SJ 16121 Car Schwein., SJ 16123 Carex complanata Torr. & Hook., a 6279, 6529; C 17335 Carex corrugata Fernald, SJ 16124; LB 27840 Carex crebriflora Wieg., SJ 16041;LB 27836 Carex debilis Michx.var. debilis, SJ 16117;LB 28181 Carex digitalis Willd. var. asymetrica Fernald, SJ Carex festucacea schkuhr ex ane SJ 16044 Carex floridana Schw Carex frankii Kunth, a ay Carex glaucescens Elliott, LB a 28266 Carex intumescens Rudge, LB 27150 Carex joorii Bailey, MM 6697; LB 28959a Carex leavenworthii Dewey, SJ 16103; Carex Seen Wahlenb. subsp. h i(F Stone, SJ 161 es = 2826/, 28201 Carex one Mack., SJ Carex louisianica le 17352;S) 16212;LB 28610, 2815 Carex lupulina “en ex Willd.,SJ 17252;LB 28272, 28209, 30563 (SB oe Carex lurida Wahlenb., LB Carex muehlenbergii ee ex Willd. var. enervis Boott, SJ 16043 Carex oxylepis Torr.& Hook., SJ 16153; MM 6337 Carex planispicata R. F.C. Naczi.SJ 16217, 16122 Carex reniformis Small, SJ 16154 5262 Carex retroflexa Muhl. ex Willd. SJ 16284 iatula Michx., LB 30538 (SBSC) Face texensis Bailey, SJ 16104; MM 6130 Carex triangularis Boeck., SJ 16118 Carex tribuloides Wahlenb. var. a aa Clokey, SJ 16228, 16270;LB 2824 Cyperus croceus Vahl, MM 6555;LB — 28387 Cyperus echinatus Britton , SJ 16400 Cyperus grayioides Mohlenbr. WRC 17705b Cyperus haspan L., LB 29015, 28729 IAW ITlalQ) BRIT.ORG/SIDA 22(1) Cyperus hystricinus Fernald, MM 6756; DR 2364 Cyperus plukenetii Fernald, LB 28671; MM 6720, 6556; DR 2363 Cyperus pseudovegetus Steud., SJ 16365; MM 6698, 660 Cyperus retroflexus Buckley, LB 28666; MM 6554; DR 2371 Cyperus retrorsus Chapm., LB 28486, 28358 Cyperus strigosus L., LB 28386, 28358 Cyperus virens Michx., SJ 16337;LB 28155 Fleocharis montevidensis Kunth, SJ 16349 Eleocharis obtusa (Willd.) aoe 16424 Eleocharis tortilis Schult., S Eleocharis tuberculosa a ens & JA. ultes, MM 6245,6578;LB 28323; =: 16363 sie squarrosa Michx., MM 6596, 662 ee odorata ve LB 28604; MM ae ica Elliott, SJ 16357 sel eae corniculata (Lam.) A. Gray, MM 6939; LB 28617 Rhynchospora globularis (Chapm.) Small, var. globularis, SJ 16382;LB 28677;MM 6530,6599 Rhynchospora glomerata (L.) Vahl, MM 6735; LB 28430, 28418,28624 Rhynchospora gracilenta A. Gray, MM 6887; LB 28413; DR 2375 Rhynchospora harveyi W. Boott, LB 28220; SJ 16397 Rhynchospora inexpansa (Michx.) Vahl, LB 28432, 28324 Rhynchospora mixta Britton, LB 28961, 28270, 28340, 28248 Rhynchospora pusilla Chapm. ex M.A. Curt. MM 6574, 6576 Rhynchospora rariflora Elliott, MM 6627;LB 28412 5J 16 Rhynchospora scirpoides (Torr.) A. Gray, DR 2376 Scirpus cyperinus L. Kunth, LB 28745, 27848; MM 6540 Scleria ciliata Michx. var. ciliata, SJ 16201, MM 6292: LB 28237 Scleria oligantha Michx., SJ 16280, 16126; MM 16126; 6325 Scleria reticularis Michx., MM 6885 Scleria triglomerata Michx., SJ 16423, 16224; LB 28 28177 CYRILLACEA Cyrilla oe L., LB 28179, 28241;MM 6591; SJ 16368 BROWN ET AI PRESERVE 713 DENNSTAEDTIACEAE Pteridium aquilinum (L.) Kuhn, MM 6139; LB DIOSCOREACEAE Dioscorea villosa L., LB 27833; SJ 16389 DROSERACEAE Drosera brevifolia Pursh, MM 6094 Drosera capillaris Poir., MM 6883,6575;LB 28416 DRYOPTERIDACEAE Athyrium filix-femina (L.) Roth var. eae (Michx.) Farw. SJ 16275; MM 624 Dryopteris ludoviciana (Kuntze) at GW 2600 Onoclea sensibilis L., MM 6345; LB 28246, 27842 arena Bees rode (Michx.) Schott, SJ 16232; LB 28274, 27808 EBENACEAE Diospyros virginiana LMM 6281;LB 28966, 28159 ERICACEAE Rhododendron canescens (Michx.) Sweet, SJ 16329 Rhododendron viscosum (L.) Torr., SJ 16289; LB 2895, 28279,27850 Vaccinium arboreum Marshall, SJ 16214; LB 28664, 28145; MM 6291 Vaccinium corymbosum L., LB 28428; 28129; MM 6137; GW 201 Vaccinium stamineum L., SJ 16066; MM 6320 ERIOCAULACEAE Er iocaulon te Croke Koern., MM 6592: SJ 16372 EUPHORBIACEAE Acalypha oe A. Gray, MM 6872, 6757; LB 28953; GW 2768 Acalyph cane idea Raf., GW 2603 Chamaesyce cordifolia (Elliott) Small, LB 28675, 28768; MM 6618 Cnidoscolus texanus (Muell.-Arg.) Small, MM 6269; SJ 16 Croton argyranthemus Michx., MM 6309; LB 28224; GW 2576 Croton Michx., MM 6718 Croton g Danahiostis L. var. septentrionalis Muell.- Arg. “MM 6695, GW 2602; LB 28722, 28370 Croton michauxii G. L. Webster, GW 2687; MM 6619 Euphorbia corollata L., LB 27770; MM 6544; GW 2045 pede ou jensis Walter, LB 28607; GW pre ra urinaria L.,LB 27927; MM 6945 Sebastiania fruticosa (W. pee Fernald, SJ 16152; MM 6327; LB 28187, 27830 Stillingia sylvatica L., SJ 16367; a 6711 Tragia smallii Shinners, GW 2577 Tragia urticifolia Michx., MM 6306,6714,6548;LB 28379 *Triadica sebifera (L.) Small, SJ 16446 FABACEAE *Albizia julibrissin Durazz., SJ) 16445; LB 29006, 27771; MM 6524 Baptisia bracteata Muhl. ex Elliott var laevicaulis (A. Gray ex Canby) D. Isley, SJ 16106 Baptisia bracteata Muhl.ex Elliott var.leucophaea (Nutt.) Kartesz & Gandhi, MM 6132 pe nuttalliana Small, MM 6316, 6133 entrosema virginianum (L.) Benth., LB 28381 a is canadensis L.var. canadensis, MM 6150;LB 28681; SJ 28381 Chamaecrista fasciculata (Michx.) Greene, MM 751; LB 28681 Chamaecrista nictitans (L.) Moench, LB 28711 28658 Clitoria mariana L.,LB 28590, 28757, 28339: MM 6553 Crotalaria sagittalis L.,SJ 16220; MM 6693, 6551; B 28327 Dalea phleoides (Torr. & A. Gray) Shinners, MM 6748 Desmodium ciliare a ex Willd.) DC.,LB 29032, 28376; GW 2714 Desmodium ie Hi ens DC., LB 28734, GW 2773 ae\ ae m yk as LB 29030 DC,,LB 2 bt, OUlUSUT Ti Demoaon paniculatum DC,, a 0;GW 2422 Desmodium rotundifolium DC., LB 28668; GW 2803 Erythrina herbacea L., SJ 16322; MM 6288; LB 28223 Galactia volubilis (L.) Britton, LB 28769; MM 6526, 6552; GW 2578 Gleditsia aquatica Marshall, LB pe Indigofera suffruticosa Mill. GW *Kummerowia striata Mill., LB oars 28925 *Lathyrus hirsutus L.,GW 2011 Lespedeza hirta Elliott, MM 6925;LB 29000 Lespedeza procumbens Michx., LB 28397, 28380, 440 Lespedeza repens (L.) Barton, LB 28758, 28231a; Lespedeza stuevei Nutt., LB 28767, 29019; GW 2/94 Lespedeza virginica (L.) Britton, LB 28751, 28232; G 6 Mimosa hystricina (Small ex Britton & Rose) B.L. Turner, SJ 16258;LB 28216 Rhynchosia difformis (Elliott) DC. LB 28750, 28667,2 GW 2413 Rhynchosia latifolia Nutt. ex. Torr. & A. Gray, GW 2565, 2/33 Rhynchosia reniformis DC.,LB 28406, 28235; MM 6270,6314 pil mari joloes (L.) a LB 28642; GW 2049 (L.) Irwin & Barneby, GW 2715 (L.) eae GW nee +Sophora affinis Torr. & A. Gray, G Strophostyles umbellata (Muhl. ex vie Britton, 8781, 28754 Stylosanthes biflora (L.) Britton, Sterns, Poggenb., FC it tol Tephrosia onobrychoides Nutt., MM 6283; LB eas GW 2463 Tep! iniana (L.) Pers.,SJ 16391;LB 28661, 2821; 7 MM 6301 He ea ium eee Schreb., MM 6146;SJ 16354 Sibthrop, LB 27845b *Trifoli um incarnatum L., SJ 16062; MM 6323 *Trifolium repens L., SJ 1 —_ *Vicia sativa L. MM 614 Wisteria frutescens (L.) ae SJ 16207; LB 28174, 28171, 28175 FAGACEAE Castanea pumila (L.) Mill. SJ 16314 Fagus grandifolia Ehrh., MM 6101; SJ 16279; LB = 27781 Quercus alba L., LB 28125; MM 6602, 6127; SJ 16075 Quercus falcata Michx., MM 6348, 6523; LB 28600; SJ 16304 Quercus hemisphaerica W. Bartram ex Willd., MM 6532; SJ 16110; LB 28777, 28713 ] artram, MM 6274,6568,6567; Quercus laurifolia Michx., LB 27844, 27827a, 30560 (SBSC) BRIT.ORG/SIDA 22(1) Quercus lyrata Walter, LB 28748 ercus margaretta Ashe ex Small, MM 6318 6311 Quercus marilandica Muench., MM 6090; SJ LB 27778, 28665 Quercus michauxii Nutt., LB 28649: MM 6078, + SJ 16084 Quercus nigra L., MM 6077, 6261; SJ 16098; LB 28183 Quercus pagoda Raf., SJ 16169 Quercus phellos L.,LB 28117,27733;MM 6268; S5 16215 Quercus stellata Wangenh., SJ 16108;LB 29005 GENTIANACEAE ne ice, s(L ae CP t.MM ae . 28239 GQUaUG CANIp GERANIACEAE Geranium carolinianum L., SJ 16068 GROSSULARIACEAE Itea virginica L., SJ 16068; LB 29013, 28133; MM 6243 HALORAGACEAE *Myriophyllum aquaticum (Vell.) Verdc. SJ 16292; GW 3345 Id Cri HIG OOS, Proserpinaca palustris L.var.crebra F LB 28741 Proserpinaca palustris L.var.amblyogona Fernald, LB 28158 HAMAMELIDACEAE Hamamelis virginiana L., SJ 16307; LB 28176 Liquidambar styraciflua L., SJ 16073, 16441; MM 6120 HIPPOCASTANACEAE Aesculus pavia L., MM 5234 HYPERICACEAE st drummondii (Grev, & Hook.) Torr. & A. B 28922, 28765; MM 6613; GW 2793 eee galioides Lam., LB 28422; MM 6587 Hypericum gentianoides (L.) Britton, Sterns, Poggenb., MM 6892, 6621; . 28988, 28764 z,MM 6098 ae uit LLB 28417, ae 60; MM 6625 Triadenum walteri (J.G.Gmel.) Gleason, MM 6931, 6953; LB 29016; GW 2821 IRIDACEAE Alophia drummondii (Graham) R. C. Foster, MM 6516; LB 28392, 28234; SJ 16416 H\ peri wim Ay peri oides (L.) Cra PRESERVE 715 BROWN ET AL., PLUAA VE TTT Herbertia lahue (Molina) Goldblatt, MM 6253; SJ 16190 Iris ssid icaulis Raf., SJ 16251 hium anaustifolium Mi IL, G Sahm etiam Bae eat ae 6263; GW 2495 Sisyrinchium sagittiferum E. P. Bicknell, MM 614 WRC 17353 JUGLANDACEAE Carya cordiformis (Wangenh.) K. Koch, SJ 16282 Carya glabra (Mill.) Sweet, LB 28623, 28679,28373; 7 Carya texana Buckley, LB 28391; MM 6271 Carya tomentosa (Lam. ex Poir.) Nutt., SJ 16287; LB 27772; GW 2447, 2448 Juglans nigra L., SJ 16393; LB 28609; MM 6349 JUNCACEAE Juncus acuminatus Michx., LB 30561 (SBSC) Juncus capitatus Weigel, SJ 1604 Juncus coriaceus Mack., MM 6935,6528;LB 28632, 28967 Juncus dichotomus Elliott, SJ 16385; MM 6604 Juncus diffusissimus Buckley, SJ 16296 Juncus effusus L., SJ 16138; MM 6254; LB 28207, Juncus marginatus Rostk, SJ 16346 Juncus repens Michx., MM 6944; LB 27851 Juncus scirpoides Lam., LB 28421; MM 6607 Juncus tenuis Willd., LB 28161; SJ 16405 Juncus trigonocarpus Steud., LB 28161;DR 2366 Juncus validus Coville, SJ 16352, 16406;LB 28328 LAMIACEAE Hedeoma hispida Pursh, MM 6624 Hyptis alata (Raf.) Shinners, MM ak cae rubellus Moench, MM 694 COPUS VIFGINICUS ls GW oe a 2966 Moe fistulosa L.GW 2 Monarda punctata L. MM i 7,6705;LB 28618 GW 2 *Perilla frutescens (L.) Britton, LB 28643 Prunella vulgaris L., MM 6305 Pycnanthemum albescens Torr. & A. Gray, LB 28736, 28755; MM 6308, 6707 Pycnanthemum tenuifolium Schrad., SJ 16432; MM 6703 Salvia azurea Michx. var. grandiflora Benth., MM 6926; LB 29023 Salvia lyrata L., MM 6117;LB 28645, 27809 — Scutellaria ees Engelm. & A. Gray, LB 2 (e lla Ili i; Muhl.e “LB 28365, cee Scutellaria integrifolia L.,LB i Scutellaria parvula Michx., GW ast 17333 hed fenuONG Willd., GW 272 L.,LB — GW 2775 iichosteme setaceum Houtt., MM 6924a; LB 29022 ve SJ 16276; LAURACEAE Persea borbonia (L.) Spreng., MM 6273, 6087; LB 28126, 28390 Persea palustris (Raf) Sarg., LB 27847, 28325; SJ 16397 Sassafras albidum (Nutt.) Nees, LB 28639; MM poate LARIACEAE inguicula pumila Michx., GW uu cornuta Michx., LB ee (SBSC) tricularia gibba L., LB 28717 icularia inflata Walter, PR s.n. icularia juncea Vahl, MM 6585 Utricularia subulata L., GW 2487 LILIACEAE Alli d L.var. canadense, MM 6282;GW 2537 Allium ae L. var. mobilense (Regel) Ownbey, SJ 16197 Hypoxis curtissil ee MM 6343; GW 2522; LB 28175a; SJ 16235 Nothoscordum bivalve (L.) Britton, MM 6095 LINACEAE Linum medium (Planch.) Britton var. texanum (Planch.) Fernald, MM 6533;LB 28424, 28218 2574 Linum striatum Walter, GW 2598, 2719; MM 6519 LOGANIACEAE Gelsemium sempervirens (L.) Aiton f, MM 6113 SEWP 2005 Mitreola petiolata (J.F. Gmel.) Torr. & A. Gray, LB 29017, 28739; GW 2780 Mitreola sessilifolia (J.F. Gmel.) G.Don, LB 28431; MM 6626; 2695 LYCOPODIACEAE Lycopodiella appressa (Chapm.) Cranfill, MM 6890, 6579 Pseudolycopodiella caroliniana (L.) Holub var. caroliniana, MM 6582; GW 2712 LYGODIACEAE *Lygodium japonicum (Thunb. ex Murray) Sw MM 6317;SJ 16211;WRC 10845 LYTHRACEAE *Cuphea pele le 6298, 674 3 *Cuphea ss Cham.& Schlecht., MM 6266; LB 28393; GW 2512 otala ramosior (L.) Koehne, GW 2782 MAGNOLIACEAE Magnolia grandiflora L., LB 27818,28196; MM 6149; SJ 16070 (Jacq.) J.F. Macbr., MM Magnolia virginiana L., MM 6248; LB 27813; SJ 1637] MALVACEAE Mal j t Dill.ex.Cav.var.drummondii (Torr.ex A. Gray) Schery, LB 28345; MM 6719 Sida rhombifolia L., MM 6752;LB 28637 Sida spinosa L., GW 2725 MAYACACEAE Mayaca fluviatilis Aubl.,, GW 2507, 2788 MELASTOMATACEAE Rhexia mariana L., MM 6608;LB 28330;GW 2597, 2969 Rhexia virginica L., LB 28728; MM 6733 MELIACEAE *Melia azedarach L., LB 28964 MENISPERMACEAE Cocculus carolinus (L.) DC., LB 28203 MOLLUGINACEAE Mollugo verticillata L., MM 6606; SJ 16377 MONOTROPACEAE Monotropa uniflora L., LB 28372 MORACEAE Morus rubra L., MM 6123;LB 28776;SJ16438;GW 3335 MYRICACEAE Morella caroliniensis (Mill.) Small, MM 6237, LB 28956, 28784, 28182 Morella cerifera (L.) Small, MM 6340; LB 28130, 28247; GW 2707 BRIT.ORG/SIDA 22(1) NYMPHAEACEAE bi lutea (L.) Sm. subsp. advena (Aiton) artesz & Gandhi, LB 26168 OLEACEAE Chionanthus virgin SJ 16089 Forestiera ligustrina (Michx.) Poir., GW 1944, 2421, 2515; LB 27817 Fraxinus americana L., MM 6319; GW 2708; LB 28106; SJ 16365 Fraxinus caroliniana Mill., (SBSC) icus L., MM 6118; LB 28137, SJ 16203; LB 30565 LGN ll lait Marshall., MM 6076 r., LB 28945; SJ 16251 CNAGRACERE a alternifolia L.,LB 28331; GW 2799 oc a pl Walter GW 2798; LB 28724; eae are Walter, GW 2720, 2967; LB 28744 Ludwigia hirtella Raf; LB 28328; MM 6585, 6734 Ludwigia leptocarpa (Nutt.) Hara, GW 2968 Ludwigia linearis Walter, MM 6897 Ludwigia pilosa Walter, LB ae ie Oenothera laciniata Hill, GW Oenothera linifolia Nutt., MM pe Oenothera speciosa Nutt., MM 6260; SJ 16344 OPHIOGLOSSACEAE Botrychium biternatum (Sav.) Underwood, MM 6910; LB 27831 Botrychium virginianum (L.) Sw., MM 6331,6147 GW 1945 ORCHIDACEAE Corallorhiza wisteriana oe aa 2007 Listera australis Lindl., SEWP Platanthera clavellata ne ne LB 28268;GW 2563 Plata th ta (Michx.) Lindl., GW 2701 Spiranthes cernua (L.) L.C. Rich., MM 6952; GW 3449 Spiranthes lacera (Raf.) Raf. var. gracilis (Bigelow) Luer, MM 6620 Spiranthes praecox (Walter) S.Watson, SJ 16259 Spiranthes tuberosa Raf., LB 28382;MM 6573 OROBANCHACEAE Epifagus virginiana (L.) W. Bart., LB 28271 BROWN ET AL., FLAA VET OSMUNDACEAE Osmunda cinnamonea L., MM 6242;LB 28190 Osmunda regalis L.var spectabilis (Willd.) A.Gray, MM 6259; LB 28123, 28206, 27855 OXALIDACEAE Oxalis corniculata L. var. wrightii (A. Gray) B. L. Turner, MM 6096 *Oxalis debilis Kunth var. corymbosa (DC.) Lourteig, $J 16231 Oxalis lyonii Pursh, (T) MM 6135;WRC 17337 PASSIFLORACEAE Passiflora lutea L., MM 6293, 6742; LB 28939, 28338 — PHYTOLACCACEAE Phytol icana L., LB 28140 PINACEAE a ead Mill. MM 6893, 6082; LB 28949; SJ q I, MM 6070; SJ 1631 ate taeda L., MM 6083; SJ 16257 PLANTAGINACEAE Plantago aristata Michx.,SJ 16295;LB 28236; MM 6623 PLATANACEAE Platanus occidentalis L., MM 6246; SJ 16434; LB 27816; GW 2046 POACEAE pie is elliottiana J.A.Schultes, GW 2501 tis hyemalis (Walter) Britton, Sterns, *Poggenb, A legans inth, MM 6235;LB 28212 aneronocen gerardii Vitman, LB 29033, 28999 Andropogon glomeratus (Walter) Britton, Sterns, Poggenb. LB 28965; GW 2784 Andropogon gyrans Ashe var.gyrans, WRC 17339 Andropogon ternarius Michx., MM 6874,6916;GW 2807; LB 28982 Andropogon virginicus L.var. virginicus, LB 29027, 28997, 28941, 28936 Aristida lanosa Muhl. ex Willd., MM 6927; LB 29002; WRC 10829; DR 2367 Aristida longispica Poir. var. geniculata (Raf.) rnald, MM 6904 Aristida se Poir. var. longispica, LB 28926; GW 2789, 2 ristida ah aes GW 2791 PRESERVE 717 Aristida purpurascens Poir. var. purpurascens, MM 6871; LB 29004, 28991, 28931 Arundinaria gigantea (Walter) Muhl., MM 6072; LB 28650, SJ 16120; SEWP 2001 Axonopus fissifolius (Raddi) Kuhlm., LB 28359, 28611; GW 2705 *Briza minor L.,MM 6079; GW 2611 *Bromus japonicus Thunb. ex Murr., SJ 16359 Cenchrus spinifex Cav., LB 28363; MM 6615; GW 2/04 Chasmanthium lfatifolium (Michx.) Yates, SJ B 28350, 28151; MM 6743 Chasmanthium laxum (L.) Yates var. laxum, LB 30 Chasmanthium laxum (L.) Yates var. sessiliflorum (Poir.) Wipff & S. Jones, MM 6722; LB 28714, 28348 Dichanthelium aciculare (Desv. ex Poir.) Gould & Clark SJ 16060, 16107;LB 28238 Dichanthelium aciculare (Desv. ex Poir.) Gould & Clark subsp. angustifolium (Elliott) Freckman & Lelong, LB 28367 Dichanthelium acuminatum (Sw.) Gould & Clark var.acuminatum, SJ 16427;LB 28335 Dichanthelium boscii (Poir.) Gould & Clark, LB 28273, 28663, 28351, 28149 Dichanthelium commutatum (Schultes) Gould subsp , MM 6239; SJ 16064, i i a ent (SBSC) Dichanthelium commutatum (Schultes) Gould ubsp. equilateral (Scribn.) Freckman & Lelong, LB 28589 Dichanthelium dichotomum (L.) Gould var. dichotomum, MM 6299; LB 28975, 28628; SJ 16171 Dichanthelium dichotomum (L.) Gould subsp. microcarpon (Elliott) Freckman & Lelong, LB 28166, 28164 Dichanthelium ensifolium (Baldw. ex Elliott) ould subsp. ensifolium, LB 29038; SJ 16308 ohana laxiflorum (Lam.) Gould, LB 27788; WRC 1 See Sein oligosanthes (Schult.) Gould, var. oligosanthes, SJ 16184 Dichanthelium oligosanthes (Schult.) Gould var. scribnerianum (Nash) Gould, LB 28369 Dichanthelium oligosanthes (Schult.) Gould, WRC 17340 Dichanthelium ovale (Elliott) Gould & C.A. Clark subsp. villosissimum (Nash) Freckman & Lelong, SJ 16263, 16195; MM 6277; WRC 17341 Dichanthelium polyanthes (Schult.) Mohlenbr,, LB 28384, 30567 (SBSC), 30573 (SBSC); SJ 16404 Dichanthelium ravenelii (Scribner & Merr.) Gould & Clark, MM 6290; SJ 16435 Dichanthelium scoparium (Lam.) Gould, SJ 16407; LB 28405; MM 6612 Dic. pe ium sphaerocarpon (Elliott) Gould, MM 6 3; SJ 16265 LB 28630 See ane wrightianum (Scribn.) Freckmann, LB 28433; MM 6583 Digitaria ciliaris (Retz.) Koel., LB 28933, 28712 Digitaria cognata (J.A. Schultes) Pilger var. LGW 2815 Digitaria filiformis (L.) Koel. var. filiformis GW 2806 Digitaria filiformis (L.) Koel. var villosa (Walter) Fernald, MM 6876; LB 29026; GW 2418 *Digitaria violascens Link, LB 29009 Elymus virginicus L., LB 28208, 28732; MM 6535; SJ 16444 Eragrostis elliottii S.Watson, GW 2812 Eragrostis hypnoides (Lam.) Britton, Sterns, Poggenb 19b — Eragrostis intermedia (Lam.) Britton, Sterns, g 894 Eragrostis refracta (Muhl.) Scribn., LB 28989 — sree (UJ. Presl) subsp. oxylepis (Tor Koch, MM 6696;LB 28774;GW 2797 (Pursh) Steud., MM 6915, 983 oe spectabilis 6869, 6902; LB 28 Eustachys petraea (Sw.) Desv., MM 6903 Gymnopogon ambiguus (Michx.) Britton, Sterns nb., MM 6865; GW 2808; DR 2368 Leersia virginica Willd., LB 28971, 28627 *Lolium perenne L., MM 6322;SJ 16350 Melica mutica Walter, MM 6108;LB 28173, 27832 Oplismenus hirtellus (L.) Beauv., LB 28648: MM 6938 Panicum anceps Michx., MM 5757, 5646; LB 29008, 28974 Panicum brachyanthum Steud., MM 6912; LB 68980, 68929; WRC 10832 Panicum gymnocarpon Elliott, LB 28973, 28740 Panicum rigidulum Bosc ex Nees, LB 2874 Panicum verrucosum Muhl., MM 6881,6950; LB 29037, 2901 | *Paspalum notatum Fluegge, MM 6875, 6537;LB 28603; GW 2585a BRIT.ORG/SIDA 22(1) Paspalum plicatulum Michx., MM 6864, 6731;LB 28199; SJ 16356 Paspalum praecox Walter, MM 3933, 3719, 3937 Paspalum setaceum Michx., GW 2419: LB 2403,28928; MM 6724 *Paspalum urvillei Steud., MM 6895, 6541; LB 28221, 29010 Phalaris angusta Nees ex Trin., SJ 16255 Piptochaetium avenaceum (L.) Parodi, MM 6329; SJ 16378 *Poa annua L., MM 6142; GW 2497, 2498 Poa autumnalis Muhl. ex Elliott, LB 27829 Saccharum ‘sania (Walter) Pers., MM 6880 Se ne scoparium (Michx.) Nash var. eee Gould, LB 28998; GW 2814 setae paraiora (Poir.) Kerguelen, LB 28361 sh, MM 6919 Sphenophols filiformis (Chapm.) Scribn., WRC 7336 nes Sorghastn || ttii (Mo hr) Na Sph en resect atnyes) Rydb., SJ 16229 Sphenopholis obtu hx.) Scribn., MM 6300 Sporobolus niet inus ee A.S.Hitche.,, MM peed LB ian gis GW 2813 j R.Br, LB 28930 Sorat us junceus (Beauv) Kunth, LB 28993:MM 0; DR 2369 —_ Steinchisma hians (Elliott) Nash, SJ 16298 Tridens flavus (L.) ead var. chapmanii (Small) Shinners, WRC 1082 Tridens flavus (L.) Tee flavus, LB 28634;GW 2426, 2819/2783 Tridens strictus (Nutt.) Nash, LB 28904 POLEMONIACEAE Phlox nivalis Lodd. ex Sweet subsp. texensis Lundell, MM s.n.;WRC 17332 POLYGALACEAE olygala mariana Mill, LB 28215; SJ 16206 Polygala polygama Walter, MM 6296;LB 28116 POLYGONACEAE Brunnichia ovata (Walter) Shinners, MM 6951:LB 28138 Polygonum punctatum Elliott, MM 6930; LB 28656; GW 2883, 2778a Polygonum eral L., LB 28656a POLYPODIACEA Pleopeltis por ypodioide ANTAIEWs & rhe oi sub sp ndrews & pARSER tds Windham, LB 28407, 37795 BROWN ET AL., FLORA OF TH C DIN SANUT POTAMOGETONACEAE Potamogeton diversifolius Raf., SJ) 16273; LB 28682; Potamogeton pulcher Tuckerman, GW 2561 PRIMULACEAE An Ili is L., MM 6089 Anagallis minima (L.) Krause, LB 277 Samolus valerandi L. subsp. a iss (Raf.) Hultén, LB 29040; SJ 1624 RANUNCULACEAE Clematis reticulata Walter, GW 2734; MM 6610 Ranunculus hispidus Michx. var. nitidus (Chapm. T.Duncan, . 27841; GW 2734 *Ranunculus parviflorus L., GW 2517 Ranunculus pusillus Poir., GW 2480 ecurvatus Poir., GW 3351 RHAMNACEAE Berchemia scandens (Hill) K. Koch, MM 6103; LB 278615) 16159 Rhamnus caroliniana Walter, MM 6122:LB 28633, 27799, 30572 (SBSC) ROSACEAE Agrimonia microcarpa Wallr., GW 2769,3425; MM 6721 ro Agrimonia rostellata Wallr., LB 28732; MM 6334 Crataegus marshallii Egglest., MM 6726;LB 28970, 27825a; SJ 16074 Crataegus opaca Hook. & Arn., SJ 16078 Crataegus spathulata Michx., SJ 16096;LB 28644, 2/ ] Crataegus uniflora Muench., GW 2014 Crataegus viridis L. GW 1947a Prunus caroliniana (Mill.) Aiton, LB 28951 Prunus gracilis Engelman & A.Gray, MM 6284; LB 27785 Prunus serotina Ehrh., MM 6119;LB 28785, 28269; SJ 16067 Prunus umbellata Elliott, LB 27800; SJ 16172 Rubus argutus Link, SJ 16155;LB 28788; GW 2706, 2457 Rubus trivialis Michx., MM 6116 RUBIACEAE Cephalanthus occidentalis L., MM 6932, 6593 Diodia teres Walter, MM 6694; LB 2840 Galium pilosum Aiton, GW 2573; MM 6326 Galium tinctorium (L.) Scop., LB 27806; GW 2483 Galium triflorum Michx., SJ 16283; LB 28789 BIG THICKET NATIONAL PRESERVE 719 Galium uniflorum Michx., SJ 16278; LB 30460a BSC) Houstonia micrantha (Shinners) Terrell, WRC 17334 Houstonia pusilla Schoepf, MM 611 Mitchella repens L., MM 6247, 6189; LB 27803; SJ 161 ee Oldenlar L., GW 2713,2790 penroden pentane (K.Schurm.) Vatke, LB 28608; 2374 — W aL, LB 28924 *Sherardia arvensis L., MM 6136 Stenaria nigricans (Lam.) Terrell, LB 28368, 28222; RUTACEAE *Poncirus trifoliata (L.) Raf.,SJ 16246;LB 28652 Zanth herculis L.,LB 28185 | 'PaiVial ylun SALICACEAE Salix nigra Marshall. SJ 16142 SAPOTACEAE Sideroxylum lanuginosum Michx.subsp.oblongi- folium (Nutt.) T.D. Pennington, SJ 16306 SARRACENIACEAE Sarracenia alata Wood, MM 6589 SAURURACEAE Saururus cernuus L., LB 28620, 28142a; MM 6943 SAXIFRAGACEAE Lepuropetalon spathulatum Elliott, LB 27768 SCROPHULARIACEAE Agalinis fasciculata (Elliott) Raf, LB 28948, 28923 +Agalinis filifolia (Nutt.) Raf, MM 6921,6922,6901 Agalinis purpurea (L.) Pennell, LB 29029; GW 2879 Aureolaria flava (L.) Farw.var macrantha Pennell, 639 Aureolaria grandiflora (Benth.) Pennell, MM 6751; LB 28591; GW 2736 Aureolaria pectinata (Nutt.) Pennell, GW 2880 Bacopa caroliniana (Walter) Robins. LB 30562 (SBSC) Gratiola neglecta Torr, GW 2526 Gratiola pilosa Michx., GW 2696; LB 29036; MM 6581 Gratiola Virgil al 1d L, GW 2485, 2525), 2604 Lindernia dubia (L.) Pennell, SJ 16146; LB 28720; GW 2605 Mecardonia acuminata (Walter) Small,GW 2781; LB 29028, 28230 720 Mecardonia procumbens (Mill.) Small, GW 809 Micranthemum umbrosum (J.G.Gmel.) Blake, SJ 16269; 28592 Pe nctemon iflary) 16077; LB 28227 Scoparia dulcis L., MM 6692; LB 28357, 28225 SELAGINELLACEA Selaginella apoda (L ‘ Spring, MM 6255;GW 2508; Pennell, MM 6307, 6088; SJ SMILACACEAE Smilax one: nox L., MM 6691 Smila ca Walter, LB 28977 in Lunplia L., MM 6250, se : 28193:SJ Smilax pumila Walter, MM 6086; SJ 1616 Smilax rotundifolia b., MM 6238: SJ 16436; LB 27779 Smilax smallii Morong, MM 6240; LB 28938, 28233; SJ 16158 Smilax tamnoides L.,SJ 16202 SOUEN RSENS Physali phylla Nees, i 6286; GW 2801 Physalis pubescens L., GW 2599 al pau Nutt. MM 5369, 5404; GW 2660 M 6723;LB 28337 *Solanum capsicastrum Link ex Schauer, GW inense L 2047 Solanum ptycanthum Dunal, MM 6937;SJ 16248 SPARGANIACEAE Sparganium americanum Nutt., SJ 16293: LB 30564 (SBSC) STYRACACEAE Halesia diptera Ellis, SJ 16412;LB 27825 Styrax grandifolius Aiton, LB 28128, 28124,28127 SYMPLOCACEAE Symplocos tinctoria (L.) L.Her, SJ 16318;LB 28783, 28277 THELYPTERIDACEAE Thelypteris kunthii (Desv.) Morton, MM 6339; SJ 16227; LB 26244, 26244 TILIACEAE Tilia americana L. var. caroliniana (Mill) Castigl., LB 27828; GW 2012, 2048 ULMACEAE Celtis laevigata Willd., GW 2513b,2514c Celtis tenuifolia Nutt,, MM 6522:LB 30534 (SBSO), 30560 (SBSC) — at BRIT.ORG/SIDA 22(1) Planera aquatica J.F. Gmel., LB 28180; SJ 16204,6414; GW 2042 Ulmus alata Michx., MM 6706, 6092; GW 2415:S) 16439 Ulmus americana L., SJ 16132,5J 16140;LB 28211; GW 2425 URTICACEAE Boehmeria cylindrica (L.) Sw., LB 28344, 28136; MM 6744 VALERIANACEAE Valerianella radiata (L.) Dufr., MM 6333 VERBENACEAE Callicarpa americana L., SJ 16442;LB 28347;MM a 6 9 Glandul sg canadensis (L.) Nutt. MM 6115; LB 2840 es orem ) Greene, SJ 16431 Vell., SJ 6908 Veibene halei Small MM 6308; LB 28236a; GW 2689 na brasili *Verbena rigida Spreng., MM 6542 *Vitex agnus-castus L., MM 6542 VIOLACEAE we lanceolata L.,LB 27859 iola palmata L., MM 6129; LB 27812; GW 2519 2542 Viola pedata L., MM 6138;SJ 16092 Viola primulifolia L.,SJ 16134;LB 29014;MM 6252: SEWP 200 Viola sororia Willd. var missouriensis (Greene) cKinney, LB 28135, 27819; GW 1941, 2443 Viola villosa Walter, MM 6109;LB 27787;GW 2481, 2931 VITACEAE Ampelopsis arborea (L.) Koehne, MM 6728, 6728; Parthenocissus quinquefolia (L.) Planch., MM 6126; SJ 16094 Vitis aestivalis Michx., GW 2580; MM 6558, 6287; 16274 Vitis cinerea (Engelm) Millard. var. cinerea, LB 605, 28132 Vitis rotundifolia Michx., MM 6124; LB 28120; SJ 16223 XYRIDACEAE Xyris ambigua Bey. ex Kunth, LB 28715 Xyris baldwiniana Schult., MM 6580; GW 2711;LB 30459 BROWN ET AL., FLORA OF THE BIG SANDY PRESERVE 721 Xyris ee 5 Chapm. var. curtissii (Malme) Kral, Xyris difformis Chapm. var. difformis, GW 2710 B 28425; MM 6732 Xyris jupicai L.C. Rich., MM 6888, 6700 Notes Agalinis filifolia— We are following Wunderlin (1998) in merging A.. pulchella with A.. filifolia. Cerastium pumilum.—Turner et al (2003) mapped this exotic species in six northeastern Texas counties with Kaufman County the most southern. Our Polk County record and another one from San Jacinto County (both at SBSC) are the most southern Texas collections known to us. Croptilon hookerianum.—The Polk County collection is the first for the pineywoods region of east Texas. The most eastern collection in Turner et al. (2003) is from Fayette County. It is distinguished from C. divaricatum and C. rigidifolium (the only other Croptilon taxa in Texas) by the lack of non-glan- dular trichomes on the distal portions of the stems. Diaperia candida.—In the Flora of North America Asteraceae volumes (ined.), our Evax taxa are treated in the genus Diaperia. Loeflingia squarrosa.—This western taxa appears to be new to Polk County (Turner et al. 2003). This, Prunus eracilis,and Croptilon hookerianum are the only strictly western sy found on the Big Sandy Creek Unit. These are char- acteristic species of xeric sandylands. Minuartia muscorum.—For the curious nomen clatural history of this taxon see Rabeler (1992). Phlox nivalis subsp. texensis.—1T his endemic to southeast Texas anda federally and state listed endangered species (Poole & Riskind 1987; Texas Parks and Wildlife 2004), was located on the Big Sandy Creek Unit during the study. The species has been planted in several locations in the unit by the USS. Park Service, but the population located during this study was a naturally occurring one which had been known for some years. Ranunculus recurvatus.—Our Polk County collection is one of only four coun- ties mapped in Turner et al. (2003). Silene subciliata.—In addition to Phlox nivalis ssp. texensis, Silene subcilata is the only other Texas rare species found on the Big Sandy Creek Unit. This West Gulf Coastal Plain endemic is rare in both Texas and Louisiana, but is not fed- erally listed. In addition to these two rare West Gulf Coastal Plain endemics, the Big Sandy Creek Unit has several other West Gulf Coastal Plain endemics (MacRoberts et al. 2002a). These are Baptisia bracteata var. leucophaea, Bapti- sia nuttalliana, Berlandiera pumila var. scabrella, Dalea phleoides var. phleoides, Echinacea pallida var. sanguinea, Hymenopappus artemistifolius var. artemisiifolius, Lobelia puberula, Mimosa hystricina, Oligoneuron nitidum, Panicum brachyanthum, Scutellaria cardiophylla, Sisyrinchium sagittiferum, Solidago ludoviciana, Tradescantia reverchonii, and Vernonia texana. 722 BRIT.ORG/SIDA 22(1) Sophora affinis—Our Polk County collection is one of only three specimens known to us from the pineywoods region in east Texas. The other two collec- tions are from the Trinity River bottomlands in Liberty County. DISCUSSION There are 126 families and 358 genera for the 693 taxa (648 native) on the Big Sandy Creek Unit list. Also, 50.9 % of all taxa belong to eight families; they are Poaceae (88 taxa), Asteraceae (85 taxa), Cyperaceae (69 taxa), Fabaceae (46 taxa), Lamiaceae (19 taxa), Euphorbiaceae (16 taxa), Scrophulariaceae (16 taxa), and Fagaceae (15 taxa). The largest genera are Carex (35 taxa), Dichanthelium (19 taxa), Quercus (13 taxa) Cyperus (11 taxa), and Rhynchospora (11 taxa). We compared our list of taxa for Big Sandy Creek Unit with the Polk County taxa reported by Turner et al. (2003) in their Atlas of the vascular plants of Texas and found that only 70 percent of our taxa are mapped for Polk County in that source. This is not surprising considering that until recently little focused col- lecting has been undertaken in the Big Thicket region and indicates that basic floristic collecting and documentation in the Big Thicket region are still needed. Thus, while itis the case that our list isincomplete and that a few taxa may no longer exist on the unit, most taxa that have grown in the Big Sandy Creek Unit during the past quarter century have been collected (probably 85+ per- cent). We estimate that the Big Sandy Creek Unit has about 660 to 750 native species/taxa. ACKNOWLEDGMENTS We thank Guy Nesom (BRIT) and David Riskind (Texas Parks and Wildlife De- partment) for their careful review and resulting improvement of this paper and the staff of TEX/LL, largely Tom Wendt and Lindsay Woodruff, for facilitating the senior authors’s examination of the William Carr specimens. This study was supported in part by a National Park Service Cooperative Agreement No. CA 14001004 to Paul Harcombe who also provided funds for the page charges. REFERENCES Autvscl, G. 1979. Wild flowers of the Big Thicket, east Texas, and western Louisiana. Texas A&M Press, College Station. Brown, L.E.,B.R. MacRoserts, M.H.MacRoserts, PA. HAaRcomBe, W.W. Pruess, |S. Ecsik, and D. JOHNSON. 2005. Annotated checklist of the vascular flora of the Turkey Creek Unit of the Big Thicket National Preserve, Tyler and Hardin counties, Texas. Sida 21:1807-2005. Desuorets, J.D. 1978. Soil survey for the Big Thicket National Preserve, Texas. U.S.D.A. Soil Conservation Service, College Station, Texas. Harcomee, P.A.and P.L. Marks. 1979. Forest vegetation of the Big Thicket National Preserve. Unpublished report: U.S. Park Service, Santa Fe, NM. BROWN ET AL., FLORA OF PRESERVE 723 Harcomee, PA., J.S. Guzenstein, R.G. Knox, $.L. Orzett, and E.L. Brioces. 1993. Vegetation of the longleaf pine region of the West Gulf Coastal Plain. Proc. Ann. Tall Timbers Fire Ecol. Conf. 18:83-103. Jones, S.D., J.K. Wipre, and P.M. Montcomery. 1997. Vascular plants of Texas. Univ. Texas Press, Austin. Kartesz, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vas- cular flora of the United States, Canada, and Greenland. First edition. |n:J.T.Kartesz and C.A.Meacham. Synthesis of North American flora.Version 1.0.North Carolina Botanical Garden. Chapel Hill. MacRoserts, M.H., B.R. MacRoserts, B.A. Sorric, and R.E. Evans. 2002a. Endemism in the West Gulf Coastal Plan: importance of xeric habitats. Sida 20:767-780. MacRoserts, B.R., M.H. MacRoseets, and L.E. Brown. 2002b. Annotated checklist of the vascu- lar flora of the Hickory Creek Unit of the Big Thicket National Preserve, Tyler County, Texas. Sida 20:781-795. Marks, PL. and PA. Harcomee. 1981. Forest vegetation of the Big Thicket, southeast Texas. Ecol. Monogr. 51:287-305. McLeop, C.A. 1971. The Big Thicket forest of east Texas. Texas J. Sci. 23:221-233. Parks, H.B. and V.L. Cory. 1936. Biological survey of the east Texas Big Thicket area. Texas Agricultural Experiment Station, College Station. Peacock, H.H. 1994. Nature lover's guide to the Big Thicket. Texas A&M. Press, College Station. Poote, JM. and DH. Riskino. 1987. Endangered, threatened, or protected native plants of Texas. Texas Parks and Wildlife Dept., Austin. TEXAS PARKS AND WILDLIFE & THE NATURE CONSERVANCY, AusTIN, Texas. 2004. List of the rare plants of Texas. http://nature.org/whwerwework/northamerica/states/texas/files/ listofrareplants/pdf. RaBeLer, R.K. 1992. A new combination in Minuartia (Caryophyllaceae). Sida 15:95—96. Turner, B.L., H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Sida Bot. Misc. 24: 1-888. Watson, G.E. 1979. Big Thicket plant ecology: an introduction. Big Thicket Mus. Publ. Ser, No. 5, Saratoga, Texas. Watson, G.E. 1982. Vegetational survey of Big Thicket National Preserve. Unpublished re- port. Big Thicket National Preserve, Beaumont, TX. WunNbeERLIN, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida. Tampa. 724 BRIT.ORG/SIDA 22(1) THE NEW YORK BOTANICAL GARDEN PRESS Flora Genérica de los Paramos Guia Ilustrada de las Plantas Vasculares Petr Sklendr, James L. Luteyn, Carmen Ulloa Ulloa, Peter M. Jorgensen and Michael O. Dillon Memoirs of The New York Botanical Garden Volume 92 ISBN-0-89327-468-2 Order no. MEM 9 $85.00 520 pages n Spanish with an English abstract =a P4ramo is one of the richest nce elevation mountain ecosystems in the world with high biological diversity and endemism, but it is ecologically fragile. This book provides keys, desc a era distrib- utional information, and illustrations for the dence ation of the 127 families and 540 genera of vascular plan le | i s de alta montafa mas rico del Mundo con una diversidad es ramo es bioldgica y un endemismo altos, pero es frégil ecolégicamente. Este libro proporciona claves, descripciones, informacién acerca de la distribucién e ilustraciones para la identificacién de las 127 familias y 540 géneros de plantas vasculares “three-digit r Manca urd nani on ont : ofA rie Ez Ord ath P sed withour this securi : instirudon, Se Tee ee ie eee eee SIDA 22(1): 724. 2006 THE NORTH-SOUTH TRANSITION OF FLORA ACROSS ARKANSAS: A PRELIMINARY PHY TOGEOGRAPHICAL ANALYSIS Michael H. MacRoberts and Barbara R. MacRoberts Bog Research 740 Columbia Shreveport, Louisiana 71104, U.S.A. and Herbarium, Museum of Life Sciences Louisiana State University in Shreveport Shreveport, Louisiana 71115, U.S.A. — ABSTRACT 1 ] Ae A : hi i | 1 Using quan- titative nethods we examined the north- south f floristic feaneiaga ACTOSS Denese to see if phyto- geographic regions are detectable by abrupt changes in the flora. We found only gradual change even though there are abrupt and major physiographical discontinuities. RESUMEN Los BleeeOssaes han dividido Arkansas en provincias fisiogeograficas y ecorregiones. Usando uantitativos, examinamos las transiciones norte-sur a través de Arkansas para ver si las regiones fitogeograficas son detectabes por Cambios peeEuDios en la flora. Hemos encontrado solo y mayores. }: cambio gradual aunque hay 3 INTRODUCTION Virtually all ¢ hic and ecoregional mappers treat the Interior High- lands of Arkansas, Illinois, Kansas, Missouri, and Oklahoma as being distinct from surrounding areas. While most appear to agree on a physiogeographic map, they present conflicting interpretations of the biotic provinces and ecoregions (e.g., Dice 1943; Braun 1950; Gleason & Cronquist 1964, Kuchler 1964; Foti 1974; Pell 1983; Omernik 1986; Takhtajan 1986; Thorne 1993; Bailey et al. 1994; Keyset al. 1995; Weakley et al. 1998; Foti & Bukenhofer 1998; Ricketts 1999). For example, Dice (1943), Takhtajan (1986), and Thorne (1993) consider the In- terior Highlands and Gulf Coastal Plain to be different provinces, Bailey et al. (1994) and Keys et al. (1995) break the Interior Highlands into two provinces, Omernik (1986) divides the area into six ecoregions, and Weakley et al. (1998) divide it into four ecoregions. While it is unquestionably the case that the Ouachita Mountains and the Ozark Plateau are physiogeographically distinct from surrounding areas, no- tably the West Gulf Coastal Plain to the south, it is not so certain that the area is particularly distinct floristically. Zollner et al. (2005:1788), speaking of the In- SIDA 22(1): 725 — 734. 2006 726 BRIT.ORG/SIDA 22(1) terior Highlands in general, question its uniqueness: “for all of its physiographic uniqueness, including age, long-term isolation from its moiety—the Appala- chian region—, and its reputation as a ‘well-known refugium’ (Meyer 1997:364) the Interior Highlands] shows surprisingly little floristic uniquity from sur- rounding regions.” This was also recognized years ago by Palmer (1921) when he pointed out that about 90 percent of the woody flora of the Ozark region also occurs in the Gulf Coastal Plain. This point was brought home to us forcefully when we conducted a floris- tic inventory of the Ouachita Mountains Biological Station in Polk County, Ar- kansas (MacRoberts et al. 2005). Having extensive experience with the flora of the West Gulf Coastal Plain, but none with the Ouachita Mountains, we ex- pected, since the physiogeography was so different and since biogeographers have separated the Interior Highlands from the Coastal Plain, that the two ar- eas would be quite floristically distinct. Surprisingly they are not. The flora of the Ouachita Mountains Biological Station presented some species unknown to us, but not many. Ina previous paper, we documented the east-west floristic transition across central Texas in which the eastern flora precipitously drops out and is replaced by a western flora and vice versa (MacRoberts & MacRoberts 2003). In this study, using similar methods we examine the north-south floristic transition across Arkansas, i.e., from the West Gulf Coastal Plain in the south into the In- terior Highlands in the north (Fig. 1). Western Louisiana and southern Arkan- sas are physiographically the Gulf Coastal Plain (Fenneman 1938; Brouillet & Whetstone 1993) and the Atlantic and Gulf Coastal Plain floristic province (Takhtajan 1986; Thorne 1993). The Interior Highlands, on the other hand, are part of the Ozark and Ouachita physiographic region (Fenneman 1938; Brouillet S Whetstone 1993) and the Appalachian floristic province (Takhtajan 1986; Thorne 1993). The purpose of this paper is to provide data on 1) how different floristi- cally the Interior Highlands is from the West Gulf Coastal Plain, 2) the effect of the physiogeographic differences on the north-south floristic transition of flora from the Coastal Plain into the Interior Highlands, and 3) the floristic relation- ship between the Interior Highlands and the regions that surround it. — — STUDY SITES The Interior Highlands and the Gulf Coastal Plain have been described in numer- ous publications (Fenneman 1938; Foti 1974; Pell 1983: Bryant et al. 1993; Skeen et al. 1993; Dale & Ware 1999; Delcourt & Delcourt 2000) and will not be redescribed here except to say that the former region consists of moderate elevation moun- tains of Paleozoic sedimentary rock with numerous swift clear streams and riv- ers and both deciduous and pine-hardwood forests; the latter region consists of low to gently rolling topography of Cretaceous and Tertiary silts and clays char- NEGCeae / mae” 2 Ouachita ae ae nu Reece ors, 3 or ae A Fic. 1. Location of Interior Highlands and Gulf ¢ | and Baxter (B), Garland (G), Newton (N), and Polk (P) counties. 728 BRIT.ORG/SIDA 22(1 => acterized by forests of pine and hardwoods, with swamps, and sluggish muddy streams and rivers, and oxbow lakes. Precipitation is about the same in both areas, with temperatures slightly cooler in the Interior Highlands. METHODS 1) As we pointed out in the introduction, we have recently conducted a floristic survey of the 211 hectare Ouachita Mountains Biological Station in Polk County, Arkansas (MacRoberts et al. 2005). It is located 174 km north of Louisiana in the Ouachita Mountains and has an elevation range of between 395 and 622 meters. It is dominated by pine-hardwood forests on shale, chert, novaculite, siltstone, and sandstone. It could not be more different edaphically and topo- graphically from the West Gulf Coastal Plain. We have documented 315 native species on the station. We compared this list to floristic lists for Louisiana (Tho- mas & Allen 1993-1998; Kartesz & Meacham 1999) to determine the percent- age of species in common. While the Ouachita Mountains Biological Station is a very small area given the size and diversity of the Ouachita Mountains, it pro- vided a first comparison of the flora between the West Gulf Coastal Plain and the Ouachita Mountains. 2) In order to obtain a more detailed understanding of the possible effect of the Interior Highlands on plant distribution, using the Thomas and Allen (1993-1998) atlas we randomly selected 254 native species occurring in north- ern Louisiana (north of TIN) that also occur in Arkansas (Smith 1988, 1994). Using Kartesz and Meacham (1999), we determined how many of these occur in Missouri, approximately 380 km to the north. The sample size is only about 10 percent of the native species but sufficient to compare the flora between the regions. 3) As an independent test of Method 2, using Hyatt (1993) we determined the percentage of native species occurring in Baxter County, Arkansas, on the Missouri border in the Ozark Plateau, that also occur in the Gulf Coastal Plain of Texas and Louisiana (Thomas & Allen 1993-1998: Kartesz &@ Meacham 1999; Turner et al. 2003). Naturally, some of the species in this sample are the same as those in Method 2. 4) Using the data in Method 2, we plotted the Arkansas county distribu- tion of the 254 native species. Because all counties are not collected equally (e.g., Jackson County has 275 reported taxa while Washington County has 1,301 reported taxa), we expressed the results as percentages; that is, we divided the number of species in our sample reported for each county by the total species reported for each county. Thus, of the 254 species in the sample, 99 are reported for Miller County, and Smith (1988) reports 544 species for Miller County (99/ 544 = 18%). 5) Having found that Method 4 did not reveal sharp distributional discontinuities in the flora across Arkansas (see “Results”), using Kartesz and , BIOGEOGRAPHICAL ANALYSIS OF THE ARKANSAS FLORA 729 Meacham (1999) we surveyed the entire flora of North America and found all of the native species that were reported to occur in Missouri and Arkansas (but not in Louisiana) and in Louisiana and Arkansas (but not in Missouri) irrespec- tive of where else they occurred. Using Smith (1988), we then plotted these spe- cies by Arkansas counties according to whether they were Missouri/ Arkansas species or Louisiana/ Arkansas species (i.e, coming from the north or the south). Our sample consisted of 499 species: 272 Missouri/Arkansas and 227 Louisi- ana/ Arkansas. Our Buea es) procedure was the same as in Method 4. Our hope was to magnify the curve b inating species that occurred in all three states. 6) In order to determine the overall North American floristic association of the Interior Highlands, using Smith (1988) we randomly selected 296 native species from Garland County, Arkansas, which is entirely within the Ouachita Mountain region, and 293 native species from Newton County, Arkansas, which is entirely within the Ozark Plateau region (both counties being well collected) and, using Kartesz and Meacham (1999), plotted their state/regional occurrence expressed as percentage of species in common. All of these measures are of presence/absence, not abundance. The data necessary to investigate abundance are not yet available on a large scale. Thus, we recognize that some taxa may be widely distributed across the region while others may not. Also, while we recognize the limitation of the main data sources for this paper, we have made every effort to adjust to those limitations. But since the study is a preliminary analysis, it can absorb some error without compro- mising the general conclusions. The paper provides a problem and an analysis that should be followed up by a more extensive sample of species and more points along the continuum. RESULTS 1) Of the 315 species on our Ouachita Mountain Biological Station list, 287 or 91% occur in Louisiana entirely within the West Gulf Coastal Plain. 2) Of the 254 species in our north Louisiana sample, 209 or 82% reached Missouri and 45 or 18% stopped in Arkansas. 3) Eighty-two percent of the native species that occur in Baxter County on the Missouri border also occur in the West Gulf Coastal Plain of Louisiana and Texas. 4) Figure 2 summarizes the results of Method 4. Counties on the southern tier of Arkansas averaged around 18% and 19% of the sample, and counties on the northern tier averaged between 12% and 16% of the sample. No marked or abrupt shifts in flora were detected that correlated with the natural physiogeographic divisions of Arkansas. 5) Figure 3 summarizes the results of Method 5. No marked or abrupt shifts in flora were detected that correlated with the natural physiogeographic divi- sions of Arkansas. 730 BRIT.ORG/SIDA 22(1) * 12 [12] 13] 46 Lali eaey" © 12 12 . 4 ere a je tial fel "18! f ag oe Pen ae ae Fic. 2. P t by £7CA] L ] . \ y t \ TUPLE CAPIdHaUlOit). 6) Figure + shows that the Ouachita Mountains has about 91% of its flora in common with the West Gulf Coastal Plain and the Ozark Plateau has about 82% of its flora in common with the West Gulf Coastal Plain. It also shows that the Ouachita and Ozark mountain/plateau regions are not only continuous flo- ristically with the surrounding regions but virtually indistinguishable from them. Laer | — DISCUSSION As noted in the introduction, biogeographers have presented a number of schemes for subdividing the Interior Highlands. These range from putting the Interior Highlands and Gulf Coastal Plain into separate biotic provinces or ecoregions to dividing the Interior Highlands into two provinces, and so on. MACROBERTS AND MACROBERTS, BIOGEOGRAPHICAL ANALYSIS OF THE ARKANSAS FLORA 731 Pee 2 | I TUPUTTET CAPId- Our analysis shows that the vast majority of plants occurring in north Louisi- ana and east Texas (West Gulf Coastal Plain) are also found in central and north- ern Arkansas. There is a gradual transition of species from the West Gulf Coastal Plain into the Interior Highlands and vice versa, and there are no marked or abrupt shifts correlated with the natural divisions of Arkansas, indicating that there are no major physiographical barriers. The only place we may have de- tected a possibly sharper transition is between the Ozarks and the Mississippi floodplain, but even here the transition is not precipitous. The Interior Highlands are not floristically distinct from adjacent regions. The Ouachita flora as represented by Garland County, Arkansas, is closely re- lated to the flora of all surrounding states especially Oklahoma, Louisiana, Mississippi, Tennessee, Kentucky, Alabama, Georgia, and the Carolinas (Fig. 4). The Ozark flora as represented by Newton County, Arkansas, is very similar to the Ouachita flora but, as would be expected, has a more northern affiliation. The Ozark region has its strongest floristic affinity with Oklahoma, Missouri, Illinois, Indiana, Kentucky, Tennessee, Ohio, Mississippi, Alabama, and Geor- gia (Fig. 4). If any generalization is necessary, the Interior Highlands as Zollner et al. (2005: 1788) have pointed out “shows surprisingly little floristic uniquity from surrounding regions.” Perhaps some or most of this lack of uniqueness is the result of movernents of species north-south for the last two million years of 20 glaciations, with strag- glers both north and south. This pattern of plant movement has been docu- mented extensively, notably by pollen analysis but also by fossil plants (Givens & Givens 1987, Delcourt & Delcourt 2000). Also, edaphic conditions are not entirely dissimilar along this north-south continuum, with high rainfall and generally mild climates. 732 BRIT.ORG/SIDA 22(1) . flnrice: Lins? fh) la (lnft) 1 far] Wa {eiahe) Ct 1c.4.N +h A of species in common. What these results mean for biogeographic classification and mapping is not entirely clear. Takhtajan (1986) has defined “province” differences as involv- ing low levels of generic endemism and high levels of species endemism. “Ecoregion” is generally defined as a large area consisting of a distinct assem- blage of natural communities and species with endemism being fairly high at the species and subspecies level (MacRoberts & MacRoberts 2004). The West Gulf Coastal Plain and the Atlantic and Gulf Coastal Plain have been the sub- ject of recent endemic studies (Sorrie @ Weakley 2001; MacRoberts et al. 2002). The West Gulf Coastal Plain has approximately 100 endemic taxa, three at the generic level. The Interior Highlands has only about 37 endemics, none at the generic level (Zollner et al. 2005). Setting aside the fact that no precise “index” has been proposed to test for provincial or ecoregional status, it seems from our analysis that the Interior Highlands and Gulf Coastal Plain are too similar flo- ristically to be classified as different provinces or ecoregions. Perhaps some lower chorionomic category might better classify these two areas. ACKNOWLEDGMENTS Douglas Zollner, The Nature Conservancy, commented on an earlier version of this paper. George P. Johnson and an anonymous referee commented on the paper. REFERENCES Baivey, R.G., PE. Avers, T. Kine, and W.H. McNas. (eds.). 1994. 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HIGH LEVELS OF SEED INVIABILITY AMONG SEVEN POPULATIONS OF THE ENDANGERED SHORTS GOLDENROD (SOLIDAGO SHORTII ASTERACEAE) Patrick J.Calie Robert F.C. Naczi Dept. of Life Sciences The Claude Phillips Herbarium 235 Mone selene Bldg. Delaware State University Eastern Kentucky University Dover, Delaware 19901, U.S.A. Biemend: Kentucky 40475, U.S.A. pat.calie@eku.edu Christina Shackleford Jessica Caicedo Dept. orenvionmenta! Health Science ries one Sciences Eastern ea Vary East y University Richmond, Kent 40475, U.S.A, eae Kentucky 40475, U.S.A, ABSTRACT The physical conditions of mature seeds If x Kentucky popu- lations and one Indiana population of the endangered species ‘Short’s eaieeed. (Solidago shortii, Asteraceae) A sadn ee bsenes oe eae Reutheny populations levealed the majority (71%- OOO | 6) I 1 insect preda- tor or to unsuccessful fertilization or aie ryo tion. Successful seedling ee from phe- notypically normal seeds eet rom 71% to 96% in the ed Kentucky populations. The majority of the damaged seeds were necrotic, ranging from 62%- 90%, The Indiana popwlanon exiubited the lowest ee of herbivory ee) ebbeived for the ESS ae congener gold anadensis ‘5 1andS or adjacent to thre lati f Short’s ae exhibited lower levels of necrosis Ae higher eee of herbivory. A ee congener species (S. gigantea) exhibited equal levels of the two seed conditions (herbivory and necrosis). The influ- ence of seed inviability upon the persistence of natural populations of Short’s goldenrod is at present een RESUMEN dici Ficicg | mil] maduras de 2 ope amenazada “Short’s goldenrod” (Solidago shortii, A yi inad i 1 Kentucky y una poblacion de Indiana. La ayer a Hae sepals de poblaci las al azar en ERentiesy Pisce arOne un numero reducido ] herbivoria, infertilidad estas ebenvabiones Semillas normales encontradas en las poblaciones de Kent oe dene ron plantulas que se desarrollaron con éxito en un 71% a 96%. En la mayoria de las semillas danadas los niveles de necrosis fluctuaron en un 62%-90%. La poblacion de Indiana presenté el menor nivel de herbivoria (2%) ob vado ena epee En dos Spee: de “goldenrod” (S. canadensis var. altissima y S lis) distribuid | “Short’s goldenrod” se observaron bajos ivelesd isyal iveles de herbivoria. E le “goldenrod” (S. gigantea) se ena Sonetone similares de mopivens y necrosis. Se deseoncte is insluencte de ae a persl SIDA 22(1): 735 — 748. 2006 736 BRIT.ORG/SIDA 22(1) INTRODUCTION Short’s goldenrod (Solidago shortii, Asteraceae) isan endangered species extant in the Blue Licks area of north central Kentucky and in one locality in Indiana (Homoya & Abrell 2005). A history of the discovery of this species, a descrip- tion of the Kentucky localities in which this plant is currently found, and gen- eral features of the species’ ecological life history are summarized in Baskin et al. (2000). As with many endangered species, a key question remains unanswered: Why does this species have such a limited geographical distribution? In an ef- fort to answer this question, an extensive body of knowledge has emerged re- garding the autecology of Short’s goldenrod (e.g. Buchele et al. 1989), involving investigations of both abiotic and biotic factors and their possible impact on the occurrence of this rare species. A later series of investigations implicated interspecific competition, primarily from introduced species, as being a lead- ing candidate greatly interfering with establishment of Short’s goldenrod seed- lings (Walck et al. 1999a, 1999b). One possible biotic component contributing to the rar 2 of the species is its reproductive biology, e.g. successful production of viable offspring. The first studies focusing on S. shortii seed germination were conducted by Buchele et al. (1991), that were later greatly expanded by Walck et al. 1997a, 1997b, 1997c, 1997d, 1997e, 1997f, 1999c). The primary (but not sole) emphasis in these stud- ies was to determine the biotic and abiotic factors most responsible for success- ful seed germination and seedling survival and the range of conditions (e.g. light, temperature, moisture, cold stratification, etc.) required for optimum germina- tion. In the first set of germination experiments (Buchele et al. 1991) seeds were obtained from two populations of Short’s goldenrod; in the second set (Walck et al. 1997a) one population was sampled. In our investigations involving the biology of Short’s goldenrod, we noticed high levels of seed damage in individuals from several Kentucky populations and asked whether differences existed among populations in reproductive po- tential, ie. the production of phenotypically normal seeds and developmen- tally normal seedlings. We posed the following questions regarding the repro- ductive biology of S. shortii. 1) What types of seed damage are present? What types of variation, if any, exists at the population level? Are some populations more prone to particular damages than others? 2) What quantitative variation, if any, exists both within and among selected Kentucky populations of Short’s goldenrod in the levels of different types of seed damage and the production of normal seedlings? 3) Do the common congeners of S. shortii in the Blue Licks vicinity (ie. S. canadensis var. altissima and S. nemoralis) and at the Indiana site (S. gigantea and S. nemoralis) exhibit similar levels of normal seed produc- tion as S. shortii? Are there statistically significant differences between the Ken- CALIE ET AL., SEED INVIABILITY IN SOLIDAGO SHORTII 737 tucky and Indiana populations of Short’s goldenrod with respect to normal seed production? MATERIALS AND METHODS To address the first two questions, matur 1s (cypselae) of S. shortii were col- lected in mid-December 2001 from six Kentucky populations designated as #1, 2,5, 7,8, and 12 (Evans 1987; Smith et al. 2004). This time was chosen to ensure the collection of mature seeds, as noted in earlier studies (Buchele et al. 1991a; Walck et al. 1997a). The sampled populations are found within a 2.2-km area of Blue Licks Battlefield State Park in northern Kentucky, and were chosen due to the prevalence of flowering stems and physical accessibility. Ten individual stems were non-randomly sampled from each population as follows. The ini- tial sampled plant stem was a minimum of 10 meters from the eg ae of that specific population (based on the GIS ing data from Smith et al. 2004). Due tothe non-random distribution of these plants, the cluster of plants in clos- est proximity to the first sampled stem was then sampled, this process being repeated until ten plants were sampled. To maximize the probability of sam- pling separate genets, each sampled stem was separated from any neighboring sampled stems by at least 3 meters. Portions of mature infructescences were removed from the stems, placed in a paper coin envelope, and stored at 4°C in the dark for 12-16 weeks. Following cold storage, the dev tal fate of individual seeds sampled from the Kentucky populations in 2001 was followed by the use of 96-well microtiter plates (Fig. 1) as germination chambers prepared as follows. A mini- mal mineral salts medium (Hickok & Warne 1998) containing 1% agar was au- toclaved and then aliquotted into each well (200 ml of medium/well) with a multichannel pipettor, then allowed to cool for 24 hours. Seeds were harvested by first scraping along the infructescence with forceps to dislodge the fruits and then collecting the seeds into a Petri plate. Single seeds were removed (with- out regard for condition) by grasping the fruit’s pappus with forceps until the appropriate number of seeds was obtained. Each seed (in the majority of cases 24 seeds per plant, with exceptions noted below) was then carefully imbedded half its length into the solidified agar within a single well, with the pappus extruding above the medium. The plate cover was labeled with the location and identity of each seed source, secured with parafilm, and the plates incubated under the following conditions. A daily photoperiod of 14 hours was provided by a bank of cool white fluorescent lights (2OW), that produced a photosyn- thetic photon flux (400-700 nm) at the germination plate level of approxi- mately 20-25 umol m™ sec! The dark period temperature was 20°C and the light period temperature was 25°C. A seedling was scored as exhibiting posi- tive Viable Seedling Development after emergence of the radicle and a mor- ] 738 BRIT.ORG/SIDA 22(1) WiahIAc 1: n | (MICH example, in column 8 well # (in d di ler) 2,6-10 all contain positive VSD individuals phologically normal set of cotyledons that persisted for at least one week post- emergence. All plates were incubated for 14 days. Preliminary experiments.— Two preliminary experiments were to address several experimental parameters. To determine if the damaged seeds were capable of either germination or forming normal seedlings, both necrotic aiid herbivore-damaged seeds sampled from 10 plants from each of the six popu- lations (n=379) were incubated in germination medium for 14 days. In the sec- ond experiment the effect of cold-storage on seed germination was evaluated with a sample of phenotypically normal seeds (5 populations, 24 seeds/popu- lation) incubated in germination medium. Due to an insufficient number of normal seeds, data from population #12 were not included. Intraspecific comparisons.—Out first experiment of this type involved the incubation of a random sample of seeds regardless of physical condition (n=1,440) from each of six populations in culture medium, to determine the level of viable seedling development (VSD) from a general sample. The second ondiicted Cd CALIE ET AL., SEED INVIABILITY IN SOLIDAGO SHORTII 739 analysis was conducted to determine the actual levels of normal vs. damaged seeds in a sample (n=1,199) from all six populations. For each population, 10 plants were sampled, 20 seeds per plant. For population #2, the seed sample size was n = 199. To address the issue of the level of seed damage due to either herbivory or necrosis, seeds from five populations were examined for physical condition (n=2,431). Ten plants from each population were sampled, typically 50 seeds per plant, the number of seeds varying in some cases due to availabil- ity. Due to an insufficient number of seeds, data from population #8 were not included. For the statistical analysis, sample size was reduced to 9 plants per population, due to the necessity of having equal sample size per population (all sampled populations had at least 9 plants with 50 seeds/plant, but not all popu- lations had 10 plants with 50 seeds/plant). Interspecific comparisons.—To address the third question mature infructescences were collected in mid-December 2003 in the Blue Licks vicin- ity from 10 plants of S. shortii and 10 plants of S. canadensis var. altissima from population #5, and from 10 plants of S. shortii, 5 plants of S. nemoralis and 5 plants of S. canadensis var. altissima from population #1. Seeds were collected from 10 plants of the Indiana population of Short’s goldenrod and from 5 neigh- boring plants of S. gigantea and S. nemoralis each in mid-December 2004. For each examined plant, 20 seeds were sampled. Seeds from this final phase of the investigation were examined for their physical condition alone. For the Ken- tucky populations of S. shortii data from two different collecting seasons (2001 and 2003) are pooled as no statistically significant difference exists between the two seasons (t test; population # 1: t = 1.8, p =0.081; population #2: t = 0.33, p = (0.75). For the S. canadensis population data, two different populations (#1 and #5) are pooled as no statistically significant inter populational difference exists (t test, t= 18, p= 0.12). Statistical Analyses.—A|l statistical analyses were conducted using the software program SYSTAT version 11 (SYSTAT Software, Inc.). To test hypoth- eses of no statistically significant difference in values of the mean between or among groups, we used the t-test and analysis of variance (ANOVA), respec- tively, except in those cases when assumptions of these tests were violated. When the data were not normally distributed and/or group variances were het- erogeneous, we used the Mann-Whitney Test or Kruskall-Wallis Test, which are nonparametric analogs of the t-Test and ANOVA, respectively (Zar, 1999). RESULTS Preliminary experiments.—In the first experiment in which damaged and ne- crotic seeds were incubated in germination medium, all were inviable as deter- mined by the absence of either a radicle or a set of cotyledons. The second ex- periment using phenotypically normal seeds was conducted to determine the percentages of VSD using the cold storage treatment prior to incubation in seed 740 BRIT.ORG/SIDA 22(1) germination medium. The VSD values ranged from 71% (population #2) to 96% (population #8) (Table 1. Intraspecific comparisons.—A preliminary visual examination of the Ken- tucky S. shortii seeds revealed two distinct morphological classes. Phenotypi- cally “normal” seeds had a buff to tan color, were ellipsoid in shape (Fig. 2A), and several examined individuals contained fully developed embryos as deter- mined by visual examination. The second class, consisting of physically dam- aged seeds, were of two types. One type exhibited either small holes bored into the seed/fruit coat (Fig. 2B) or a broken end. We term this group as being ex- posed to “herbivory,” since the damage appears to be insect-induced. The sec- ond group, termed “necrotic,” consisted of seeds that were either very dark brown to black, or were of typical color but deflated (Fig. 2C). An examination of sev- eral seeds of the former type revealed a shrunken, black mass assumed to be the remnants of an embryo. Our first experiment of this type involved the incubation of a random sample of seeds regardless of physical condition (n=1,440) from each of six populations in culture medium, to determine the level of viable seedling devel- opment (VSD) from a general sample (Table 2). The levels of VSD were rather low ranging from 3% for population #12 to 33% for population #8. The differ- ences in levels of VSD between populations are not significant (ANOVA, F = 2.0, p= 0.097). To determine the actual levels of normal vs. damaged seeds, a second sample of seeds (n=1,200) from all six populations was examined (Table 3). The fre- quency of normal seeds in a population sample ranged from 1% (population #12) to 29% (population #1). The next issue addressed was the level of seeds damaged by herbivory vs. necrotic seeds (Table 4; n=2,431). Five populations were sampled (all except population #8, due to an inadequate amount of seeds). The level of seeds dam- aged by herbivory ranged from 10% (population #5) to 38% (population #1). The differences between populations in terms of seeds damaged by herbivory was not statistically significantly different (ANOVA, F = L9, p = 0.12). For this analysis it should be noted that sample size was reduced to 9 plants per popu- lation, due to the necessity of having equal sample sizes per population (all populations had at least 9 plants with 50 seeds/plant, but not all populations had 10 plants with 50 seeds/plant). Interspecific comparisons.—Levels of seed herbivory were similar between S.shortii population #5 and S. canadensis var. altissima populations #1 +5 (16% vs. 18%, respectively) (Table 5). However, the levels of necrosis were significantly different—62% for the former, and 19% for the latter (Mann-Whitney Test, U=140, p<0.0001). Temporal differences for inviable seed production (combining the necrotic and physically damaged groups) between the 2001 and 2003 collec- tions (Tables 3 and 5) were not significant for either population #1 (t = 18; = CALIE ET AL., SEED INVIABILITY IN SOLIDAGO SHORTII 741 TABLE 1.Frequency of VSD (Viable fee Pere comer’ from encheyeley normal seeds sub- ject to cold storage as determined by gen a radicle and a complete set of cotyledons in a sample of phenotypically normal can Pop'n %VSD CON NN > > 8 Ss p = 0.081) or population #5 (t = 0.33; p = 0.75). Furthermore, there were no sig- nificant differences in inviable seed production between populations #1 and #5, as determined by the Mann-Whitney test (U = 130, p= 0.064). In comparisons of S. shortii with the local congeners in 2003, the percentage of normal seeds was much greater in the more common species (44% and 73% for S. canadensis var. altissima for populations #1 and #5, respectively, and 57% for S. nemoralis from population #1) (Table 5). Among the damaged seed sets, there was again significant S. shortii interpopulation variation for 2003: 36% of the total seed sample from population #1 being necrotic but 62% of the population #5 sample exhibiting similar necrosis. The single Indiana population of Short’s goldenrod exhibited a very high level of seed necrosis (74%) in comparison to the Kentucky populations (#1 - 36%, #5 - 62%). S. gigantea also had a rate of necrosis (41%) higher than that of the Kentucky congeners S. canadensis var. altissima (#1 - 21%, #5 - 9%) and S. nemoralis (14%). The percentage of normal Indiana seeds for S. shortii (25%) was higher than the overall mean among all six Kentucky populations (16%; Table 3). In comparing either of the Kentucky S. shortii populations (#1 and #5) with the Indiana S. shortii population, there was again no significant dif- ference in inviable seed production (Kruskal-Wallis Test statistic = 3.7; p = 0.16). There were no differences in inviable seed production between S. shortii and S. canadensis occurring syntopically in Kentucky for the comparison in- volving population #1 (t-Test, t = 0.14; p = 0.89) but yes for population #5 (t= 5.3; p < 0.001) (Table 6). Finally, there were no significant differences in inviable seed production between the Indiana population of S. shortii and the syntopic congeners S. gigantea and S. nemoralis (Kruskal-Wallis Test statistic = 2.6; p = 0.2 The final comparison involved the levels of seed herbivory and necrosis among the three populations of S. shortii. The levels of herbivory did signifi- cantly differ between Kentucky population #5 and the Indiana population (Mann-Whitney Test: U = 85; p = 0.005), but not for population #1 vs. #5 (U = 31; p = 0.14) and population #1 vs. the Indiana population (U = 68; p = 0.12). For BRIT.ORG/SIDA 22(1 CALIE ET AL., SEED INVIABILITY IN SOLIDAGO SHORTII 743 Fic. 2. Photomicrographs of the different seed conditions. A = phenotypically normal seeds; B ds subjected to 7 f \ and the general “shrunken” appearance). The bar represents a length of 1 mm. necrosis, populations #1 vs. #5 (U = 21; p = 0.025) and population #1 vs. the Indiana population (U = 19, p = 0.021) showed statistically significant differ- ences. The difference between population #5 and the Indiana population was not statistically significant (U = 28; p = 0.087). DISCUSSION A number of embryo-lethal and mutant developmental alleles have been cata- logued (e.g., Meinke 1994), and their existence in the homozygous state at a suf- ficient frequency within a population could have extreme consequences for the long-term persistence of that population. For that reason, in our tests for seed- ling viability, we measured the production of developmentally normal seed- lings. In past studies of seed germination (see previously cited Buchele et al. and Walck et al.) the primary criterion for determining seed germination was emergence of the radicle after two weeks of incubation, as this is the definitive physiological indicator for completion of the germination process (Bewley 1997). The germination data in these prior studies was then transformed with data 744 BRIT.ORG/SIDA 22(1) Taste 2. Levels of Viable Seed Development (VSD) from a random sample of seeds obtained from six populations of Short's goldenrod. All samples consisted of orale normal as well as physically damaged and necrotic seeds. ' = data from Smith et a Pop’n %VSD Pop’n size! Density! ] 30% 2,549 0.842 2 21% 573 0.419 5 25% 3,488 0.648 7 14% 1,000 1.91 8 33% 672 0.877 12 3% 1,846 0.378 Taste 3.Proportion of seeds from a random sample that were either phenotypically normal or dam- aged (either due to herbivory or necrosis). Pop'n % normal % damaged ] 29% 71% 2 14% 86% 5 28% 72% 7 8% 92% 8 14% 86% 12 1% 99% Taste 4. Proportion of seed damage due to either external (herbivory) or internal (embryo abor- tion?) factors in a sample of damaged seeds from six populations of Short’s goldenrod.. Damaged = seeds exhibiting a small hole in the coat, or a broken end; necrotic = seeds with a dark brown or black color,and a twisted, shrunken shape. Pop’n (sample size) % damaged % necrotic | (464) 38% 62% 2 (487) 30% 64% 5 (490) 10% 90% 7 (500) 20% 80% 12 (490) 12% 88% obtained from seeds that had not yet undergone germination, but that contained respiring embryos as determined by a histochemical stain (Cottrell 1947). Our criteria differ from the criteria previously used to determine seed germination, as we posed a different question. Our interest in determining the frequency of developmentally normal seed- lings produced from each sample of seeds was in obtaining an estimate of com- parative reproductive potential. Those populations producing a higher fre- quency of developmentally normal seedlings would have a greater potential for persistence than a population with-a low level of normal seedlings. Our CALIE ET AL., SEED INVIABILITY IN SOLIDAGO SHORTII 745 TABLE zp ECU SINO! He renotypicall normal vs.damaged seeds in two populati f Short's gold- ntaining gener species. Nor = phenotypically normal; eget cece exhibiting damage die to insect Karbon) Nee = seeds with a darkened and physically distorted evcial appear- ance. Populations #1 and 5 are from Kentucky; IN = the Indiana population. The parenthetical nu- merals refer to the number of plants sampled. Pop’n Taxon Seed condition Nor er Nec 1 S. shortii (10) 54% 10% 36% 1 S.canadensis var. altissima (5) 44% 35% 21% ] S.nemoralis (5) 57% 29% 14% 5 S. shortii (10) 22% 16% 62% 5) S.canadensis var. altissima (10) 72% 9% 19% IN S. shortii (10) 25% 1% 74% IN S. gigantea (5) 18% 41% 41% IN §.nemoralis (5) 43% 45% 12% Taste 6. Summary statistics (mean + 1 SD and range) for the number of inviable (necrotic + dam- aged) seeds from a single sample per plant for Kentucky (#1 and #5) and Indiana (IN) 5. shortii and Kentucky (#1+#5) S. canadensis populations. Pop'n Taxon Mean Range ] S. shortii 12-263 3-20 5 S. shortii 15+ 40 8-20 IN S. shortii 15 241 5-19 145 S. canadensis LAE 6.0 0-20 specific question was not one of seed germination rates among and between populations, but one of the frequency of normal seedling production. We em- ployed incubation conditions that would, based on previous studies (e.g. Walck et al. 1997a) optimize seed germination levels, then examined seedlings for nor- mal early development. During the course of this study, of considerable surprise to us were two dis- coveries: the frequencies of seed predation (herbivory) and necrotic (presum- ably embryo aborted) seeds. All seven sampled populations experienced marked levels of either mechanically damaged seeds or necrotic seeds. The number of affected seeds (either damaged or necrotic) in our samples ranged from 70% (population #1) to 99% (#12). Of the damaged seed sets, the majority were ne- crotic, ranging from 63% (population #1) to 90% (#5). Seed damage in popula- tions | and 7 was previously noted “in some infructescences” and attributed to larvae of the family Cecidomyiidae (Diptera) (Buchele et al. 1992). There were no quantitative data in this earlier report. Another possible candidate is bruchid beetle larvae (R. Kingsolver, pers. comm.). 746 BRIT.ORG/SIDA 22(1) There is no previous mention in the literature concerning Solidago regard- ing the occurrence of necrotic seeds in mature infructescences. The major threat to successtul formation of developmentally normal seedlings appears not to be an external agent, e.g.an insect depositing eggs in S. shortii seeds asa food source for developing larvae, but rather the frequency of embryo-aborted seeds. There could be three possible explanations for the phenomenon of embryo abortion. First, due to the small sizes of the populations (Smith et al. 2004) it is possible that through stochastic events lethal allele combinations have arisen (Levin 2000) possibly leading to embryo abortion. Second, there could be pollen com- petition among Short’s goldenrod plants and the more common goldenrod spe- cies. There is a temporal overlap in anthesis between the three goldenrod spe- cies at Blue Licks, and we have observed individual insects visiting inflorescences of all three goldenrod species. Likewise plants of the Indiana population of Short’s goldenrod could have been pollinated with pollen from either S. nemoralis or S. canadensis, leading in both cases to abortive embryo formation due to gametic incompatibility (Grant 1971). Third, as Solidago is an obligate outcrosser (Buchele et al. 1992) lack of suitable crosspollination would lead to unfertilized embryo sacs and arrested development of the resultant seeds (Proctor et al. 1996). At present there are no data available to distinguish among these three possibilities. Itis difficult to assess the impact of reduced seed production on the persis- tence of S. shortii, given that three more widespread congener species (S. canadensis var. altissima, S. nemoralis and S. gigantea) are likewise subject to seed predation. The presence of congener species with a lower level of necrotic seeds (and assumedly a pre-predation higher level of normal seeds) does not appear to influence the behavior of the seed predators on Short’s goldenrod, as there was no statistically significant difference between populations that con- tained (#1) or were adjacent to (#5 and the Indiana population) congener spe- cies and those without. In the absence of interspecific competition, Walck et al. (1999a) observed successful establishment of Short’s goldenrod seedlings in field plots over a three-year period. Given the large number of florets in a typical Short’s goldenrod inflorescence, there evidently are a sufficient number of vi- able seeds generated to provide for some level of sexual reproduction. — ACKNOWLEDGMENTS The able field and laboratory assistance of Ms. Tiffany Carpenter and Ms. Jackie Hucul are gratefully appreciated. Thanks are extended to Ms. Deborah White of the Kentucky State Nature Preserves Commission and the Kentucky State Parks Commission for permission to collect plant materials. Funding was pro- vided to PJC through an EKU Faculty Research Award, and from NIH/NCRR/ KBRIN Award #1-P20 RR 16481. The review and comments of Dr. Bruce Ford on an early version of this manuscript are gratefully acknowledged. CALIE ET AL., SEED INVIABILITY IN SOLIDAGO SHORTII 747 REFERENCES Baskin, J.M.,J.L.Watck, C.C. Baskin, and D.E. Buchete. 2000. Ecology and conservation biology of the endangered plant species Solidago shortii (Asteraceae). Native Pl. J. 1:35-41. Bewtey, J.D. 1997. Seed germination and dormancy. Pl. Cell 9:1055-1066. BucHete, D.E., J.M. Baskin, and C.C. Baskin. 1989. Ecology of the endangered species Solidago shortii. |. Geography, populations, and physical habitat. Bull. Torrey Bot. Club 116:344- Buchete, D.E.,J.M. Baskin, and C.C. Baskin. 1991.Ecology of the endangered species Solidago shortii. Ill. Seed germination ecology. Bull. Torrey Bot.Club 118:288-291 Buchete, D.E., J.M. Baskin, and C.C. Baskin. 1992. Ecology of the endangered species Solidago shortii. lV. Pollination ecology. Bull. Torrey Bot. Club 119:137-141. Cortrett, HJ. 1947. Tetrazolium salt as a seed germination indicator. Nature 159:748, Evans, M. 1987. Field survey for Short's goldenrod (Solidago shortii). Report for the USFWS Region 4, Atlanta, GA. Grant, V. 1971. Plant speciation. Columbia Univ., Press, NY. Hickox. L.G. and T. Warne. 1998. C-Fern manual. Carolina Biological Supply Co., Burlington, Homoya, M.A.and D.B. Asrett. 2005.A natural occurrence of the federally endangered Short's goldenrod (Solidago shortii T. & G.) [Asteraceae] in Indiana:lts discovery, habitat, and associated flora. Castanea 70:255-262. Levin, D.A. 2000. The origin, expansion, and demise of plant species. Oxford Univ. Press, New York. Meinke, D.W. 1994. Seed development in Arabidopsis thaliana. In Arabidopsis, C.R. Somer- ville ,E.M. Meyerowitz, Eds. Cold Spring Harbor Press, NY. Smith, B.D., A.J. DenHAM, J.B. Beck, and PJ. Catt. 2004. High resolution GIS mapping and cur- rent status of the ten viable populations of Short’s goldenrod (Solidago shortii — Asteraceae) in Kentucky. Sida 21:1121-1130. Proctor, M., P. Yeo, and A. Lack. 1996. The natural history of pollination. Timber Press, Port- land, OR. Watck, J.L., JM. Baskin, and C.C. Baskin. 1997a. A comparative study of the seed germination biology of a narrow endemic and two geographically-widespread species of Solidago (Asteraceae). 1. Germination phenology and effect of cold stratification on germina- tion. | Sci. Res. 7:47-58 Watck, J.L., J.M. Baskin, and C.C. Baskin. 1997b.A comparative study of the seed germination aS of anarrow endemic and two g phically-widespread species of Solidago (Asteraceae). 2. Germination responses to Ponca seeds in relation to seasona! tem- perature cycles. Seed Sci. Res. 7:209-220. Watck, J.L., JM. Baskin, and C.C. Baskin. 1997¢.A compalallve study of the seed germination Biology of a narrow endemic and two geographically-widespread species of Solidago (Asteraceae). 3. Photoecology of germination. Seed Sci. Res. 7:293-301. Watck, J.L., J.M. Baskin, and C.C. Baskin. 1997d. A comparative study of the seed germination — ee) 748 BRIT.ORG/SIDA 22(1) It biology of anarrow endemic and two geog ally-widespread species of Solidago (Asteraceae). 4. Role of soil moisture in regulating germination. Seed Sci. Res. 7:303- Watck, J.L., J.M. Baskin, and C.C. Baskin. 1997e.A comparative study of the seed germination biology of a narrow endemic and two g hically-widespread species of Solidago (Asteraceae). 5. Effect of dry storage on after-ripening and survivorship. Seed Sci. Res. 7:311-318. Watck, J.L., J.M. Baskin, and C.C. Baskin. 1997f. A comparative study of the seed germination biology of a narrow endemic and two geographically-widespread species of Solidago Asteraceae). 6. Seed bank. Seed Sci. Res. 8:65—74. Watck, J.L.,J.M. Baskin, and C.C. Baskin. 1999a. Effects of competition from introduced plants on the establishment, survival, growth and reproduction of the rare plant Solidago shortii (Asteraceae). Biol. Conserv. 88:213-219. Watck, J.L., JM. Baskin, and C.C. Baskin. 1999b. Relative competitive abilities and growth characteristics of a narrowly endemic and a geographically widespread Solidago spe- cies (Asteraceae). Amer. J. Bot. 86:820-828. Wack, J.L., J.M. Baskin, and C.C. Baskin. 1999c. Ecology of the endangered species Solidago shortii.VI. Effects of habitat type, leaf litter, and soil type on seed germination. J. Torrey Bot. Soc. 126:117-123. Zar, J.H. 1999. Biostatistical analysis, 4" ed. Prentice Hall, Upper Saddle River, NJ. — NATURALIZATION AND EXTIRPATION OF WATER HYACINTH (EICHHORNIA CRASSIPES, PONTEDERIACEAE) IN SOUTHWESTERN ARKANSAS, U.S.A. Renn Tumlison and Brett Serviss Department of Biology Henderson State University Arkadelphia, Arkansas 71999, U.S.A, ABSTRACT Southern a pcaae to be at he edge of the eee range for naturalization of the water hyacinth, Ei Ims, due to the cold intolerance exhibited by the plant. A population discovered eonine in DeGray Lake, a County, Arkansas, was examined from 2002- 2003 to determine whether the population could survive after the freezes common during winter. The population was of particular interest because it had become rooted and remained submerged with higher lake levels during the surface freezes of winter. Rosettes that fragmented from the popu- lation established new growth during the next growing season, but fluctuating lake levels eventu- ally stranded this growth and caused extirpation of the population RESUMEN Arkansas meridional parece estar en laf lel ial para | lizacion del jacinto H 4 ° ] y BES Rae | por la planta. el Lago DeGray, Condado de Clark, Arkansas, se examino de agua, Fichhornia i (M Sol de aan intolerancia Una poblacion se descubrié iend entre 2002-2003 ae ane si ls poniacon podria sobrevivir después de las heladas comunes ] durante el invierno. La } poe habia llegado a arraigar y quedaba d ae 4 lr sumergida con ie Rosetas que se fragmentaron de la poblacion tuvieron crecimiento nuevo en la siguiente estacion de de lago cme la la superficie en invierno. crecimiento, pero los niveles fluctuantes del lago desfavorecieron finalmente este crecimiento y causaron la extincion de la poblacién. INTRODUCTION Eichhornia crassipes (Martius) Solms (water hyacinth) is one of two natural- ized species of Fichhornia found in the United States (Horn 2002; Wunderlin 1998). Water hyacinth is native to tropical South America, specifically Amazonia (Barrett & Forno 1982). Water hyacinth now occurs in over 50 coun- tries across five continents (Barrett 1989). It is believed that Eichhornia was in- troduced into the United States at a Centennial Exposition at New Orleans in 1884 (Penfund & Earle 1948). It is one of the most aggressive aquatic weeds in the tropics, doubling its population as quickly as every 11-18 days (Penfund & Earle 1948). Further, Eichhornia is known to produce physiological changes to the aquatic environment (Center & Spencer 1981; Fitzsimmons & Velljos 1986; Penfund & Earle 1948; Ultsch 1973). Seed production in temperate regions is SIDA 22(1): 749 — 754. 2006 750 BRIT.ORG/SIDA 22(1) usually only about one-half of that in the tropics, primarily because of differ- ences in levels of insect visitation (Barrett 1980). Fichhornia crassipes is only marginally cold hardy and is completely de- stroyed when the rhizome is exposed to temperatures at or below -5°C for 12 or more hours (Penfund & Earle 1948). Water hyacinth should not reliably establish in areas where these temperature conditions occur annually. In southwestern Arkansas, air temperatures reach this level for several days during most winters, which likely would limit the yearly regrowth of populations of Eichhornia. However, a population of Eichhornia crassipes was discovered in a cove of DeGray Lake, Clark County, Arkansas on 28 January 2002. Living rosettes with broken stolons were discovered along the north shoreline, and a search to re- veal the source of these usually floating plants led to the discovery of the popu- lation submerged in the lake waters. We followed the fate of this population, to determine whether the submerged condition would allow survival through the winter due to protection from freezing. lt was presumed that the typically clear water would allow adequate photosynthesis to carry the population until the next growing season. — METHODS The source of the original established population was not known. Upon fur- ther search, the only locality found supporting plants was the initial discovery along the north shore of the cove. Prevailing winds from the south presumably pushed floating rosettes into the north shore of the cove and effected establish- ment at the site. We assumed that the population was established during the previous summer. Numerous old inflorescences were discovered still attached to the rosettes of the population. We returned to the site on several occasions during 2002 to determine the fate of the population. Water temperature (surface and at one meter) and depth of the population was measured when possible. During 2002, several rosettes collected from the field site were grown in the lab and in a small plastic outdoor pond to determine whether the plants would flower and produce viable seed. With the onset of winter 2003, a few of these rosettes were planted in pots and submerged to determine whether such overwintering plants would produce break-away rosettes that would establish a new population during spring. RESULTS The submerged lake population on 28 January 2002 had rosettes of 15-20 cm width. The water depth at the location of the deepest specimens (the water in the cove was perpetually clear, allowing us to see the plants) was 90 and 107 cm, but most rosettes were located at a depth of 80-85 cm. The widest portion of the population (about | m wide) was located toward the head of the cove, 751 where the slope of the substrate was more gradual and the water was shallow, but most of the population occurred in a strip about 1/2 m wide by 80 m long along the contour of the shoreline. Samples of the rosettes showed old inflores- cences. Plants toward the head of the cove exhibited morphology more typical of floating rosettes, but most other specimens had small floats and an upright elongate central or main stem. Temperature at the surface on 28 January was 12°C, and at one meter was 10°C. On 15 February, the population looked as it had previously, but the lake had risen about 25 cm so that most plants were at a depth of 105-110 cm. Sedi- ments occasionally produced by wave action were deposited as a film over the population, although sampled plants still were green. Many plants had become separated from the submerged mat and were floating near shore. Temperature was 9°C at the surface and 8°C at one meter. On 24 April, water depth had increased again, making the rosettes impos- sible to see. Several rosettes that had separated and floated now were rooting into the sediments along the elevated shoreline, and 10 of these were marked to follow their survival. Several green rosettes were trapped among debris at the head of the cove, protected from waves created by boaters. Also, eight rosettes were found in the last 15 m of the cove along the southern shoreline. Tempera- ture was 23°C at the surface and 21°C at one meter. By 9 May, water depth had increased again, placing the population at an estimated 1.5-2 m depth. The plants rooted into the sediments and marked dur- ing the previous trip now were submerged and not relocated. Along the shore- line, we counted about 40 rosettes that recently had broken free of the original population, many at the head of the cove. One of these was partially buried in the wet sandy soil about 1.5 m from water. Most of these stranded rosettes were small (3-10 cm width), but had new growth and new roots were evident. Tem- perature was 23°C both at the surface and at one meter. On 12 June, the highest water level we observed kept the original popula- tion, if still alive, submerged beyond visibility. Wind action had placed all float- ing rosettes (the 40 observed on 9 May) along the shore near the head of the cove. These plants now were larger (8-20 cm width), had new growth, and in- cluded clusters with side branches. The specimen partially buried in the sand on the previous date was not relocated, and all plants found were floating with the exception of a few stranded at the head of the cove by wave action. Tem- perature was 31°C at the surface and 29°C at one meter. On 18 July, depth of the lake had decreased about 3/4 mand the water was very clear, but the original population of Eichhornia had disappeared. Only 18 rosettes of the 40 were surviving. These were found in 8 clusters all near the head of the cove in muddy to sandy areas covered by organic debris and in almost constant shade. The roots of these plants had become anchored in sedi- 752 BRIT.ORG/SIDA 22(1 = ments, and now had rosettes up to 30 cm in diameter and side branches with upright leaves with small bladders. These plants were in wet soil located 5-9 m from the water of the lake. Temperature was 29.5°C at the surface and 29°C at one meter. On 9 August, the few remaining exposed plants were drying and dying back, with the healthiest three clusters located 5, 5, and 22 m from the lake in the sandy soil of the head of the cove. Rains fell before 6 September, which appar- ently rejuvenated the plants and permitted new growth of leaves. Additional rain fell before a 20 September visit, and had resulted in continued new growth. On 7 January 2003, the site was revisited with the expectation that the frost of 25 November would have killed the remaining plants that had been isolated in exposed locations along the cove. Of the last 18 rosettes, only two were relo- cated - dead and submerged under 20 cm of water. In the small pond experiment, the rosettes proliferated and filled the pond during the summer, and produced numerous inflorescences from 6 August through September. However, all surface plants were killed with the first frost. The frost killed only exposed portions of 5 otherwise submerged plants. The re- maining portions of these 5 plants remained green until early spring, then died. No new rosettes were able to break away, surface, and proliferate during the next spring. Seeds left in the pond and those used in greenhouse experiments failed to germinate within one year, and no germination of seeds has been discovered at the field location. It is important to note that the seeds used in our experiments were not intentionally scarified to promote laboratory germination. DISCUSSION A population of Eichhornia crassipes became established in DeGray Lake, at a northern latitude presumably marginal for this cold-intolerant plant to over- winter outdoors. The original established population survived through periods of freezing because the root system anchored the plants, not allowing them to float as the level of water in the lake rose during winter. Although the original population did not survive the entire winter, rosettes that periodically broke free and floated survived if they surfaced after the last freeze. The higher level of water during spring deposited these surviving rosettes higher up the bank than the original population. Some survived and grew dur- ing the next season, but they became exposed and isolated from the lake as water levels dropped during summer. Because none of the originally submerged plants were observed after 18 July, none was present for future re-establishment. From the available data, we believe that the original population was estab- lished during the summer of 2001 when the lake level was near its seasonal low. Plants grew along the shoreline, held there by the winds, and became an- chored by their roots. Any plants that grew across the deeper water of the cove, and were not held by the roots as the water level rose, would have died with freezing air temperatures. This scenario would have left the observed 80 m long, 0.5-1 m wide population that followed the contour of the shoreline. This por- tion of the original population actually survived the winter by being submerged, and re-established and grew the following spring. However, fluctuating lake levels led to the extirpation of the population that had established. Penfund and Earle (1948) unexpectedly observed that Eichhornia crassipes plants died while submerged during the winter. They felt that protection from the freezing sur ures would protect the plants, but argued that a lack of oxygen might have explained the death of their submerged population. Be- cause even clear water filters light, we argue that insufficient light to the sub- merged plants might have precluded adequate photosynthesis. Further, water saturates with much less oxygen and carbon dioxide than is available in air, which likely contributed to the observed death of the rooted population in our study. Eichhornia crassipes has a means of survival when a rooted population becomes inundated. Penfund and Earle (1948) found that 10 days after submer- gence, E. crassipes begins to produce an abscission layer across the rhizome just below the lowest living leaves, forms new roots just above the abscission layer, and floats to the surface. Those new rosettes could continue the existence of the population if surface conditions were within the range of tolerance. Seedlings of E. crassipes can survive submergence via the same mechanism (Penfund & Earle 1948; Robertson & Thein 1932). Seeds of E. crassipes only ger- minate when exposed to air, and thus seeds that germinate on areas where wa- ter has receded could become rooted and attached into the soil substrate, and subsequently submerged if the water level of the lake rises. We did not observe seedling development in our field or lab studies, so we found no evidence that re-establishment would occur by those means at our site. We believe that this species would be able to survive at our site if the sea- sonal high and low water levels were less different, or if the relative timing of lake fluctuation and f afforded protection. Thus, submerged plants would have a greater chance of surviving longer into the winter and spring, and sepa- rating rosettes would not meet high water and be placed too far away from the lake to undergo the rapid growth possible during summer. The present condi- tions appear to help prevent the long-term establishment of this exotic plant at this site. ACKNOWLEDGMENTS We thank two anonymous reviewers for their comments and suggestions. REFERENCES Barrett, S. 1989. Waterweed invasions. Sci. Amer. 261:90-97. Barrett, S.C. 1980. Sexual reproduction in Eichhornia crassipes (water hyacinth) ||. Seed production in natural populations. J. Appl. Ecol. 17:113-124. BRIT.ORG/SIDA 22(1) Barrett, S.C. and |W. Forno. 1982. Style morph distribution in New World populations of Eichhornia crassipes (Mart.) Solms—Laubach (water hyacinth). Aquatic Bot. 13:299-306. Center, T.D.and N.R. Spencer. 1981.The phenology and growth of water hyacinth Eichhornia crassipes (Mart.) Solms in a eutrophic north-central Florida lake. Aquatic Bot. 10:1-32. Fitzsimons, R.E.and R.H. Vattesos. 1986.Growth of water hyacinth [Eichhornia crassipes (Mart.) Solms] in the middle Parana River (Argentina). Hydrobiologia. 131:257-260. Horn, C.N. 2002. Pontederiaceae.|n: Flora of North America Association, eds. 2002. Flora of North America. Volume 26. Oxford University Press, New York, NY. Penrunb, W.T. and T.T. Earte. 1948. The biology of the water hyacinth. Ecol. Monogr. 18: 448-472. RoserTsON, H.F.and B.A. THeIn. 1932. The occurrence of water hyacinth (Eichhornia crassipes) seedling under natural conditions in Burma. Agric. and Livestock of India. 2:383-390. Uttsch, G.R. 1973. The effects of water hyacinths (Fichhornia crassipes) on the microenvi- ronment of aquatic environments. Arch. Hydrobiol. 72:460-473. WuNDERLN, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida. Gainesville. FUNGI ASSOCIATED WITH BORRICHIA FRUTESCENS (ASTERACEAE): INSECT GALLS AND ENDOPHYTES Diane Te Strake, Amy Haddock Keagy, and Peter D. Stiling Department of Biology University of South Florida Tampa, Florida 33620-5200, U.S.A. ABSTRACT This report compares fungi found in galled and non-galled plants of Borrichia frutescens (L.) DC, Asteraceae. Fungi were observed in the plant galls of the midge, Asphondylia borrichiae Rossi & Strong. A variety o endophytes unely were oe me the apic cal meristems, stems and leaves of nels eda nd non-g it Florida. Fifteen percent of midges r U | morphological and distributional information is conveyed in the keys. All taxa are a cally arranged, without authorities and publication details; common names are provided but scien- tific synonymy is minimal. T dated from predeces- sors, but concepts of families and genera remain conservative. Fore ne species coverage and nomenclature is much up cample in the sterace —~ cae groups of similar species are keyed in conglomerates (Bal hiza-Wyethia complex, Aste Machaeranthera complex, Ericameria-Pyrrocoma- Columbiadori id-Hazardia complex, ies. li \ l | lex). The user friendly keys, the Euthamia complex, Madia-Ani SOCUFPUus Hemizonella heart and soul of this volume, appear to be effective and Kozloft’s Weak undoubtedly will quickly find it nto wide use.—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, ee ee U.S.A SIDA 22(1): 764. 2006 TYPE LOCALITIES OF VASCULAR PLANTS FIRST DESCRIBED FROM OHIO: SUPPLEMENT! James S. Pringle R yal B tanical Gardens PO. Box 399. Hamilton Ontario, CANADA L8N 3H8 Jpringle@rbg.ca In 2003 Ronald L. Stuckey and I published a list of Ohio localities at which the type specimens of the names of vascular plants had been obtained, including the plant names based on specimens from each locality. Omissions have be- come apparent, hence this supplement. As in the original list, double asterisks denote names that are currently accepted for the respective taxa. Otherwise, the currently accepted names are given in brackets. Minor variants not other- wise accounted for are now generally not recognized taxonomically, but remain included within the same species. Further explanation of the format appears in our earlier paper. OHIO Carex xanthocarpa E.PBicknell, Bull. Torrey Bot. Club 23:22. 1896, not Degl. 1807: C. vulpinoidea var. xanthocarpa Kuk., Pflanzenr. (Engler) 4(20, heft 38):148. 1909, based on C. xanthocarpa E.PBicknell; C. bicknellii E.G.Camus, Notul. Syst. (Paris) 1:239. 1910 and C. brachyglossa Mack., Bull. Torrey Bot. Club 50:355. 1923, both published in substitution for C. xanthocarpa E.PBicknell [Carex annectens (E.PBicknell) E.PBicknell] Laciniaria shortii Alexander ex Small, Man. S.E. Fl. 1335.1933|Liatris squarrulosa Michx]]. Vernonia maxima Small, Bull. Torrey Bot. Club 27:280. 1900 [Vernonia gigantea (Walt.) Trel. var. gigantea. Maumee RIVER Baptisia leucantha Torr. & A. Gray, Fl. N. Amer. 1:385. 1840 [Baptisia alba var. macrophylla (Larisey) Isely]. Miami River, Miami Country Geum franckii [sic] Steud., Nomencl. Bot., ed. 2, 1:683. 1840, not Sm. 1810: pub- lished in substitution for Geum parviflorum Hochst. & Steud., that name not validly published [Geum vernum (Raf.) Torr. & A. Gray], R ‘Contribution No. 128 from the Royal Botanical lens, Hamilton, Ontario, Canada, SIDA 22(1): 765 - 767. 2006 766 BRIT.ORG/SIDA 22(1) FRANKLIN COUNTY Columbus. Trillium erectum var. declinatum A. Gray, Man., ed 5, 523. 1867, T. declinatum (A. Gray) Gleason, Bull. Torrey Bot. Club 33:389. 1906, not Raf. 1840, and T. gleasonii Fernald, Rhodora 34:21. 1932, published in substitution for T. declinatum (A. Gray) Gleason [Trillium flexipes Raf]. HAMILTON COUNTY Cincinnati. Aira controversa Steud, Syn. Fl. Glumac. 1:224. 1854 [Sphenopholis intermedia (Rydb.) Rydbl]. Crataegus coccinea [var] e? [sic] mollis Torr. & A. Gray, Fl. N. Amer. 1:465. 1840 [Crataegus coccinea L., not Crataegus mollis Scheele]. Lippia nodiflora var. acutifolia Kuntze, Revis. Gen. Pl. 2:508.1891[Phyla nodiflora (L.) Greene; often retained in Lippial. Stachys cincinnatensis Kuntze, Revis. Gen. Pl. 2:521. 1891 [Stachys tenuifolia Willd! Volvulus sepium var. biangulo-sagittatus [sic] Kuntze, Revis. Gen. Pl. 2:447. 1891 [Calystegia sepium (L.) R.Br. subsp. angulata Brummitt]. Cincinnati, Lawrence Bay. Sida spinosa var. flava Kuntze, Revis. Gen. Pl. 1:73. 1891. Lawrenceburg. Iriha [sic] autumnalis var. caespitosa Kuntze, Rev. 2:752. 1891 [Fimbristylis autumnalis (L.) Roem. & Schult]. HockinG County Cedar Falls. **Vittaria appalachiana Farrar & Mickel, Amer. Fern J. 81:69. 1991. LoraAIN COUNTY North Amherst. Geranium carolinianum var. confertiflorum Fernald, Rhodora 37:300. 1935 [Geranium carolinianum L. var. carolinianuml. Lucas County Alexis [interpreted as the railroad junction in present-day Toledo rather than the community in Monroe County]. Cardamine douglasii forma albidula Farw,, Amer. Midl. Nat. 9:261. 1925. Toledo, Maumee River bank, Walbridge Park. Malus coronaria var. dasycalyx Rehd., J. Arnold Arbor. 2:52. 1920. Meics County Slope above Leading Creek, Co. Rt. 10, 0.25 mi SW of Twp. Rt. 27, N of Dexter, Sect. 6, Salem Twp. Polypodium x incognitum Cusick, Amer. Fern J.92:241. 2002 [= Polypodium appalachianum Hauller & Windham ~ P.virginianum Ll. RICHLAND COUNTY Mansfield. Convolvulus sepium var. communis R.M. Tryon, Rhodora 41:419. 1939 [Calystegia sepium (L.) R.Br. subsp. sepium. PRINGLE OHIO PLANTS 767 Scioto COUNTY Friendship, Shawnee State Forest, Camp Gordon C.C.C. Disporum schaffneri Moldenke, Phytologia 1:167. 1935 [Prosartes maculata (Buckley) A. Gray]. In addition, the names *Oxalis grandis Small, based on O. recurva Elliott sensu Trelease, not as originally applied by Elliott, and **Sanicula trifoliata E.PBicknell were originally based on syntypes that included specimens respec- tively from “Ohio” and “Franklin County, Ohio.” Elymus canadensis var. inter- media Vasey ex S.Watson @ J.M.Coult. was originally described with no speci- mens cited but with the range said to include Ohio [now included in E. virginicus L. var. virginicus]. For all of these mames later authors have designated speci- mens from other states as the lectotypes. REFERENCE Stuckey, R.L. and J.S. Princte. 2003. Type localities of vascular plants first described from Ohio. Sida 20:1677-1692. 768 BRIT.ORG/SIDA 22(1) Book NOTICE Prrer Gregory. 2000. Plant Roots: Growth Activity and Interaction With Soils. (ISBN 1-4051-1906-3; 978-1-4051-1906-1, hbk.). Blackwell Publishing Pro- fessional, 2121 State Street, Ames, IA 50014-8300, U.S.A. (Orders: www black wellplantsci.com, 800-862-6657, 515-292-3348 fax). $199.99 hbk., 328 pp., illustrations, 7" x 10' As suggested by the cover notes, “this book will be a vital tool for plant, crop, soil, and agricultural I scientists, ecologists, and hydrologists.” It’s expensive, scientists, plant physiologists but perhaps “essential” as well as “vital.” There are nine chapters, densely packed with technical in- formation but easily readable, as follow. . Plants, Roots and the Soil (evolution, roots and shoots, roots and soi Architecture of Root Systems (structure, extension ae branching, root tip, system . Roots and th ees socal men a Growth of Root Systems (measurement of root systems, system development, ng) size, n, longevity and turnover, modelli 4. The Functioning Root System (anchorage, water ae nutrient uptake). 5. Roots and the Physico-chemical Environment (temperature, tropistic responses, soil pores anc mechanical properties, atmospheric CO2). Environment (soil organisms, symbioses, pathogens and parasites, root . Roots and the Biologica herbivory 7. The Rhizosphere (rhizodeposition, chemical changes affecting nutrient acquisition, physical changes in the rhizosphere) 8. Genetic Control of Root System Properties (genotypic differences, genetics, breeding). 9. Root Systems as Management Tools (optimal root systems, intercropping and agroforestry, crop rotations, phytoremediation). —Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A SIDA 22(1): 768. 2006 NOTES ON SOUTHWESTERN MORACEAE Alan T.Whittemore 1/S National Arh retum 1 New York Ave NE ie. DC 20002-1958 U.S.A. ABSTRACT Specimens forming the basis of reports of Ficus palmata (F pseudo-carica) in North America are misidentified and actually belong to F. carica, so F palmata must be excluded from the adventive ‘lora of North America. Morus nigra is present as a rare waif in ee nia and Nevada. It is best distinguished from other Morus species in the area by its deeply cordate leaves and densely pubes- cent styles 3-5 mm long RESUMEN Los especimenes en que se basan las citas de Ficus palmata (E pseudo-carica) en Norte América estan mal identificados y pertenecen a F carica, por lo que E palmata debe excluirse de la flora adventicia de Norte América. Morus nigra esta presente como una planta rara abandonada en California y Ne- vada. Se diferencia de otras especies de Morus en el area por sus hojas profundamente cordadas y estilos densamente pubescentes de 3-5 mm de longitud. Moraceae are primarily trees and shrubs of tropical and subtropical areas, with few herbaceous species and few taxa found in areas with cooler climates. The family is poorly represented in the flora of the southwestern United States: re- cent floras list only two native species, Morus microphylla Buckl. (which oc- curs from central Texas west to Arizona and across northern Mexico) and M. rubra L. (an eastern species that crosses the Great Plains in Texas), and three introduced species, M. alba L., Ficus carica L., and Maclura pomifera (Raf.) Schneid., from the western half of the country (McClintock 1993; Wunderlin 1997). Two other exotic species not mentioned in these floras, Ficus palmata Forrsk. and Morus nigra L., have been recorded as escapes in California and Nevada. In the course of revising McClintock’s treatment for the second edition of the Jepson manual of the higher plants of California, specimens vouchering these reports were examined in order to clarify the status of these species as possible escapes in North America. California reports of Ficus palmata Ficus palmata is a small tree or shrub native from Iran east to India and Nepal, and south to Ethiopia and Somalia. It is closely related to the commercial fig, Ficus carica L., a native of the Near East that is now naturalized at scattered sites across much of the southern United States (Wunderlin 1997), but F palmata differs in having smaller, often unlobed, leaves and smaller fruit (Browicz 1982: Friis 1993; Ghafoor 1985). Ficus palmata (=F. pseudo-carica Miq.) has been re- SIDA 22(1): 769 - 775. 2006 770 BRIT.ORG/SIDA 22(1) ported from southern California, first by Munz (1974). Munz used the newer synonym E pseudo-carica Mig. and described it as similar to F carica, but with leaves more deeply divided, a description that is at odds with descriptions and keys in recent floras of areas where F palmata is native (Browicz 1982; Friis 1993; Ghafoor 1985). Munz listed the species as “Reported as occasional escape in Santa Barbara region,” the phrasing suggesting that Munz picked up his iden- tification from another source. Smith (1976, 1998) mentioned two collections by Henry M. Pollard from “west fork of Cold Springs [sic] Canyon, Montecito.” The species was not treated for California in the most recent state flora (Mc- Clintock 1993) or the Flora of North America (Wunderlin 1997), but it was re- ported again (this time under the older name F palmata) by Hrusa et al. 2002), who cited a single collection from Santa Barbara County, California, made by Henry Pollard in 1958. In view of the irregular treatment of this species (reported by several local floras, but not the most recent state and national floras) and the mismatch be- tween the keys and descriptions of F palmata in the North American and Old World literature, it seemed best to examine the specimens that the California reports are based on and compare them with specimens collected from the na- tive range of E palmata in the Old World, as well as the full range of variation in cultivated F carica. Variation in Ficus palmata and FE. carica Thirty-one specimens from three herbaria (MO, NA, and US), representing Fi- cus palmata from throughout its native range, were examined and coded for morphological characters that potentially distinguish the species from F carica. The results are shown in Table 1, along with characters of cultivated F. carica, taken from 70 sheets of cultivated figs, and characters of the vouchers for the California oS of F. palmata (see below). Characters taken from herbarium s agree well with descriptions and illustrations of the two species from Este sources (Browicz 1982; Condit 1947; Friis 1993; Ghafoor 1985). Representative specimens examined: ses aaa INDIA: Botanical Garden, University of Delhi, Rodin 8049 (US). Setabani, Hem Raj s.n. 28 Feb 1931 (MO). Koraput district, Jaypore Estate, Orissa, 3000-3500 ft, H.E Mooney 3893 (MO). ee a W. Koelz 20496 (NA). Almora, United Prov- inces, 6000 ft, W. Koelz 19996 (NA). PAKISTAN: Peshawar, W. Koelz 8206 (NA); 3 mi from Abbotabad on road to Havelion, M. Qaiser & A. Ghafoor 1999 (NA), Karakar, M. Shah & Dilawar 1001 (MO). NE- PAL: Wada, 6600 ft, Mr Hoh and Rajbhandari 1157 (US). SAUDI ARABIA: Al Karn, Dwyer & El Sheikh 13676 (MO). YEMEN: Attara, 1900 m, Deflers 402 (US). ERITREA. Amasen: Ad Rassi, eis 4925 (US). ETHIOPIA. Kaffa Prov.: 7 km E of oe Burger 1158, Meyer 7820 (US). Harar Proy.: 22 km SE of Harar, ca. 1350 m, Burger Ae 1622 (US). Ficus carica (all cultivated, a f depos at ay CHILE. Juan Fernandez Islands: Mas Afuera, EG. Meyer 9461 IRAN. Tehran, W. Koelz 16085. U.S.A. Arkansas. Drew Co.: Monticello, E. Sundell 91LL California. Alameda Co.: es 23041. Butte Co.: Chico, WF Wight 1279. Riverside Co.: Citrus Experiment Station, Riverside, LJ. Condit 1. Santa Barbara Co.: Franceschi Nursery, Santa Barbara, W.E Wight 4879. Sonoma Co.: Sonoma, H. Grinstead s.n. 4 Jun 1978. Florida. Alachua Co.: Gainesville, WHITTEMORE, NOTES ON SOUTHWESTERN MORACEAE 771 EG. Meyer & PM. Mazzeo 14365. Georgia. Morgan Co.: Madison, FG. Meyer & PM. Mazzeo 20440. Illinois. Cook Co.: Morton Arboretum, Lisle, N. Gavlak 4098V90. South Carolina: Columbia, J.B. Nel- son 19580. ZIMBABWE: Salisbury, H. M. Bigel 4444. California material referred to Ficus palmata The collections cited as Ficus palmata and F. pseudo-carica by Smith (1976) and Hrusa et al. (2002) are part of a series of four collections made by Henry Pollard (Pollard s n. 20 November 1955, 29 September 1956, 23 December 1958, and 28 November 1959 [all CAS]). All bear the same locality data (bed of west fork of Cold Spring Canyon, Santa Barbara Co.) and evidently came from the same colony (perhaps the same plant), described by Pollard on the herbarium labels as a clump of tall, suckering stems growing among boulders in the streambed. The plant was evidently immature when Pollard first collected it, and he revis- ited it several times to get more mature material. The first three specimens (1955, 1956, and 1958) are all sterile, and the 1956 label specifically describes the plants as “Adventive seedlings,” while the 1959 collection is the only fertile specimen in the series, and the label says: “Largest shoots, heretofore sterile, beginning to bear fruit.” The 1955 collection was annotated, apparently in Pollard’s hand- writing, as F. carica; the others were named “Ficus pseudo-carica Miq.?,” all three with the question mark. Morphological characters of these specimens are given in Table L. — RESULTS The Pollard collections from Cold Spring Canyon match F carica, not F palmata, in most of their characters (the thickness and dark coloration of the twigs, the large, deeply lobed leaves with obtuse or acute apices, strongly cordate bases, and shallowly crenate margins, and the turbinate fruit). Only the density of the indumentum on the twigs and petioles is in any way unusual for F carica, and this cl is rather variable in F carica. The indumentum is dense only in the earlier Cold Spring Canyon collections (1955 and 1956, made when the plants were quite immature); later collections, made as they approached and reached sexual maturity (1958 and 1959), show sparser indumentum, within the normal range of variation of F carica. The plant collected by Pollard and reported by several authors as F. pseudo-carica or F palmata is clearly F. carica; the unusual indumentum characters may be related to juvenility. Specimens of F. palmata have occasionally been grown horticulturally in the United States (WE Wight 4870, Franceschi nursery, Santa Barbara California, NA), but there is no evidence that the species has ever escaped from cultivation. It should be excluded from the adventive flora of North America. Morus nigra in the southwest Black mulberry (Morus nigra) is a small tree native to the mountains of south- west Asia (probably Iran and the Caucasus). It isa very old cultigen—mulberry Taste 1.Comparison of morphological characters of Asian Ficus palm reports of F palmata.The last column notes for each morphological character which species is the best match for the California specimen ata, cultivated F.carica,and California specimens forming the basis of the North American Ficus palmata Ficus carica Basis of CA F. palmata report Material coded Twig color diameter mm wig indumentum ee length mm Petiole indumentum Blade length mm Blade width mm Blade shape Blade lobing Blade/lobe margin Blade base Blade apex Fruit shape Fruit width mm 31 sheets from India, Nepal, Pakistan, Saudi Arabia Yemen, Eritrea and Ethiopia tan or brown, sometimes, greenish, usually with a + ashy cast from the white indumentum 1.5-4 moderately to densely hirsute 16-70 sparsely to densely hirsute, seldom puberulent 57-142 50-140 ovate to triangular usually unlobed, seldom 3(-5)- lobed to 0.4(-0.7) of length shallowly to strongly crenate- toothed or bluntly toothed broadly obtuse to truncate or shallowly cordate rounded, apiculate or acuminate globose, often from a linear stipe-like base 9-15 mm ca. 70 sheets, cultivated worldwide brown, usually dark 4-5 glabrous or sparsely hirsute 40-119 glabrous or sparsely hirsute 114-211 129-210 ovate to very broadly ovate in outline 3-5-lobed 0.5-0.85 of length rarely shallowly lobed) crenate or irregularly wavy —a shallowly to deeply cordate, rarely broadly obtuse rounded or obtuse, ver rarely acute but never acuminate turbinate more than 15 mm 4 Pollard collections, W fork Cold Spring a Santa Barbara Co. purplish brown sparsely to moderately hirsute 110-125 sparsely to moderately 240-370 ovate to very broadly ovate in outline 5- se 0.75-0.8 of cordate obtuse or acute turbinate too immature to measure Best match for Pollard figs F.carica F.carica intermediate F.carica F. palmata F.carica F.carica either F.carica F.carica F.carica F.carica F.carica (L)@2 VaIs/9¥O'LIYS WHITTEMORE, NOTES ON SOUTHWESTERN MORACEAE 773 seeds are known from early archaeological sites in Mesopotamia and Egypt (Brothwell & Brothwell 1969)—and black mulberry is now widespread in cen- tral and western Asia and the Mediterranean basin. In the North American botanical literature, the name M. nigra has been confused since Small (1903, 1933) misapplied it to dark-fruited forms of M. alba, an east Asian species that is widely naturalized in the United States (Wunderlin 1997). Wunderlin’s dis- cussion seems to suggest that the two taxa may be conspecific, but M. alba and M. nigra are actually quite distinct, differing (among other features) in their chromosome numbers, M. alba being diploid with 2n=28 whilea variety of poly- ploid or high aneuploid numbers, varying from 2n= 89-308, have been reported for M. nigra (Voltattorni 1947; Darlington & Wylie 1955; Hans 1972). Morus nigra was reported to escape occasionally in Beatty Townsite, Nye Co., Nevada, by Beatley (1976). This report has not been mentioned in subse- quent publications, including the Flora of North America Moraceae treatment (Wunderlin 1997). A check of Beatley’s hers showed that they are correctly determined and also revealed specimens of apparent waifs collected by Henry M. Pollard at a site in southern California (see below for specimen data). These collections indicate that black mulberry is able to reproduce as a waif at widely separated sites in the southwestern United States. It is only known asa waif in disturbed areas close to settlements, and there is no evidence that it can persist for long periods or disperse into intact natural vegetation. The species is not correctly described or keyed in any North American flora (except McMinn & Maino 1947, who included it as a cultivated species), soa brief description of M. nigra and an emended key for the region are provided. In order to enlarge the sample size and capture the normal range of variation in the Species the de scription below was supplemented with cultivated material from the south- western United States. Illustrations of M. nigra are provided by McMinn and Maino (1947, fig. 203) and Browicz (1982, fig. 1). KEY TO SPECIES OF MORUS IN THE WESTERN UNITED STATES 1. Leaves reniform-triangular to almost circular, base deeply cordate. Styles 3-5 mm aieee densely hairy all over. eae purple-black or black M. nigra . Leaves ovate or triangular-ovate, base rounded, truncate, or shall rdate. Styles 1-2 mm long, glabrous ioe with a few hairs). Syncarp red, adiele or white. 2. Upper surface of leaf smooth, often very shiny, glabrous or with a few hairs on the main veins; underside glabrous except for scattered hairs on the major veins or in vein axils. Syncarp short-cylindrical to almost spherical, 0.6-1.8 X 0.5-0.7 M. alba 2. Upper surface of leaf dull, often scabrous, with hairs (very inconspicuous in M. rubra) evenly scattered over the blade; underside pubescent with hairs evenly scattered over the veins and blade. Syncarp various 3. Trees to 20 m tall. Leaves ovate to subcordate or almost circular, 7-28 * 7-25 cm, if lobed then lateral lobes abruptly acuminate. Syncarp short-cylindrical, 1-1.9 X 0.6-0.8 cm M. rubra a 774 BRIT.ORG/SIDA 22(1 = 3. Low shrubs to small trees 2-5 m tall. Leaves ovate or triangular-ovate, 2.8-8.3 1.6-5.3 cm, apex abruptly acuminate, if lobed then lateral lobes acute, ob- tuse, or occasionally rounded. Syncarp more or less spherical, 0.5-1.2 * 0.6- 13cm M. microphylla Morus nigra L., Sp. Pl. 2:986. 1753. BLACK MULBERRY Small trees 3-5 m tall. Twigs brown, 2-4 mm thick, puberulent, pilose, or gla- brous; buds ovoid, 5-7 mim long. Petiole 1.2-3.0 cm long, pubescent with long soft hairs, at least ventrally. Blade reniform-triangular to almost circular, unlobed or rarely 3-lobed, 7-16 cm long and wide, (3-)5-veined from base; base deeply cordate, apex abruptly short-acuminate, margins dentate, teeth rounded- obtuse, 2-5 mm long; upper surface green, smooth or very weakly scabrous, gla- brous or with a few scattered hairs mostly confined to the major veins; under- side lighter green, soft-pubescent on veins and blade. Unisexual. Styles 3-5 mm long, densely hairy all over. Fruiting peduncle densely pubescent; syncarp short- cylindrical, 1.4-2.2 cm es 10-16 cm wide, purple-black or black. Specimens examined. U.S.A. NEV, e Co eee) escaped, in thicket with Screw-bean, Beatty townsite, eo ee USA. E.C’s Nevada Test Site, J. Beatle y 13438 (US). CALI- FORNIA, Ventura Co.: two small trees in low waste ground in dense brake of Rubus procerus, S of railroad and Fox St. drain, Ojai, origin unknown, location suggesting escape from cultivation, H.M. Pollard s.n., 21 Oct 1965 (NA); same locality, H.M. Pollard s.n., 20 Jun 1968 (NA). ACKNOWLEDGMENTS I would like to thank the curators at CAS, MO, UC and US for providing facili- ties for work at their herbaria, and Guy Nesom, Steve Boyd, Dieter Wilken, and Margriet Weatherwax for helpful comments. REFERENCES Beatey, J.C. 1976. Vascular plants of the Nevada Test Site and central-southern Nevada. Springfield, VA, National Technical Information Service, 1D-26881. BrotHWwett, D.and P. BrotHwelt. 1969. Food in antiquity. Ed. 2. Baltimore: Johns Hopkins Uni- versity Press. Browicz, K. 1982. Moraceae. Flora lranica fascicle 153. Graz: Akademische Druck- u. Verlagsanstalt. Conor, |.J. 1947. The fig. Waltham, Mass.: Chronica Botanica Co. DaruncTon, C.D, and A.P. Wyue.1955.Chromosome atlas of flowering plants. Ed. 2. London: Allen & Unwin Ltd. Friis, |. 1993. Moraceae. In: M. Thulin, ed. Flora of Somalia, vol. 2. London: Royal Botanic Gardens, Kew. Pp. 91-104. GuHaroor, A. 1985.Moraceae. Flora of Pakistan fascicle 171.Islamabad: Pakistan Agricultural Research Council; and Karachi: Dept. of Botany, University of Karachi. Hans, A.S. 1972. Cytomorphology of arborescent Moraceae. J. Arnold Arbor. 53:216-225. Hrusa, F., B. Ertrer, A. SAnbers, G. Lepric, and E. Dean. 2002. Catalogue of non-native vascular WHITTEMORE, NOTES ON SOUTHWESTERN MORACEAE 775 plants occurring spontaneously in California beyond those addressed in The Jepson manual—part |. Madrono 49:61-98. McCuntock, E. 1993. Moraceae. In: J.C. Hickman, ed. The Jepson manual: higher plants of California. Berkeley: University of California Press. Pp. 764-765. McMinn, H.E.and E. Maino. 1947. An illustrated manual of Pacific Coast trees. Ed. 2. Berkeley: University of California Press. Munz, PA. 1974.A flora of southern California. Berkeley: University of California Press. Smatt, JK. 1903. Flora of the southeastern United States. New York: Published by the author. Sma, J.K. 1933. Manual of the southeastern flora. New York: Published by the author. Smito, C.F. 1976. A flora of the Santa Barbara region, California. Santa Barbara: Santa Barbara Museum of Natural History. Smit, C.F. 1998. A flora of the Santa Barbara region, California. Ed. 2. Santa Barbara: Santa Barbara Botanic Garden. Voutattorni, S. 1947. Embriologia e cariologia di Morus nigra L. Annali Sperim. Agrar. N.S. Wunoertin, R.P. 1997. Moraceae. In: Flora of North America Editorial Committee, Flora of North America, volume 3. New York: Oxford University Press. Pp. 388-399. 776 BRIT.ORG/SIDA 22(1) Book REVIEW CHET VAN Duzer. 2004. Floating Islands: A global bibliography with an Edition and Translation of G. C. Munz’s Exertacio academica de insulis natantibus (1711). USBN 0-9755424-0-0, hbk.). Cantor Press, Los Altos Hills, Califor- nia 94024, US.A. (Orders: www.cantorpress.com). $44.95, 400 pp., illus- trated, 6 1/8" x 9", To an average versed and educated reader the word floating island evokes association to Jules Verne and Jonathan Swift, to the description of explorations of swamps along the upper course of the Nile River and to the floating gardens of the Titicaca Lake. These do not seem entirely concrete objects anime in the aie life. This is we case in most countries without such islands. Floating Islands | review of 2500 years of publications on floating islands, cov- ering all plpeevile items from floating bogs to pumice islands and from bird nest rafts to floating air port o Duzer’s book consists of two main parts. The first (33 pp.) isa reprint of the full Latin text and English translation of Georgius Christophorus Munz’s rare 1711 Exercitatio acade mica de Insulis natantibus. Thus the basic work on the topic is easily accessible to the reader, both in its original form as well as its translation. Comprehensive and clear notes by Van Duzer follow (pp. 35-67) the Munz’s description. The second of the book is Global Bibliography on Floating Islands. On 280 pages there are gathered more than 1800 references of books and articles in twenty languages treat- ing this subject. The entries are annotated and cross- ened and thet are e both h thematic and geographic indices. All aspects of floatin f floating islands, the causes of their buoyancy, ree role in the ecology of lakes aad wetlands, their flora and auna, their role in Bime eepeteal: st pate ana nee and methods for controlling and managing them. Worl ial {1 1 for agriculture, human habitation, wild- life habitat, and imp The book melndes the text al an sees aouapuuae Ww iui asiaied notes, of G.C. Munz’s rare 1711 fl islands, E .as well as photographs of sev- eral floating aoa Van Duzer’s book is not only a complete (or one of the ae eiPiloarapiy on floating islands, f water ake: and floating ade in literature, myth, and legend. but also an indispensable source of literature for everyone { float- ing islands. Floating Islands succeeds in condensing a large body of historical ne and thought, in addition, it contains summaries of the recent research on floating wetlands. Essentially every feature of floating islands is s exarined—their formation, causes of their puCyaney, dominan 1 } flora and fauna, and tt leir interactions with the wetland, lake, and river habitats with w hich they are 1. Th ee ea associatec I d if ing islands, to their uses, including human habitation, and into classical inenanine and mythology. Van Duzer has not only provided a masterful treatment of the subject but has also provided an excellent sets fee ing ‘Islands should serve asa pasts for ne sc OlaeP for years tocome. Recommended fo ¢, botanical, herbarium Larger public librar- ies would be well cee to include it in their collection.—Gary Jennings, ee Research Insti- tute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A. SIDA 22(1): 776. 2006 EXOTIC PLANT INTRODUCTION IN KANSAS, TWO NEW SPECIES lralee Barnard 962 300 Avenue Hope, Kansas 67451, U.S.A. findiralee@yahoo.com ABSTRACT Described in this paper are two new records for Kansas, Myagrum Bae coals Li ~ (> 1000 - 2500 ~ WM >2500 ba Way =" - La | rd Pi pee ay Ret iri L COON Pp Fic. 1. Peri Eleocharis liesneri S. Gonzalez & Reznicek, Novon 6:361. 1996. (Fig. 2a—c). Tyre: NEZUELA. BOLIVAR: Gran Sabana, ca. 10 km SW of Karaurin Tepui at junction of Rio Karaurin and Rio Asadon (Rio Sanpa), gallery forest, 519°N, 61 03’W, 900-1000 m, riverbank, 23 Apr 1988, R. Liesner 23698 (HOLOTYPE: MO} ISOTYPE: NY}). Perennial herb; roots coarse, fibrous, dark drab brown; rhizomes elongated, to 784 BRIT.ORG/SIDA 22(1) ig & Nees) Steud. d—e. Detail of achene, perianth bristles, and it ! Inf 1 ~ a] t yt “39 7N0 MAN ie ie ae Cea ORME PRT 5 eee! | tubercle. f. Detail of spil 7634 (C) by Neva Mikulicz. ROSEN, RECORDS OF ELEOCHARIS LIESNERI FROM SOUTH AMERICA 785 ca. 1.8 mm thick, scales not seen, tubers absent; culms (when dry) terete or dis- tally obscurely 3-angled, 40-65 cm x 2-4.4 mm, rigid, internally spongy, with incomplete transverse septa, finely longitudinally striate, olive-brown; leaves reduced to sheaths, sheaths 2, apically notched, membranous, loose, friable, variable in color proximally from dark brown to maroon, distally cinnamon to rufous, apex acute; spikelet cylindric, narrowly-lanceolate, 1.6-2.4.cm X 3.9-4.5 mm, acute, proximal scale empty, obtuse, amplexicaul, appearing asa continu- ation of the culm; floral scales appressed to weakly spreading, oblong, 5.5-6.2 mm X 2.3-3.2 mm, distal and medial 0.2-0.5 mm hyaline-erose, cartilaginous, darkish centrally or subapical dark splotch, reddish-stramineous marginally, apex acute (the distally hyaline region obtuse-rounded if present), central area nearly flat, coarsely many veined, mid-vein not distinguishable abaxially, con- spicuous in adaxial view; perianth bristles 6-8, sub-equal, exceeding the tu- bercle, ascending, broad and strap-shaped proximally, coarsely retrorsely spinu- lose nearly to the base, the spinules coarse distally, becoming finer and more closely spaced proximally, stramineous; stamens 3; style trifid; achene bicon- vex, more or less obovoid to broadly obpyriform, 2.8-3.2 mm (including tubercle) x 13-19 mm, with 23-26 longitudinal rows of deeply concave transversely oblong polygonal cells visible through transparent periclinal layer on each achene face, dull, buff, maturing to shiny light brown, apex constricted to about 0.4 times the width of the achene to a distinct neck (this sometimes obscured by the style base); tubercle dorsoventrally compressed, triangular, 0.5-1.2 mm x 0.7-0.9(-1) mm, dark brown. Specimen citations. BRAZIL. Mato Grosso do Sul: Central Brazilian Plateau, ca. 1-3 km W of Km 261 Xavantina-Cachimbo road, in standing water in wet forest marsh, 12° 49'S, 51° 46’W, 19 Jan 1968, Philcox & Fereira 4124 (NY). Sau Paulo: Gesammelt in Sud-Brasilien um Sao Paulo, without date aaa eae? annotation a. C. B. Clarke dated 24 Apr 1906), Usteri s.n. (ZT). COLOMBIA. o. de Sonson, km 4 of road Sonson-La Union (52 km from La Union), dictated roadside 8 - 44'N, 75° 18 W, 2480 m, 04 Oct 1987, Zarucchi et al. 6240(MO). ACKNOWLEDGMENTS lam very grateful to the herbaria indicated for loan of specimens, to Socorro Gonzalez-Elizondo for advice, encouragement, and helpful comments, to Neva Mikulicz for preparing the excellent illustrations, and Tony Reznicek for re- viewing this manuscript. Thanks to Larry Brown (SBSC) for requesting loans of the type specimens. Financial support for research was provided by the Frank W. Gould Award for Graduate Student Research in Plant Systematics and the Dr. Harry Wayne Springfield Graduate Student Endowment. REFERENCES Barros, M. 1960.Las ciperaceas del estado de Santa Catalina. Sellowia 12:262. Ciarke, C.B. 1898. Cyperaceae. In: R. Chodat, Plantae Hasslerianae soit €numération des 786 BRIT.ORG/SIDA 22(1) nlantes récoltées par le Dr.Emile Hassler, d’Aarau (Suisse) de 1885-1895 et déterminées par le Prof. Dr. R. Chodat avec l'aide de plusiers collaborateurs. Bull. Herb. Boissier 6, Appendix 1:19-22. GonzAez -Euizonpo, M.S.and A.A. Reznicek. 1996. New Eleocharis (Cyperaceae) from Venezu- ela. Novon 6:356-365. GONZALEZ-ELIZONDO, M.S.and P.M.Peterson.1997.A classification of and key to the supraspecific taxa in Eleocharis (Cyperaceae). Taxon 46:433-449. GonzAtez-Evizonpo, M.S.and J.A.Tena-Fiores. 2000. Eleocharis (Cyperaceae) in the New World. In: K.L.Wilson and D.A. Morrison, eds. Monocots: systematics and evolution.CSIRO Pub- lishing. Australia. Pp.637-643. GonzALez-EuIzoNnbo, M.S., M. GONZALEZ-EvIZONDO, and S.G. SmitH. 2002. Eleocharis obtusetrigona (Cyperaceae) new to North and Central America. Acta Botanica Mexicana 60:7-11. Svenson, H.K. 1939. Monographic studies in the genus Eleocharis V. Rhodora 41:13-19; 95-104. NEW AND SIGNIFICANT REGORDS.OF VASCULAR PLANTS FOR FLORIDA AND FOR COLEIFE COUNTYAND LEE COUN FYE RORIDA George J.Wilder Martha R.McCombs Division of Ecological and Social Sciences 28260 Pine Haven Way, Unit 85 Florida Gulf Coast University Bonita Springs, Florida 34135, U.S.A. 10501 FGCU Boulevard South mecombsmarthar@aol.com Fort Myers, Florida 33965-6565, U.S.A. gwilder@fgcu.edu ABSTRACT Documented records are reported for 89 taxa (81 species, 7 varieties, and one hybrid) from Collier County (31 taxa) and from Lee County (71 taxa), Florida (102 records altogether). A newly discovered hybrid, Carex gigantea x Carex lupuliformis, is discussed. Key Worps: Florida, native species, alien species, new records, hybrid RESUMEN Se realizan las citas documentadas de 89 taxa (81 especies, 7 variedades, y un hibrido) de Collier Sony iO taxa) y de Lee County (71 taxa), Florida (102 citas en total). Se discute un hibrido , Carex gigantea X Carex lupuliformis. INTRODUCTION Herein we present new documented records ont ian of vascular plants from Collier County and/or Lee County, Florida, and wv discuss a Carex hybrid here reported for the first time in the literature. Collier County and Lee County occur within the southwestern portion of the Florida Peninsula and border one another from south to north, respectively. These counties encompass 2,026 and 804 square miles, respectively (Southwest Florida Regional Planning Council 2002). They merit especial attention for five reasons. (1) Gann et al. (2002), who grouped Collier and Lee counties among South Florida’s counties, characterized South Florida as “.. one of the most biologi- cally diverse regions in North America.” They reported over 2,200 species of native and naturalized plants from South Florida, including over 1,400 native species. By contrast, Wunderlin and Hansen (2003) recognized less than twice as many native and nonnative taxa (species, infraspecific taxa, and hybrids) for Florida’s 67 counties, overall (.e., 4,145 taxa). Wunderlin and Hansen (2004) listed the following numbers of taxa for the ten South Florida counties recog- nized by Gann et al. (2002): Broward Co., 1001; Charlotte Co., 709; Collier Co., 1194; Glades Co., 485, Hendry Co., +93; Lee Co.,, 1172; Martin Co., 988; Miami- SIDA 22(1): 787 -— 799. 2006 788 BRIT.ORG/SIDA 22(1) Dade Co., 1655; Monroe Co., 575, 776 (the mainland portion of Monroe Co. and the Florida Keys, respectively); and Palm Beach Co., 1036. (2) South Florida extends into the subtropics, thus exhibiting impressive overlap between typically temperate, and characteristically tropical vegetation. Coordinate with their subtropical milieu, at all times of year Collier and Lee counties exhibit particular species in flower. (3) Both counties manifest intense residential and commercial develop- ment. Within extensive areas their floras are being decimated, and increasing numbers of taxa face extirpation or rarity. The Southwest Florida Regional Plan- ning Council (2002) defined Southwest Florida as consisting of six counties, including Collier and Lee counties. From 1950 to 2000, Southwest Florida’s share of total U.S. population increased over nine-fold, from 0.047% to 0.429%. Simi- larly, from 1950 to 2000 Southwest Florida’s share of Florida’s population in- creased nearly three-fold, from 2.571% to 7.550%, the latter increase transpired despite the circumstance that Florida’s population growth exceeded that of the U.S., overall. (4) Together with Hawaii, Florida ranks among the two states “..with the most severe nonnative species problems...” (Simberloff 1997). One-third of the South Florida flora consists of escaped, nonnative plants (Gann et al. 2002). Simberloff (1997) stated that two primary factors predispose Florida, particu- larly southern Florida, to invasion by, and damage from nonnative species: (a) “destruction and disturbance of native habitats and their replacement by novel habitats...” and (pb) *..the geographic features of tropicality, insularity, and the great expanse of aquatic habitats.” Simberloff (1997) considered the problem of nonnatives to be aggravated by the considerable tourism and transportation into Florida. (5) No flora has been published for Collier County or Lee County. “ Climate Annually, Collier and Lee counties exhibit a warm rainy season and cooler dry season. During the dry season shallow, rain-fed pools dry up and the land ap- pears drier, overall. Liudahl et al. 1998) cited data gathered at Fort Myers (Lee Co.)—20 miles north of Collier County—to characterize the climate of Collier County. Thus, those data represent both counties. They indicated annual average tempera- ture of nearly 74° F and temperatures ranging from an average within the low sixties in January to the low eighties during summer. Average daily maximum temperatures vary from 74.7° F January) to 91.1° F (August); corresponding daily minimum temperatures for the two months are 53.6° F and 74.1° F Frost develops few times, yearly (Liudahl et al. 1998). Collier and Lee coun- ties each exhibit USDA plant hardiness zone nos. 10a and 10b, and Lee County also exhibits zone Ob ( th ESE ZOTIES represent average ann ual m inim umtem pera- tures of 30°-35" F 35°-40° F and 25°-30° F respectively; Anonymous 2005). WILDER AND MCCOMBS, FLORIDA PLANT RECORDS 789 Average annual rainfall exceeds 50 inches. Approximately, two-thirds of rain falls from June to September, when average rainfall is nearly eight inches per month. Monthly average rainfall is less than two inches from November through January and is a little over two inches from February through April (Liudahl et al. 1998). Geology and soils Abutting the Gulf of Mexico and belonging to the Atlantic Coastal Plain, both counties exhibit low, flat terrain. They are part of a larger, unnamed, primarily southern and peripheral region of Florida exhibiting maximum elevation of 70 feet above sea level (Schmidt 1997). Physiographic regions prominent within Collier and Lee counties are the Big Cypress Spur, Caloosahatchee Valley, Gulf Barrier Chain and Gulf Coastal Lagoons, Immokalee Rise, Reticulate Coastal Swamps, and the Southwestern Slope (Liudahl et al. 1998; Schmidt 1997). Within each county, outcrops and shallow subcrop rocks are either Pliocene or Pleis- tocene-Holocene in age (Randazzo & Jones 1997). Henderson, (1984) and Liudahlet al. (1998) surveyed the soils of both coun- ties. For Collier Co., Liudahl et al. (1998) grouped general soil map units under either of three main headings: (1) urban land and soils in urban areas, (2) soils on the flatwoods and hammocks and in sloughs, and (3) soils on prairies and in swamps and freshwater marshes. For Lee Co., Henderson (1984) grouped major units similarly, but under four main headings: (1) soils of the manmade areas, (2) soils of the flatwoods and sloughs, (3) soils of the swamps and sloughs, and (4) soils of the tidal areas and barrier islands. Ecosystems and land-use history Despite their minimal ranges of elevations, both counties, collectively, exhibit diverse ecosystems, e.g., pine flatwoods, scrub, diverse kinds of hammocks, swamps, freshwater marshes, saltwater marshes, mangrove forests, and ruderal sites (Myers & Ewel 1990; Taylor 1998; G. Wilder, pers. obs.). In addition to development, land use in Collier and Lee counties has en- tailed these extensive human-induced modifications: construction of drainage canals and of ditches for mosquito control; elimination of, and various alter- ations of wetlands; cattle ranching; row-crop agriculture; and lumbering. Both counties also maintain major preserves or portions of preserves: Big Cypress National Preserve, Collier Seminole State Park, Corkscrew Regional Ecosystem Watershed, Corkscrew Swamp Sanctuary, Fakahatchee Strand State Preserve, and Everglades National Park. Furthermore, Everglades restoration, currently underway partly within Collier and Lee counties, ranks among the largest of efforts for ecosystem restoration, worldwide. MATERIALS AND METHODS We made most collections between and including 2002 and 2005, however, we collected Limnobium spongia in 1994 and Leptochloa nealleyi, Paspalum 790 BRIT.ORG/SIDA 22(1) distichum, and Scirpus pungens in 1998; Appendix). All collections but one were prepared as dried herbarium specimens; Landoltia punctata was fixed in aque- ous formalin-propionic acid-ethanol solution and stored in aqueous glycerin- ethanol solution. Specimens cited are deposited in the Herbarium of Southwest- ern Florida (SWF), housed at Florida Gulf Coast University (Fort Myers, FL). We characterize taxa as native, alien, and endemic, according to Wunderlin and Hansen (2003, 2004). Nomenclature for species and infraspecific taxa follows Wunderlin and Hansen (2003), but for several taxa we also provide commonly used synonyms. Family circumscriptions and common names also follow Wunderlin and Hansen (2003), except for Lemnaceae, Nyssaceae, and Asclepiada- ceae, which we keep separate from Araceae, Cornaceae, and Apocynaceae, re- spectively. The Atlas of Florida Vascular Plants Wunderlin @ Hansen 2004) was our principal source for determining whether taxa represented new county or state records. That database does not reference voucher specimens, however, Wunderlin (2002) specified that “Each distribution record on the atlas website is documented by an herbarium specimen or a reliable published source, such asa monograph or revision.” We also recognized documented plant records from Austin et al. 1990) and Gann et al. (2002). There exists a plethora of additional reports and inventories pertaining to the floras of Collier County and Lee County, which do not reference voucher specimens, which are unpublished, and/or which have insufficient circulation. Because of these shortcomings, we do not consider records cited within them as adequately established. As Wunderlin (2002) stated in regard to voucher specimens, if a “..species is not documented..there is no record that the plant ever existed...” RESULTS AND DISCUSSION Reported herein are 89 taxa (81 species, 7 varieties, and one hybrid) that repre- sent 31 new documented records for Collier Co. and 71 for Lee Co., Florida (02 records altogether). The hybrid, Carex gigantea < Carex lupuliformis, is newly discovered and thus a new Florida record. Asclepias feayi is included here, although it was documented previously. That species was originally collected near Immokalee (Collier Co.) in 1967 (Gann etal. 2002). It is cited here, because those workers designated A. feayi as histori- cal in South Florida Gincluding Collier Co.), a designation implying that the species might have been extirpated there. Muss et al. (2003) published a plant-species list for the Big Cypress Na- tional Preserve, which includes portions of Collier, Miami-Dade, and Monroe counties. They listed nine species cited in this article: Cyperus involucratus, Dalbergia sissoo, Eleocharis flavescens, Ixora coccinea, Limnophila sessiliflora, WILDER AND MCCOMBS, FLORIDA PLANT RECORDS 791 Melilotus albus, Paspalum distichum, Reimarochloa oligostachya, and Senna alata, but did not indicate which, if any, records represented Collier County. The Institute for Regional Conservation (IRC) cites plant records for South Florida, from many sources (Gann et al. 2005). These records as presented by Gann etal.(2005) are undocumented and, therefore, not accepted here. Gann et al. (2005) cited four taxa that we report for Collier Co. (Eleocharis flavescens, Limnophila sessiliflora, Melilotus albus, Scirpus californicus) and eight taxa presently listed for Lee Co. (Asclepias feayi, Cirsium nuttallii, Commelina gambiae, Palafoxia integrifolia, Paspalidium geminatum, Paspalum conjugatum, Phyllan- thus amarus, Quercus chapmanii). Taxonomic data The 89 taxa represent pteridophytes (1 taxon; 1.1 % of taxa), monocotyledons (44 taxa; 49.4%), and dicotyledons (44 taxa; 49.4%). Numbers of families in each group are: | (pteridophytes), 10 (monocotyledons), and 24 (dicotyledons; Appendix). Families of monocotyledons with the most taxa are Poaceae (21 taxa) and Cyperaceae (15 taxa). Remaining monocotyledonous families each include one taxon. The 36 taxa of Poaceae and Cyperaceae, collectively, constitute 40.4 % of presently reported taxa. This disproportionately high percentage suggests that previous collectors within Collier and Lee counties documented these families insufficiently. Families of dicotyledons with the most taxa are Fabaceae (8), Asteraceae (5), Veronicaceae (5), Euphorbiaceae (4), Brassicaceae (2), and Urticaceae (2). Remaining dicotyledonous families each include one taxon. Native and Endemic Taxa Wunderlin and Hansen (2003) classified as native to Florida 48 (53.9%) of the 89 taxa reported here (Appendix). They listed two of our taxa as endemic to Florida (Asclepias feayi and Linum carteri; Appendix). We consider Carex gigantea x Carex lupuliformis endemic, as well. In the Poaceae and Cyperaceae, the families most prominently represented in this study, 12 and 10 taxa are native, respec- tively (ie. 57.1% and 66.7% of taxa of these families). The present report of a large proportion (ca. one-half) of nonnative taxa accords fairly well with the consider- able proportion (ca. one-third) of such taxa indicated previously for South Florida (Gann et al. 2002) and for Florida, overall (Wunderlin & Hansen 2003). Rare Florida Taxa The eighty-nine records include a considerable number of rare taxa. Foremost, is Carex gigantea X Carex lupuliformis, first collected by George Wilder on April 22, 2003. We found that hybrid solely at one station, in Six Mile Cypress Slough Preserve (Lee County). Two years ago, Dr. Anton Reznicek (MICH) annotated three herbarium specimens of this hybrid as “a beautiful intermediate collec- tion” (Wilder & McCombs [W & M] 17258, W & M 17259, and W & M 17260). 792 BRIT.ORG/SIDA 22(1) Preliminary study revealed three differences between both parental spe- cies and the hybrid. () Perigynia exhibit achenes (both parental species), but are sterile (the hybrid); indeed, sterility isa common feature of hybrids, overall. (2) Relative to spike axes, perigynia extend at, or approximately at right angles (C. gigantea), are ascending (C. lupuliformis), and have generally intermediate orientations (the hybrid). (3) Spike diameters (excluding perigynium beaks) are narrow (C. gigantea), broad (C. lupuliformis), and intermediate (the hybrid). Cayouette and Catling (1992), in their comprehensive review of Carex hy- brids, referred to “Carex gigantea Rudge x lupulina Willd. @ lupuliformis Sartwell). Possible hybrid origin according to Correll and Johnston (1970).” That language is interpretable in various ways—e.g., to mean the existence of C. gigantea X C. lupuliformis, however, the latter interpretation is incorrect. Correll and Johnston (1970) (who construed C. lupuliformisasa form of Carex lupulina Muhl., rather than as a discrete species) actually said of C. lupuliformis, that “ ..these plants seem to show the introgressive influence of C. lupulina by C. gigantea.” Thus, Correll and Johnston (1970) suggested hybrid character of C. lupuliformis, overall, not a definite C. gigantea/C. lupuliformis hybrid. Carex gigantea X Carex lupuliformis was not reported or recorded in Flora of North America (Reznicek 2002), in section Lupulinae of the genus Carex, nor was it indicated in floristic manuals listing both parental species (Fernald 1950; Gleason 1968: Gleason & Cronquist 1991; Godfrey & Wooten 1979; Mohlenbrock 1999; Small 1933; Wunderlin & Hansen 2003). Some of these work- ers submerged, or apparently submerged Carex lupuliformis into Carex lupulind Mull. Leptochloa nealleyi and Kyllinga hyalina are also unusual in Florida. George Wilder collected L. nealleyi on Sanibel Island (Lee Co.), on Aug, 27,1998 (W & M 9774). We know of one other Florida collection of L. nealleyi, from Miami-Dade County (Keith A. Bradley 2583, Aug. 19, 2004). Leptochloa nealleyi was listed by Wunderlin and Hansen (2004), but not by Wunderlin and Hansen (2003). We collected Kyllinga hyalina in Collier County and Lee County, where it isa lawn weed and grows in disturbed areas. Two years ago, Dr. Anton Reznice (MICH) annotated six of our specimens as “Cyperus hyalinus Vahl 2nd US. col- lection” (W & M 17261 to W & M 17266, all of Sept. 8, 2002). (Wunderlin & Hansen [2002] listed Cyperus hyalinus Vahl and Kyllinga hyalina (Vahl) T. Koyama as synonyms). Wunderlin and Hansen (2003) ranked 17 presently reported taxa as rare in Florida. We increase that number by two, by including Carex giantea X Carex lupuliformisand Leptochloa nealleyi (taxa which those workers did not consider), Based on data from Wunderlin and Hansen (2003) and on our assessments of these two taxa, the 89 taxa considered here are ranked for Florida, as follows: rare, 19 taxa (21.3% of all 89 taxa); occasional, 48 taxa (53.9%); common, 4 taxa (4.5%): and frequent, 19 taxa (21.3%). Rare and occasional taxa predominate among our —— N WILDER AND MCCOMBS, FLORIDA PLANT RECORDS 793 collections—composing, collectively, three-quarters of taxa presently listed. Com- mon and frequent taxa represent, collectively, ca. one quarter of taxa. Wunderlin and Hansen (2003) listed as rare in Florida the following of our taxa: Acalypha arvensis, Acalypha setosa, Achyranthes aspera var. pubescens, Alysicarpus vaginalis, Callitriche pedunculosa, Chamaesyce lasiocarpa, Commelina gambiae, Dalbergia sissoo, Heteropogon contortus, Ixora coccinea, Kyllinga hyalina, Leptochloa virgata, Linum carteri, Paspalum nicorae, Senna alata, Spigelia anthelmia, and Zoysia tenuifolia. Five of the nineteen taxa des- ignated as rare by Wunderlin and Hansen (2003) and by ourselves, collectively, are native to Florida (Appendix). For Florida, Coile and Garland (2003) classified Linum carteri as endan- gered and Tillandsia variabilis[Tillandsia valenzuelana A. Rich.Jas threatened. Gann et al. (2002) considered as rare in South Florida eight species pres- ently listed. They indicated two of these species as extirpated (Lipocarpha maculata [“collected once in 1965 near Immokalee”], Scirpus californicus [“last collected in 1965 near South Bay in Palm Beach County”), two species as his- torical (Asclepias feayi [“last collected in 1967 near Immokalee”|, Reimarochloa oligostachya [“last collected in 1977"]), and four species as critically imperiled (Leptochloa virgata, Nyssa sylvatica, Rhynchospora baldwinii, Rhynchospora wrightiana). We documented Gann et al’s (2002) eight taxa from various localities: Lipocarpha maculata (Lee Co,, 2 localities), Scirpus californicus (Collier Co., | locality), Asclepias feayi (Collier Co., 1 locality; Lee Co., 6 localities), Reimarochloa oligostachya (Collier Co., 1 locality), Leptochloa virgata (Lee Co., 3 localities), Nyssa sylvatica (Lee Co., 1 locality), Rhynchospora baldwinii (Col- lier Co., 1 locality; Lee Co., 1 locality), and Rhynchospora wrightiana (Collier Co., 1 locality). ~— — Directions and extents of Range Extensions Data from Wunderlin and Hansen (2004) indicate that 24 of our taxa repre- sent range extensions southward in Florida: Acalypha setosa, Anthaenantia villosa, Callitriche pedunculosa, Chasmanthium nitidum, Cyperus lanceolatus, Dalea carnea var. albida, Dichanthelium strigosum var. leucoblepharis, Hemarthria altissima, Iris virginica, Lindernia dubia var. dubia, Poa annua, Monotropa uniflora, Nyssa sylvatica, Portulaca amilis, Rhynchospora baldwinii, Rhyncho-spora fil vou id, Rite vere aaa Run: obovatus, Scirpus — californicus, Solida sis var. scabra, Sp} _Stylosanthes biflora, Urochloa texana, and Urtica namdedy oides. Fewer within Florida are northward (Leptochloa virgata), westward (Gonce coccinea, Spigelia anthelmia), both southward and eastward (Sida santaramensis), and both northward and westward (Kyllinga hyalina, Leptochloa nealleyi). 794 BRIT.ORG/SIDA 22(1) For each of our taxa except the Carex hybrid, we gauged the relative dis- tance between the county (or two counties, collectively) newly represented here and the nearest county indicated by Wunderlin and Hansen (2004). For 60 taxa the newly reported county (or counties, collectively) bordered one or more of the indicated counties. For 2] taxa one or two counties intervened between the newly reported county (or counties, collectively) and the nearest county. For five taxa the number of intervening counties was 3 or 4 (Iris virginica, Callitriche pedunculosa, Dalea carnea var. albida, Stylosanthes biflora, Sida santaremensis). For three taxa between 6 and 8 counties intervened (Dichanthelium strigosum var. leucoblepharis, Solidago canadensis var. scabra, Lindernia dubia var. dubia). Prognosis for presently discussed taxa Development and habitat destruction are explosive within Collier County and Lee County, and we anticipate accelerated destruction of taxa and habitat in these counties. Certain localities represented herein have already been destroyed, and the following taxa are now extirpated from one or more localities repre- sented in this paper: Asparagus aethiopicus, Cirsium nuttallii, Cyperus sphacelatus, Melilotus albus, Melilotus indicus, Paspalum conjugatum, Paspalum notdatum var. notatum, Penstemon multiflorus, Polygala verticillata, and Urochloa texana. APPENDIX The following list of species, varieties, and a hybrid represent new documented records for Collier County and Lee County, Florida. Data are presented in the following order after the Latin name of a species, hybrid, or variety: relevant synonym, if any (between brackets); common name; the Wilder and McCombs collection number(s) of voucher specimen(s); habitat(s) where the species was collected; Wunderlin and Hansen’s (2003) as- sessment of frequency of the taxon in Florida; and county(ies) for which the taxon is presently documented. For individual taxa documented for both Col- lier Co.and Lee Co., collection numbers are indicated in the order listed of these counties. * = alien to Florida; © = endemic to Florida. PTERIDOPHYTE THELYPTERIDACEAE Mariana Maiden Fern—22,158;damp forest; “Macrothelypteris torresiana (Gaudich.) Ching, occasional; Lee MONOCOTYLEDONS ALISMATACEAE ASPARAGACEAE ss ittaria latifolia Willd, Broadleaf Arrowhead— — *Asparagus aethiopicus L. [Asparagus sprengeri 22697; damp soil; occasional: Lee Regel], Sprenger’s Asparagus-Fern— 20731: scrub; occasional: Collier WILDER AND MCCOMBS, FLORIDA PLANT RECORDS BROMELIACEAE Tillandsia HaMaoys is Schltdl., Airplant vamp; COMMELINACEAE *Commelina gambiae C.B. Clarke, Gambian Day- flower—22522; highly insolated, disturbed land; edge of field; rare; Lee CYPERACEAE xX Carex gigantea Rudge X Carex lupuliformis Sartwell ex Dewey— 19688; swamp; rare; Le *Cyperus involucratus Rottb. [Cyperus alternifolius L.J, Umbrella Plant—18666; insolated land beneath power lines; occasional; Collier oe lanceolatus Poir.,Epiphytic Flatsedge— etland within insolated, cleared ne iaeoleted land bordering trail; occa- sional; Lee *Cyperus sphacelatus Rottb., Roadside Flatsedge—19715, 18686; bulldozed, insolated land; insolated lawn; occasional; Collier, Lee Eleocharis flavescens (Poir.) Urb., Yellow ae 20749; along/on shaded trail ugh damp lowland; frequent; Collier ie inga aie (Vahl) T. Koyama [Cyperus hyalinus Vahl], Peduncled Spikesedge—dis- : Nise land; lawn; 18498, 17265; rare; Collier, Leatherleaf ccasional: Lee bie nga ane Michx.,L sedge—22033; disturbed land; oa Lee *Kyllinga squamulata Thonn. Ex Vahl, Asian ee pine flatwoods; occa- nal; Lee ree maculata (Michx.) Torr, American Halfchaff Sedge—18030; insolated wetland beneath power lines; disturbed land (for- merly either a flatwoods or scrub);on trail; occasional; L Rhynchospora i dwinii A. Gray, Baldwin's Beaksedge—19780; wet portion of pine flatwoods; occasional; Collier Rhynchospora filifolia A. Gray, Threadleaf eaksedge— 19084; periphery of pond; Fe eee llier Rhynchospora grayi Kunth, Gray’s Beaksedge— 17911; pine flatwoods; occasional; Lee Rhynchospora wrightiana Boeck., Wright's Beaksedge—19812;along firebreak through pine flatwoods; occasional; Collier C irp r aif (C.A.Mey.) Steud., Giant Bul- tush—16350. edge of pond; occasional; Collier Scirpus pungens Vahl, Threesquare Bulrush— 9790, 19209; insolated wetland; roadside; fre- quent; Collier, Lee HYDROCHARITACEAE Limnobium spongia (Bosc) Rich.ex Steud., Ameri- can Spongeplant—7392; occasional: in ca- nal; Lee HYPOXIDACEAE Hypoxis curtissii Rose, Common Yellow Star- grass—20142; swamp; common; Lee IRIDACEAE Iris virginica L., Virginia Iris—18207; insolated depression along road; occasional; Lee LEMNACEAE ti eee . oe Les & D. J. Craw- d Du | 1] JIIAHOVY weter; fre- ope ne POACEAE Anthaenantia villosa ane) P. Beauv., Green Cilkevweral Loew cei J ye ie JIIKYStdl I i.e) 2; pine fi ' | Lee *Bothriochloa ischaemum (L.) Keng, King Ranch Bluestem—19118; pasture; occasional: Lee Cenchrus gracillimus Nash, Slender Sandbur— 19178, 21335; along dirt road through bull- dozed land; strand vegetation by Gulf of Mexico; frequent; Collier, Lee Chasmanthium nitidum (Baldwin) Yates, Shiny Woodoats—22291; Hammock; occasional; Lee Dichanthelium strigosum var. leucoblepharis (Trin.) Freckmann—19364; edge of trail through open woodland; occasional; Lee *Hemarthria altissima (Poir.) Stapf & C. E. Hubb, Limpograss—20728; edge of pond; occa- sional; Collier Heteropogon contortus (L.) P. Beauv. ex Roem. & Schult., Tanglehead— 19467; pine flatwoods; along railroad tracks; rare; Lee *Leptochloa nealleyi Vasey, Nealley’s Sprangletop—97/74; disturbed land on Sanibel Island; rare; Lee Leptochloa virgata (L.) P. Beauv., Tropical Sprangletop—22364; damp depression along road; rare; Lee Panicum anceps Michx., Beaked Panicu 26; wet prairie; pine flatwoods; on ai common; Lee doe idium - vatum (Forssk.) Stapf, Poyelal palidium—22686; insolated wetlan fell e Paspalum conjugatum PJ. Bergius, Sour spalum—21578;| k- recently bull- dozed, ae insolated land; along dirt road; occasional; L sc dsc mL, ANeghesS: 9787; dis- and; occasional; Collier enh nicorae Parodi, Brunswickgrass— 21583:pine flatwoods; rare; Lee *Paspalum notatum Flugge var. notatum, Bahiagrass—213/70; ao highly insolated land; occasional: Le oO \O nn ea tS *Poa annua L.,Ann baile Olas “19550 lawn:shaded,b 1erj frequent; Collier, Lee Reimarochloa oligostachya (Munro ex Benth.) Hitchc., Florida Reimargrass—21136; meadow; occasional; Collier *Secale cereale L., Cultivated Rye—22847; bull- dozed, highly insolated land; occasional; Lee eat olis obtusata (Michx.) Scribn., Prairie Wedgescale— 19594; field; frequent; Lee *Urochloa texana (Buckley) R. D. Webster, Texas Signalgrass - 19633; bulldozed, highly insolated sane a Lee — tenuifolia Willd. Thiele, Manila emplegrass— 19638, ok disturbed land; a Lee DICOTYLEDONS ACANTHACEAE *Thunbergia fragrans Roxb., Whitelady—22253; disturbed hammock, edge of field; occa- sional; Lee AIZOACEAE Sesuvium maritimum (Walter) Britton et al., Slen- der Seapurslane—21161;insolated trail; oc- casional; Coll. AMARANTHACEAE bcc aspera var. pubescens (Maq.) C.C. evil's Horsewhip—21102; oak- hee within Caloosahatchee River Regional Park; rare; Lee BRIT.ORG/SIDA 22(1) ASCLEPIADACEAE = Asclepias feayi Chapm.ex A.Gray , Florida Milk- weed—1/78/74; scrub, on/along trails through pine flatwoods, grassy portion of pineland remnant, disturbed land (formerly, apparently pine flatwoods); occasional; Lee ASTERACEAE Cirsium nuttallii DC., Nuttall’s Lae ae lawns (including tree lawn); frequen *Cyanthillium cinereum (L.) H. Rob. ae ci- nerea (L.) Less.]—18454; along/on dirt road; naliColller ae ae (Nutt.) Torr. & A. Gray, Coastalplain Palafox— 22101; field; pineland; frequent; Lee Solidago canadensis L.var.scabra (Mubhl.ex Willd.) orr. & A, Gray—18432, 17315; median strip of | 75; along road; wet von shallow wa- ter; frequent; Collier, Le *Taraxacum officinale ee ex F. H. Wigg. [in- cluding Taraxacum laevigatum (Willd.) DC]—19682, 18787 (red-fruited and brown- fruited formas, respectively); occasional; Lee BRASSICACEAE *Coronopus (L.) Sm., Lesser winecress 20397; highl val insolated dirt nian ae occasional; Collier, Lee Descurainia pinnata (Walter) Britton, Western Tansymustard—20401; tree lawns along main roads; common; Lee CALLITRICHACEAE *Callitriche pecunes losa Nutt., Ua Wraterctan rt phere) depression situated ee sen land be- neath power lines; rare; Lee ERICACEAE uniflora L., Indianpipe—16699; ha portion of scrub; occasional; Collier py ne REIACERE ilypha arvensis pp., Threeseed Mercury - 17225; disturbed land by creek; rare; Lee *Acalypha setosa A. Rich., Cuban Copperleaf— 18821;shrub bed within supermarket park- g lot; rare; Lee *Chamaesyce lasiocarpa (Klotzsch) Arthur, Road- side Sandmat—19225: field; tree lawn; insolated wasteland;crack in pavement; rare; Lee in WILDER AND MCCOMBS, FLORIDA PLANT RECORDS ee amarus Schumach. & Thonn., Gale- — 21947; insolated, recently bull- a land of median of road; occasional; FABACEAE *“Alysicarpus vaginalis (L.) DC., White Money- wort 9; insolated, dry substrate of median of road;land by intersection; rare;Lee “Dalbergia sissoo Roxb. ex DC., Indian Rose- wood—21427; along railroad tracks; rare; Collier Dalea carnea var.albida (Tort.& A.Gray) Barneby, Whitetassels— 17934; pine flatwoods; occa- sional; Lee *Melilotus albus Medik., White Sweetclover— 16471; field; frequent; Collier *Melilotus indicus (L.) All, Indian Sweetclover— 905; bulldozed land; insolated dirt pile; occasional; Collier, Lee Mimosa quadrivalvis L., Sensitive Brier—17883; field; frequent; Lee *Senna alata (L.) roxb., Candlestick Plant— 19040, yee cea land; by lines; con- n site; rare; Collier, ene biflora (L.) se et on 22524; bor- dering, and extending into ditch; occasional; FAGACEAE Quercus chapmanii Sarg., Chapman's Oak.— 17198; scrub; pineland; edge of disturbed forest; frequent; Lee ITEACEAE tea virginica L., Virginia Willo 20580; forest by creek; common; Lee LINACEAE 2 Linum carteri Small, Carter's Flax—16958 field; rare; Lee MALVACEAE ae cise Monteiro, Moth Fanpetals— 1; edge of field; occasional; Collier MORACEAE Morus rubra L.,Red Mulberry—19757; hammock, disturbed land; frequent; Lee NYSSACEAE Nyssa sylvatica Marshall, Tupelo—20834; shore f pond; frequent; Lee i 797 POLYGALACEAE Po oe a verticillata L., Whorled Milkwort— ,edge of field; occasional; Lee POLYGONACEAE *Rumex obovatus Danser, Tropical Dock-—16909, insolated land by canal; dried-up ditch; weedy sod; occasional; Lee PORTULACACEAE “Portulaca amilis Speg., Paraguayan Purslane— 20769, 21705; edge of insolated, bulldozed land; lawn weed; frequent; Collier, Lee RUBIACEAE “Ixora coccinea L., Scarlet Jungleflame—21085; natural land; rare; Collier, Lee STRYCHNACEAE Spigelia anthelmia L., West Indian Pinkroot— 21232; insolated gravel and bulldozed land; tree lawn; meadow; rare; Collier; Lee URTICACEAE *Pouzolzia zeylanica (L.) Benn., Pouzolz’s Bush— oe land along road; insolated mp; occasional; Lee ae ica cramer Pursh, Heartleaf Nettle — 3; lawn weed; occasional; Lee SS NOS ~S VERBENACEAE *Verbena brasiliensis Vell., Brazilian Vervain— 19764; depression by intersection; dried-up, barren portion of lawn; occasional; Lee VERONICACEAE *Limnophila — sessiliflora Blume, Marshweed—21181;damp substrate along ee Lindernia dubia var. anagallidea (Michx.) Cooperr., ee False Pimpernel.— 21196;damp trench through insolated, bull- dozed land (growing together with the fol- lowing variety); occasional; Lee Lindernia dubia (L.) Pennell var. dubia— 21199; damp trench through insolated, bulldozed land; occasional; Lee Penstemon multiflorus (Benth.) Chapm.ex Small, Manyflower Beardtongue—22283; scrub; field; frequent; Lee *Russelia CACC OTS ae 2 nel, sional; Collier 798 BRIT.ORG/SIDA 22(1) ACKNOWLEDGMENTS We thank Edwin Everham III, Aswani Volety, and Florida Gulf Coast Univer- sity for providing space, equipment, and herbarium supplies; the Coccoloba Chapter of the Florida Native Plant Society for financial support; Roger Clark, John Yarbrough, and the Lee County Dept. of Parks and Recreation for permis- sion to collect specimens in County parks and conservation lands; Bruce Hansen, John Kunzer, and Richard Wunderlin for identifying/confirming our identifications of virtually all taxa considered here; Anton Reznicek for con- firming our identification of Carex gigantea x Carex lupuliformis and for iden- tifying Kyllinga hyalina; Anik Smith, Barbara Conelly, and Richard Workman for directing us to single localities for Panicum anceps, Achyranthes aspera var. pubescens, and Sagittaria latifolia, respectively; Edwin Everham Ill, George Gann, Mark Garland, Bruce Hansen, Richard Wunderlin, and an anonymous reviewer for making suggestions about the manuscript; and Jordana Bitner, Brian Gilbert, Kathleen Gonzalez-Sanz, Tristan Gonzalez-Sanz, Debbie Preston, Jean Roche, Graham Walker, and Jennifer White for assistance in curating the Her- barium of Southwestern Florida. REFERENCES Anonymous, 2005. Florida Landscape. http://www.wgcu.org/fllandscape/zonemap.html. Florida Gulf Coast University, Fort Myers. Austin, D.F., J.L. Jones, and B.C. Benner. 1990. Vascular plants of Fakahatchee Strand State Preserve. Florida Sci. 53:89-122. Cayouerte, J.-and PM. Carina. 1992. Hybridization in the genus Carex with special reference to North America. Bot. Rev. 58:35 1-438. Cote, N.C.and M.A. Gartanp. 2003. Notes on Florida's endangered and threatened plants. Bot. Contr. No. 38, 4" ed. Florida Dept. Agric. & Cons. Serv., Div. Plant Industry, Gainesville. Corrett, D.S. and M.C. JoHNsTon. 1970. Manual of the vascular plants of Texas. Texas Re- search Foundation, Renner Fernato, M.L. 1950. Gray's manual of botany, 8" ed. D. Van Nostrand Co., New York. Gann, G.D., K.A. Braotey, and S.W. Woopmansee. 2002. Rare plants of South Florida: their his- tory, conservation, and restoration. The Institute for Regional Conservation, Miami, FL. Gann, G.D., K.A. Braotey, S.W.Woopomansee, and J.L. Sante. 2005. The floristic inventory of South Iconservation.org. The Institute for Regional Con- Florida database. http://ww servation, Miami, FL. GLEASON, H.A. 1968. The new Britton and Brown illustrated flora of the northeastern United States and adjacent Canada. Vol. 1. Hafner Publishing Co,, NY. Gteason, H.A. and A. Cronauist. 1991, Manual of vascular plants of northeastern United States and adjacent Canada, 2" ed. The New York Botanical Garden, Bronx. Goorrey, R.K. and J.W. Wooten. 1979. Aquatic and wetland plants of southeastern United a7 WILDER AND MCCOMBS, FLORIDA PLANT RECORDS 799 States. Vol. 1. Univ. of Georgia Press, Athens. HENDERSON, W.G. Jr. 1984. Soil survey of Lee County, Florida. U.S. Dept. Agriculture Soil Con- servation Service. LIUDAHL, K.eT AL. 1998. Soil survey of Collier County area, Florida. U.S. Dept. Agriculture Natu- ral Resources Conservation Service. Mon ensrock, R.H. 1999. The illustrated flora of Illinois. Sedges: Carex. Southern Illinois Uni- versity Press, Carbondale. Muss, J.D., D.F. Austin, and J.R. Snyper. 2003. Plants of the Big Cypress National Preserve, Florida. J. Torrey Bot. Soc. 130:119-142. Myers, R.L. and J.J. Ewer (eds.). 1990. Ecosystems of Florida. University of Central Florida Press, Orlando. Ranpazzo, A.F.and D.S. Jones (eds.). 1997. The geology of Florida. University Press of Florida, Gainesville. Reznicek, A.A. 2002. Carex Linnaeus sect. Lupulinge Tuckerman ex J.Carey.In: Flora of North America Editorial Committee, ed., Flora of North America north of Mexico. Vol. 23. Ox- ford University Press, NY. Scumiot, W. 1997.Geomorphology and physiography of Florida. |In:A.F. Randazzo and DS. Jones, eds. The geology of Florida. University Press of Florida, Gainesville. Simpertorr, D. 1997. The biology of invasions. In: D. Simberloff, D.C. Schmitz, and T.C. Brown, eds., Strangers in paradise. Island Press, Washington, DC. Smatt, J.K. 1933.Manual of the southeastern flora. Published by the author, New York. SOUTHWEST FLORIDA REGIONAL PLANNING COUNCIL. 2002. Strategic Regional Policy Plan.Vol.1.South- west Florida Regional Planning Council, Fort Myers. Taytor, W.K. 1998. Florida wildflowers in their natural communities. University Press of Florida, Gainesville. WunNDERLIN, R.P.2002.Where's the proof? The importance of herbarium specimens. Palmetto. 21:6, 8. WUNDERLIN, R.P. and B.F. Hansen. 2003. Guide to the vascular plants of Florida, 2"? ed. Univer- sity Press of Florida, Gainesville. WUNDERLIN, R.P.and B.F. Hansen. 2004. Atlas of Florida vascular plants. http://www.plantatlas. usf.edu/isb/. Institute for Systematic Botany, University of South Florida, Tampa 800 BRIT.ORG/SIDA 22(1) Books RECEIVED Martua J. GRoom, Gary K. Merve, and C. RONALD CARROLL. 2005. Principles of Con- servation Biology, Third Edition. (SBN 0-87893-518-5, hbk.). Sinauer As- sociates, Inc., 23 Plumtree Road, Sunderland, MA 01375-0407; U.S.A. (Or- ders: orders@sinauer.com, wwwsinauer.com, 413-549-4300, 413-549-1118 fax). $92.95, 699 pp., 369 illustrations, 8 1/2" x 11" From the publisher—*Principles of Conservation Biology, Third Edition, features a wholly revised or- aie oe eae ing analyses of different categories of threat and approaches to conservation. Coverage has been expanded to incorporate both terrestrial and marine conservation issues, and ee the US. and across the globe. INDERJIT CEd.). 2005. Invasive Plants: Ecological and Agricultural Aspects. (ISBN 3-7643-7137-4; 978-3-7643-7137-1, hbk.). Birkhauser Verlag, PO. Box 133, CH-4010 Basel, SWITZERLAND. (Orders: wwwspringer.com). $159.00, 283 pp., illustrations, 6 1/2" x 91/4", From the publisher.—The aim ol this book is to discuss fundamental questions of invasion ecology, such as why particular communities become more invasible than others, what the mechanisms of exclusion of native species by invaders are, and whether invasion can be predicted. In addition, agricultural practices influencing invasion, the environmental and economic costs of invasion as w el as ao ee management strategies are discussed. Readers will get a unique per spective on inv gy through employing general principles of ecology to plant invasions. SIDA 22(1): 800, 2006 CAREX BICKNELLII (CYPERACEAE) NEW TO ARKANSAS Paul M.McKenzie C. Theo Witsell U.S. Fish and Wildlife Service Arkansas Natural Heritage Commission 101 Park DeVille Dr, Suite A 1500 Tower Building Columbia, Missouri 65203-0057, U.S.A. 323 Center St. paul_mckenzie@fws.gov Little Rock, Arkansas 72201, U.S.A. theo@arkansasheritage.org Joe Woolbright Ozark Ecological Restorations Inc. 04 Skilern Siloam Springs, Arkansas 72761, U.S.A. joewoolbright@cox-internet.com ABSTRACT We report the first record of Carex bicknellii Britt. tor Arkansas. The species was collected by the authors in 2003 and 2005 from Baker Prairie Natural Area, an unplowed tallgrass prairie remnant in Boone County, Arkansas. RESUMEN Se realiza | le Carex bicknellii Britt para Arkansas. | | lecto por los autores en 2003 y 2005 en Baker Prairie Natural Area | | Boone County, Arkansas. The range of Carex bicknellii Britt. extends from Maine and Manitoba south to Kansas and South Carolina, with the bulk of the distribution extending from Missouri and southeastern Kansas northward to southeastern Manitoba and the western end of Lake Erie (Rothrock & Reznicek 2001; Mastrogiuseppe et al. 2002). Disjunct populations occur in Maine, western Kansas, South Carolina, and the eastern Lake Ontario Region (Rothrock & Reznicek 2001). Habitat for the species includes dry to mesic prairies, rock, sand or serpentine barrens, open woodlands, openings in mesic to dry upland forests and bluff ledges, dry sandy or rocky fields, thickets, and less commonly along margins of ponds and fens (Yatskievych 1999; Rothrock & Reznicek 2001; Mastrogiuseppe et al. 2002). Orzell and Bridges (1987) and Smith (1988) reported C. bicknellii (as C. bicknellii var. bicknellii) for Franklin County, Arkansas but the specimen (Barber 861, UARK), which was the basis for the reports, was subsequently annotated by A.A. Reznicek as Carex bicknellii Britt. var. opaca FJ. Herm. J. Gentry, UARK, in litt. 21 October 2005). Hyatt (1988), Rothrock and Reznicek (2001), and Mastrogi et al. (2002) did not include Arkansas within the range of Carex bicknel Hii (either as C. bicknellii or as C. bicknellii var. bicknellii). Rothrock and SIDA 22(1): 801 — 804. 2006 802 BRIT.ORG/SIDA 22(1) Reznicek (2001) referenced all specimens within the C. bicknellii group col- lected in Arkansas through 2000 and determined that all prior reports of C. bicknellii (as C. bicknellii var. bicknellii) in Arkansas were based on other species. On 9 May 2005, the senior author discovered this species scattered in ir- regular colonies on dry mounds of Baker Prairie Natural Area,a 28 ha remnant of native tallgrass prairie in Boone County in north central Arkansas located near the western edge of the town of Harrison. On 10 May 2005, McKenzie and Witsell returned to the site to assess the overall distribution and abundance of the species on the area and to record plant associates. Carex bicknellii was asso- ciated with Baptisia bracteata Muhl. ex Elliott, Berlandiera texana DC, Carex bushii Mack., Ceanothus americanus L., Commandra umbellata (L.) Nutt., Dodecatheon meadii L., Euphorbia corollata L., Hypoxis hirsuta (L.) Cov, Luzula bulbosa (Alph. Wood) Rydb., Physalis sp., Schizachyrium scoparium (Michx.) Nash, Silphium integrifolium Michaux, S. laciniatum L., and Tradescantia ohiensis Raf. Plants were irregularly scattered over much of the drier portions of the prairie and extended to a few of the swales that drain the site. Carex bicknellii was initially collected on Baker Prairie in a wet swale by Witsell on ] June 2003 but the specimen was incorrectly identified as the closely related C. opaca (FJ. Herm.) PE. Rothrock & Reznicek. With the exception of a small (approximately + ha) former pasture along its eastern edge, Baker Prairie Natural Area is a high quality unplowed prairie remnant, and is jointly owned by the Arkansas Natural Heritage Commission and the Arkansas Field Office of The Nature Conservancy. It is the only rem- nant of what was once an approximately 2000 ha tallgrass prairie located west and south of present day downtown Harrison and is the largest known rem- nant of Ozark prairie that occurs on a chert substrate in Arkansas (Arkansas Natural Heritage Commission 2004). Baker Prairie Natural Area occurs on the Boone Formation of the Springfield Plateau which is characterized by lime- stone with embedded chert (Harper et al. 1981, Arkansas Natural Heritage Com- mission 2004). The Natural Area is underlain by Nixa and Noark very cherty silt loams which are deep, well-drained, moderately permeable soils that formed in residuum from cherty limestone (Harper et al. 1981; Arkansas Natural Heri- tage Commission 2004). Elevation on the natural area ranges from 354 to 378 m above mean sea level on slopes of 3 to 20 % (Arkansas Natural Heritage Com- mission 2004). Much of the natural area is gently rolling with scattered “pimple” mounds (Arkansas Natural Heritage Commission 2004). Baker Prairie includes dry-mesic chert prairie, mesic chert prairie (restricted to swales that bisect the area), and non-native grassland (Arkansas Natural Heritage Commission 2004). Major threats to the prairie prior to state ownership and restoration included lack of prescribed fire, excessive haying, the introduction of tall fescue (Festuca arundinacea Schreber) for grazing livestock, development pressure from adja- and — MCKENZIE ET AL., CAREX BICKNELLII NEW TO ARKANSAS 803 cent communities, and the encroachment of woody vegetation (Arkansas Natu- ral Heritage Commission 2004). Due to the prairie’ ownership and such current management practices as regular use of prescribed fire and the control of exotics, most historical threats have been eliminated. One possible threat to plant species inhabiting Baker Prairie, however, is the potential for reduced ge- netic fitness due to the geographic isolation of this prairie from other high qual- ity prairies in Arkansas and adjacent states. Following European settlement and prior to state ownership, haying and grazing were apparently the main management tools used to maintain the prai- rie (Arkansas Natural Heritage Commission 2004). Beginning in 1992, Baker Prairie Natural Area has been the site of aggressive management efforts that include the removal of encroaching woody vegetation and exotic species; seed collection and planting; brush hogging; herbicide treatment of Festuca arundinacea, Lespedeza cuneata (Dum.-Cours.) Don, Lonicera japonica Thunb,, Carduus nutans L., Albizia julibrissin Durazz, Daucus carota L., woody sprouts and stumps; and the use of prescribed fire (Arkansas Natural Heritage Com- mission 2004). These intensive management efforts have been successful in maintaining the rich botanical diversity of the area and restoring the ecologi- cal integrity of the prairie. In addition to Carex bicknellii, Baker Prairie pro- vides habitat for other rare plant species tracked as elements of conservation concern by the Arkansas Natural Heritage Commission. These include Antennaria neglecta E. Greene, Aster sericeus Vent., Calopogon oklahomensis D.H. Goldman, Carex gravida L. Bailey, Gentiana puberulenta J. Pringle, Nemastylis nuttallii Picker ex R. Foster, Silene regia Sims, Trillium pusillum Michaux var. ozarkanum (EJ. Palmer & Steyerm.) Steyerm., Viola pedatifida G. Don, and Zizia aptera (A. Gray) Fern.. Rare animals of conservation concern at the site include ornate box turtle [Terrapene ornata ornata (Agassiz)], grass- hopper sparrow [Ammodramus savannarum (Gmelin)], willow flycatcher [Empidonax traillii (Audubon)], and prairie mole cricket (Gryllotapa major Saussure). Voucher Spec imens. U.S.A. ARKANSAS. Boone Co.: Ar} N l Heritage Commission and TNC’s Baker Prairie Natural Area in western Harrison, ca. 0.4 km $ of the inter. of Industrial Dr. and Goblin Dr; TI8N,R20WS5 SW1/4NE1/4N W1/4; Gaither 7.5' Quad.; N 360° 14'40.2", W 93° 08'01.6" (Datum = NAD83 Zone 15); infrequent in scattered colonies on dry mounds of native prairie; in association with Baptisia bracteata, Berlandiera texana, Carex bushii, Ceanothus americanus, Commandra umbellata, Euphorbia corollata, Hypoxis hirsuta, Luzula bulbosa, Physalis sp., Schizachyrium scoparium, Silphium integrifolium, S. laciniatum, and Tradescantia ohiensis,; 9 May 2005 Paul M. McKenzie 2187 (MO, MICH, UMO); 10 May 2005 Theo Witsell 05-0330 (anhe-Arkansas Natural Heri- tage Commission, UARK, peh-pers. herb. of Philip E. Hyatt); 1 Jun 2003 Theo Witsell 03-0437 (anhc). Botanical nomenclature listed herein follows Yatskievych and Turner (1990) except for Calypogon oklahomensis that follows Yatskievych (1999), and Carex bicknellii and C. opaca that follow Rothrock and Reznicek (2001). BRIT.ORG/SIDA 22(1) ACKNOWLEDGMENTS We are grateful to Anton Reznicek of the ey of Michigan Herbarium, Ann Arbor, Michigan, andanan ny for their assistance with this report. REFERENCES ARKANSAS NaturAL Heritace Commission. 2004. Baker Prairie Natural Area management plan. Arkansas Natural Heritage Commission. Little Rock. Harper, M.D., D.H. Fowckes, and D.A. Howard. 1981. Soil survey of Boone County, Arkansas. U.S. Dept. of Agric. Soil Conservation Service, in cooperation with the Arkansas Agri- cultural Experiment Station. Hyart, PE. 1998. Arkansas Carex (Cyperaceae): a briefly annotated list. Sida 18:535-554. Mastroaiusepre, J., PE. RoTHROCK, A.C. Digste, and A.A. Reznicek. 2002. Carex Linneaus sect. Ovales Kunth, Enum. PI. 2:394. 1837.|n: PW. Ball, K. Gandhi, R.W. Kiger, D. Murray, J.L.Zarucchi, A.A. Reznicek, J.L.Strother, eds.Flora of North America North of Mexico, vol.23 Magnoliophyta: Commelinidae: Cyperaceae. Oxford Univ. Press, NY. Pp. 332-378. Orzett, S.L.and E.L.Brioces. 1987.Further additions and noteworthy collections in the flora of Arkansas, with historical, ecological, and phyteographical notes. Phytologia 64: 81-144 RorHrock, PE.and A.A.Reznicek. 2001.The taxonomy of the Carex bicknellii group (Cyperaceae) and new species for Central North America. Novon 11:205-228. Smith, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas. Second Ed. Published by the author. Fayetteville, AR. YATSKIEVYCH, G. 1999. Steyermark’s flora of Missouri- Vol. 1- Revised Ed. Missouri Depart- ment of Conservation, Jefferson City, MO. YatskievycH, G. and J. Turner. 1990. Catalogue of the flora of Missouri. Missouri Bot. Gard. Monogr. Syst. Bot. No. 37. Braun-Brumfield, Inc, Ann Arbor, MI. GREEN FRINGED ORCHID (PLA TANTHERA LACERA, ORCHIDACEAE) IN SOUTHERN LOUISIANA Charles Allen, Sara Thames, ward Anderson lll, Bill Newton Ill,and Rhonda Hampton George Fisher Colorado State Univ., Fort Polk 509 Bon Ami 1645 23° Street DeRidder, Louisiana 70634, U.S.A. Fort Polk, Louisiana 71459, U.S.A. ABSTRACT A new population of Platanthera lacera is reported for Vernon Parish, southern Louisiana. RESUMEN Se cita una nueva poblacion de Platanthera lacera de Vernon Parish, Sur de Louisiana. Green fringed orchid (Platanthera lacera (Michx.) G. Don) was reported for Loui- siana as early as 1852 by Riddell (1852). A specimen was collected in Shreveport in Caddo Parish by MacRoberts in 1976 (MacRoberts 1977). Apparently, this specimen was misidentified and not included in the orchid flora of Louisiana by Prigeon and Urbatsch (1977) and excluded by Thomas and Allen (1993). This species was reinstated to the Louisiana flora by MacRoberts and MacRoberts (1998): the authors also stated that the site for the collection in Caddo Parish was now a housing development. Green fringed orchid is reported from all of the eastern United States except for Florida (USDA, NRCS 2005). In Texas, this orchid is reported only from Bowie County (Correll 1947; Liggio & Liggio 1999). The habitats reported for this orchid include open sedge swamps and marshes, bogs, meadows, and glades of open woods, swampy woods and wet or occasionally dry open fields and prairies, and in thickets (Correll 1978; Liggio & Liggio 1999; Radlord et al. 1968). In Flora of North America Editorial Com- mittee (2002), this sp is reported from sphagnum bogs, alluvial and swamp forests, stream banks, riparian meadows, sand flats, moist and seeping slopes, prairies, roadside banks, ditches, old fields, and borrow pits. According to Liggio and Liggio (1999), this orchid has a preference for acidic soils but is indifferent to varying conditions of moisture, sunlight, and shade. On May 6, 2005, a new population of green fringed orchid was discovered in Vernon Parish, Louisiana which is ca 150 miles south of the Caddo Parish location and apparently the southernmost location in the United States. The collection data are: SIDA 22(1): 805 — 809. 2006 806 BRIT.ORG/SIDA 22(1 = Voucher specimen: LOUISIANA. Vernon Parish: along edge of mowed hunting trail, edge of pasture and beech/white oak forest in Section 14 T2S R7W, GPS N30.942 W93.053, ca. 1/4 mi SW of Talbert- Pierson Cemetery (Pine Grove Church) and ca. 5 mi NW of Sugartown in drainage of unnamed creek that drains into Whiskey Chitto Creek, 24 May 2005, Allen et al 19415, (BRIT, Fort Polk Herb., ULM). The habitat varied from an open mowed trail to the dense edge of the beech/ white oak forest of the creek edge and from moderately well drained to moist soil. The soil type is Gore which is a very fine sandy loam with 5 to 12 percent slopes (Soil Survey Staff 2003). The elevation in the area ranges from 150 to 160 feet. The plants had just begun to flower on May 6, seemed to peak around May 20, and just a very few flowers were noted on June 11, 2005. During May 2005, the site was revisited several times and 35 clumps of the orchid were noted; most clumps had a single flowering stem but a few had two flowering stems. The area surrounding each of the 35 clumps was examined and the five nearest individuals were identified including herbaceous, woody vine, shrub (woody non-vines shorter than six feet), shrubs/saplings (woody non-vines taller than six feet and five inch or less dbh), and trees (woody non- vines taller than six feet and larger than 5 inch dbh) species (Table 1). The per- cent of each species out of the 175 total individuals (five times 35) for each cat- egory is listed in Table |; for the herbaceous and woody vines only one percent is reported for each species and for the woody non-vine species, the first per- cent listed is the shrub, the second is the shrubs/saplings, and third is the trees. The surrounding area was examined for a distance of twenty feet and all asso- ciated species were identified and listed in Table 1. A total of 124 species from 90 genera and 56 families were identified as a nearest individual or associated species. Densiometer readings were taken at each clump and the average cover percent was 61.04% and ranged from 43.03% to 75.00%. The most common occurring nearest herbaceous species was Chasmanthium sessiliflorum (Poir.) Yates (17.71%) and was followed b Dichanthelium dichotomum (L.) Gould (12.57%), Sanicula canadensis L. (8.57%), Allium canaddense L. (8.00%), and Dichanthelium laxiflorum (Lam.) Gould (5.71%). The woody vine species that was most commonly nearest the orchid was loxicodendron radicans(L.) Kuntze (38.29%) and was followed by Lonicera japonica Thunb, (37.14%) and Parthenocissus quinquefolia (L.) Planch. 12.57%). The most common nearest shrub species was Rubus argutus Link (25.71%) and was followed by Quercus alba L. (8.57%), Callicarpa americana L. (7.43%), and Quercus hemisphaerica Bartr. ex Willd. (6.86%). The most common nearest tree species with smaller dbh (5 inch or less) were Pinus taeda L. (30.86%) and Ilex vomitoria Ait. 12.00%) and with the larger dbh (> 5inch dbh) were Liquidam- bar styraciflua L. (23.43%), Triadica sebifera (L.) Small (20.00%), and Quercus alba L. 17.14%). ares ALLEN ET AL PLATANTHERA Taste 1. List of species associated with 35 clumps of green fringed orchid in Vernon Parish, Louisi- ana, during May 2005 with percent! of nearest individual records. ACANTHACEAE Ruelli liniensis (J.F.Gmel.) Steud. (1.14%) ACERACEAE Acer rubrum L. (5.71%, 1.71%, ANACARDIACEAE Toxi ron radicans (L.) Kuntze (38.29%) 1.71%) APIACEAE Ptilimni apillaceum (Michx.) Raf. Sanicul densis L.(8.57%) AQUIFOLIACEAE llex opaca Ait. (1.14%, 0, 0) Ilex vornitoria Ait. (3.43%, 12.00%, 0) ARACEAE Arisaema triphyllum (L.) Schott ARALIACEAE Aralia spinosa L. (2.29%, 3.43%, 0.57%) ARISTOLOCHIACEAE Aristolochia serpentaria L. ASPLENIACEAE Asplenium platyneuron (L. ASTERACEAE Elephantopus carolinianus Raeusch. (2.86%) Helenium flexuosum Raf. / t aAnnaden B.S.P. er lid anadensi L. 8 So dace rugosa P.Mill. (1. Symphyotrichum eae (L.) A.& D. Love (2.29%) BETULACEAE Carpinus caroliniana Walt. (1.71%, 3.43%, 1.71%) Ostrya virginiana (P. Mill.) K. Koch (0, 1.14%, 0) a aan Big 1L. (3.43%) CAMPANULACEAE | icpilata heliaq annen ak CAPRIFOLIACEAE Lonicera japonica Thunb. (37.14%) Sambucus mye , cai: 2.29%, 0) Vib rife Vib dep ertune lt 14%, 2.29%, 0) CELASTRACEAE Euonymus americana L. (0.57%, 0, 0) CLUSIACEAE Hypericum hyperi oldes (L.) Crantz Hynericum mutilum L P , CORNA Cornus pa [ L.(0.57%, 0, 0) CUPRESSACEAE Juniperus virginiana ee (0, 0.57%, 0) CYPERACEAE Carex complanata Torr. & Hook. (2.86%) Carex debilis Michx. Carex digitalis Willd. Carex festucacea Schkuhr ex Willd. Carex flaccosperma Dewey (0.57%) Carex longii Mack. (0.57%) Carex rosea Willd. (1.14%) ge ola ot: sh ea iao laantha ree DIOSCOREACEAE Dioscorea villosa L. DRYOPTERIDACEAE Onoclea sensibilis L. Palystichum acrostichoides (Michx.) Schott. EBENACEAE Diospyros virginiana L. (0.57%, 0, 0) ERICACEAE eee eee Marsh. tii Chapman (1. 14%, 0, 0) EUPHORBIACEAE Triadica sebifera (L.) Small (1.14%, 8.00%, 20.00%) FABACEAE Galactia volubilis (L.) Britton Lespedeza cuneata (Dum.-Cours.) G.Don (0.57%) Trifolium repens L. (0.57%) Vicia sativa L.ssp. nigra (L.) Ehrh. (1.71%) FAGACEAE Fagus grandifolia Ehrh. (0, 0.57%, 10.29% Quercus alba L. (8.57%, 0.57%, 17.14%) Quercus hemisphaerica Bartr. ex Willd. (6.86%, 6.86%, 0) —s Quercus nigra L. (4.00%, 2.29%, 1.14%) HAMAMELIDACEAE Hamamelis virginiana L. (0, 1.71%, 0) Tage 1.continued BRIT.ORG/SIDA 22(1) Liquidambar styraciflua L.(3.43%, 5.14%, 23.43%) JUGLANDACEAE Carya alba (L.) Nutt. ex Ell. (0, 0, 0.57%) Carya glabra (P. Mill.) Sweet var. hirsuta (Ashe) Ashe (5.14%, 0, 3.43%) Carya illinoinensis (Wangenh.) K. Koch JUNCACEAE alice coriaceus Mack. ia _ 57%) epha I) Lincs nol JUNCUS 16 uis Willd. : 7 ne LAMIACEAE Pycnanthemum tenuifolium Schrad. Salvia lyrata L.(0.57% Scutellaria elliptica Muhl. ex Spreng. (0.57%) LAURACEAE Sassafras albidum (Nutt.) Nees (0.57%, 0, 0) LILIACEAE inad ie a 00%) im michauxii Polygonatum bi oe (Walt.) Ell. LINACEAE Linum striatum Walt. LOGANIACEAE Gelsemium sempervirens (L. YCOPODIACEAE ma St. Hil. (0.57%) i | lygodium japonicum (Thunb. ex Murr.) Sw. (0.57%) MELIACEAE Melia azedarach L. (0.57%, 2.29%, 0.57%) MORACEAE Morus rubra L. (0.57%, MYRICACEAE Morella cerifera (L.) Small (0.57%, 0.57%, 0) NYSSACEAE, Nyssa sylvatica Marsh. (4.57%, 0, 4.00%) OLEACEAE een saat us L.(0,0,0.57%) Ir. (4.57%, 8.00%, 0) 1.71%, Q) ONAGRACEAE liiduyinia ite ifaly L. Ludwigia palustris (L.) Ell. OROBANCHACEAE Epifagus virginiana (L) W. Bart. OSMUNDACEAE Osmijnda ci Osmunda regalis L. OXALIDACEAE Oxalis stricta L. PINACEAE Pinus taeda L. (0, 30.86%, 10.29%) POACEAE Andr dii Vitman (1.14% =< Briza minor L. hasmanthium laxum Chasmanthium sessi Worun (Poir.) Yates (17.71%) Dichanthelium acuminatum (Sw.) Gould & C.A Clark var. lindheimeri (Nash) Gould & CA. Clark Dichanthelium boscii (Poir.) Gould & C.A. Clark (4.00%) Dichanthelium commutatum (J.A. Schultes) Gould (4.57%) hanthelium dichot 7 (L.) Gould (12.57%) rum (Lam.) Gould (5.71%) Leersia virginica Willd. Lolium perenne L. (2.29%) Melica mutica Walt. is (L.) Beauv. (2.29% Paspalum u urvi ie Steud. Dich nth lium la if Oplist AEN IC Hirtell oe a (Michx.) Scribn. (0.57%) RHAMNACEAE Berchemia scandens (Hill) K. Koch (1.71%) edna Crataegus marshallii Eqglest. (0.57%, 0, 0) ae serotina Ehrh. (0.57%, 0, 0) Rubus argutus Link (25.71%, 0, 0) Rubus trivialis Michx. (3.43%, 0, 0) RUBIACEAE Galium pilosum Ait. oa ium tinctori um i. (4.00%) ichx. Mitchella repens L. . 86%) SMILACACEAE Smilax glauca Wall (2.29%) mila \ Walt. Smilax pu Smilax rotundifolia L. ALLEN ET Al 809 Tas_e 1.continued Smilax smallii Morong VERBENACEAE Smilax tamnoides L. Callicarpa americana L. (7.43%, 2.29%, 0) le salaries VIOLACEAE hii (Desv.) Morton Viola sororia Willd. (1.14%) ULMACEAE VITACEAE Ulmus alata Michx. (1.14%, 2.29%, 2.86%) Parthenocissus quinquefolia (L.) Planch. (12.57%) Ulmus americana L. (0,0, 1.71%) Vitis rotundifolia Michx. (4.00%) URTICACEAE Boehmeria cylindrica (L.) Sw. ' Percent of nearest individual records = number of nearest ae records divided by 175 and mul- tiplied by 100 for each peels Cae The categories aceous, woody vine, shrub (woody non-vines shorter than six ft t percent), ie ee non-vines taller than six feet and five inch or less dbh) eee percent), and trees (woody non-vines taller than six feet and larger than 5 inch dbh) (third percent). ACKNOWLEDGMENTS We would like to thank Larry Magrath and Charles J. Sheviak for reviewing our article. REFERENCES Correll, D.S. 1947. Additions to the orchids of Texas. Wrightia 1:166-181. Corrett, D.S. 1978. Native orchids of North America north of Mexico. Stanford University Press, Stanford, California. Fiora of NortH America EprtoriaL Committee. 2002. Flora of North America: Volume 26; Magnoliophyta: Liliidae: Liliales and Orchidales. Oxford Univ. Press, New York. Liccio, J.and A.O. Licaio. 1999. Wild orchids of Texas. Univ. of Texas Press, Austin. MacRosents, D.T. 1977. Additions to the Louisiana flora. Sida 7:220-222. MacRoserts, M.H. and B.R. MacRoserts. 1998. A note on Platanthera lacera (Michx.) G. Don (Orchidaceae) in Louisiana. Phytologia 84:38-39. Priccon, A.M.and L.R. Ursatscu. 1977. Contributions to the flora of Louisiana. ||: Distribution and identification of Orchidaceae. Castanea 42:293-304. Ravroro, A.E., H.E. Antes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolina. Univ. North Carolina Press, Chapel Hill. Riovett, J.L. 1852. Catalogus Florae Ludovicianae. New Orleans Med. Surg. J. 8:734—754. THomas, R.D. and C.M. Auten. 1993. Atlas of the vascular flora of Louisiana, Vol. 1:ferns and ferns allies, conifers, and monocotyledons. Louisiana Department of Wildlife and Fish- eries, Baton Rouge. Soi Survey Starr. 2003. Soil survey of Vernon Parish, Louisiana. United States Department of Agriculture, Natural Resources Conservation Service, Washington, D.C USDA, NRCS. 2005. The PLANTS Database, Version 3.5 (http://plants.usda.gov). National Plant Data Center, Baton Rouge, LA 70874-4490 USA. BRIT.ORG/SIDA 22(1) Book Notices Timber Press Books (Reviews Forthcoming) Davip Fross and Dirter WILKEN. 2006. Ceanothus. (ISBN 0-88192-762-7, 978-0- 88192-762-7, hbk.). Timber Press Inc, 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax.). $39.95, 272 pp., color photos, b/w illustrations, 7" x 9". Contents.—Pretace, Acknowledgments. 1) Ceanothus in the Garden and Landscape (Natural Distri- bution, History of Cultivation, Growing Ceanothus Disease and Pests, Cultivar Development, Propa- gation, Ceanothus pecs ane Cultivars tox the Gi ina and L peeaes Ceanothus Selection Guide. 2) i inthe I C ; stribution, Form and Function in Ceanothus. s, Other Ceanodus Names). C onversion Tables, Glossary, Bibliography, Index. aS Davin D. Stuart. 2006. Buddlejas. (ISBN 0-88192-688-4, 978-0-88192-688-0, hbk). Royal Horticultural Society Plant Collector Guide. Royal Horticultural Society and Timber Press Inc, 133 S.W. Second Avenue, Suite 450, Portland, OR 97204- 3527, US.A. (Orders: www.timberpress.com, mail@timberpress.com, 503- 227-2878, 1-800-327-5680, 503-227-3070 fax.). $34.95, 192 pp., color pho- tos. a. Contents.—Foreword by Michael A. Dirr, Preface, Introduction. 1) Species. 2) Hybrids and Cultivars. 3) Care and Cultivation. 4) Propagation. 5) Wildlife Attracted to Buddlejas. Where to See Buddlejas, Where to Buy Buddlejas, Bibliography, Index. RoBIN Wuite. 20060. Daphnes: A Practical Guide for Gardeners. (ISBN 0-88192- 752-X, 978-0-88192-752-8, hbk.). Timber Press Inc, 133 S.W. Second Avenue. Suite +50, Portland, OR 97204-3527, U.S.A. (Orders: wwwtimbet press.com mail@timberpress.com, 503-227-2878, l-800-327-5680, 503-227-3070 fax.). $34.95, 232 pp., color photos, 7" x 9". Contents.—Foreword, Preface, 1) History, Taxonomy and Mor = hology. 2) Daphne Species. 3) Daphne Hybrids. 4) Propagation. 5) Cultivation. 6) Pests and Diseases. 7) Garden Use. Useful Addresses. Glos- sary, Bibliography, Index of Daphnes, Conversion Tables. SIDA 22(1): 810. 2006 SEYMERIA FALCATA (SCROPHULARIACEAE), A NEW RECORD FOR TEXAS AND THE UNITED STATES Joselyn Fenstermacher Sul Ross State University Alpine Texas 79832, U.S.A. josfenster@alumni.duke.edu ABSTRACT Upon review of herbari i | Hlecti itis clear that Seymeria falcata B.L. Turner irst report of occurs in the Dead Horse Mouncats of southern Brewster County, Texas. This is the f this species for Texas and the United States. RESUMEN Basandonos en la revision de especimenes herbario y colecciones recientes esta claro que Seymeria falcata B.L. Turner ocurre en las Montanas Caballos Muertos en el condado meridional de Brewster, Tejas. Este es ne primera cita de esta especie para Tejas y los Estados Unidos Turner (1982) provided a revisional study of the largely Mexican genus Seymeria. In this he described S. falcata B.L. Turner, a new species then known only from north-central Mexico, partially sympatric with its closest ally S. scabra Gray. The latter is typified by material collected by Charles Wright in the Davis Mountains of Jeff Davis County, Texas, and is distinguished by mostly symmetrical, semi-falcate fruits and markedly scabrous stems and foliage. In contrast, S. falcata has weakly to decidedly falcate fruits and minutely puberulous or glabrous stems and foliage, and glabrous fruits. Turner et al. (2003) recognized only a single species of Seymeria, S. scabra, as occurring in Trans-Pecos Texas. Recent collections from the Dead Horse Mountains in the Sierra Del Carmen of Big Bend National Park have revealed the presence there of S. falcata. Voucher specimens: TEXAS. Brewster Co.: Dead Horse Mountains, on rocky ridge above Brushy Can- yon, ca. 1/2 mi N of lodge, 1220 m, 15 Aug 1994, Talbot 137 (SRSC). Dead Horse Mountains, north sister of Sue Peaks at crest, 1740 m, 16 Sep 1994, Talbot 255 (SRSC); western slopes of Dead Horse Mountains below Sue Peaks, 1420 m, 3 Sep 2005, Fenstermacher 1187 (SRSC); Dead Horse Mountains, 5 miinon Telephone Canyon Trail from Old Ore Road, 1310 m, 8 Mar 2005, Fenstermacher 571(SRSC). Seymeria falcata was first collected by Miller Talbot in the Dead Horse Moun- tains but his 1994 specimen was identified as Seymeria scabra. My recent col- lections of S. falcata from the Dead Horse Mountains spurred a review of the Seymeria specimens in the SRSC herbarium, thereby discovering and correctly identifying Talbot's previous collection. My S. falcata collections were made at SIDA 22(1): 811 — 812. 2006 812 BRIT.ORG/SIDA 22(1) two locations in relative proximity to Talbot's collection sites, at elevations be- tween 1310 m and 1430 m (Fenstermacher 571, 1187, SRSC). | also collected S. scabra in a range of elevations in proximity to all collections mentioned here (Fenstermacher 744D, 1018, 1050, SRSC). The two species are clearly sympatric and hybridization between the two is suspected (Fenstermacher 1018, 1050, SRSC) ACKNOWLEDGMENTS | would like to acknowledge the generosity of Billie L. Turner and thank him for bringing this to my attention. Thanks also go to the Sul Ross State Univer- sity herbarium for research space, holdings, and technical expertise, without which this discovery would not have been made. The two reviewers, Guy L. Nesom and Richard D. Worthington, made helpful suggestions. REFERENCES Turner, B.L. 1982. Revisional treatment of the Mexican species of Seymeria (Scrophulariaceae). Phytologia 51:403-422. Turner, B.L., H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Sida Bot. Misc. 24, Volume 1.Botanical Research Institute of Texas, Fort Worth. CAYRATIA JAPONICA (VITACEAE) NEW TO NORTH CAROLINA AND AN UPDATED KEY TO THE GENERA OF VITACEAE IN THE CAROLINAS Alexander Krings Robert J. Richardson Herbarium, Department of Botany Department of Crop Science North Carolina State University North Carolina State University Raleigh, North Carolina 27695-7612, U.S.A. Raleigh, North Carolina 27695-7620, U.S.A. Alexander_Krings@ncsu.edu Rob_Richard ABSTRACT A population of the introduced and expanding Cayratia japonica (Thunb.) Gagnep. (Vitaceae) is reported for North Carolina. Voucher specimens are cited and an updated key to the genera of Vitaceae in the Carolinas is presented. RESUMEN Se cita de Carolina del N blacién de | 1 nexpansion Cayratid japonica (Thunb.) Gagnep. (Vitaceae). Se ae los pliegos tstigs y se presenta una clave puesta al dia de los géneros de Vitaceae en las Carolinas. The Asian vine Cayratia japonica (Thunb.) Gagnep. var. japonica (Vitaceae) is known from Louisiana, Mississippi, and Texas, but has not been previously re- ported escaped elsewhere in the United States (Shinners 1964; Brown 1992; Al- ford 2003; USDA, NRCS 2005). The species was not previously reported for North Carolina by Radford et al. (1968), Pittillo et al. (1969), Pittillo et al. 1972), Kral (1981), Pittillo and Brown (1988), or USDA, NRCS (2005). However, an aggres- sive and persistent population of the plant was recently encountered on pri- vate property in a suburban area near Winston-Salem (Forsyth Co., NC). Vines were growing up trees (reaching heights of 9-12 m high), along a fence, and generally throughout the landscape—covering about 0.5 acre. When visited in late August, the vines were in full flower and well-attended by honeybees and wasps. Based on conversations with the landscaper, individual ramets aggres- sively re-sprouted following handweeding. Apparently, a vine of > 2.5cm diam at the base had been removed. Remaining roots at the site were ca. 0.6 cm diam. It remains unclear how the species arrived. Both previous and current homeowners, the latter an avid birdwatcher, indicated that the species had not been planted. However, intentional planting or contamination through other ornamental plantings cannot be ruled out. Bird dispersal may be a possibility, although unlikely as additional populations between North Carolina and the Gulf Coast would be expected under this scenario and reproduction is reported to be primarily vegetative in North America—at least in Texas (Brown 1992). SIDA 22(1): 813 - 815. 2006 BRIT.ORG/SIDA 22(1) Although the vector of arrival remains unknown, the persistence of the North Carolina population is of concern. Alford (2003) reported a Mississippi popu- lation tripling in size within LO months. Little information is currently avail- able regarding control measures. Voucher specimens: U.S.A. NORTH CAROLINA. Forsyth Co.: Wesleyan Lane, Winston-Salem, 1] Aug 2005, Jonathan Todd s.n.ONCSC, USCH). Wesleyan Lane, Winston-Salem, 29 Aug 2005, Robert]. Rich- ardson s.n. (FLAS, NCSC, USCH). Cayratia japonica (Fig. 1) can be distinguished from other Vitaceae taxa in the Carolinas by the pedate quinqueloliate leaves. Tendrils are bifurcating and not disk-tipped. An updated key to the genera of Carolina Vitaceae follows. Cissus is included here based on the listing of C. trifoliata (L.) L. by Weakley (2005). Cissus trifoliata is cultivated in North Carolina, but-has not been observed to escape (Krings, pers. obs.). 1. Tendrils, at least some, terminating in adhesive disks 1. Tendrils lacking adhesive disks 2. Leaves pedate quinquefoliate 2. Leaves simple, palmately, or pinnately compound, but never pedate quinquefoliate. 3. Inflorescence a thyrse; petals connate at their apices Vitis L. 3. Inflorescence a dichotomous or umbelliform cyme; petals free at their apices. 4. Cyme dichotomous; flowers 5-merous 4. Cyme umbelliform; flowers 4-merous Parthenocissus Planch. Cayratia Juss. OQ. Ampelopsis Michx. Cissus L. ACKNOWLEDGMENTS We thank Jonathan Todd for bringing the Forsyth population to our attention and the curators and staff of DUKE and NCU for access to their collections. Guy Nesom offered helpful review comments. REFERENCES Atrorb, M.H. 2003. Noteworthy collections Mississippi Castanea 68:93. Brown, L.E. 1992. Cayratia japonica (Vitaceae) and Paederia foetida (Rubiaceae) adventive in Texas. Phytologia 72(1): 45-47. Kral, R.1981.Some distributional reports of weedy or naturalized foreign species of vas- cular plants for the southern states, particularly Alabama and middle Tennessee. Cas- tanea 46:334-339. Pittito, J.D., JH. Horton, and K.W. Greentee. 1969. Additions to the vascular flora of the Caro- linas. |. J. Elisha Mitchell Sci. Soc. 85:18-22. Prrmitto, J.D., JH. Horton, and K.E. Herman. 1972. Additions to the vascular flora of the Caroli- nas. ll. J. Elisha Mitchell Sci. Soc. 88:144-152 Pirtito, J.D.and A.E. Brown. 1988. Additions to the vascular flora of the Carolinas. Ill.J. Elisha Mitchell Sci. Soc. 104:1-18. Raprorod, A.E., H.E. AHves, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. The University of North Carolina Press, Chapel Hill. KRINGS AND RICHARDSON, CAYRATIA JAPONICA NEW TO NORTH CAROLINA foli leaf 1 bif Jril: B. inflorescence: C. detail Hy ec 14:-L t 1 dol} / | 1) ond €] hid-M dick (ctl 1). Leaf, flower bud, on? (NCU); 4ril 1 inf] I d S fell 1580 (NCU). SHinners, L.H. 1964. Cayratia japonica (Vitaceae) in southeastern Louisiana: new to the United States. Sida 1:384. USDA, NRCS. 2005. The PLANTS Database, Version 3.5 (http://plants.usda.gov). Data com- piled from various sources by Mark W. Skinner. National Plant Data Center, Baton Rouge. Weaxtey, A.S.2005.Flora of the Carolinas, Virginia,and Georgia:Working draft, 10 June 2005. North Carolina Botanical Garden, Chapel Hill. BRIT.ORG/SIDA 22(1) Book NOTICES Blackwell Publishing Kenpaut R. LaMkKey and Micnaet Ler. (Eds.). 2006. Plant Breeding: The Arnel R. Hallauer International Symposium. (ISBN 0-8138-2824-4, 978-0-8138-2824- 4, hbk.). Black well Publishing, 2121 State Ave., Ames, [A 50014-8300, U.S.A. and 9600 Garsington Road, Oxford, OX4 2DQ, UK. (Orders: 515-292-0140, 515-292-3348 tax 1-800- oe 6657, www.blackwellprofessional.com, lers@ blackwell hing.com). $149.99, 379 pp., numerous illus- trations, 7" < LO", Douc-as D. Stokke and Lestir H. Groom. 2006. Characterization of the Cellulosic Cell Wall. (ISBN 0-8138-04 39-6, 978-0-81 38-04 39-2. hbk.). Black well Pub- lishing. 2121 State Ave., Ames, [A 50014-8300, U.S.A. and 9600 Garsington Road, Oxford, OX4 2DQ, UK. ue 515- fatal 0140, 515-292-3348 fax l- 800-862-6657, www.black well ional.com, ord blackwell- publishing.com). $149.99, 274 Dp, color photos, numerous b/w illustrations, 7" X 10" JouN F Lesuir and Brett A. SUMMERELL. 2006. The Fusarium Laboratory Manual. (ISBN 0-8138-1919-9, 978-0-8138-1919-8, spiral pbk.). Blackwell Publishing. 2121 State Ave., Ames, IA 50014-8300, U.S.A.and 9600 Garsington Road, Oxford, OAT 2ZDO.U - LOnderss ee 292-0140, 515-292-3348 fax 1-800-862-6657, blacl ional.com, orders@ blackwellpublishing.com). VV VV $124.99, 388 * b/w Tiveeetione Ole From the publisher—For the first time in over 20 years, a comprehensive collection of photographs and descriptions of species in the fungal genus Fusarium is available. This laboratory manual pro- vides an overview of the biology of Fusarium and the techniques involved in the isolation, identi cation and characterization of individual species and the populations in which they occur. It is the ogical and molecular approaches have been incorporated into a vol first time that genetic, morphol h new and ume devoted to Fusarium identification. The authors include descriptions of species, bot old, and provide protocols for genetic, morphological and molecular identification techniques. SIDA 22(1): 816. 2006 NEW AND NOTEWORTHY COLLECTIONS FOR ARKANSAS James H. Peck Brett E.Serviss Department of Biology Department of Biology University of Arkansas at Little Rock Henderson State University Little Rock, Arkansas 72204, U.S.A. bags ia, Arkansas 71999, U.S.A. jhpeck@ualredu visb@hsu.edu ABSTRACT Ten species of vascular plants, all native or naturalized in the soutaeastern United States, are re- ported as new or noteworthy for Arkansas. Of these, fi -native. Seven are reported as state records, and the accounts for the other three species provide clarifications, includ- ing updates on the status of two aquatic weeds, one federally listed as an invasive non-native weed (Hydrilla verticillata) and another erroneously thought to be native to the United States (Salvinia ma). Species reported as a state record are: Amaranthus blitum, Chamaesyce hypercifolia, Chamaesyce opthalmica, Euphorbia graminea, Hedyotis corymbosa, Ipomoea quamoclit, and 4 {] | FAFriLClare AYLUT LUM RESUMEN Se citan diez especies de plantas vasculares lizad te de los Estados dos bl Ark Uni i rkansas. De estas, Ci i y cl i Siete se acts : snes ] lds citan para el estado, y par oe estatus de dos ee acuaticas, una ade ™ as del listado federal de plantas 1 invasoras no nativas minima). Las especies citadas como nuevas ve ara el estado son: Amaranthus blitum, Chamaesyce hypercifolia, Cl : 1, Hedyotis corymbosa, Ipomoea quamoclit, [ fo) y Panevan oridana. INTRODUCTION The Arkansas Vascular Flora Project is an on-going collective effort to produce a complete and modern treatment to the vascular flora of Arkansas in the form of achecklist, atlas, and manual (Peck 2003). Toward that end, urban and rural parks, transportation routes, and other ruderal areas in Pulaski Co. and else- where in central Arkansas were surveyed to test the completeness of the floris- tic record. The state occurrence and state county distribution of species col- lected was determined from dot-maps kept at Herbarium LRU, University of Arkansas at Little Rock. The national occurrence and distribution of the new state records was determined from the national flora database kept by NRCS (1999). Records at the state and county level were documented with herbarium specimens. The records reported here support the notion that lesser quality habitats, such as urban and rural disturbed or ruderal habitats, whether xeric ones along railroad tracks or aquatic ones such as mudflats along impound- ments, are under-collected. Although less pristine and aesthetic than non-an- SIDA 22(1): 817 — 820. 2006 818 BRIT.ORG/SIDA 22(1) thropogenic habitats, ruderal environments warrant further survey in Arkan- sas to ensure completeness of the floristic record. SPECIES NEW OR NOTEWORTHY FOR ARKANSAS Amaranthus blitum L. (Amaranthaceae). Purple amaranth, a US non-native, occurs across the coastal states from Massachusetts south to Florida and west to Louisiana and Texas. This is the first documentation of this species in Ar- kansas. Voucher specimen: Pulaski Co.: urban ruderal habitat, along railroad tracks at University Ave. and 65" St, Little Rock, 13 Oct 2001, Peck 2001231 (LRU). Chamaesyce hirta (L.) Millsp. (Euphorbiaceae). Pillpod sandmat occurs across the coastal states from South Carolina south to Florida and west to Louisiana and Texas. Known from Bradley and Pope Cos., Arkansas. This is the third docu- mentation of this species in Arkansas. Voucher specimens: Pulaski Co.: urban ruderal habitat along railroad tracks at University Ave. and 65" St. Little Rock, Ll Nov 2001, Peck 20014892, 2001499, 20014895, 2001503 (LRU). Chamaesyce hypercifolia (L.) Millsp. Euphorbiaceae). Graceful sandmat occurs across the southeastern United States west to Louisiana and Texas. This is the first documentation of this species in Arkansas. Voucher specimens: Pulaski Co.: urban ruderal habitat along railroad tracks at University Ave. anc 65" St, Little Rock, 10 Nov 2001, Peck 2001490, 2001496, 2001500, 2001504 (LRU). Chamaesyce opthalmica (Pers.) Burch (Euphorbiaceae). Florida hammock sandmat occurs in Georgia and Florida west to Louisiana. This is the first docu- mentation of this species in Arkansas. Voucher specimens: Pulaski Co.: urban ruderal habitat along railroad tracks at University Ave. and 65" St. Little Rock, 10 Nov 2001, Peck 2001488, 2001497, 2001502 (LRU Euphorbia graminea Jacq. (Euphorbiaceae). Grassleal spurge, a US non-native, occurs as escapes in Florida and Hawaii. This is the first documentation of this species in Arkansas. — Voucher specimens: Pulaski Co.: urban ruderal habitat, along railroad tracks at Barton state fair grounds, Roosevelt Rd., Little Rock, 1] Nov 2001, Peck 2001567 (LRU): ruderal Saita near railroad tracks, east of University Ave. at 65" St, Little Rock, 4 Sep 1999, Peck 99764 (LRL Hedyotis corymbosa (L.) Lam. [Oldenlandia corymbosa L_] eee: Flat- top mille graines is a coastal species from South Carolina south to Florida and west to Louisiana and Texas. This is the first documentation of this species in Arkansas. Voucher specimens: Jefferson Co.: urban ruderal habitat, along railroad tracks at Old 167 in Pine Bluff near Arkansas River, 14 Oct 2001, Peck 2001243. Pulaski Co.: ruderal habitat, along railroad tracks at University Ave. and 65" St., Little Rock, 13 Oct 2001, Peck 2001232 (LRU). Saline Co.: ruderal ditat, along railroad tracks, S side of Benton, 13 Oct 2001, Peck 2001236 (LRU). — ha PECK AND SERVISS, NOTEWORTHY COLLECTIONS FOR ARKANSAS 819 Hydrilla verticillata (L. f.) Royle (Hydrocharitaceae). Waterthyme, United States non-native and an aggressive invasive species in the southeastern United States, especially from Florida west to Texas, was first documented from Arkansas by Parker and Serviss (2003). The present collections further document the spread of this species downstream along the Ouachita River impoundments to other counties in central Arkansas. Voucher specimens: Garland Co.: stranded on mudflat by drawdown event of Ouachita River, shore- line at boat ramp park at N side of Carpenter Dam, Lake Hamilton, 23 Nov 2004, Peck 2004248 (LRU); stranded on mudflat by drawdown event of Ouachita River, shoreline at Spencer’s Bay, mouth of Gulpha Creek, Lake Catherine, 23 Nov 2004, Peck 2004240 (LRU); stranded on mudf a by draw- down event on Ouachita River, Lake Ouachita, shoreline at Brady Mountain campground, end of CR 119, 9 Dec 2004, Davis s.n. (LRU). Hot Spring Co.: stranded on mudflat by Eerie n event of Oua- chita River, shoreline of Lake Catherine, Lake Catherine State Park, 23 Nov 2004, Peck 2004243 (LRU). Montgomery Co.: stranded on mudflat by drawdown event of Ouachita River, shoreline at Big Fir Campground, Lake Ouachita, end of CR 56, 23 Nov 2004, Peck 2004262 (LRU). Ipomoea quamoclit L. (Convolvulaceae). Cypressvine is native to tropical America, but is known to escape from cultivation across the southeastern United States, and is known from all states surrounding Arkansas. This collec- tion documents the first occurrence of this species in Arkansas. Voucher specimen: Pulaski Co.: urban ruderal habitat, present for three years as escape at informal ae grounds along railroad tracks at Barton state fair grounds, Roosevelt Rd., Little Rock, 14 Nov 2005, Peck 2005009 (LRU). Parietaria floridana Nutt. (Urticaceae). Florida pellitory is a coastal species that occurs from Delaware south to Florida and west to Louisiana and Texas. This is the first documentation of this species in Arkansas. Voucher specimens: Garland Co.: ruderal habitat along roadside, 4 mi E Hot Springs on 270, 22 May 2002, Peck 2002043 (LRU). Grant Co.: ruderal habitat along roadside, 3 mi E Poyen, 22 May 2002, Peck 2002032 (LRU). Hot Spring Co.: ruderal habitat along roadside, 4 mi S Malvern on 9, 22 May 2002, Peck 2002043 (LRU). Pulaski Co.: urban ruderal habitat along roadside in Boyle Park, Little Rock, 22 May 2002, Pec : 2002018 (LRU). Saline Co.: ruderal habitat along roadside, 4 mi S Benton on 25, 22 May 2002, Peck 2002054 (LRU). Salvinia minima Baker (Salvinieaceae). Least waterspangles, a non-native US aquatic fern, occurs in the southeastern United States from South Carolina south to Florida and west to Louisiana and Texas (Peck, 2002). The species was first documented from Arkansas by Peck (1999, 2002). These recent collections docu- ment a westerly range extension to three counties in west-central Arkansas. Voucher specimens: Garland Co.: stranded on mudflat by drawdown event of Ouachita River, shore- line at boat ramp park at N side of Carpenter Dam, Lake Hamilton, 23 Nov 2004, Peck 2004250 (LRU); stranded on mudflat by drawdown event of Ouachita River, shoreline at Spencer's Bay, mouth of Gulpha Creek, Lake Catherine, 23 Nov 2004, Peck 2004242 (LRU); stranded on mudflat by draw- down event on Ouachita River, Lake Ouachita, shoreline at Brady Mountain Campground, end of CR 119, 23 Nov 2004, Peck 2004260 (LRU). Hot Spring Co.: stranded on mudflat by drawdown event of Ouachita River, shoreline of Lake Catherine, Lake Catherine State Park, 23 Nov 2004, Peck 2004245 820 BRIT.ORG/SIDA 22 _~ 1) (LRU). Montgomery Co.: stranded on mudflat by drawdown event of Ouachita River, shoreline at Big Fir Campground, Lake Ouachita, end of CR 56, 23 Nov 2004, Peck 2004264 (LRU). ACKNOWLEDGMENTS The authors thank Theo Witsell and one anonymous reviewer for making im- provements to the manuscript. REFERENCES NRCS, Nationat Resources CONseRVATION Service. 1999. The PLANTS database. [http:// plants.usda.gov/plants]. USDA, National Plant Data Center, Baton Rouge, LA . Parker, M. and B.E. Seaviss. 2003. Occurrence and status of Hydrilla verticillata (L. f.) Royle Hydrocharitaceae) in Arkansas. J. Arkansas Acad. Sci. 57:15. Peck, J.H. 1999. Salvinia minima in Arkansas. Amer. Fern. J.89:215-216. Peck, JH. 2002. Survey of Salvinia (Salviniaceae) in eastern Arkansas. J. Arkansas Acad. Sci. 55:115-118. Peck, J.H.2003. Arkansas fl iditi instat ts,exclusions, and re-exclusions. Sida 20:1737-1757. — CLEMATIS MOREFIELDII (RANUNCULACEAE) NEW TO TENNESSEE Dwayne Estes Chris Fleming The Herbarium Breedlove, Dennis, Young suean Inc. Dept. of Ecology & i cee Biology 133 Holiday Court / Suite 2 The University of Tennessee Franklin, Tennessee 3706/7, 7 A. Knoxville, Tennessee 3 USA cfleming@bda-inc.com tnplants@yahoo.com ABSTRACT Clematis Cue (Morefield’s leather flower, Huntsville vasevine), a ieedetaly endanseree ape Ta nted for the flora of Tennessee for the first time a small ae ravine ee on the lower slopes of the western seionied of the Guabatiad Plateau in Franklin County, Tennessee. A general description of the habitat and associated flora is included. RESUMEN Clematis morefieldii (flor de cuero de Moretield,), especie en peligro, se documenta para la !lora de Sarasa BOE Paine. vez. Se descubri ieron dieciocho plantas a lo largo de un pequeno barranco en lental del Cumberland Plateau en Franklin County, Tennessee. Se incluye una descripcion pencil del habitat y de la flora asociada. INTRODUCTION Clematis morefieldii Kral (Morefield’s leather flower, Huntsville vasevine) is a federally endangered species previously thought to be endemic to Madison and Jackson counties, Alabama (Kral 1987; USFWS 1992; Pringle 1997). This rela- tively recently described species (Kral 1987) is similar to the widespread C. viorna but differs from all variants of that species in the cobwebby tomentose to villous pubescence of its stems, and in having bracts at or very near the base of the peduncle rather than well above the base (Pringle 1997). Clematis morefieldii also usually differs from C. viorna in having more flowers clustered in the leaf axils on shorter peduncles, in its sepals more pinkish and greenish- tinged, and abaxial leaf surfaces more densely pubescent (Kral 1987). As of 2003, C. morefieldii had been documented from ca. 10 populations, and currently at least two of these (including the type locality) are considered extirpated. All known C. morefieldii sites occur along the dissected western es- carpment of the Cumberland Plateau in rocky limestone woods on mostly south or southwest-facing slopes. These sites are typically dominated by Juniperus virginiana L. and various hardwood species characteristic of basic soils, includ- ing Cotinus obovatus Raf., which isa key indicator species for C. morefieldii (Kral SIDA 22(1): 821 — 824. 2006 822 BRIT.ORG/SIDA 22(1) 1987). This habitat type, referred to by Kral (1987) as the Cotinus association, is restricted to the lower slopes of the western escarpment of the Cumberland Plateau in northeastern Alabama and southeastern middle Tennessee. Kral (1987) suggested that further field exploration of suitable habitats within this community type in Alabama and Tennessee could result in the discovery of additional populations. In spring 2003, we began searching lor C. morefieldiiin southeastern middle Tennessee close to the nearest known Alabama population of C. morefieldii.On 5 Jun 2003, we discovered a small population of C. morefieldii in Franklin County, Tennessee ca. 12.5 km northeast of the nearest Alabama population. This is the first report lor C. morefieldii from Tennessee. Voucher specimen: Franklin Co.: ca. 4.2 mi SE of Huntland (Beans Creek Quad), SE of Motlow Cove in headwaters of Wilhite Creek, N side of Blooin Hollow, cela Fee small seasonal stream in rocky limestone woods, 5 Jun 2003, D. Estes & C. Fleming 04795 N). DISCUSSION The Franklin County, Tennessee site is on private property approximately 6 km ESE from the town of Huntland on the dissected western escarpment of the Cumberland Plateau in the Elk River watershed of the Tennessee River Ba- sin. The population is located along the lower portion of a SSW-facing slope over Monteagle limestone at an elevation of ca. 347 m (1140 ft). Eighteen plants, one in flower and one with immature fruit, were found. Most of the vines were 0.5 m or less in length and apparently damaged by insect herbivory. The indi- viduals were scattered along a 20 m reach of a wet weather conveyance 1-2 m in width, growing both along the banks and among the boulder substrate within the channel. The surrounding forest contained the following associates: Acer saccharum Marsh., Carya carolinae-septentrionalis (Ashe) Engl. & Graebn., Celtis tenuifolia Nutt, Fraxinus americana L., F quadrangulata Michx., Quercus muehlenbergii Engelm., Juniperus virginiana L., Cotinus obovatus, Cercis canadensis |. Ostrya virginiana (P. Mill.) K.Koch, Forestiera ligustrina (Michx.) Poir, Hamamelis virginiana L., Hypericum frondosum Michx., Rhus aromatica Ait., Bumelia lycioides (L.) Pers., Symphoricarpos orbiculatus Moench, and the rare Neviusia dlabamensis Gray and Viburnum bracteatum Rehd. The most common herbaceous taxa observed were Dasistoma macrophylla (Nutt.) Raf. Polymnia canadensis L., Scutellaria ovata Hill, and Solidago auriculata Shuttlw ex Blake. Several species considered rare in Tennessee were discovered growing with or adjacent to the Clematis population, including a small population of the state- endangered (TNHP 2004) Viburnum bracteatum (Estes & Fleming 04788 TENN). One shrub was growing at the edge of the wet weather conveyance con- taining C. morefieldii and another group of 15-20 individuals were ca. 100 m TO TENNESSEE 823 away along the rocky banks of a nearby stream. This species is extremely rare throughout its range and is endemic toa small area centered on the Cumberland Plateau of northeastern Alabama, northwestern Georgia, and southeastern Middle Tennessee. This is the second reported Tennessee occurrence for V. bracteatum(C. Bailey, Tennessee Natural Heritage Program, pers. comm.). Out- side of Tennessee, the species is only known from ca. eight extant populations. There is one each in Floyd and Walker counties in Georgia J. Allison, Georgia Natural Heritage Program, pers. comm.), and six documented populations in Alabama (Etowah, Jackson, and Madison counties), although one of these has not been observed in more than 50 years (M. Barbour, Alabama Natural Heri- tage Program, pers. comm.). Another significant find at the site was a small population of the state- threatened (TNHP 2004) Neviusia alabamensis (Estes & Fleming04790 TENN). Approximately 30-40 stems were located along a rocky streambank in asso- ciation with C. morefieldii. The Neviusia, like the Clematis, was visibly impacted by herbivory. Our collection represents a new county record for N. alabamensis in the state and the first report for the species from the Tennessee portion of the Cumberland Plateau; however, other populations are known fror the Cumberland Plateau of Alabama (Long 1989; D. Estes, pers. obs.) and Georgia (Long 1989). There are nine previously reported Tennessee populations (C. Bailey, Tennessee Natural Heritage Program, pers. comm.) within the Central Basin Section or at the transition zone between the Central Basin and Highland Rim Sections of the Interior Low Plateaus (ILP) Physiographic Province (Horn & Somers 1981; Long 1989). The Franklin County station is ca. 18 km southeast of the nearest previously known Tennessee population in Moore County. Other rare taxa tracked by the Tennessee Natural Heritage Program (2004) found within 100 m of the C. morefieldii site were Juglans cinerea L. Tennessee threatened), Lonicera dioica L. Tennessee threatened), Solidago auriculata (Ten- nessee threatened), and Cotinus obovatus (Tennessee special concern). The Cotinus and Solidago were common components of the forest while only one individual was found of J. cinerea and L. dioica, respectively. SUMMARY The recent discovery of C. morefieldii in Tennessee not only represents a state record but also extends the range of the species by some 12.5 km to the north- east. Recent fieldwork in the area has revealed several sites with suitable habi- tat for C. morefieldii on the western escarpment of Tennessee’s Cumberland Pla- teau from the Alabama state line northeastward to southern Warren County. Systematic surveying in this area might result in the discovery of more popula- tions of the federally endangered Morefield’s leather flower as well asa number of other significant rare taxa. 824 BRIT.ORG/SIDA 22(1) ACKNOWLEDGMENTS Our sincere gratitude is extended to Drs. Robert Kral and Michael Dennis for confirming our identification of C. morefieldii. Furthermore, we appreciate the information provided to us by Steve Threlkeld of the Alabama Department of Conservation and Natural Resources, Al Schotz and Michael Barbour of the Alabama Natural Heritage Program, James Allison of the Georgia Natural Heri- tage — and Claude Bailey of the Tennessee Division of Natural Heritage. Thanks are also due to Dan Spaulding, curator of collections at the Anniston Museum of Natural History, who loaned us specimens of Morefield’s leather flower from the nearest Alabama population. Dr. B. Eugene Wofford, curator of the University of Tennessee Herbarium, was very helpful throughout the project and his editorial comments improved the manuscript. We wish to thank the Department of Botany at the University of Tennessee for supporting the travel costs associated with this project. REFERENCES SS Horn, D.D.and P. Somers. 1981. Neviusia alabamensis (Rosaceae) in Tennessee. Sida 9:90-91, Krat, R. 1987. A new “Viorna” Clematis from northern Alabama. Ann. Missouri Bot. Gard. 74:665-669, Lona, A.A. 1989. Disjunct populations of the rare shrub, Neviusia alabamensis Gray (Ro- saceae). Castanea 54:29-39, Prinate, J. 1997. In: Flora of North America Editorial Committee, eds. Flora of Nort North of Mexico. Vol. 3. Oxford University Press, New York. TENNESSEE NATURAL HERITAGE PROGRAM (TNHP). 2004. Tennessee rare plant list. Tennessee Divi- sion of Natural Heritage, Dept. Environ. Conserv., Nashville. United States Fish and Witouire Service (USFWS). 1992. Endangered and threatened wildlife and plants; determination of Clematis morefieldii (Morefield’s leather flower) to be an endangered species. Fed. Register 57(98):21562-34420. 1 America GRATIOLA BREVIFOLIA (PLANTAGINACEAE) NEW TO THE FLORA OF DELAWARE, THE DELMARVA PENINSULA, AND THE MID-ATLANTIC Wesley M. Knapp Dwayne Estes Maryl ag aie ‘ment of Natural Resources Univers! ee Mee Idlife & Heritage Service a utionary Biology PO Box 68 xville Se 37996, USA. Wye Mills, Maryland 21679, U.S.A. tnplants@yahoo.com wknapp@dnr.state.md.us ABSTRACT Gratiola brevifolia (Pl i d | native additi he fl f Delaware, the Delmarva Peninsula, arid tarhe Mid- Atlantic This species is dis} PI ly 835 km (520 mi) from the closest known population in Burke Co., Georgia. RESUMEN Gratiola brevifolia (Plantagina e cita como una gone native rara ea anadir a la flora de 835 km (520 Delaware, la Peninsula de Delmarva, y peuane Minds Atla mi) aproximadamente de la poblaci cida cana €1 Burke Co; Georgia, The genus Gratiola L. (Plantaginaceae) consists of ca. 30 species (Estes, unpub- lished data) widely distributed through the North and South Temperate zones and on mountains within the Tropics (Pennell 1935). Six species of Gratiola (s.) have been reported on the Delmarva Peninsula (Tatnall 1946, McAvoy 2001), an area that lies entirely within the Atlantic Coastal Plain Physiographic Prov- ince of the eastern United States and consists of the Eastern Shores of Mary- land and Virginia and the majority of the state of Delaware. The northern por- tion of Delaware lies within the Piedmont Physiographic Province (Plank & Schenck 1998). These six species are G. aurea Muh. G. neglecta Torr, G. pilosa Michx., G. ramosa Walt., G. virginiana L.,and G. viscidula Pennell. On Delmarva, G. pilosa and G. virginiana are considered to be common, G. aurea and G. neglecta infrequent, and G. ramosaand G. viscidula are thought to be historical, having not been reported for 20 or more years (McAvoy, pers. comm.). Gratiola ramosa was last collected from Wicomico Co., Maryland, near the town of Sal- isbury (Canby s.n. PH), and G. viscidula was last collected from New Castle Co., Delaware, near the city of Wilmington (Tatnall s.n. DOV; Commons s.n. DOV). On 23 June 1992, Frank Hirst and Ron Wilson reportedly rediscovered Gratiola ramosa in Sussex Co, Delaware (R. Wilson 0719941, pers. herbarium). Recently, this population was visited by the first author and specimens were collected and sent to the second author for verification. Subsequently, the speci- SIDA 21): ): 825 — 829. 2006 826 BRIT.ORG/SIDA 22(1) mens were determined not to be G. ramosa but instead were identified as G. brevifolia Ral., a species not previously reported from Delaware, the Delmarva Peninsula, or the Mid-Atlantic. Though G. brevifolia and G. ramosa are easily distinguished by the presence or absence of two bracteoles at the base of the calyx (Godt rey @ Wooten 1981), Hirst and Wilson’s misidentilication could be expected because G. brevifolia is not included in any manual commonly used in the Mid-Atlantic Ge., Fernald 1950; Radford et. al 1968; Brown & Brown 1984: Gleason & Cronquist 1991). Voucher Specimen: U.S.A, DELAWARE. Sussex Co.: Coastal Plain Physiographic Province, ditched portions of Tussocky Branch paralleling Piney Branch Rd., northwest of the town of Delmar, 29 Jul 2005, Knapp 1549 (DOV, TENN, Maryland Natural Heritage Program Herbarium). Gratiola brevifolid is an erect perennial of wet, sandy pinelands (Pennell 1935), oak barrens (Tennessee), and sandy riverbanks (Arkansas and Oklahoma). The species ranges from Tennessee and Florida west to Oklahoma and Texas (Pennell 1935) and is found in four centers of distribution, with two occurring east and west of the Mississippi (Fig. L). Eastward, G. brevifolia is centered on the Coastal Plain of northern Florida and southern Georgia with scattered populations west tosoutheastern Alabama. The species is also significantly disjunct to the High- land Rim and Cumberland Plateau of central Tennessee (Chester et al. 1997). To the west, G. brevifolia is most frequent in the West Gulf Coastal Plain of south- eastern Texas and southwestern Louisiana, with disjunct populations in the West Gulf Coastal Plain of southeastern Oklahoma and the Ouachita Province of Oklahoma and Arkansas (Estes, unpublished data). Despite the broad geo- graphic distribution, it is considered imperiled in Arkansas (NatureServe Ex- plorer 2005) and probably should be considered a species of conservation con- cern in Alabama, Oklahoma, and Tennessee (Estes, unpublished data). ld ramosdasa species of conservation concern pa: for Delaware based upon the discovery and identification by Hirst and Wilson. With this publication, G. ramosd should no longer be considered a component of Delaware's native flora; however, it was historically a component of Delmarva’s native flora based on Canby’s collection cited above. The population of Gratiola brevifolia in Delaware consists of ca. 1500 in- dividuals restricted to the lower banks and bottoms of a ditched perennial stream. Interestingly, several other species in the vicinity of the G. brevifolia population are considered rare in Delaware, based on McAvoy (2003). These species include: Amphicarpum purshii Kunth, Hypericum adpressum Raf. ex WeBart., H. denticulatum Walt., Sabatia difformis (L. Druce, and Utricularia rddiata Small. The discovery of Gratiola brevifolia in Delaware marks a new addition to the flora of Delaware, the Delmarva Peninsula, and the Mid-Atlantic region, and is significantly disjunct from all other known populations in the South- east. The closest population to the Delaware occurrence is ca. 835 km (520 mi) KNAPP AND ESTES PNR WHERES 997 Fic. 1. Range of Gratiola brevifolia. Each @ represents a specimen examined. to the southwest in Burke Co., Georgia (E.T. Wherry s.n. PH). Various sources have attributed South Carolina to the range of G. brevifolia (NatureServe Ex- plorer 2005; South Carolina Plant Atlas 2005; USDA 2005; Weakley 2005), based upon specimens from Clarendon (J.F Townsend 384 CLEMS), Lee (C.A. Aulbach- Smith 4070 and 4077 with S. Hutto USCH), and York counties {f.B. Nelson 821] with L. Lundquist USCH), South Carolina, but these specimens, however, have been annotated by either Deborah Lewis or the second author as G. ramosa (Clarendon County) or G. viscidula (Lee and York counties). The nativity of discoveries, such as this, are highly debatable, and entire papers have been devoted to the topic (Lamont & Young 2005). We consider G. brevifolia a native component of the flora of Delaware, the Delmarva, and the Mid-Atlantic region for several reasons. First, though the habitat supporting the species in Delaware is highly degraded, G. brevifolia is found in similar habi- tats in other portions of its range. Second, two other species of Gratiola, G. ramosa and G. viscidula, once reached their northern range limits on the Delmarva Peninsula. The historic Maryland population of G. ramosa was also significantly disjunct, ca. 500 km (300 mi), from its closest known occurrence in Hoke Co., North Carolina (Radford et al. 1968). Third, the genus Gratiold is known for having strange and highly disjunct distributional patterns. For in- stance, De Lange (1997) reported G. pedunculata R.Br. as a new addition to the 828 BRIT.ORG/SIDA 22(1) flora of New Zealand and considered the species to be a recent natural migrant likely distributed by vagrant birds from mainland Australia, a disjunction of ca. 2150 km (1332 mi). Furthermore, G. virginiana, a species primarily distrib- uted in the southeastern United States, is disjunct to the states of Veracruz and Puebla, Mexico from central Texas, a distance of more than 900 km (525 mi) where it occurs in association with other species characteristic of the temper- ate southeastern United States (Miranda & Sharp 1950). Even G. brevifolia has a fragmented range characterized by wide disjunctions. For example, on the Gulf Coastal Plain, the easternmost population of G. brevifolia west of the Mis- sissippi River in Rapides Parish, Louisiana is separated by ca. 623 km (387 mi) from the westernmost population east of the Mississippi River in Pike Co., Ala- bama. Lastly, G. brevifolia is not known to be cultivated or grown for horticul- tural purposes; therefore, it is highly unlikely t introduced from nearby cultivated plants. a rat this species would have been ACKNOWLEDGMENTS Appreciation is extended to several people who contributed information or com- ments to this project: Curtis Hansen, Auburn University; Frank Hirst; James Macklin, Philadelphia Academy of Natural History; Bill McAvoy, Delaware Natural Heritage Program; Lucile McCook, University of Mississippi; Robert Naczi, Delaware State University; John Nelson, University of South Carolina; Heather Sullivan, Mississippi Department of Wildlife, Fisheries and Parks; Chris Frye of the Maryland Department of Natural Resources, Wildlife and Heritage Service; and Ron Wilson. Additional thanks are given to the following herbaria where specimens were examined: AUA, BRIT, CLEMS, DOV, LSU, MISS, NCU, NO, PH, TENN, USCH. REFERENCES Brown, M.L.and R.G. Brown. 1984. Herbaceous plants of Maryland. University of Maryland, College Park. CHesrer, E.W., B.E. Worrorp, and R. Krat. 1997. Atlas of Tennessee vascular plants. Volume 2. Angiosperms: dicots. Misc. Publ. No. 13. The Center for Field Biology, Austin Peay State University, Clarksville, TN. De Lance, PJ. 1997. Gratiola pedunculata (Scrophulariaceae): a new addition to the New Zealand flora. New Zealand J. Bot. 35:317-322. FerNatb, M.L. 1950. Gray’s manual of botany (ed. 8). Dioscorides Press, Portland, OR. GteAson, H.A. and A. Cronouist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada (ed. 2). The New York Botanical Garden, Bronx. Goprrey R.K. and J.W. Wooten. 1981. Aquatic and wetland plants of southeastern United States: Dichotyledons. Univ. of Georgia Press, Athens. Lamont, E.E.and S.M.Youns. 2005. Juncus diffusissi an addition to the flora of New York, with notes on its recent spread in the United States. J. Torrey Bot. Soc. 132:635-643. KNAPP AND ESTES 829 McAvoy, W.A. and K.A. Bennett. 2001. The flora of Delaware: an annotated checklist. Dela- ware Natural Heritage Program, Delaware Division of Fish and Wildlife (Document No. 40-05/01/01), Smyrna, DE. McAvoy, W.A. 2003. Rare vascular plants of Delaware. Delaware Natural Heritage Program, Delaware Div. of Fish and Wildlife, Smyrna, DE, unpublished report. Miranpa, F.and A.J. SHare. 1950. Characteristics of the vegetation in certain temperate re- gions of eastern Mexico. Ecology 31:313-333. NatureServe Explorer: An online encyclopedia of life [web application]. 2005. Version3.1. NatureServe Arlington, Virginia. Available .org/explorer.(Accessed: 17 Dec 2005). Pennett, EW. 1935.The Scrophulariaceae of Eastern Temperate North America.Wickersham Printing Company. Lancaster, PA. Pp. 78-80 Plank, M.O.and WS. ScHENck. 1998, Delaware Piedmont Geology. Delaware Geological Sur- vey, University of Delaware, Newark, Delaware, Special Publication No.20, 69 pages. Raproro, A.E., H.E. AHLEs, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. University of North Carolina Press, Chapel Hill, NC. Sorrie, B.A.and A.S.Weakiey. 2001. Coastal plain vascular plant endemics: Phytogeographic patterns. Castanea 66:50-82. SouTH CAROLINA PLANT ATLAS. 2005. University of South Carolina. Available http:// cricket.biol.sc.edu/herb/. Tatnatt, R.R. 1946, Flora of Delaware and the Eastern Shore. Soc. Nat. Hist. of Delaware. Lancaster, PA. Weaktey, A.S. 2005. Flora of the Carolinas, Virginia, and Georgia, Working Draft: June 10, 2005. Available: http:// herbarium.unc.edu USDA (United States Department of Agriculture). 2005. Natural Resources Conservation Service. The PLANTS database (plants.usda.gov/plants/). National Plant Data Center, Baton Rouge Louisiana (Accessed: 17 Dec 2005). 830 BRIT.ORG/SIDA 22(1) Book Notices University of California MELINDA A. Zeper, DANIEL G. BRADLEY, Eve EmsHwitter, and Bruce D. Smitu. 2006. Documenting Domestication: New Genetic and Archaeological Paradigms. (ISBN 978-0-520-24638-6, hbk.). University of California Press, California/ Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, US.A.(Orders: www.ucpress.edu/, 609-883-1759, 609-883-7413 fax). $70.00, 361 pp., b/w photos, illustrations, 8 1/2" x 11" The book is divided into four sections: 1) Archaeological Documentation of Plant Domestication. 2) Genetic Documentation of Plant Domestication. 3) Archaeological Documentation of Animal Do- mestication. 4) Genetic Documentation of Animal Domestication. From the Publisher—Agriculture is the lever with which humans transformed the earth over the the last 10,000 years and created new lorms of plant and animal species that have forever alterec face of the planet. In the last decade, significant technological and methodological advances in both molecular biology and archaeology have revolutionized the study of plant and animal domestica- tion and are reshaping our understanding of the transition from foraging to farming, one of the ma- a turning g points in human history. This groundbreaking volume for the first time brings together eading veologists and biologists working on the domestication of both plants ve ani uc to pee a wide variety of archaeological and genetic approaches to tracing the of domesticates. It provides a comprehensive overview of the state of the art in this ene cals recent findings on specific crop and livestock species in the Americas, field as well as reviews of 1 Africa. Offering a unique global perspective, it explores common challenges and poten- Eurasia, anc tial avenues for future progress in documenting domestication. M. Nevin Smit. 2006. Native Treasures: Gardening with the Plants of Califor- nia. ISBN 0-520-24425-7, pbk.). University of California Press, California/ Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, US.A.(Orders: www.ucpress.edu/, 609-883-1759, 609-883-7413 fax). $24.95, 278 pp., color photos, 7" x 10" From the Publisher—The author explains how California's diverse terrain, climate, and geology sup- port a wealth of plant species—more than 6000—and offers suggestions for designing with most ol cultivation, as well as with some more obscure but garden-worthy groups. With sity celebrates = the major né an engaging narrative and a wealth of illustrations, this ode to beauty and dive -alifornia’s rich store of native plants and encourages readers to visit them in rn native haunts and invite them into their gardens. SIDA 22(1): 830. 2006 HEDYOTIS AUSTRALIS (RUBIACEAE) NEW TO MISSOURI AND: FLORIDA AND RELATED SPECIES IN THE SOUTEL-CENTRAL UNITED STATES Walter H. Lewis Department of Biology Washington University St. Louts, Missouri 63130, U.S.A. ABSTRACT The distribution of white-flowered Hedyotis australis is extended to Missouri and Florida as well as within Alabama, Arkansas, Mississippi, and Tennessee. This distribution is compared to allied homostylous species H. crassifolia and H. rosea which also flower as winter annuals in the south- central United States. Hedyotis crassifolia is a diploid species having a flowering peak earlier than the tetraploid H. australis. An example of aaa out- eee within H. crassifolia via bee pollina- =a) tion is discussed. Hedyotis rosea is now ¢ ‘tirpated in Missouri RESUMEN Hedyoti tralisde fl bl distribuida por Missouri y Florida asi como también en Alabama, Arkansas, Mississippi, y Tennessee. Su distribucion se copa la de las otras especies homostilas H. oS ia y H. rosea la region centro- sur de Estados Unido ide Hedyoti ifolia ti le f] 5 — a especie ene H. aera Se Sale un emul de posible entrecruzamiento dente de H.c ae via polinizacion por abejas. H ra extinguida en Missouri. INTRODUCTION Three winter annual and homostylous bluets (Hedyotis or Houstonia) flower and fruit in the south-central U.S. largely between January and April. The most common and widely distributed of these is the purple- or violet-colored Hedyotis crassifolia Raf. (Houstonia pusilla Schépf). Less well-known is a smaller an- nual with tiny white corollas which is easily overlooked even when flowering in the same habitat and at the same time as H. crassifolia. Consequently, this southern white-flowered Hedyotis australis WH. Lewis & D.M. Moore [Houstonia micrantha (Shinners) Terrell] is infrequently collected even when common. A third very small bluet with large pink- or rose-colored corollas, Hedyotis rosea Raf.{Houstonia rosea (Raf.) Terrell], is more narrowly distributed and occurs in eastern Texas, Louisiana, central and southern Arkansas, west- ern Mississippi, and southeastern Oklahoma, with outlying populations in Ala- bama (Tuscaloosa Co.) and in 1931 southeastern Missouri (Dunklin Co.). During March 2005, collections were made of these species concentrating on areas where H. australis was unreported but could be expected to occur. Find- SIDA 22(1): 831 — 836. 2006 832 BRIT.ORG/SIDA 22(1) ing these small bluets by their distinct flower colors of purple-violet, white, and pink-rose is one of the best ways to identify them. In addition, corolla tubes of H. australis are short with calyx lobes as long as or longer than the tubes (Fig. 1), while lobes of H. crassifolia are much shorter than its longer tubes. How- ever, without these floral characters their discovery is greatly limited even when growing side by side as they often do, and particularly since anthesis is con- fined toa few weeks at most. All collections were made by Walter H. Lewis and Memory Elvin-Lewis; their numbers and herbaria of deposit are provided for each collection cited. Many additional ones are at MO. Cytological differences also exist between these species: H. crassifolia is diploid, 2n = 2x = 16, x = 8, H. australis is tetraploid, 2n = 4x = 32, x = 8, and H. rosea is diploid, 2n = 2x = 14, x =7,a reduced base number from the others. That eos a uctions, and to some extent polyploidy, have played impor- tant roles in the evolution of these and other North American species of Hedyotis (Lewis 1965) is supported by recent molecular phylogenetic analyses by Church (2003) who ei found that “Houstonia is not distinct from either North American Hedyotis or Stenaria.” Additional research is needed lor North Ameri- can (including Mexican) taxa and others worldwide before generic- and tribal- level complexes of both the Hedyotideae and Spermacoceae are satisfactorily resolved. Clearly, this paper does not provide additional data regarding classili- cation and phylogeny, for it is intended only to extend certain species distribu- tions and comment briefly on specific ecologic and reproductive observations. To aid the reader in identifying the three species, as well as two forms of H. crassifolia, f. crassifolia and [. albiflora (Standley) W.H. Lewis, characters were obtained from Lewis (1970), Smith (988), Terrell (1996), and herbarium collec- tions at BRIT and MO: — 1. Corollas white, lobes and tube each 1.5-2.5 mm long; calyx lobes as long as or longer than corolla tube (Fig. 1) australis 1. Corollas purple-violet or pink-rose, rarely white, tube exceeding length of calyx lobes. 2. Stems 3 eles cm long; corollas purple-violet, occasionally pale purple to white, lobes 2.5-3.5 mm long, tube 3.5-4.5 mm long; calyx lobes about half as long as corolla eee nmon ____ crassifolia 3. Corollas purple-violet throughout f. crassifolia 3. Corolla lobes white to pale purple-violet, tube purple-violet to reddish f. albiflora 2. Stems 1-3 cm long, often clumped; corollas pink-rose fading to oe yes tol nearly white lobes 2.5-4.5 mm long, tube 5—7 mm long; calyx lobes 1.2-2.5 mm long; occasional rosea Hedyotis australis Prior to 2005 the known continuous distribution of H. australis was limited to eastern Texas, Louisiana, Arkansas (excluding the northeast), southeastern Oklahoma (only McCurtain Co.), western and central Mississippi, and south- western Tennessee (only Shelby Co.), in addition to outlying populations in cen LEWIS AND FLORIDA 833 Fic. 1. Hedvoti tralis W.H. Lewis & D.M.M inl f t y, Natchitoches, N hitoches Parish, Loui- siana, WH, Lewis 15867 (MOQ).A h g calyx lot long Ila tube ( imes longer). Bar equivalent to about 1 cm tral Alabama (Tuscaloosa Co.), and central and eastern Georgia (Bibb and Co- lumbia Co.’s) (Chambers 1965, Lewis 1968, Terrell 1996). Its current distribu- tion is now extended to include: (1) southeastern Missouri.—northern Dunklin Co., N of Campbell, 15968 F MO, UMO, US; northern Dunklin Co., NW of Campbell, 15969 MO; Butler Co., Poplar Bluff, 15970 MO, US. These are the first reports for Missouri. The Poplar Bluff collection at 36'76'N is the most northerly record of the species, although this distribution is similar to the new record at Paris, Tennessee (36'17’N). Hedyotis australis was found in Missouri’s Bootheel region and adjacent Butler County in low foothills at the southeastern edge of Ozark Plateau where a three collections grew in grassy cemeteries and where herbicides were not used. None were found in nearby pastures or meadows or in lower-lying areas where cotton is now grown in large farms and where herbicides are widely used. The species was also absent from flood-plains and other riparian habitats to the east along the Mississippi River valley. (2) northeastern Arkansas.—Greene Co., Paragould, 15921 MO, 15964 BRIT, 834 BRIT.ORG/SIDA 22(1) MO. As in Missouri H. australis was not found in the Mississippi River valley, but only along a low ridge just west of the Missouri Bootheel among grass in a hilly cemetery. This is its most northeasterly location in Arkansas which par- allels its northwestern distribution (Smith 1988). (3) western panhandle of Florida.—Escambia Co., Hwy 4 at Sandy Hollow Rd, 15936 FSU, MO, USE This new outlying record was found in a hilly region of extreme northwestern Florida growing with H. crassifolia in grass by a road- side corner. Both species were in flower 22 March, although flowering had just begun for H. australis (no mature capsules) compared with numerous flowers and maturing capsules of H. crassifolia. Tetraploids like H. australis often have peak flowering later than their diploid relatives (as H. crassifolia) and this gen- eralization applies here (Lewis 1980). Hedyotis australis was not seen elsewhere in Escambia County, adjacent Santa Rosa County, or nearby Mississippi and Alabama, although H. crassifolia was found with regularity. (4) eastern Mississippi.-Hinds Co., Hwy 27 at Bear Creek Rd, 15930 BRIT, MO; Smith Co., Taylorsville, 15932 MISS, MO, US; Itawamba Co., Fulton, 15950 DAO, MO; Lee Co., Tupilo, 15951 MISS, MO; Union Co., New Albany, 15953 MO, US; Marshall Co., Holly Springs, 15954 MISS, MO. In Mississippi's southeastern area H. australis was only occasionally found along grassy roadside verges, but where found it was common while H. crassifolia was less so (Hinds, Smith Co.’s); in the northeast it was widespread in the more frequent upland areas (Itawamba Co., the most easterly Mississippi collection, Lee, Union, Marshall Co.’s) where H. crassifolia was also less common. Observation of fewer flowering plants sug- gests that {lowering of H. crassifolia had peaked some time earlier rather than limited frequencies of plants. (5) western Alabama.—Marion Co, Hamilton, 15949 BRIT, MO, US. Plants of this second collection known for Alabama were growing in abundance among other invaders in lawns. This new extension eastward in Alabama from those found in nearby northeastern Mississippi is about 80 miles northwest of the only other known Alabama collection at Tuscaloosa in Tuscaloosa Co. (Terrell 1996) where H. australis was re-collected (15945 BRIT, MO, US) growing with H. crassifolia 15946 BRIT, MO) and nearby (ca. 20 m) H. rosea 15947 BRIT, MO, US), the most easterly record for this species. (6) western Lennessee.—Tipton Co., Covington, 15926 MO; Fayette Co., Hwy 57 at Hwy 18, 15956 MO, US; Hardeman Co., near Bolivar City, 15957 MO, TENN; Gibson Co., Humboldt 15960 MO, TENN, Bradford 15961 MO, TENN; Henry Co., Paris, 15962 MO, TENN. Tothe one collection of H. australis previously reported in the extreme southwestern part of the state at Memphis (Shelby Co.) (Terrell 1996) are added numerous collections throughout western Tennessee as far east as 88° 15°W in Fayette, Gibson, Hardeman, Henry, and Tipton Counties found in lawns, hillsides, ruderal areas, and cemeteries. As in Missouri, the species was not found in the Mississippi River valley, but did occur with some frequency on LEWIS. 835 the eastern ridge and uplands eastward and as far north as near the Kentucky state line to Mississippi in the south. Although the species was not found in western Kentucky, it might occur there. Hedyotis crassifolia Many collections were made of H. crassifolia whose range, including H. minima (Beck) Torrey & Gray, extends from Maryland to Georgia west to Wisconsin and Iowa, and south from Kansas to central Texas along the Gulf Coast states to the panhandle of Florida (Terrell 1996). Often it occurs with H. australis and less commonly also with H. rosea, particularly in eastern Texas and Louisiana where all three species can be found in flower together. All are homostylous with different chromosome numbers; recently Church (2003) found H. crassifolia with cleistogamous flowers among greenhouse-grown plants. In- breeding is clearly predominant and no hybrids between the three species have been found. Thus, I was surprised to observe foraging in a population of H. crassifolia a bee, Agapostemon virescens (Halictidae), extending its proboscis into the corolla tube while wings beat and rapidly flying from one flower to another after a few seconds stay. It visited about 30 flowers (occasionally re- turning to the same one) of 12 or so plants during 3-4 minutes and then flew off. This occurred near the Appalachian ridge in Alabama, Clay Co., at the Bap- tist Church cemetery adjacent Highway 49, 23 March. No other species of Hedyotis was found in the vicinity; in fact few species were in flower. How com- mon such visits might be in the early spring (few pollinators seen) is unknown, but that it was observed even once suggests a means of at least limited out- crossing for this homostylous species. One could venture that its distributional success covering a wide range of habitats compared to other related winter an- nuals might be due to the success of even secondary out-crossing in combina- tion with in-breeding via homostyly and possibly cleistogamy. Hedyotis rosea In representative specimens for H. rosea Terrell (1996) includes a collection from Missouri in Dunklin Co., near Campbell, 5 April 1931, collected by A.L. Grant s.n.(MO). Recently George Yatskievych located a second sheet collected by Julian Steyermark 406 (MO) on the same day, both undoubtedly made during the same collecting trip. Steyermark’s label locality is more precise at “ca. 5 miles north- west of Campbell” in open flat ground surrounded by Quercus phellos woods. This is the same area of northern Dunklin Co. where new records of H. austra- lis were found only in “protected” (from herbicides) cemeteries on 28 March 2005. Searches around Steyermark’s locality, in much of the northern part of the county, and in neighboring counties proved fruitless. I suggest that this single outlying population collected in 1931 is extinct and that H. rosea is now extirpated from the Missouri flora. No collection has been found in neighbor- 836 BRIT.ORG/SIDA 22(1) ing Arkansas or Tennessee and the nearest H. rosea in central Arkansas (Terrell 1996) is about 175 miles southwest of the original Missouri find. Indeed the loss of habitat and heavy use of herbicides for many years in the Bootheel counties may have contributed to the loss of H. rosea and perhaps other species whose ranges once extended to this region. ACKNOWLEDGMENTS | thank Memory Elvin-Lewis, Washington University, who assisted me in the field during both 2005 collecting trips, Richard Clinebell, entomologist with the Missouri Botanical Garden, who determined the bee discussed under Hedyotis crassifolia, and N. Rogerio Castro for translating the Abstract into Spanish. lalso acknowledge the use of important collections of Hedyotis at BRIT and MO. REFERENCES CHameers, K.L. 1963. Hedyotis australis in Georgia. Rhodora 65:271-273. CHurcH, S.A. 2003. Molecular phylogenetics of Houstonia (Rubiaceae): descending aneup- loidy and breeding system evolution in the radiation of the lineage across North America. Molec. Phylogenetic Evol. 27:223-238. Lewis, W.H. 1965. Pollen morphology and evolution in Hedyotis subgenus Edrisia (Rubiaceae). Amer. J. Bot. 52:257-264. Lewis, W.H. 1968. Notes on Hedyotis (Rubiaceae) in North America. Ann. Missouri Bot. Gard. 50188) Lewis, W.H. 1970. Hedyotis L.In:D.S. Correll and M.C. Johnston. Manual of the vascular plants of Texas, Texas Research Foundation, Renner. Pp. 1487-1490. Lewis, W.H. 1980. Polyploidy in species populations. In W.H. Lewis, ed. Polyploidy: Biological relevance. Plenum Press, New York. Pp. 103-147. SuitH, E.B. 1988.An atlas and annotated list of the vascular plants of Arkansas, ed. 2. Pub- lished by the author, Fayettesville, AR. TerreLt, E.E. 1996. Revision of Houstonia (Rubiaceae-Hedyotideae). Syst. Bot. Monogr. 48. The American Society of Plant Taxonomists. Book REVIEWS TosHio YosHipa. 2005. Himalayan Plants Illustrated Hi hokubutsu daizukan (ISBN 4-635-58031-8, pbk.). YAMA-KEI Publishers, Co., Ltd., 1-9-13 Akasaka, Minato-ku, Tokyo, JAPAN. Price: ¥13,000 (ca. $115), 800 pp., (773 pp.in color), 18.5 x 26 cm, in Japanese. When I returned from field work in Asia in September of 2005, a book of beautiful photographs of sree wildflowers was among the items in Ge mail. The book, Himalayan Plants Illustrated, is by Toshio Yoshida, one of the most talented, knowledgeable and intrepid photographers of alpine Bee in eastern ee ae Yoshida has traveled ne ly to capture images of the characteristic aes of the Greater Himalayan region. Having also visited some of the areas where he has worked, I wa amazed to see that he | g p eray of the species in one of the world ee areas of plant pages The photog bool he f nine auEnOrs = ia of field studies in the Hima- ] di China and from leadi The photos are richly su upplemen tedb iled f h dy of herbarium specimens and fror the literature. The introductory material inc ludes a fresh discussion of floristic zones and their de- limitation in the Greater Himalaya, southern Xizang (Tibet) and the Hengduan Mountain region of China, based on the author’s observations and interpretation. The nt contains 2,739 color photos covering 1,771 plant taxa, of which 1,586 are at species rank, 40 are subspecies, 61 are varieties, 2 are forms, and the remaining 82 are unidentified. The pho- tos were taken in all parts of the Himalayan range; northern Pakistan, Kashmir, northwestern India, — Sikkim, Bhutan and Tibet, as discussed in more detail in the front pages of the book. The book compared with Flowers of the Himalayas by Oleg Polunin and Adam Stainton, published in 1964 by Oxford University Press. Polunin one Stainton’ s book covers 1495 plant taxa, with 694 color 1 in the last 128 pages, and 3 gs. Because of the politics of those days, the field sites covered by Polunin and Stainton are in about half the Himalayan range, excluding Bhutan, Tibet and most parts of Pakistan and Sikkim n Mr. Yoshida’s book the photos and descriptions/discussions are on the same page, crossed Gus by number. Plants exhibiting a wide range of variation, a likely phenomenon in the malaya, are represented by two or more photos to give a sense of eens variation. Below a photo is the plant name in Japanese, which is often the Japanese transliteration of the scientific me. Also preva if bce is ou icient space, are details of the photograph, Hues date, place, i pie and i the eine The letters A-Z3 preceding the place name below each photograph ee to the 34 maps on the front pages. A brief guide to the ee is on the inside flap of the sae ine and can be read without turning pages ach photograph is keyed to : nding text entry. The text entry gives the Japanese name, the scientific name and synonyms, if anys fe Sliawed ee flowering period, altitudi- nal distribution and details of the habitat and life form. The morphological description in the text matches the plants in the corresponding photo(s), since it is basically derived from author's field notes at the time the photo was taken and from dried specimens of the photographed a? sif they were obtained. The oa are further enriched with information from the literature. The 82 Dice meine’ cen which may represent new species or varieties, also text with data about th ts. They are provided with the name of the species that the author } bs ] result Del C1 Plant morphology, especially of species dwelling in the alpine zone, is strongly affected by the environment. With that thought i in mind, the author has tried to include photos that show not only the plant, but its habit and its place within the habitat. SIDA 22(1): 837. 2006 838 BRIT.ORG/SIDA 22(1) Che introductory material includes articles on the history of research on Himalayan plants written by Prof. Hideaki Ohba on pages 4-7, geobotany of the Himalaya on pages 8-1], and a map of the whole Himalaya. On pages 12-25 are 34 topographical maps. Botanical guides to the areas of the aya are on pages 26-32. An analysis of the horizontal and alecidinal: distribution patterns of —_ Hima the plants covered in the book is on pages 33-39 and adaptive strategies of alpine plants in the aya are covered on oo 40-43. Himalayan plants in the oe mountains of southwest —_— Hima China are discussed on pages though the book is ae the high quality and detail in the photos make the diversity of the flora and vegetation in this unique and highly diverse area easy to comprehend. The number of individual taxa included for each genus provides a good sense of the range . shel within large genera. Some examples: Arisaema, 17; Astragalus, 24; Gentiana, 33; Impatiens, 22; Leontopodium, 10; Meconopsis, 21; Pedicularis, 66; Rhododendron, 58; Saxifraga, 06; Sdussurea, 40. book is beautifully bound on high quality paper and one of the nice features is t pages lie flat when the book is opened at any page. Rumor has it that an English language edition is being considered.—David E. Boufford, Harvard University Herbaria, 22 Divinity Avenue, Cambridge, MA 02138-2020, U.S.A, david_ boufford@harvard.edu. rat the Q. Two British Orchid Books ANNE and Simon Harrap. 2005. Orchids of Britain and Ireland: A field and site guide. (ISBN 0-7136-6956-X, pbk., durable field guide binding). A & C Black, 38 Soho Square, London, W1D 3HB, U.K. (Orders: +44 (0)20 7758 0200 customerservices@acblack.com; http://www.acblack.com/search.asp) £29.99 ($51.95 US), 480 pp, 360 color plates, numerous watercolors, line art, 49 maps, glossary, bibliography, index, 21.6 x 13.8.cm. MicHaet Fotey and Sipney Clarke. 2005. Orchids of the British Isles. SBN 0-9541916- 1-7, hbk.). Griffin Press Publishing Limited, 35 Wessex Way, Cox Green, Maidenhead, SLO 3BP, UK (Orders: http://www.griffinpress.co.uk/ orchids.htm). £45 ($78 US), 390 pp., extensively illustrated with color photographs; 50 maps, anaes bibliography, index, 27.6 x 20.5 cm. It is not often a reviewer has t] ity to review two eae that cover the same subject and are pubis! hedat the same time. eThisis the situation with both books on the orchids of the British Isles. Each | from the British Isles, Both books are packed with full- color photographs, maps, eeebnical details, and similar information. Yet they are as different from her as apples and oranges. While Foley and Clarke’s large format book (published in associa- tion the Royal Botanic Garden, Edinburgh), contains the more technical information, the Harraps’ work is designed as a field guide—and that it very much is. Asa field guide the binding is a durable Flexibind type that is sown and open tat at any given pom Is eal) ita Pabst binding i is an injus- dare \ ry detail. A i= a tice. The photographs in Orchids of Britain though both books use the maps oe Ne Atlas of the British and Irish Flora ioe: reproduced in the Harraps’ book are much easier to read owing to the fact that the distributional dots representing county records for specific time segments are in red, black, and green as opposed to those in Foley and Clark that are in graduated shades from black to grey to red to pale pink. The wealth of informa- tion that appears in both books is very complete and covers every possible aspect of the orchids in- cluding their on, W ae extensive notes on more than 0 supspccics, varieties, and forms; dis- tributional i and status. $ although ina slightly different format in each book. In Orchids ide and Ireland the Harraps go even further and detail th SIDA 22(1): 838. 2006 BOOK REVIEWS 839 history of the known populations and their current status. Both books treat the genera Dactylorhiza and Epipactis in great detail and make every effort to sort out the species, subspecies, and varieties— no simple task! It is not surprising that the bibliography and glossary are similar in each work. Given that both books cover much of the same information what does one book have that the other does not? Foley and Clarke @ £45 Harrap @ £29.99 covers 55 ee covers 56 species historical information extensive specific information for seeking the superior chapter on orchid taxonomy orchids (44 page site guide to 330+ places to checklist of species see orchids authors names with the species easier to read distribution ma photographic hints chart for each species ae eine over the keys to the genera and species of selected genera years chapters by guest authors on cultivation and — convenient field guide format conservation growth pattern gr ae of several genera comparative plate of the bee orchids, Ophrys, of Europe Both books use the term variety is the sense of forma—color and growth forms and both are cio of the misuse of the term albino. A true albino plant = no oe other than white-stems, leave and flowers. What is usually meant is white-fl fte hthe name albiflora. aioe it is the only publication of the two with keys for identification, be and Clarke is by no means a field guide and the size limits its use as such. Having a copy of both books would be ideal. Use the Harraps in the field and then when returning home consult gens and cl arke for confirming or addi- ae information. The photographs in both books at, but in Foley and Clarke many of ‘ull-page images have blurred. I feel certain the er ee were sharp but they were en ee just a ou beyond a limits. The me) momceavie enor found in HOY and Clans The first, under Spi simply piranthes {x}steigeri being a hybrid ee i 5 romanzoffiana and S. cernua. It is not been viewed as such for over 25 years. Spiranthes {x}steigeri is a synonym for S. ochroleuca. Information under Pseudorchis albida stating that it occurs in North America from the east to Alaska is incorrect. The species, as P. straminea, is restricted to northwestern Newfoundland and a single site in western Quebec on the shore of Hudson's Bay. Both facts could have been easily found in Flora of North America. ographical information is given for both Foley and Clarke and their guest authors but none for the ieee It would have been nice to know a bit acon: them and their other work. All four ae authors should be congratulated on their worl contribution to the 21" Century's knowledge of wild orchid in the British Isles and are heartily recommend d fo both i: novice and Byorsionala alike. If youare visiting the British Isles for an orchid hunting expe- dition the An Simon Harraps'’ field guide will be especially useful —Paul Martin Brown, au- thor, Wild meee ‘of Florida, The Wild Orchids of North America, Wild Orchids of the Southeastern United States, 10896 SW 90" Terrace, Ocala, FL 34481, U.S.A. naorchid@aol.com. 5; rt — Orchids of Mexico E. HAcsater, M.A. Soto ARENAS, G.A. SALAZAR CHAVEZ, R. JIMENEZ MACHORRO, M.A. Lopez Rosas. and RL. Dresser. 2005. Orchids of Mexico. (ISBN 9-6878-8908- X, hbk.). Published by Instituto Chinoin, A.C. Mexico City. Spanish and English versions available. (Orders: redactamex@yahoo.com.mx; Redacta, S.A. de SIDA 22(1): 839. 2006 840 BRIT.ORG/SIDA 22(1 CV, Avenida 10 de Mayo # 249, Col. San Pedro de los Pinos, C.P.03800 Mexico, D.F MEXICO). $100.00, plus shipping, 304 pp., 650 color photographs, 13.25" mR IO(3I 9 R25 5c) Initial disappointment may often give way to surprised delight. Such is the case of Orchids of Mexico. Given the publication and research history of the estimable authors | had expected a detail taxo- nomic oe the orchids of Mexico. Imagine my surprise when a large format-33.5 x 25.5cm- oO ‘coffee table’ book arrived! This sumptuously illustrated volume is much more orchids ‘in Mexico’ than ‘of Mexico’ and wastes no time in luring the reader to that country. | soon discovered that the | a searchable CD that will cover the 164 genera and 1200+ species with over 1500 color photogr aphe a checklist, and cross-referenced taxonomic treatment | was so looking forward to is fort synonymy. The current publication mentions 162 of those genera and 450 of the species. But back to the volume at hand; sitting before me is one of the lushest books on orchids | have ever seen. Apart from the text, which we will get to later, it is lavishly illustrated with the highest quality piptogs ae and es ae oduction I have seen in many years and the printing on heavy | =p stock only y of the book. The authors have provided us with not just an overview of the oi tha nat grow w eh Mexico, but with details of the several climatic and geological regions of the country shown at first in a two-page layout of a satellite image of Mexico. After relating the history of orchids, and orchid research in Mexico, with many reproductions of plates from well- known historic volumes, it concludes with the ethobotanical history featuring Vanilla planifolia. A brief chapter on orchid biology is fae illustrated with photographs of plants from flower to fruit to flower with the many stages in between. ow to our adventure searching ne wilds of Mexico. As we visit the eight various habitats the narrative highlights the many species of orchids that are present and and numerous vies of the habi- tats and illustrates them with color nes Sai At this a few synonyms are given and occa- sionally the reader may be temporarily confused with some of the newer g c concepts for famil- iar species such as Prosthechea citrina for Cililes: citrinad and Tamayorkis for a few species traditionally known as Malaxis. Many species found in Mexico are also found in the bordering re- gions of the southwestern United States, Texas, and Florida. It is always welcome to see photographs of these more southerly sire that may be very rare in the United States. Because of the arrange- ment of chapters by habitat it req sulting the index to find information on specific species. But that is not the apparent aim of this book. It is to immerse the reader in the diversity of the orchids of Mexico and the verdant and prolific variec habitats. A final chapter on Conservation efforts and successes is followed by detailed photo creits and an extensive bibliography. es nks to the team of authors and photographers the text is scientifically accurate and this reviewer did not note any mipurectly ae deled or positioned graphics (a feat not many books of this scope can boast!). There few i with a missing letter Le, seudobulb for pseudobulb, but nothing that could | essibly distract? from the intense text. Many photographers were credited in the book and the meticulous editing of these photographs has provided the finished book with a continuity not often seen in similar publications. Is it expensive?—yes, at $100—but worth every penny of the price. Keep in mind that two books were really written; one in Spanish and then one in English. Were Orchids of Mexico not sponsored by Chinoin lam sure it would have to sell for much more or not even have ever gone into production. If you have any interest in North American ore ae Finca rom the southern regions of that rl} t . 1 s volume. And remember, that the interac- tive CD will soon be available to satisfy those a us sas thrive on taxonomic treatments!.—Paul Mar- tin Brown, author, Wild Orchids of Florida, The Wild Orchids of North America, Wild Orchid sof the Southeastern United States, 10896 SW 90" Terrace, Ocala, FL 34481, U.S.A. naorchid@aol.com. O ‘e) — ~ => = =) io) +) la } S 1) = a | s y gy i= C = 7 ~_ iS i a £ = 7 SIDA22( (1): 840. 2006 BOOK REVIEWS 841 Orchids of Cuba Juan LLAMACHO (text) and JuLio A. LARRAMENDI (photographs). 2005. The Orchids Cuba—Las Orquideas de Cuba. (Cloth. ISBN 84-933615-2-6, hbk.). Greta see Plaza Mayor, 24 25340 VERDU, SPAIN. (Orders: +34 690 733 O11, http://www.gretaeditores.com inlo@gretaeditores.com). +9.00 Euros [$58.19 2/13/06], 287 pp., Spanish-English in parallel columns, color photographs, interpretive graphics, table of species, and glossary, 9.6" x 7" Following a recent trend for ae orchid books to be both scientifically accurate and written such a manner that they are popular with the non-academic enthusiast, The Orchids of Cuba fulfills a geographic niche ee has ie been void. This work of two enthusiastic Cubans has produced not only the first full-color book on the orchids of the island archipelago but the first text in English (Spanish/English in parallel columns) that provides an alluring enticement to the island nation. Following an imprimatur by Carlyle A. Luer, M.D. and foreword by James Ackerman, Ph.D. the books starts with the basic biology of orchids and biogeography of Cuba and proceeds to merge the two, describing and illustrating the several regions ol the r nation. These ieee and the orchids found within them are then treated in detail. Of the 305 sr ted from Cuba at the writing of the book, more than 150 are treated in detail and, for the most spate have excellent color photographs. Both the author's and photographer's love of the pleurothallids (Lepanthes and related genera) is me in the spectacular close-up photographs of many of these tiny species. Following the species treatments is a section on undescribed (those areas that have not yet been identified and/or de- cribed to science) species, with the same high-quality photographs. The book cana: witha table of all 305 ly found in Cuba, their flowering times an g n,a brief biniosoeny: and an ‘excellent glossary. What is unfortunately ae isan ee Because of the arrangement of species it makes it difficult to find s ickly. A listing of syn- onyms for the species treated in full Id also have been nihelpfa ul. No oe to ee are given as this book is, ey just the prelude to more publications covering all of the orchids of Cuba. F e reviewer's standpoint The Orchids of Cuba has the same proble her works that treat orchids from the Caribbean and Central America/Mexico. W ee Ee are stated the occurrence a the Spaces in the United States, usually Florida, is omitted. In part, tk € in several works. Rather than Ph: on pation fr rom a quarter century or more ago, simply consulting Flora of North America or Wild Orchids of Florida would have solved this problem. Those species included in es or and also documented lata, Bulbophyllum Florida include lonopsis utriculariodes, Prosthechea boothiana,S Pe ee Epidendrum nocturnum, Liparis nervosa, Maxillaria crassifolia, Brassia caudata, Eulophia elata, Malaxis pane neal distans, Phaius tank eae avert! amo ee drum radicans, Prescottia opligantha, and Pelexia adnata. Govenia trib- uted to Florida (Govenia floridana is - correct eae From a taxonomic standpoint ane issue can be taken other than to note that the treatment and photographs of Habenaria quinqueseta are oe H. ues yn. H. quinqueseta var. macroceratits) ane the curious treatment of Sa inceolata and Stenor hy nchos squamulosum. The a tw | ld be treated as Sacoila ee - | I lif { | Sacoila and Steno or absence of a mentum (in the latter). The English translation is excellent and very readable and the only error noted would be the incomplete name, or omission of a period after the abbreviation, in one of the authors’ names: Gal for Galeotti. The book is pamiede on very cies Blew heavy paper and durably bound. It is to be highly recommended toall i f the Caribbean and south Florida as well.—Paul Mar- SIDA 22(1): 841. 2006 BRIT.ORG/SIDA 22(1) tin Brown, author, Wild Orchids of Florida, The Wild Orchids of North America, i d oe rids of the Southeastern United States, 10896 SW 90" Terrace, Ocala, FL 34481, U.S.A. na Splendor in the Hill Country Grass BRIAN and Suir_ey Lorin. 2006. Grasses of the Texas Hill Country. 2006. (ISBN 1-58544-467-7, flexbound with flaps). Texas A&M University Press, 4354 TAMU, College Station, TX 77843-4354, U.S.A. (Orders: 979-458-3982, 979- 847-8752 fax; http://www.tamuedu/upress/). $23.00, 208 pp., 170 color photos, 7 lin art, 2 maps, black and white figures, glossary, bibliography, index, 53/4" x 81/2". Grasses of the Texas Hill Country is a pictorial guide to grasses of the Hill Country region of Texas. Designed for the layperson, this book is full of crisp color photographs on a black background that clearly show the inflorescences (seed heads) of grasses common in the Hill Country as well as many other areas of Texas. Authors Brian and Shirley Loflin have created a guide that is easy to use with grasses grouped by pan in inflorescence structure. The front and back book flaps serve as ler types and the icons used throughout the text. handy The ope ag chapter of the book introduces readers to the Hill Country region and other veg- etation regions of Texas. The introduction discusses the basic structures 5 of grass and includes color d t e different inflores- images. Within the “how to use this book” section is a detail cence structure types used to organize the grasses in the book. This ene section includes other items of interest such as a discussion on how . are ey scientific names, the meaning of those names, and a list of both the genus and commo es of grasses included in this book. here are seventy-seven grasses included in the or Each grass has an associated profile that eo includes features of the species, plant uses, growing season information, and habitat information in- cluding soil, location and distribution. The features section of each profile includes important infor- mation such as typical height, description of grass inflorescence, some vegetation characteristics, whether the grass is cool or warm season, native or introduced and whether the grass is annual or perennial. Much of the feature information is included in a bar under the common name of the grass ina rather intuitive icon form. The economic usage sections describe a species’ use as a forage grass, pace grass, aneseane grass, or value tow beanies ce grass profile has at least one large image of the int photographs of growth habit in nature. There are a few items that would have made this book more useful to both beginners and more experienced grass speenva tnete are ® fantastic pee ar as of the inflorescences included with each grass, but unfortur ately pho tographs of the vegetative structur ies (ligule and auricle) needed to nee identify grasses when the inflorescence is not available. However, the au- nin the “features” of the plant descrip- an thors have included some vegetative structure information wit tion. It would have also been beneficial to have the growth habit photographs th lat accompany many of the plant descriptions for every grass included in the text. There were a couple of typos/mix-ups no- ticed; an example is the aioe is p hoa in Figure 5 where the auricle was incorrectly labeled as the ligule. Some other small t roticed. Despite these few items, there is no doubt that this is a very useable book for persons with an interest in learning grasses of the Hill Country area If you are a beginner to grass identification or are looking for a good color photograph guide to grasses tor the Hill Country (and surrounding area) of Texas, Grasses of the Texas Hill Country by Brian and Shirley Loflin would be a terrific book to start with. The glossary, writing style and icon quick guide add to make te ee ial eos an easy to reference and use. This book ise sa) recom- of: mended for use b planti cation enthusiasts.—Lee Lucke ydoo, *y Herbarium, Botanical Research Thsti tute afie xas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. SIDA 22(1): 842. 2006 BOOK REVIEWS FREDERICK CAMPBELL and RICHARD Dube. 1997. Landscaping Makes Cents. (Paper- back Edition). (ISBN 0-88266-948-6.). Storey Publishing, 210 MASS MoCA Way, North Adams, MA, 01247. U.S.A. (Orders: # 66948, 1-800-441-5700, wwwstorey.com). $16.95, 170 pp., color and b/w figures, appendices, glos- sary, further reading guide, index, 81/2" x 11" If you want to improve the look of your homes’ landscape and increase its’ resale value at the same time, pe sure to pick ap a copy of i Landscaping Makes Cents yy ceapod and Dube. The book is | who is either Ul landscape by themselves or with the help of an outside contractor. The book is div ided ne three main sections, the first of which isa discussion of how landscaping can add or detract value from a home. The second section is how to plan for hardscape elements and plants as well as on thoughts on HOuRUIBUnE a — The final section contains all the info you need to know when hiring an The first section is subdivided into four chapters that can help readers recognize which ele- ments of ier a add phe) most value to EheDE property; these ee include several tip boxes pecial | tics. Some of the more specific topics include how to deteumine the values of trees, pl “int igen removal, adaition, environmental value, historic value and aesthetic value. The final chapter of the first section s] addresses landscape consider- ations when buying, puree and sell ing a home. ine second section of tl hi I hei dit land- id e anninga pices: anda landscape scaping considerations for ine with various periods of time until oe ele and finally the idea of financing help. Readers will need to investigate into budgets for their own situations, as the inlomtation included i in cl hapter five on budgeting is ie generalized. Chapter six of this sec- as well as multiple assessment tools to help readers think through and de- sign their ee For eck one of the first tools er in this chapter is “What do you want in your landscape?” with seventy-plus | uae epien: you ee want, including patios, stor- hie ies S. ane authors and an age sheds, small fruit trees, hot tubs, benches, gaze melee a short discussion of aesthetic landscape He p ve lities for outline for plotting the actual design. Chapter seven discusses investing in cee elements with inclusion of wall types fences patios, we alkways arbors drip edges ee and others. Once again, the authors include a number of helpful tip and eonsideradon boxes. Chapter eight provides the reader ideas for planting in their landscape. There are discussions of different plant types, growth forms and some helpful hints and te for planting plants. One would need to do further research into the m well in their ea Hula areas (the Anno’ nae provided a Usbe2 zone al _Th fj lch rot plants that would ae diness map inan scape. Readers are provided with many aoe and ideas for planting a Suse able Saas mee an excellent assessment tool for performing a self-environmental audit of ones’ own rty. The final section of the book contains three chapters, which cover all aspects of hiring a pro- ee ce. contractor. For those readers interested in hiring such a person, these chapters will be especially helpful for you. This section's first chapter discusses how to hire a contractor be- ginning with the titles, definition and services offered by various landscape professionals; it includes things to ask about prior to hiring the person. The second chapter of this section focuses on negotiat- chapter is a very in-depth chapter on what questions to ask of contractor ing contracts. The final during the planning sae and how to Rian specific concerns. There is also an included appendix on how to evaluate the aesthetic value of ones’ landscape. ee ng Ma i Cents by Frederick Campbell and Richard Dube is easy to read and includes helpful tip eeu aes cies uae topics for nae in planning and designing a home land- Lands ng as a fantastic resource for readers who have an in- SIDA 22(1): 843. 2006 844 BRIT.ORG/SIDA 22(1) Es terest in, either personally or through a landscape contractor, designing their home's landscape to improve both its financial and aesthetic value.—Lee Lucke ate - rbarium, Botanical Research In- stitute of lexas, 509 Pecan Street, Fort Worth, TX 76102-4060, U. Ropert H. MOHLENBROCK. 2006. Aquatic and Standing Water Plants of the Cen- tral Midwest: Filicineae, Gymnospermae and Other Monocots, Excluding Cyperaceae. (ISBN 0-8093-2670-1, hbk.). Southern Hlinois University Press, Carbondale, IL, U.S.A. (Orders: 618-453-6633, http://wwwsiuedu/‘siupress). $65.00, +00 pages, b&w line drawings, glossary and indexes to the com- mon names and genus and species, 6" X 9", Aquatic and Standing Water Plants of the Central Midwest: Filicineae, Gymnospermae and other monocots, excluding Cyperaceae by Mohlenbrock is a flora, which consists of the ferns, conifers, and other monocots, excluding the sedges portion of the larger Aquatic and Standing Water Plants of the Central Midwest series. The Cyperaceae family is included in another book by the same author within the same series title. The book’s Central Midwest range includes: Ohio, Indiana, Kentucky, Illinois, lowa, Missouri, Nebraska, and Kansas The flora includes information on 221 species, including members of the genera Azolla, Wood wardid, Equisetum, lsoetes, Lycopodiella, Osmunda, Acorus, Sagittaria, Elodea, Iris, Juncus, Lemna, Najas, Platanthera, Spiranthes, Calamagrostis, Echinochloa, Glyceria, Leersia, Zizania, Heteranthera Potamogeton, Sparganium, Lypha, Xyris, and others. There is no overall key to ie mine family, but = the individual families have keys to determine genus, as well as within genus keys to determine spe- cies. Each species is presented in the book by black and white drawings, which provides images in- cluding: plant habit, leaves, inflorescence, achene, fruit, and/or sheath/ligules a many grasses, as las an accompanying detailed description. he description for each species contains a great deal of information, including the current we accepted specific epithet, synonyms, as well as the authority and references for publications on each specific epithet. Each species’ description includes plant habit information, descriptions and mea- surements of leaves and flowers (when applicable), root types, inflorescence, notable colors and, where ap] plicable, details on parts unique to the family or genus, such as root collar information in Poaceae. The description eeecases information, witha listing of which states the species have wetland designations (ie., FACW) within the U.S. Fish and Wildlife Wetlands Inventory. The author has also included handy oe field traits within the plant description to hel p determine that species from similar looking others. Aquatic and Standing Water Plantsof the Central Midwest: Filicineae, Gymnospermae and other monocots, excluding Cyperaceae by Mohlenbrock would be a nice addition to the library of any per- ee Mohlenbrock’s book on Cyperaceae within the son who works with aquatic plants, right a same series. Although this series focuses on the Midwest, a number of the species included in this flora are found in various regions of North America. The black and white drawings are crisp, easy to read images that show major Saran traits for the species. Some familiarity with family traits will be necessary to know which family key to use to identify a plant. The keys are straightforward, but the flora does contain a glossary for any unfamiliar terms, and is user friendly. The description for each species is thorough, which can help you conclude that you have determined the correct spe- ] ] cies or not; and includes very helpful “in the field” identification hints. If you need to identify mono- cots and ferns in aquatic regions, be sure to pick up this helpful book. —Lee Luckeydoo, Herbarium, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A SIDA 22(1): 844. 2006 BOOK REVIEWS 845 French/English Review Dov FE Sax, JOHN J. StAcHOWwicz, and STEVEN D. Gaines (eds.). 2005. Les invasions despéces: Considérées dans le cadre de l’écologie, de Pévolution et de la Biogéographie. Sinauer Associates, 23 Plumtree Rd, Sunderland »MA oe 0407, US.A.(Orders: 413-549-1118 fax; ord om; Ww $51.95, 495 pp., b&w figures and tables, 7"x 91/4". ee —Ce sate constitue une vue a ensemble de Hompueuet> pECHEICH ayant été menées sur les éc ologique | 1 l’évolution a ws x a o >) a L | ee hinl De sept chapitres ont leurs peopes illustrés grace a des exemples tirés de publications i bien le milieu aquatique, aérien que E eae es grands noms de la recherche se sont rassemblés pour mettre leur savoir en commun. La contribution ee ae Soles a peums de prendre en considoauog un large éventail de eal 6 AUX iaires, en passant par le role des maladies infectieuses sur les communautés ace on exemple. rés les auteurs, ce livre peut-étre lu dans sa globalité afin de connaitre les tenants et aboutissants de nombreuses études antérieures, mais peut également étre abordé par chapitre same selon les préoccupations de chacun. e lecteur peut Come Nae e comment Petude des espéces invasives est devenue une discipline res dé s. Par exehipis Petude des BSpeces | invasives permet sient phare depuis | ‘appréhender la facon dont : ver les pro- cessus en temps réel, plutét que de déduire les processus ayant eu lieu dans | e passé seulement grace aux modéles quils ont pu engendré Ce livre est le dernier ouvrage sorti et constitue le premier de référence pour toute personne voulant orienter ses recherches vers les théories les plus récentes et vers les méthodologies dediées a létude des invasions d’espéces. English.—This book is an overview of previous research as well as current investigations about spe- cies invasions. The study of non-native mere their invasions can provide insights into ecology, evolution, and biogeography. ere are seventeen chapters that cover everything from — to aerial and terrestria inva- sions; each chapter ends with its own set of supporting references. Leading researchers were brought together to review the field of invasion Pioloay. an contri bution of 45 scientists has provided the editors with a diverse array of i communities, avian populations on islands, to the role of the infectious diseases on natural communities. The editors note, “Individu- ally, we believe that each of these chapters has something significant to offer. Collectively, we ho that this book has much to offer to both invasion biology and to our fundamental understanding _ ecology, evolution, and biogeography.” reader wil — quickly gain an understanding of why the study of invasive species has be- come an important scientific discipline in the last few decades. For instance, the study of invasive ae helps to better understand the function of an ecosystem. “Looking closely at invasions allow s to observe processes in real time, rather than having to infer the operation of processes that oc- see in vihe past solely from the aus they eave s the most recent book on species invasions and is recommend it to anyone inter isis in the ee of invasive species . isa source o oD to odologies and will t king in this discipli Virginie SIDA 22(1): 845. 2006 846 BRIT.ORG/SIDA 22(1) H. Raquet, Graduate Student, Botany Curatorial Assistant, ELL. d Herbarium Department of Bio- logical Sciences Texas Tech University Flint & Main ae nae , TX 79410-3131, U.S.A vireinie raquet@ttuedu Oo tT ANNA Pavorb. 2005. The Naming of Names. (ISBN 1-59691-071-2; hbk.). Bloomsbury Publishing, 175 Fifth Ave., New York, NY 10010, U.S.A. (Orders: http:// www.bloomsbur y.com/). $45.00, 384 pp., illustrated, 8" x 9 5/8" T animals, germs, stars, storms, rocks, and other huge kingdoms have eventually ee broken down into types and grouped so that we could begin to understand them. The Naming of Names traces t — ae natural world presents innumerable objects which humans have needed t i | he search for order in the natural world. Such a process has taken many centuries, and we have gotten better at it with a scientific understanding of the world, but the impulse has been there for as long as we have been thinking about the things around us. Pavord, author of the The Tulip (2001), Flower Power: the Meaning of Flowers in Art (2003), and an expert gardener, details the history of plant tax- onomy from the ancient Greeks to 17th-century British botanist John Ray. She reveals the history of plant classification and shows how the process was affected by intellectual, political and cultural cues The journey, traced here in detail for the first time, involves the culture of Islam, the first the Indies and the first eons in the New World. In Athens, Aristotle's pupil Theophrastus was the first man ever to write a book about plants. How can we name, sort, and order them? He asked. 7 The lebat s still, two thousand years later. The Naming of Names gives a compelling nsigne intoa wen in oF intr igus and patensely sonrpeHtye egos. She has gone back to the ancient Greeks sance tried to get a grasp on the disorderly a kingdom, with eventual success even bel fore the taxo- nomic standards see down by Linnaeus which we still follow. It is the pre-Li in efforts that Pavord has chr icled. 1 reisalsoa histo OTY O of plant illustration W ithin the se pages. The eventual woodcuts did not have to Be = with many meprodiices Here S107 ae swirling masses of plants or delicate leaves in fine detail. 1 t books could show enough useful detail to be excellent field g ike dithaush for centuries authors neliea on previous works of folklore. Along with Theophrastus, Pavord’s highest praise goes to Englishman John Ray, who in 1696 coined the term “botany.” He provided six rules by which to categorize plants, not only the ones familiar to him in England, but the spectacular finds being brought from distant lands. Others had previously insisted on classifying plants by use, which was entirely artificial, or more helpfully by leaf or seed form, but it was Ray who ue pokes on its ou irst real nec oi nouns the distinction of seeds that sprout with one leaf ort ). He kne W he Was pal 18 ol an ongoing process, predicting hae future botanists would look back and “our proudest discoveries will seem slight, obvious, almost worthless.” He might have been right, but seen as a tribute to their efforts, Fhe Naming of Names shows how fe discoveries, achieved over eas nine ao curious, ia voted, and fallible plantsmen, have brought us to our current und ane ith Ray, barely mentioning the recent advances that have been ee with DNA testing; euch ests have confirmed much of what was eventually realized as o speauae es tree, but have upset ac Pere as well. It nas peeH a a long botanical trip 5», and P. ‘ ps shi 1 inclusion of rmous fun. The book eee illustrated, with a third of the | pages being taken up with ileaniees (most in color) nicely keyed to the text Int he best sections, she slows down to draw detailed portraits of researchers and describe how each contributed to the ay evolv ing sand until Me ate 1000s, Enmenee) science of botany. The ] story makes ] { 1] | } Lue i HIAaAGE LIC ee Ol comprehensible. It would be an eelledt choice for Sublic libraries w ‘ith al ipi sted in the history of botany.—Gary Jennings, Botanical Research Institute of Texas, 509 ee Street, Fort Worth, TX 76102-4068, U.S.A. SIDA 22(1): 846. 2006 an - 1d i in si A ais - 7 ; JUL TANA | BAM ae 1 ( yay * el i, rey Pub A ANS fA! PUORIDA TOVVA ee gS A 7. VIRGINIA OREGON ‘NE -BRASK eS A NEW Me NICO. C J PUPP? ORTH CAROLINA SOUTH CAROLINA MONTAR Vv olf the Hlustrated Flora of East Texas is the first fully illustrated flora for East Texas, a species-rich area that contains roughly two-thirds of all the plant species known for lexas. The book covers all the native and naturalized ferns and similar plants, gymnosperms, and monocotyledons (1,060 species) known to occur in East Texas Published by Botanical Research Institute of Texas Pars NG Texas Floras Pro ojec £7, proj Texas (B and the Austin College Center for Environmental Studies. Pr f| od x10 //" 4 pages 1; ee bw sia 5 ] ,0004 Sida, oe 426 Illustrated Flora es of East Texas isbn 1-889878-12-X, X, hbk. George Diggs, Barney Lipscomb, May 2006 Monique Reed, Robert J. O’Kennon $89.95 Texas residents add 8.25% tax ($7.42) From Big “D" ' ' of beau prone iDisecken we groves of timber, please inquire) ( rchids nay ads, azaleas, and Texas Pride! Available from Botanical Research Institute of Texas Press 50 9 Pecan Street Fort Worth, TX 76102-4060, U.S.A ‘I E-mail: sida@brit.org Fax: 1-8 2-4112 Fost: AO www.brit. Lae a/sbm/ BRIT his photographic guide to the Wild Flowers of ; - ; WILD FLOWERS OF MOMBACHO Mombacho is color-coded and arranged by Family NICARAGUA within the color sections. It covers flowering plants found Sie alee ee MOM on the whole of the Volcano; which includes both tropical cloud forest and dry, deciduous forest, at lower altitudes. The book covers 200 plants, each of which has two or three photographs and a short description in both English and Spanish giving common name(s); a short description of the plant; height; flowering period; habitat and any known use of the plant. The vast majority of plants are likely to be found in similar habitats throughout tropical HELEN PICKERING America and nearly one third are pan tropical, making this book of interest to a wide audience throughout the tropics. WILD FLOWERS OF MOMBACHO Sida, Bot. Misc. No 28, 2006 pisiieck issn 0823-1475 FLORES SILVESTRES DEL MOMBACHO isbn 1-889878-14-6 BY HELEN PICKERING 6.5” x 8.5", x + 217pp. over 500 photographs @ $15 + p&p" ‘International: $7 BI 211 Texas residents add $1.24 sales tax PRESS : RRANSAS FLORIDA IOWA MiISSISS Barney L. Lipscoms, EDIToR Botanical Research Institute of Texas 509 Pecan Street Fort Worth, Texas 76102-4060, USA 2-444] / 817 332-4112 FAX Electronic mail: sida@brit.org Home page at the URL: http://www.brit.org/sida/ S1DA CONTRIBUTIONS TO BOTANY WAS FOUNDED BY 7 JOHN W. THIERET, ASSOCIATE EDITOR Lovo H. SHINNERS (LEFT) Dept. of Biological Sciences Northern Kentucky Universit IN 1962. INHERITED BY Perel epee ea Ta Highland Heights, KY 41099 USA WILLIAM F. MAHLER (RIGHT), DiRecTOR EMERITUS FELIX LLAMAS, CONTRIBUTING SPANISH EDITOR oF BRIT IN 1971, AND Dpto. de Botanica, Facultad de Biologia l niversidad de Leon SINCE 1993, IT HAS BEEN E-2471 Leon, SPAIN PUBLISHED BY Brit PRESS. . os The views expressed in this journal do not necessarily ate reflect those of the editors or of the Botanica Research Institute of Texas Guidelines for contributors are available upon request and on our SIDA home page as well as the last pages of each volume. Subscriptions for year 2006: $39. Individual $75.USA Institutions $85. Outside USA numbers issued twice a year OSIDA, CONTRIBUTIONS TO BOTANY VOLUME 22, NUMBER 2, PAGES 847-1274, COPYRIGHT 2006 BOTANICAL RESEARCH INSTITUTE OF TEXAS PRINTED IN THE UNITED STATES OF AMERICA SSN 0036-1488 TABLE OF CONTENTS A new name for Sida, Contributions to Botany (1962-2006) BARNEY LipscomB SYSTEMATICS A new species of witch-hazel (Hamamelis: Hamamelidaceae) apparently endemic to southern Mississippi S.W. LEONARD Eriogonum callistum (Polygonaceae), a new species from the Tehachapi Mountains of California JAMES L. REVEAL Six new combinations in Dodecatheon (Primulaceae) JAMES L. REVEAL Ericameria ubsp phila (Asteraceae), a new rabbitbrush from the San Luis Valley of Colorado LorAN C. ANDERSON A new species of Boltonia (Asteraceae) from the Ridge and Valley physi- ographic province, U.S.A. JOHN F TOWNSEND AND VESNA KARAMAN-CASTRO A new species of waterlily (Nymphaea minuta: Nymphaeaceae) from Madagascar KENNETH LANDON, RICHARD A. EDWARDS, AND P. IVAN NOZAIC Two new combinations in Peyritschia (Poaceae: Pooideae: Aveninae) Victor L. FINOT, PAUL M. PETERSON, AND FERNANDO O. ZULOAGA Poa matri-occidentalis (Poaceae: Pooideae: Poeae: Poinae), a new species from Mexico PAuL M. PETERSON, ROBERT J. SORENG, AND YOLANDA HERRERA ARRIETA Bromus ayacuchensis (Poaceae: Pooideae: Bromeae), a new species from Peru, witha key to Bromus in Peru JEFFERY M. SAARELA, PAUL M. PETERSON, AND NANCY F. REFULIO-RODRIGUEZ Psidium cauliflorum (Myrtaceae), a new species from Bahia, Brazil Lesiie R. LANDRUM AND MARCOS SOBRAL New species of Myrtaceae from Ecuador and Peru Bruce K. Hoist AND MariA LUCIA KAWASAKI Zeuxine pantlingii, sp. nov. (Orchidaceae), a new species from India AVISHEK BHATTACHARJEE AND HJ. CHOWDHERY Four novelties and a lectotypification in Matelea (Apocynaceae: Asclepiadoideae) from Hispaniola ALEXANDER KRINGS New combinations in the genus Cymopterus (Apiaceae) of the southwestern United States RONALD L. HARTMAN 873 Six new species of Sphagnum (Bryophyta: Sphagnaceae) from North America RICHARD E. ANDRUS Review of Crataegus series Pulcherrimae (Rosaceae) J.B. Puipes, RJ. OTKENNON, AND K.A. Dvorsky Crataegus series Bracteatae and Triflorae (Rosaceae) J.B. Pipes, R. LANCE, AND K.A. Dvorsky Morphometric analysis of an Amelanchier (Rosaceae: Maloideae) complex on the Delmarva Peninsula (Delaware, Maryland and eastern Virginia) re- solves the taxonomic identities of Amelanchier obovalis and A. canadensis CHRISTOPHER T. FRYE Carex reznicekii, a new widespread species of Carex section Acrocystis (Cyperaceae) from eastern North America Davip A. WERIER A new species of Eugenia (Myrtaceae) from the Monteverde region, Costa Rica FRreD R. BARRIE Taxonomic review of Symphyotrichum patens (Asteraceae: Astereae) Guy L. Nesom Drepanostachyum falcatum var. sengteeanum identity and origins (Poaceae: Bambusoideae C.M.A. STAPLETON Rejection of lectotypification of Aster amellus (Asteraceae: Ast ) and se- lection of a new lectotype JOHN C. SEMPLE ANATOMY AND MORPHOLOGY Trichome morphology in selected Mexican red oak species (Quercus section Lobatae) M. Lucia VAZQUEZ BOTANICAL HISTORY Thomas Walter Typification Project, I. Observations on the John Fraser folio DANIEL B. WarD FLORISTICS, ECOLOGY, AND CONSERVATION Evidence for hybridization between two sympatric violet species, Viola grahamii and V. hookeriana (Violaceae), in central Mexico AuREA C. CorTES-PALOMEC AND Harvey E. BALLARD, JR. Herbivory of feral goats on Espiritu Santo Island, Gulf of California, Mexico José Luis LFON-pDE LA LUZ AND REYMUNDO DOMINGUEZ-CADENA A demographic and ecological analysis of Geocarpon minimum (Caryophyllaceae): a federally threatened species in southwest Missouri TIM SMITH AND JOSEPH S. ELy 1009 1027 1049 1071 1075 1081 1087 1091 1111 1119 1135 1145 A quantitative study of styles and achenes of terminal and basal flowers of Schoenoplectus hallii (Cyperaceae), a rare plant species of transient wetland habitats MaRrIAN SMITH, SARA AMMANN, NANCY PARKER, AND PAIGE METTLER-CHERRY Annotated checklist of the vascular flora of the Lance Rosier Unit, of the Big Thicket National Preserve, Hardin County, Texas Larry E. BROWN, BARBARA R. MACRopserts, MICHAEL H. MACROBERTS, PauL A. HARCOMBE, WARREN W. PRuEss, I. SANDRA ELSIK, AND STANLEY D. JONES An updated, annotated vascular flora of Caddo Parish, Louisiana, with notes on regional phytogeography and ecology BARBARA R. MACROBERTS AND MICHAEL H. MACROBERTS Carex stipata (Cyperaceae), Chamaesyce geyeri (Euphorbiaceae), Eurytaenia texana (Apiaceae), Pediomelum esculentum (Fabaceae), and Talinum calycinum (Portulacaceae) deleted from the Louisiana flora BARBARA R. MACROoBERTS AND MICHAEL H. MACROBERTS New vascular plant records for New Mexico RONALD L. HARTMAN, BRIAN REIF, B.E. NELSON, AND BRIAN JACOBS We . (Cc 1 7 f£ Galium tricornutum (Rubiaceae) and P iaceae) new to Oklahoma Amy K. BUTHOD AND BRucE W. HOAGLAND Capraria mexicana (Scrophulariaceae) in Cameron County, Texas: rediscov- ered in the United States ALFRED RICHARDSON AND KEN KING Range expansion of Hexalectris grandiflora (Orchidaceae) in Texas MARGARET BROWN-MARSDEN AND ANNE B. COLLINS Erodium malacoides (Geraniaceae), new to Texas Davip E. LEMKE AND JED L. APLACA New distribution records for Eupatorieae (Asteraceae) in the United States Guy L. Nesom AND BILLIE L. TURNER 1159 1175 1191 1221 1225 1235 1237 1239 1245 1249 Book reviews and notices 862, 866, 894, 904, 930, 954, 958, 1008, 1026, 1074, 1090, 1134, 1144, 1158, 1174, 1190, 1220, 1224, 1234, 1254 Announcements 847, 1236 Sida, Contributions to Botany update—1257 Reviewers for Volume 22, 2006-1258 Index to volume 22, 2006 Titles of Articles With Authors—1260 Authors—1264 Botanical Names and Subject—1266 New Names and New Combinations—1272 Index to new names and new combinations in Sida 22(2), 2006 Boltonia montana J.F Townsend & V. Karaman-Castro, sp. nov—874 Bromus ayacuchensis Saarela & P.M. Peterson, sp. nov—919 Calyptranthes manuensis B. Holst @ M.L. Kawasaki, sp. nov—931 Carex reznicekii Werier, sp. nov—1052 Cymopterus breviradiatus (WL. Theob. & CC. Tseng) R.L. Hartm., comb. & stat. nov— 955 Cymopterus davidsonii (J.M. Coult. & Rose) R.L. Hartm., comb. nov—956 Cymopterus glomeratus var. fendleri (A. Gray) R.L. Hartm., comb. & stat. nov—956 Cymopterus sessiliflorus (WL. Theob. & CC. Tseng) R.L. Hartm., comb. nov—956 Dodecatheon clevelandii var. gracile (Greene) Reveal, comb. nov—863 Dodecatheon clevelandii var. insulare (HJ. Thomps.) Reveal, comb. nov—863 Dodecatheon clevelandii var. patulum (Kuntze) Reveal, comb. nov—863 Dodecatheon pulchellum var. macrocarpum (A. Gray) Reveal, comb. nov—863 Dodecatheon pulchellum var. shoshonense (A. Nelson) Reveal, comb. nov—864 Dodecatheon utahense (N.H. Holmgren) Reveal, stat. nov—864 Ericameria nauseosa subsp. ammophila L.C. Anderson, subsp. nov—868 Eriogonum callistum Reveal, sp. nov—857 Eriogonum Sect. Lanocephala Reveal, sect. nov—857 Eugenia haberi Barrie, sp. nov—1071 Eugenia yasuniana B. Holst @ M.L. Kawasaki, sp. nov—934 Hamamelis ovalis S.W. Leonard, sp. nov—850 Matelea corynephora Krings, sp. nov—942 Matelea phainops Krings, sp. nov—948 Matelea rhynchocephala Krings, sp. nov—949 Matelea torulosa Krings, sp. nov—951 Nymphaea minuta K. Landon, R.A. Edwards & PI. Nozaic, sp. nov—887 Peyritschia howellii (Hitchc.) Finot & PM. Peterson, comb. nov—897 Peyritschia pinetorum (Swallen) Finot & P.M. Peterson, comb. nov—899 Poa matri- occidentalis P.M. Peter: son & pOune sp. nov.—906 oa matri lentalis sub I i g & PM. Peterson, subsp. nov—911 Psidium cauliflorum Landrum & Sobral, sp. nov—927 Sphagnum beothuk Andrus, sp. nov—966 Sphagnum bergianum Andrus, sp. nov—964 Sphagnum kenaiense Andrus, sp. nov—961 Sphagnum mcqueenii Andrus, sp. nov—959 Sphagnum sitchense Andrus, sp. nov.— 969 Sphagnum talbotianum Andrus, sp. nov—970 Zeuxine pantlingii Av. Bhattacharjee @ HJ. Chowdhery, sp. nov—935 A NEW NAME FOR SIDA, CONTRIBUTIONS TO BOTANY (1962-2006) Barney Lipscomb Botanical Research Institute of Texas Fort Worth, Texas 76102-4060, U.S.A. barney@brit.org Beginning in 2007, the journal Sida, Contributions to Botany will no longer be published by that name but instead will be called the Journal of Botanical Research Institute of Texas (J. Bot. Res. Inst. Texas). The new journal will have a new look but generally the same format, botanical focus, and editorial direction will remain. Reason for chang The leadership of. the Botanical Research Institute of Texas has decided the name “Sida” is offensive and has too much of a negative connotation because of its similarity to the word meaning AIDS in Spanish, French, Italian, and Portuguese. Perhaps the name change is a good thing but lam personally saddened to see Shinners’ original and long-standing name displaced. Brief History of Sida 1962—Lloyd H. Shinners, a member of the Southern Methodist University faculty anda prolific researcher and writer who wanted to edit his work and the work of others, pub- lished the first issues of Sida. He named the journal for a genus of yellow-flowered plants of the mallow family, distributed throughout the world and especially common in Texas. Shinners continued as editor and publisher until his death in 1971. 1971—Wnm. FE. Mahler, professor of botany at SMU, became editor and publisher. 1977—Barney Lipscomb became assistant editor to Wm. F Mahler. 1982—Barney Lipscomb became editor, Wm. F Mahler remained publisher. 1993—With Barney Lipscomb as editor, Sida became a publication of the BRIT Press. 2002—BRIT celebrated the 40th anniversary of Sida, Contributions to Botany. 2006—BRIT Press publishes last issue of Sida, Contributions to Botany. Thank you To all authors, we thank you for choosing Sida, Contributions to Botany. A big thank you to the hundreds and hundreds of individuals who generously supported Sida through their time and expertise in reviewing more than 1000 manuscripts published from 1962- ZOO Thanks to all authors, reviewers, subscribers (individuals, institutions, and readers for your continued interest and support. It is our plan to continue bringing you the best sources of current research in classical and modern systematics through the Journal of Botanical Research Institute of Texas with your continued support. I personally wish you the best in 2007 and beyond. Your comments are welcome. SIDA 22(2): 847 — 848. 2006 848 Publication dates of individual issues of Sida, Contributions to Botany. BRIT.ORG/SIDA 22(2) Volume Issue Pagination Publication Date Volume Issue Pagination Publication Date 22 2 847-1274 22 Nov 2006 10 4 269-341 27 Nov 1984 oy) 1-846 11 Aug 2006 10 3 195-267 8 Jun 1984 21 4 1955-2511 21 Dec 2005 10 2 95-193 28 Nov 1983 21 3 1249-1954 5 Aug 2005 19 1-93-28 Apr 1983 21 2 511-1248 28 Dec 2004 9 4 269-381 7 Dec 1982 21 1 1-150 20 Aug 2004 9 3 195-268 10 Jun 1982 20 4 1343-1836 22 Dec 2003 2 2 “194 24 Nov 1981 20 3 877-1341 29 Jul 2003 = : 1-97-29 Apr 1981 20 2 461-876 23 Nov 2002 8 4 317-372 17 Dec 1980 20 1 1-460 19 Jul 2002 8 3 218-315 11 Jun 1980 19 4 767-1233 28 Dec 2001 8 2 123-216 30 Nov 1979 19 3 445-766 23 Aug 2001 : ep eee Tee 19 D 235-444 20 Dec 2000 7 4 309-397. 13 Dec 1978 19 1 1-234 23 Sep 2000 7 3 227-308 30 Jun 1978 18 4 941-1316 23 Dec 1999 d 2 94-226 9 Dec 1977 18 : 631-940 20 Sep 1999 if 1-93 27 May 1977 18 2 377-630 28 Dec 1998 6 4 243-331 29. Sep 1976 18 1-376 15 Oct 1998 6 3 123-242 14 May 1976 17 4 645-873 31 Dec 1997 6 2 63-122 11 Nev 1375 17 3 533-643 12 Sep 1997 S ee “eaaee 17 2 305-532 30 Dec 1996 5 5 301-361 28 Feb 1975 17 1-303 30 Sep 1996 5 4 191-300 16 Sep 1974 16 4 613-809 29 Dec 1995 5 3 137-190 21 Jan 1974 16 3 389-610 28 Aug 1995 2 : pon )eo: SN IAS 16 2 225-388 30Dec 1994 5 ie A OMal Ia? 16 1-222 26 Aug 1994 4 4 279-430 28 Sep 1972 15 4 565-669 29 Dec 1993 4 3 228-278 = 22 Jul 1971 15 3 349-563 28 Sep 1993 ‘i 2 ae EBL 15 2 173-348 31 Dec 1992 2 I-36 16 Sep 1968 15 1 1-172 17 Sep 1992 3 7 451-444 3 Apr 1970 14 4 525-638 31 Dec 1991 : ° ead dade 14 3 315-524 27 Aug 199] 2 a 14 2 141-314 27 Dec 1990 3 : 191-283 21 Jun 1968 a ae. oes 3 3 123-190 24 Nov 1967 3 2 77-122 31 May 1967 13 4 393-518 29 Dec 1989 3 1 _76 31 May 1967 13 3 265-392 31 Aug 1989 13 2 125-264 30 Dec 1988 7 6 are =O Der Tee ie oe 2 5 349-394 27 Sep 1966 2 4 269-348 20 Jul 1966 12 4 Never published 2 3 177-268 Apr 1966 12 3 Never published 2 2 115-175. 31 Dec 1965 12 2 253-434 30 Dec 1987 2 1 1-113 3 Mar 1965 12 1 1-251 29 May 1987 1 é 299-417 30 Oct 1964 11 4 357-497 16 Dec 1986 1 5 957-298 25 Jun 1964 11 3 2507250 20 May 1986 1 4 187-256 25 Jun 1964 11 2 107-253 12 Dec 1985 1 3 109-186 12 Dec 1963 11 ] 1-106 19 Jun 1985 1 2 63-108 23 Nov 1962 1 1 62. 23 Nov 1962 A NEW SPECIES OF WITCH-HAZEL (HAMAMELIS: HAMAMELIDACEAE) APPARENTLY ENDEMIC TO SOUTHERN MISSISSIPPI S.W. Leonard The Nature Conservancy Camp Shelby Field Office CSTS-ENV, Bldg. 6678 Camp Shelby, Mississippi 39407, U.S.A. ABSTRACT A new species of witch-hazel, Hamamelis ovalis, is described from Perry County in southern Mississippi, US.A The red flower color, clonal habit, and late: winter prOMeNng Sugeest affinity with the Ozarkan endemic H. ve ridilis however, the large leaves, unlike a merican or A witch-hazels, readily differenti- a ate H. ovalis from its congeners. The problematic H. macrophylla is discussed. RESUMEN Se describe una nueva especie de la avellana-bruja, Hamamelis ovalis, del condado de Perry, Mississippi del sur, los Estados Unidos. El color rojo de la flor, el habito conics. y ie norcen radi en i g aunadad con el Ozarkan endémico H. vernalis. Sin embargo, | de avellanas-brujas americanas 0 asiaticas eine Poiimente H. ovalis de sus Weoneencies Se discute el problematico H. macrophylla. During surveys for proposed tracked vehicle maneuver areas at Camp Shelby Training Site in southern Mississippi, an unusual colony of witch-hazel (Hamamelis) was discovered in July 2004. Distinctive features initially observed were the large, abaxially pale whitish- green ovate leaves, densely pubescent cinnamon-brown twigs, and abundance of low clonal sprouts. The colony was periodically checked to see when anthesis would occur and if the petals would be yellow like all the known native Hamamelis plants in southern Missis- sippi. In early January 2005 the large tawny buds opened to expose brilliant red sepals and petals entirely maroon, scarlet, rose, or red with yellow tips. Subsequent surveys in the area revealed colonies with mixed flower colors as well as colonies with flowers entirely wine- red. Leaf and floral appearance as well as the clone-forming habit of the plants did not match anything known for Hamamelis from the Gulf Coast region. Witch-hazel shrubs, with pale yellow petals less than 8 mm long and occasional plants with leaves less than 6 cm long are present at Camp Shelby. These yellow-flowered plants do not fit this writer’s concept of Hamamelis virginiana L. var. virginiana of the Carolinas nor Hamamelis virginiana L. var. henryae Jenne ex C. Lane of northern Florida. However, for simplicity of discussion in this paper, they will be referred to as H. virginiana. Floristic treatments indicate a range for H. virginiana from Nova Scotia to Texas (Small 1933; Fernald 1950; Steyermark 1963; Radford et al. 1968; Godfrey 1988; Meyer 1997) and possibly Mexico Jenne 1966; Lane 2005). In Mississippi these shrubs and small under- story trees are frequent on acidic, sandy woodland slopes and bluffs, moist or dry, in full sun or shade. Flowering at Camp Shelby begins in November, peaks in December and tapers off in January. In addition to the native woodland Hamamelis virginiana in Mississippi, other witch-hazels are occasionally used as landscaping shrubs. These cultivars include SIDA 22(2): 849 — 856. 2006 850 BRIT.ORG/SIDA 22(2) the Ozarkan witch-hazel, Hamamelis vernalis Sargent, and less frequently the Oriental species, Hamamelis mollis Oliver, Hamamelis japonica Siebold and Zuccarini, and the hybrid Hamamelis x intermedia Rehder. Neither H. vernalis, nor the Asian plants are known to naturalize in Mississippi. In March 2005 the dried collection of Hamamelis at the Missouri Botanical Garden Herbarium (MO) and late flowering specimens of H. vernalis Sargent on the Garden’s grounds were examined. Although considerable variation in foliage was noted, nothing in the MO collection approximated the big-leaved, red-flowered plants found earlier at Camp Shelby, Mississippi. With no visible evidence that the Camp Shelby colony was located near a house site where cultivars might have been introduced, exotic witch-ha- zels were eliminated. Leaf size, pubescence, clonal habit, flower color, petal length, and phenology tended to rule out the common eastern witch-hazel, H. virginiana—highly variable as it is known to be. Clonal habit and infusion of red in the petals suggested an affinity between the Camp Shelby plants and the variable Hamamelis vernalis,an Ozarkan endemic that grows 600 kilometers to the northwest of the Mississippi site. Rehder (1920, 1928) and Steyermark (1963) recognized forms of H. vernalisto account for the pure red flower color and tomen- tose leaves within certain colonies of H. vernalis. Jenne (1966) proposed that the variable- flowered, clonal plants of H. vernalis originated as putative hybrids between the eastern H. virginiana and an unknown parental type (possibly no longer extant). However, Bradford and Marsh (1977), af ter ] parative studies in Arkansas of H. virginiana and H. vernalis, were inclined to disagree with Jenne’s interpretation of the hybrid origin of H. vernalis. Nevertheless, as Jenne (1966) hypothesized, a tentative Ozark relict “would be expected to be extreme in the following directions: short petal length, presence of red pigment in the petals and on the inner surface of the sepals, linear staminodia, less than 0.5 mm wide, the tips of which are not dilated, clonal habit of vegetative suckering, with 75 to 100 stems emerging from one root system.” Looking at these features only, the Camp Shelby plants come uncannily close to Jenne’s parental phenotype, but size and vestiture of leaves, branching pattern, habitat, and other characteristics are hard to reconcile the Camp Shelby shrub asa parent of H. vernalis. On the other hand, with exception of variation in petal color, nothing about the Camp Shelby plants suggests a hybrid origin. In fact, other features such as larger leaf size, ear- lier flowering, musty floral fragrance, plant habit, and dry pineland habitat point to- ward an independently derived species. As documented below the Mississippi plants are sufficiently different from the Ozark plants to merit a taxonomic designation at specific rank. Therefore the following name and diagnosis are provided: ee ovalis S.W. Leonard, sp. nov. (Figs. 1-2). Type: U.S.A. Mississippr Perry Co. shallow vine in mixed pine-hardwoods near headwaters of Garraway Creek, Camp Shelby Training — T-17, Compartment 96, DeSoto National Forest, T2N, R1W, Sec. 6, NEI/4, 7 Jul 2004, S.W. Leonard 11073 (HOLOTYPE: MO, ISOTYPES: FSU, NCU, NY). Frutex clonis, crescit ad 5 m. Differt a Hamamelis virgini ) quod scarlatinus vel rubropurpureus est, floris folia sunt coloris vinosi aut rosei; differt a H. ver nelas e0 ty folia magna ovata habet et pars obversa folii habet pilos stellatos albos. Rhizomatous shrubs to 5 m tall with twigs densely stellate pubescent, brown to brown- ish-gray on older twigs; terminal buds russet. Leaves tardily deciduous, alternate, ovate 12-24 cm long, 5-17 cm wide, dark green and scurfy above, pale green with dense stellate tomentum below initially white and aging to golden tan, margins sinuate on distal half LEONARD, A NEW SPFEAICS UF 851 Fig. 1. Powel vet) eee ovalis, ii, Cee ken on January: 2006.1 (right) j | yp ality eruleris 15 c I } jal £ dq + | hy bnit J J J and entire on proximal half; petioles 1.0-3.5 cm long; leaf bases oblique with longer lobe rounded to cordate, principal veins of leaves alternate, usually 11 (five on one side of mid- rib, six on the other); leaf apices acute to obtuse. Flowering late December to early Febru- ary with peak around January 20; calyx of 4 ovate sepals 3-4 mm long, acute, densely tawny pubescent abaxially and glossy scarlet adaxially, spreading, slightly reflexed, and ultimately strongly reflexed with age, subtended by 2 or 3 densely tawny tomentose bractlets; petals liguliform, at first uncoiling 7 mm long and continuing to lengthen to 14 mm over a period of 3 weeks, circinnately coiled in bud and irregularly revolute when expanded, various shades of red (wine-red, scarlet, rose) or infrequently tipped with yel- low; filaments purplish-red, gradually expanded toward apex and resembling gavels until anthers turn outward from the connective exposing sockets in the distal ends of the fila- ments; filaments alternating with staminodes; staminodes dilated distally and curved outward with the broader tips straddling the base of each petal, thus pushing the petal away from the stigmatic surface; ovary partly inferior, pubescent with mostly appressed hairs, fruits slightly compressed ca. 2 cm long calyx adhering to capsule except for re- curved tips; seeds 2 per locule, glossy black, 7-10 mm long, slightly obovate with each face of attachment scar (2 mm wide and 3 mm long) about twice the width and length of attachment scars on seeds of H. virginiana from local populations (Fig. 1). Ecology and phenology.—At the type locality, the plants occur in a shallow flat-bot- tom ravine and on adjacent low slopes. Soil at the site is loamy. The ravine drains into a headwater tributary of Garraway Creek that flows into the Leaf River between the City of Hattiesburg and the small municipality of New Augusta in Perry County, Mississippi. The location is in Compartment 96 of DeSoto National Forest, Training Area 17 of Camp Shelby Training Site, T2N, R11 W, Section 6. The Hamamelis occurs beneath a canopy of pine and mixed deciduous hardwoods. The main species of this vegetation are: Pinus taeda, Pinus palustris, Carya tomentosa, Nyssa sylvatica, Quercus falcata, Magnolia macrophylla, Cornus florida, Sassafras albidum, Ilex vomitoria, Ilex opaca, Liquidambar styraciflua, Vaccinium arboreum, Vaccinium elliottii, Morella cerifera, Callicarpa americana, Tridens flavus var. chapmanii, 852 BRIT.ORG/SIDA 22(2) Fic? Clockwi £ ra Late. U iB ae a (F fa MS). H yi Th (P rf. MS) H, Tj, Th. (T ‘a . L MO), Hamamelis virginiana (Perry Co., MS). Each LEONARD 853 Chasmanthium sessiliflorum, Sorghastrum elliottii, Smilax pumila, Mitchella repens, Berchemia scandens, and Vitis rotundifolia. Flowering of Hamamelis ovalis was observed in late December with the peak occur- ring around the first ten days of January in 2005. The 2006 peak (during an unusually warm January) was closer to January 20th. In addition to the taller plants developing flowers, clonal sprouts—some no more than 20 cm tall—also flower. The following observations were made in 2006 on potted plants of H. ovalis and H. vernalis: flowering of H. ovalis begins (late December) as flowering of H. virginiana is ending; flowering of H. vernalis begins (mid-February) as flowering of H. ovalis is end- ing. Leaves of H. ovalis are 30-40 percent of mature size before the leaves of H. vernalis begin to emerge from the buds. Habit and morphology.—Although Hamamelis ovalis, like H. vernalis, isa clonal shrub, the root sprouts are generally uniform in height on upper slopes where prescribed fire at 2-6 year intervals isa standard longleaf pine management procedure. These small shoots, typically 3-5 dm tall, extend outward as muchas 6-8 meters from the central plant. How- ever, the habit of the H. ovalis is markedly different from that of H. vernalis. Instead of forming a rounded bush of several sucker stems with major branches diverging in all directions from near the base of the plant as in H. vernalis, mature shrubs of H. ovalis have two growth forms—distichous branching ona geniculate central axis and a graceful arched branching on an erect straight stem. On the former, alternate branches diverge at approximately right angles in the same plane, gradually becoming longer as they progress up the main stem, thus giving the plants an espaliered look. Rhizomes of H. ovalis are 6- 10 mm in diameter and are shallow, often just beneath the leaf duff. In contrast, rhizomes of H. vernalis are deep seated among overwash stones in flood zone sands. Specimens along the Caddo River in Clark County, Arkansas were tightly bound to the substrate and not easily extracted. The type locality colony of Hamamelis ovalis comprises 92 stems greater than 1.5m tall with the tallest being 3.8 m (%2.3 m) and about 120 smaller sprouts. It is estimated these 200+ stems arise from the rhizomes of fewer than two dozen plants. Field work during 2005 in the headwaters of Garraway Creek revealed about two dozen smaller colo- nies of witch-hazel in drier habitats: upper slopes, crests of ridges, and 1-2 hectare level areas now covered with mixed longleaf and loblolly pine secondary growth forests. Calyces are consistently deep reddish-maroon adaxially with conspicuous tawny hairs from the dense abaxial pubescence protruding along the lobe margins. Calyces of some plants become tinged with green toward the base of the lobes after the petals wither. Petals were predominantly wine-red, scarlet, and reddish-pink, with at least one plant having yellow petals with the basal half deep red. Petals persist for around three weeks before they begin to turn brown from the tips progressively toward the base. Floral fra- grance of H. ovalisis at first faintly spicy or “fruity,” thereafter degrades to a musty scent similar to that of decaying pears, and eventually to a disagreeable “wet dog” odor! One of the distinctive features of Hamamelis ovalis is the abaxial appearance of ma- ture leaves in the early part of the growing season. If viewed at an angle in direct sun- light, the developing leaves have a hoary white appearance above due to closely spaced stellate hairs. At the same time the lower surface is whitish (very pale green) from closely- knit stellate pubescence. On plants of Hamamelis ovalis, even short sucker sprouts, two or three of the leaves 854 BRIT.ORG/SIDA 22(2) toward the stem tip will be greater than 20cm in length. This dimension exceeds maxima for all witch-hazel taxa currently described. In autumn the leaves of H.ovalis turn yellow, then brown, and many continue to hang downward on the branches until mid-March. Anobservation that may have some bearing on data collection from herbarium speci- mens is that witch-hazels respond vigorously after forest fires. At Camp Shelby Training Site, stems of Hamamelis virginiana in the first one or two growing seasons after fire have larger than normal leaves, usually 16-18 cm long. These post-fire leaves tend to have cordate rather than oblique leaf bases and lack conspicuous abaxial pubescence. Hamamelis ovalis vs. H. vernalis . Leaves ovate, 12-24 (%19.7) cm long, 5-17 cm wide, soft velvety green above for first half of growing season and ae becoming darker green ae SC ay above; pale green va ae dense white tomentum in spring that graduall nin autumn, sinuate on distal half, base rounded, strongly cordate-oblique, usually 11 major veins arising from midrib with 5 on one side and 6 on the other; suckers from base of main stem absent; plants strongly soboliferous with rhizome sprouts mostly less than 5 dm tall; flowering late December to early February, flowers faintly spicy becoming musty for the three-week duration ez of the petals H. ovalis . Leaves mostly obovate, 7-13 (%=8.7) cm long and 4-10 cm wide glabrous above or sparingly stellate pubescent becoming darker green and smooth above; ce ous beneath or rusty stel- late pubescent along midrib and veins, margins sinuate apically, cuneate basally or infrequently weakly rounded and oblique, usually 9 or 10 major veins arising nae midrib; plants suckering strongly from base and occasionally sending up rhizome sprouts that form dense thickets 2-3 meters tall; flowering late February throughout March, flowers sweetly aromatic H. vernalis Leaves of Hamamelis ovalis are more than twice as large as leaves of H. vernalis, and sel- dom if ever have an obovate shape. In contrast, foliage of H. vernalis observed along the Caddo River in Arkansas as well as leaves of cultivated plants of this species are consis- tently obovate, slightly glaucous, and with broad cuneate bases. Flowers of Hamamelis ovalis are similar in size and possibly as variable in color as H. vernalis, but lack the pro- nounced fragrance. Hamamelis ovalis vs. H. virginiana arge ovate leaves with pale densely stellate tomentum on abaxial surface; mostly 11 lateral veins per leaf; abundant short rhizome sprouts sometimes forming a groundcover; tawny pu- bescent twigs of current and previous season's pe flower buds and terminal buds larger, scarlet adaxial surface of calyx; maroon, scarlet, or deep pink petals and larger; flowering princi- pally in January in Mississippi; meee nat seeds with larger attachment scars H. ovalis . Leaves 3.7-16.7 cm long and 2.5-13 cm wide (Meyer 1997); elliptic, obovate, nearly rounded; margins sinuate, base broadly cuneate, ned weakly to strongly oblique with one lobe rounded the other cuneate; surface dull green above, scabridulous or smooth, lower nae pale green, glabrous or pubescent or tomentum restricted to midrib and major veins; 9 or | ateral veins arising from midrib; plants suckering from base with only a few sprouts arising from rhizomes; flowers pale yellow, smaller (6-8 mm) than published dimensions [10-20 mm see Meyer 1997)] for the species, opening in November to mid-December 1s slight srl with smaller attachment scars H. virginiana 1. cz Features that differentiate Hamamelis ovalis from the widespread eastern North American H. virginiana are the larger leaves with pale densely stellate tomentum on abaxial surface; tawny pubescent twigs, longer bud scales, scarlet adaxial surfaces of calyx, reddish petals instead of typical yellow, floral fragrance, and strong development of clonal sprouts. Flow- ering of H. ovalis begins ate December) as flowering of H. virginiana is ending. Among the nearly two dozen discrete colonies of Hamamelisovalis observed at Camp LEONARD 855 Shelby, only one has been found that contains both H. ovalis and H. virginiana (small leaf form) and at this site there are only a few plants of the latter. In areas where Hamamelis ovalis occupies drier sites at the heads of drainages, H. virginiana tends to be a stream- side plant. Identity of Hamamelis macrophylla Because the large leaf size of the new Mississippi witch-hazel is one of its most distinc- tive features, a logical name for the shrub would be Hamamelis macrophylla. This name was in fact used by John Lyon, a Scottish nurseryman working out of Philadelphia and making commercial collecting trips as far south as Georgia and Florida. Where Lyon found his new species is unknown. He offered it for sale in 1812 under the name Hamamelis macrophyllus sp. nova, but he did not publish a Latin diagnosis (Ewan and Ewan 1963). Pursh appropriated the name, provided a brief Latin diagnosis, and published the name in his Flora americae septentrionalis (Pursh 1814). Although Pursh said the type came from “river banks in the western part of Georgia,” Ewan and Ewan (1963) pointed out in a footnote to the annotated Lyon journal that some of Pursh’s locations were not “trust- worthy.” Loudon, a Lyon biographer writing in 1844, according to Ewan and Ewan (1963), said the witch-hazel was native to the “Katawba (sic) Mountains.” This location would place the type locality in western North Carolina. Regarding the Lyon/Pursh Hamamelis macrophylla, Pursh described only “large leaves punctated on their lower side, with rough tubercles and other marks...” Appar- ently he never saw flowers of H. macrophylla and if Lyon saw the plants in flower, he either withheld the information from Pursh or omitted it from his journal of 1799-1814. Pressed specimens were either lost or destroyed. A clue regarding the provenance of Hamamelis macrophylla might be ascertained from associated flora. Having seen the magnificent Magnolia macrophylla, Lyon seemed to have become obsessed with it and is said to have collected more than 4,000 seedlings from a single location in North Carolina (South Carolina Association of Naturalists news- letter June 1996). It is possible that on one of his collecting trips to Magnolia colonies in western Georgia (or western North Carolina?) he collected a large-leaved Hamamelis. According to Lyon’s notes of his Georgia travels, he was near the Ocmulgee River on June 23,1803, the Flint River on July 17, 1803, and the Chattahoochee River on October 6, 1803. However, these dates appear to be too early if the witch-hazels were not advertised for sale until 1812. A more likely site and date of collection would be western North Carolina near the Catawba River in October 1809, when Lyon packed 3,600 seedlings of Magnolia macrophylla tor shipment to Europe. agnolia macrophylla is a frequent associate of Hamamelis ovalis in the upper Garraway Creek watershed in Mississippi. If the Georgia or the Carolina plants bloomed in late winter or spring, were red-flowered and were clonal, then Pursh’s Hamamelis macrophylla might be the same species as recognized here as newly described. If the Mis- sissippi plants were a colony of a conjectured red-flowered H. macrophylla, such a dis- junction would be about 425 kilometers from a theoretical site in Georgia or more than 1100 km from a North Carolina location, based on the distribution of Magnolia macrophylla and assuming the magnolia’s range has changed little since the early nine- teenth century. Treatments of the common eastern Hamamelis virginiana have included Hamame- lismacrophylla Pursh as asynonym (e.g, Sargent 1922; Coker & Totten 1945). Unless, how- ever, John Lyon’s steps can be retraced to an extant colony of a big leaf witch-hazel, or 856 BRIT.ORG/SIDA 22 — 2) unless nursery stock from Lyon’s 1812 sale has been propagated for almost two centuries in an English garden, the true identity of H. macrophylla may never be known. ACKNOWLEDGMENTS My thanks to Alan S. Weakley, NCU Herbarium Curator, for searching references tucked away in the University of North Carolina archives and to Mary Stevens of the Mississippi Museum of Natural Science for obtaining the paper by Bradford and Marsh. Lisa Yager, Camp Shelby Field Office Coordinator, arranged interlibrary loans of two crucial docu- ments. Suggestions by reviewers, Guy Nesom, Bruce Sorrie, and Stephen Spongberg greatly improved the manuscript. | thank Fr. Peter Verhalen for corrections to the Latin diagno- sis, the Mississippi Army National Guard for funding, and the US Forest Service, DeSoto Ranger District, that has final management responsibility for conservation of the species and its habitat. REFERENCES Braprord, J.L.and D.L. Marsu. 1977.Comparative studies of the witch hazels Hamamelis virginiana and H. vernalis. Proc. Arkansas Acad. Sci. 31:29-31. Coker, W.C.and H.R. Torten. 1945. Trees of the southeastern states. Univ. of North Carolina Press, Chapel Hill, North Carolina. Pp. 195-196. Ernst, WR. 1963.The genera of Hamamelidaceae and Platanaceae in the southeastern United States. J.Arnold Arbor. 44:193-210. Ewan, J.and N. Ewan. 1963. John Lyon, nurseryman and plant hunter, and his journal, 1799-1814. Trans. Amer. Philosophical Soc. Vol. 3, pt. 2. Philadelphia, PA. Fernato, M.L. 1950. Gray’s manual of botany. American Book Company, New York. P.752. Goorrcy, R.K. 1988. Trees, shrubs and woody vines of northern Florida, and adjacent Georgia and Alabama. Univ. of Georgia Press, Athens. Pp. 372-374. Jenne, G.E. 1966. A study of variation in North American Hamamelis L. (Hamamelidaceae). Master of Arts thesis, Vanderbilt Univ., Nashville, TN. Lane, C.2005.Witch hazels. Timber Press, Inc., Portland, OR. Mever, E.G. 1997. Hamamelidaceae. In: Flora of North American Editorial Committee, eds. Fl. North Amer. Oxford Univ. Press, New York and Oxford. 3:362-36/7. Pursu, FT. 1814. Flora americae septentrionalis. London,White, Cochran and Co,, London, Great Britain. 1:116. Raprord, A.E.,H.E. Antes, and C.R. Beit. 1968. Manual of the vascular flora of the Carolinas. Univ.of North Carolina Press, Chapel Hill. Pp. 529-531. Revper, A. 1920. New species, varieties, and combinations. J. Arnold Arbor. 1:256. ReHper, A. 1928. New species, varieties, and combinations. J.Arnold Arbor. 9:30. SarGENT, C.S. 1922. Manual of the trees of North Americ a. Houghton, Mifflin and Co., Cambridge, MA. 1:368-371. Smatt, J.K. 1933. Manual of the southeastern flora (1972 facsimile of the 1933 edition). Hafner Pub- lishing Company, New York. Pp. 600-601. South CAROLINA ASSOCIATION OF NATURALISTS. 1996. June newsletter. Columbia, South Carolina. SreverMaRrK, JA. 1963. Flora of Missouri. lowa State Univ. Press, Ames, lowa. Pp. 786-789. ERIOGONUM CALLISTUM (POLYGONACEAE), A NEW SPECIES FROM THE TEHACHAPI MOUNTAINS OF CALIFORNIA James L. Reveal! ersity of Maryland College ee a and 20742-5815, U.S.A. and New } rk Botanic Cd! rde! Bronx Park, New York 10458- 5026 USA. jreveal@umd.edu ABSTRACT Eriogonum callistum, a new species of subg. Eucycla Nutt. from the Tehachapi Mountains of Kern Co., California, is the sole representative of Sect. Lanocephala, a new section allied to Sect. Lachnogyna Torr. & A. Gray, a taxon pans al ne acre ce but probably more osely revatee to Sect Pal sonia oy ot the agile peak The gla rg eC compact ensel ae new pieaves from which arise several UpHent poapoee flowering s stems terminated by a capitate inflorescence subtended by many bracts with many g hairy, bright white flowers on long, hairy pedicels, and glabrous, trigonous achenes. RESUMEN Eriogonum callistum, una nueva especie del subg. Eucycla Nutt. ede las montanas Tehachapi de Kern Co., Califor- nia, es el Unico pep ieschiante oS la Sect. Panocephala, | ite Sect. Lachnogyna Torr. & A. babl 4 a Sect Lattjola Benth. de la costa pacifica. La nueva especie ee port formar grand i lond hojosas y rosetas de hojas nuevas, de las que I iba: llos florid una inflore ia capitada subtendida por muchas bracteas, con muchos involucros ae een numerosas flores blanco brillante, sobre pedicelos largos y pelosos, y aquenios trigonos glabros. The discovery of another undescribed member of Eriogonum Michx. (Polygonaceae Juss., subf. Eriogonoideae Arn.) from California is not, in itself, a surprise, but one so distinct as to represent a new section, and of such potential as a cultivated garden plant, is at least unusual. Eriogonum Michx. Sect. Lanocephala Reveal, sect. nov. Type: Eriogonum callistum Reveal. A Eriogono sectionibus singularis floribus et pedicellis pilosis. sa ea callistum Reveal, sp. nov. (Fig. 1). Typr:U.S.A.CALIFORNIA: Kern Co: Tehachapi Mountains, y 2006, R.E. Preston 2400 (HOLOTYPE: UC; IsoryPes: NY, RSA, U A Eriogono lachnogyno Torr. ex Benth. foliis latioribus et longioribus, involucris numerosis (10-35 nec 2-5), et floribus albis (nec flavis) et E. latifolio Sm. floribus pilosis differt Plants perennial forming densely white-tomentose, rounded, compact mounds, 3-10(-11) dm across and (0.5-)1-3.5 dm high, arising from a stout, woody taproot with numerous, spreading, tightly compact caudex branches covered with marcescent leaves ultimately terminated by a tight leafy rosette of numerous, newly-formed basal leaves; leaves fas- ciculate in terminal tufts, persistent, the petiole 1.5-5 cm long, grayish-white tomentose, the petiole-base narrowly triangular, glabrous, 0.5-3 cm long, 0.3-0.8 cm wide, becoming 'Mailing address: 18625 Spring Canyon Road, Montrose, Colorado 81401-7906, U.S.A. SIDA 22(2): 857 — 861. 2006 858 BRIT.ORG/SIDA 22(2) a7 G § \ » { yy a \ ‘A !y y x \ We oe a wy Mbsuwey Fic. 1. Illustration (lef ight) of a leaf, an involucre, and a flower with its pedicel and bractlets. ~s Pp papery and reddish-brown with age, the blade elliptic, flattened or slightly folded longi- tudinally, (1-)2-5 cm long, 0.8-2 cm wide, silky grayish-white tomentose on both sur- faces, the margins entire, flat; flowering stem scapose, arising centrally from rosette, erect or nearly so, slender, solid, not fistulose, (1.5-)2-4(-4.5) dm long, greenish to grayish, be- coming reddish with age, thinly tomentose; inflorescences capitate, 2-4 cm across, bracts 8-16, subfoliaceous, linear to narrowly lanceolate, 4-12(-15) mm long, 1-3 mm wide, to- mentose on both surfaces except glabrous basally on adaxial surface; peduncles absent, involucres 10-25(-35) per cluster, turbinate, (5-)6-8(-9) mm long, 2-4 mm wide, rigid, tomentose abaxially, glabrous adaxially, the teeth 5, erect, 2-3.5 mm long, acute apically; flowers bisexual, not attenuated basally, 2-5 mm long, the pedicels pilose, (5-)6-10(-12) mm long, the bractlets linear, 4-8(-10) mm long, pilose throughout, the perianth pink- ish-white in bud, becoming bright white, densely white tomentose with long, soft hairs abaxially, glabrous adaxially except for glands along the rosy to yellowish-green mid- ribs, the tepals connate |/4-1/3 their length, essentially monomorphic, oblong to ellip- tic, the outer three slightly broader than the inner three; stamens exserted, 3-6 mm long, the filaments pilose proximally, the anthers oblong, pale yellow with a tinge of pink to light rose, 0.4-0.5 mm long; achenes trigonous, 3-4 mm long, glabrous. Other specimens examined: U.S.A. Kern Co.: Tehachapi Mountains, 13 Jun 2006, B.D. Schafer 214 (DAV, NY, RSA, UC, US); 25 Jul 2006, [-L. Reveal et al. 8781 (NY, RSA, UC, US). ~ Eriogonum callistum (from kalliston Gr. most beautiful; Figs. 2, 3) represents a new sec- tion within subg. Eucycla Nutt. The Sect. Lanocephala (from lenos Gr. wooly hairs, and REVEAL, A NEW SPECIES OF ERIOGONUM FROM CALIFORNIA 859 cae srbete tae A ae “Hee > 860 BRIT.ORG/SIDA 22(2) kephale, head, as to the pilose flowers on hairy pedicels arranged in a capitate inflores- cence) is similar to Sect. Lachnogyna Torr. & A. Gray, a taxon of two species encountered from eastern Arizona to western Texas northward onto the southern Great Plains to Colo- rado and Kansas. Although the type, E. lachnogynum, has hairy achenes, the other spe- cies, E. havardii S. Watson, has glabrous fruits. The soft, silky, grayish-white tomentum of the leaves of E. callistum is similar to that seen in both species, and while both mem- bers of Sect. Lachnogyna have yellow flowers, those of E. callistum are a bright white. The monospecific section is probably more closely allied to Sect. Latifolia Benth., and specifi- cally the coastal element of the taxon, E. latifolium Sm. Both form rounded mounds of leaves and have numerous involucres in capitate heads. Still, E. latifolium is best charac- terized asa subshrub or shrub with long, aboveground, woody stems. Eriogonumcallistum is an herbaceous perennial with short, stout, caudex branches buried in a mixture of soil and marcescent leaves. Nonetheless, Sect. Lanoce phala is the only taxon in the genus with hairy pedicels. The flowers of E. callistum are individually small but numerous in nu- merous involucres that together form a capitate cluster atop long, slender scapes. The achene is trigonous, glabrous, and contains a curved embryo in abundant mealy en- dosperm; as such it is typical of most species of subg. Eucycla. Ongoing molecular stud- ies should resolve an exact placement of Sect. Lanocephala. At present, Eriogonum callistum is known only from a few scattered locations that altogether contain some 2000 individuals in a variety of age classes. The new species grows mainly in open places among manzanita on rock outcrops of pre-Cretaceous lime- stone (Jennings & Strand 1969), on open ridges within mixed chaparral. Associated spe- cies include Adenostoma fasciculatum Hook. & Arn., Amelanchier utahensis Koehne, Arctostaphylos glauca Lindl, Arctostaphylos parryana Lemmon, Hesperoyucca whipplei (Torr) Trel., Pinus monophylla Torr. & Frém., Ceanothus cuneatus (Hook.) Nutt., Quercus berberidifolia Liebm., Tauschia parishii J.M. Coult. & Rose) J.F Macbr, Castilleja martinii Abrams, and Corydanthus eremicus (Coville & C. V. Morton) Munz. In the key to Eriogonumin Flora of North America (Reveal 2005), the new wild buck- wheat will key to subg. Eucycla, Key 1-California (p. 225) and will proceed to lead num- ber 12 (p. 226). That may be modified as follows: 12. Perianth glabrous; s Sierra Nevada, Transverse Ranges, and desert ranges to the east from ono County s to Los Angeles and San Bernardino counties 73.E pias kennedyi 12. Perianth pubescent; Last Change Range, Inyo and Mono counties, or Tehachapi coe Kern County. 12a. Achenes pubescent; pedicels glabrous; scapes up to 0.3 dm long; Last Change Range, Inyo and Mono counties 68.E dee shockleyi 12a. oe glabrous; pedicels pilose; scapes 2—3.5 dm long; Tehachapi wen Ker County 92a — Mm Ilictiim The author has agreed not to publish at this time detailed information as to the exact location and distribution of Eriogonum callistum. Eriogonum callistum should make an attractive addition to the rock garden flora. The large, roundish mats are composed of numerous tight rosettes of persistent leaves that remain attractive long after flowering. The fresh leaves, plus those of previous years that remain on the caudex branches, allow the mats to accumulate soil and plant debris and thus enlarge as roundish mounds with each growing season. The numerous, closely arranged caudex branches mean that the plants can be grown as tight, compact clusters along borders or in patterns. The plant’s natural habitat suggests that it requires little REVEAL, A NEW SPECIES OF ERIOGONUM FROM CALIFORNIA 861 watering, prefers gravelly soil in open, sunny places, and should survive with little or no maintenance. ACKNOWLEDGMENTS I wish to thank Hazel Pollard for preparing the line drawing. REFERENCES JENNINGS, C.W. and R.G. STRAND. 1969. Geologic map of California, Los Angeles Sheet. California Division Mines and Geology, Sacramento. Scale: 1:250,000. Reveal, J.L. 2005. 44a. Polygonaceae Jussieu subfam. Eriogonoideae Arnott, Encycl. Brittannica (ed. 7), 5:126. 1832. Wild buckwheat. Fl.N. Amer. 5:218-478. BRIT.ORG/SIDA 22(2) Book REVIEW Guy GusMAN and LILLIAN GusMAN. 2006, The Genus Arisaema, a Monograph for Botanists and Nature Lovers, Second Revised and Enlarged Edition. (ISBN 3-906166-37-6. Publisher: ARG Gantner Verlag Kommaditgellschaft FL 9491 Ruggell. Distributor: Koeltz Sci- entific Books, POB 1360, D-61453 Konigstein/Germany. (Orders: Timber Press Inc, 133S.W Second Avenue, Suite +50, Portland, OR 97204-3527, U S.A; wwwtimber press.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). US$79.95 and C$109.95. +74 pp., 400 +color ee 29 figures, and 18 tables, 6 3/8" x 91/2". ribed since ene First Edition was published, many This Second Edition includes more than ten new s] new photographs, geographical distribution of species by country, d Sevelopaicatid in Arisaema taxonomy, types of acai taxa 4 ie d piles aphical references. ‘The genus Arisaema consists of about 200 species, subspecies and varieties whose extraordinarily wide range extends from Central and East Africa towards Southeast Asia, via Yemen, Oman, Pakistan, Afghanistan, the whole Himalayan range, India, China, Korea, Japan and Siberia and auras to North America. Arisaema is the third largest of the 105 genera that make up the Araceae (Mayo, Bogner & Boyce), 1997), a remarkable fam- ily of monocotyledons that contains some of the most extraordinary ee in the world.” (pg. 12) This, a second, revised and enlarged edition, is a comprehensive book on the genus, written by two collec- tors and growers, who are themselves scientists. Commonly called Cobra-Liles, most do not resemble a snake and the genus is unrelated to lilies. The initial chapters include ees ee use in chemistry and medicine, occurrence in the wild and in cultivation, Beas ation and systematics. Most of the book is taken up in the de- ections, which includes photographs, drawings, tables, etc. Chapters on tables for key species, recent scription of species by each of the doubtful species, invalid names and then hy ae follows. The final three chapters include a ee bibliogra- phy, and an index to species names. The book is a must have for botanists, horticulturalists, plant collectors, ee growers, and anyone interested in the fascinating genus, Arisaema.—David Leedy, Volunteer, Botanical Research Institute of Texas, Fort Worth, TX 76102-4060, U.S.A. SIDA 22(2): 862. 2006 SIX NEW COMBINATIONS IN DODECATHEON (PRIMULACEAE) James L. Reveal! Professor Emeritus, University of Maryland College Park, Maryland 20742-5815, U.S.A. Honorary Curator, New York Botanical Garden Bronx, New York 10458-5026, U.S.A. jreveal@umd.edu ABSTRACT Six new combinations are required in Dodecatheon L. for a forthcoming treatment of the genus in Flora of North America north of Mexico: D. clevelandii Greene var. gracile (Greene) Reveal, D. clevelandii var. insulare (H Thomps.) Reveal, D. clevelandii var. patulum (Kuntze) Reveal, D. iene var. macrocarpum (A. Gray) Reveal D. pulchellum var. shoshonense (A. Nelson) Reveal, and D. utahense (N.H. Holmgren) Reveal. RESUMEN Se necesitan seis combinaciones nuevas en Dodecatheon L. en el tratamiento del género en la Flora of North America north of Mexico: D. clevelandii Greene var. gracile (Greene) Reveal, D. clevelandii var. insulare (HJ. Thomps.) Reveal, D. clevelandii var. patulum (Kuntze) Reveal, D. pulchellum vz (A. Gray) Reveal, D. pulchellum var. shoshonense (A. Nelson) Reveal, y D. utahense (N.H. ares Reveal. The following new combinations are required for a forthcoming treatment of Dodecatheon L.(Primulaceae) in volume 8 of Flora of North America north of Mexico. The rank of vari- ety is used throughout following Holmgren (1994). Dodecatheon clevelandii Greene var. gracile (Greene) Reveal, comb. nov. Dodecatheon See var. gracile Greene, Erythea 3:72. 1895. Type: U.S.A. CALIFORNIA. Santa Clara Co. gar specimen obtained from root collected at Loma Prieta by J.B. Davy (Not found). um f= Dodecatheon clevelandii Greene var. insulare (HJ. Thomps.) Reveal, comb. nov. Dodecatheon clevelandii subsp. insulare HJ. Thomps., Contr. Dudley Herb. 4:134. 1953 (as “insularis”). Type: U.S.A. CALI- FORNIA. Santa Barbara Co.: in chaparral along road to La Cumbre Lookout, 2.3 mi E of San Marcos Pass, Santa Inez Mts., 3000 ft., 25 Mar 1950, HJ. Thompson 1079 (HOLOTYPE: DS; ISOTYPE: OSC). pace clevelandii Greene var. patulum (Kuntze) Reveal, comb. nov. Meadia patula Kuntze, Revis. Gen. PL. 2:398. 1894. Dodecatheon patulum (Kuntze) Greene, Erythea 3:72. 1895. The earlier a name (Pittonia 1:211. 1888) is not valid as it was provisional when published. Type: U.S.A. CALIFOR- Co.: Livermore Valley, 23 Mar 1888, E.L. Greene s.n. (LECTOTYPE: NDG; ISOLECTOTYPE: DS), ae by HJ. Thompson, Contr. Dudley Herb. 4:135. 1953. The variation within Dodecatheon clevelandii, as defined by Thompson (1953), is main- tained with only the rank altered. The epithet var. gracile is equivalent to subsp. sanctarum (Greene) Abrams. Dodecatheon pulchellum (Raf.) Merr. var. macrocarpum (A. Gray) Reveal, oe nov. Do theon meadia L. var. macrocarpum A. Gray, Bot. Calif. 1:467. 1876. a U.S.A. ALASKA. Valdez- ordova Co.: Middleton Island, 2 Jun 1874, WH. Dall s.n. (LECTOTYPE: GH). N E: The ee pe was in- correctly termed a “neotype” by Beamish, Bull. Torrey Bot. Club 82:364. 1955. pee HJ. Thompson (1953) fad considered a “ BISve? specimen ( 27, GI Df rom the Santa Susana Mts., Ventura Co., Califor- considered t1 orm” of his var. macrocarpum, and nia, y annotated the prac eee “macrocarpum, igen atum.” This phase of Dodecatheon pulchellum has long been known as subsp. alas] (Hultén) ‘Mailing address: 18625 Spring Canyon Road, Montrose, Colorado 81401-7906, U.S.A. SIDA 22(2): 863 — 865. 2006 864 BRIT.ORG/SIDA 22(2) Hultén or subsp. superbum (Pennell & Stair) Hultén. As treated by Thompson (1953) and by Taylor and MacBryde (1978), it is the tetraploid, coastal expression that occurs from southern Alaska to Oregon. Dodecatheon pulchellum (Raf.) Merr. var. shoshonense (A. Nelson) Reveal, comb. nov. Dodecatheon pauciflorum (Durand) Greene var. shoshonense A. Nelson, Bot. Gaz. 54:143. 1912 (as shoshonensis). TYPE: U.S.A. IDAHO. Twin Falls Co. Shoshone Falls, 26 Jul 1911, A. Nelson & J. F Macbride 1362 (HOLOTYPE: RM). This variety of Dodecatheon pulchellum is found mainly in alkaline meadows on the foot- hills and valley bottoms of the northern Mojave Desert and Intermountain Region. In the latter area, var. pulchellum is a plant of the mountains. Only in the Pacific Northwest will one find var. pulchellum at lower elevations. The distinctive yellow pollen sacs of var. shoshonense are seen in populations of var. pulchellum in the Black Hills area of north- eastern Wyoming and adjacent South Dakota. Welsh (2003) proposed two entities which are synonyms of var. shoshonense: subvar. atwoodii and subvar. coriaceum. Dodecatheon utahense (N.H. Holmgren) Reveal, stat. nov. Dodecatl tatum Hook. var. utahense N.H. Holmgren, Brittonia 46:91. 1994. Type: U.S.A. UTAH. Salt Lake Co.: Moss Falls, Big Cottonwood Can- yon, Wasatch Range, 13 Jun 1977, N.H.H l g & PK. Holmgren 8412 (HOLOTYPE: NY, ISOTYPES: BRY, UTC). The work by Mastet al. (2001, 2004) demonstrated that varieties of Dodecatheon dentatum Hook., recognized by Holmgren (1994), should be regarded as distinct species. Accord- ingly three widely disjunct entities are now accepted: D. dentatum mainly on the eastern slope of the Cascade Range from south-central British Columbia to central Washington, with disjunct populations along the western portion of the Columbia River, the Colum- bia River Gorge of both Oregon and Washington, northeastern Oregon (Wallowa Co.), and in northern Idaho; D. ellisiae Standley of north central New Mexico in the moun- tains of Bernalillo, Sandoval and Torrance cos., and in southeastern Arizona in southern Apache, Greenlee, southern Graham, and northeastern Santa Rita cos.; and D. utahense,a narrow endemic confined to Big Cottonwood Canyon, Salt Lake Co., Utah. The recent molecular work on Primuldceae by Mast et al. (2001, 2004) and others (Kallersjé et al. 2000; Martins et al. 2003; Trift et al. 2002) has shown that recognition of Dodecatheon creates a paraphyletic Primula. Accordingly, Mast and Reveal (2006) trans- ferred all of the entities assigned to Dodecatheon to Primula. ACKNOWLEDGMENTS lam grateful to the Flora of North America Committee for providing funding to study Dodecatheon in the field and herbarium. To the several curators who made available their collections for study lam in your debt. In particular | must mention Kenton L. Chambers and Harry L. Thompson, both well acquainted with the genus, who willingly shared their decades of experiences. As always, the assistance of Kanchi Gandhi is indispensable. The Holmgrens, both Noel and Pat, provided me with information both from the literature and their experience with the genus. Both Kanchi and Noel formally reviewed the manu- script and lam grateful for their comments and timely corrections. Finally, my wife, C. Rose Broome, managed to survive the three months of getting Dodecatheon done; her help in herbaria and in the field is acknowledged with thanks. To the FNA editors and several reviewers who examined the “first draft” and provided comments and corrections, I give my thanks as we REVEAL, NEW COMBINATIONS IN DODECATHEON 865 REFERENCES Beamish, K.. 1955. Studies in the genus Dodecatheon of northwestern America. Bull. Torrey Bot. Club 82:357-366. HitcHcock, C.L. 1959. Dodecatheon L. Shooting star. Univ.Wash. Publ. Biol. 17(4):40-46. Hoimaren, N.H. 1994. Redefinition of Dodecatheon dentatum (Primulaceae) and rationale for use of varietial rank. Brittonia 46:87-94. KALLERSJO, M., G. Bercavist, and A.A. Anbersers. 2000. Generic realignment in primuloid families of the Ericales s.l: A phylogenetic analysis based on DNA sequences from three chloroplast genes and morphology. Amer. J. Bot.87:1325-1341 Martins, L., C. OBeRPRIELER, aNd FH. Hetiwic. 2003. A phylogenetic analysis of Primulaceae s.l. based on internal transcribed spacer (ITS) DNA sequence data. Pl. Syst. Evol. 237:75-85. Mast, A.R., S. Ketso, A.J. RicHaros, D.J. LANG, D.M.S. Fetter, and E. Conti. 2001. Phylogenetic relationships in Primula L.and related genera (Primulaceae) based on noncoding chloroplast DNA. Int. J. PI. Sci. 162:1381-1400. Mast, A.R., D.M.S. Fever, S. KeLso, and E. Conti. 2004. Buzz-pollinated Dodecath igi d from within the heterostylous Primula subgenus Auriculastrum (Primulaceae): A 7-region BcoENA phylogeny and its implications for floral evolution. Amer. J. Bot. 91:926-942. Mast, A.R. and J.L. Reveat. 2006. Transfer of Dodecatheon to Primula (Primulaceae). Brittonia, in press. THompson, H.J. 1953. The biosystematics of Dodecatheon. Contr. Dudley Herb. 4:73-154. Taytor, R.L. and B. MacBryoe. 1978. New taxa and nomenclatural changes with respect to Vascular plants of British Columbia:a descriptive resource inventory. Canad. J. Bot.56:184-195. Tritt, |, M. KALLERSJO, and A.A. AnperBerG. 2002. The monophyly of Primula (Primulaceae) evaluated by analysis of sequences from the chloroplast gene rbcL. Syst. Bot. 27:396-407. WELSH, S.L. 2003. Dodecatheon L. A Utah flora, ed. 3:526-527. 866 BRIT.ORG/SIDA 22(2) Book REVIEW SUSANNA Ly_r. 2006. Fruit and Nuts. (ISBN 0-88192-759-7, hbk.). Timber Press Inc, 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, US.A.(Orders: wwwtimberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax.). $59.95, 480 pp., color photos, figures, glossary, bibliography and index, 81/4" x 111/4". Have you ever looked at strange food in the grocery produce aisle and wondered which part you can eat? Or what preparation is needed to eat it? The new book Fruit and Nuts by Susanna Lyle will be a handy book to reference. Over two hundred food plants are presented in Fruit and Nuts along with details on floral structures, cultivation and uses. To help visualize the ae the book also includes over three hundred fantastic color photographs and ee Fruit and Nuts opens with a very informative section that introduces the readers to the various aspects of fruit and nuts and their production. Plant classification and re aan are covered in such a way that any reader can understand, and more detail can be found for bolded words in the glossary of the book. lyle gives readers botanical definitions of fruit and nut types with Sane examples and accented by color illustrations de- picting nut and fruit structures. Additional information in the introductory s by aa region, cultivation, propagation and the health benefits of nuts and fruits. e majority of the text is the fruit and nut “encyclopedia” in eer order by specific epithet from Acer sacc ae um (sugar my) to nics Pam as aebet should be noted that a few non-fruit nut items are included I t views of plant types 1 fruit or nut entry contains a crisp color image of e edible portion of the plant along with descri oe structure, harvesting methods, cultivation, prop apation a other relative species. A side box for each ry lists uses such as food, nutritional value, alternative uses and ornamental value. The descriptions are very thorough and easy to read. An appendix covers plants for differing growth conditions/uses such as ornamental and health benefits. There is also a glossary, bibliography by topic and an index. Fruit and Nuts by Susanna Lyle is a fun and informative book with great value to those in food sciences, horticulture, gardening food plants or anyone with a general interest in learning more about the fruits and nuts consumed around the world. The text is thorough, easy to read, and is well illustrated with both color illustra- tions and color photographs. These illustrations and photographs are fantastic in helping readers to determine the edible portions of many Beale unknown fruits and nuts. This could be a wonderful reference book or gilt for cooks, horticultural students or “foodies” in general.—Lee Luckeydoo, Herbarium, Botanical Research In- stitute of Texas, 509 Pecan on Fort Worth, TX 76102-4060, U.S.A. SIDA 22(2): 866. 2006 ERICAMERIA NAUSEOSA SUBSP. AMMOPHILA (ASTERACEAB), A NEW RABBITBRUSH FROM THE SAN LUIS VALLEY OF COLORADO Loran C. Anderson Professor Emeritus, ee ca ological Science tate Univer ihiestes FI fen pene U.S.A. ABSTRACT A previously undescribed, distinctive subspecies of rabbitbrush has been recently discovered in south-central Colotads ane it is ene dominant plant on eandenect = is e eequent on low sand aad It is = Bete natied ammophila. A r and a key is provided. RESUMEN Se ha descubierto recientemente una nueva subespecie de chamiso en el centro-sur de Colorado dinde es la planta commtanite en el arene) y eae en oe bajas. Se ha nombrado como Ered merida nauseosd subsp. ammophila p i y se aporta una Slave: One of the most widespread species in western North America is the rabbitbrush Ericameria nauseosa (Pallas ex Pursh) Nesom & Baird. It ranges from southern Canada south to northern Mexico and from the Dakotas, Kansas, and Texas west to Washington and California. This rabbitbrush was long known as Chrysothamnus nduseosus (Pallas ex Pursh) Britton. H.M. Hall (with F E. Clements) monographed Chrysothamnus in 1923 and Haplopappus in 1928, and his views held for many years. However, when the North American species of Haplopappus were demonstrated not to be closely related to the South American species (which included the type species for the genus), several sections of Hall's North American Haplopappus were recognized as separate genera. One such genus is Ericameria. Some species of Chrysothamnus, such as C. nauseosus, have long been noted for their close similarity to Ericameria. When DNA data suggested that the traditional Chrysothamnus was not monophyletic, Nesom and Baird (1993) transferred four of its species, including C. nauseosus, to Ericameria. Anderson (1986a) recognized 22 subspecies of C. nauseosus, whereas Nesom and Baird (1993) recognized two subspecies of E. nauseosa (ie., subsp. nauseosa and subsp. consimilis) and several varieties. Now, a new infraspecific taxon of E. nauseosa is to be described. Should it be a variety or a subspecies? Nesom and Baird (1993) emphasized the category of variety, but I believe my rationale for use of subspecies (Anderson 1986b) is solid, and I therefore describe the new subspecies, Ericameria nauseosa subsp. ammophila, below. Variability in the species is great, and my extensive study of the group in the field and in the herbarium has convinced me to apply infraspecific names to only the more signifi- cant morphotypes. It is surprising that this new subspecies has escaped recognition for so long. Specimens of it were not represented in the more than 12,000 collections of C. nauseosus that I studied from 56 herbaria (including COLO, CS, NCM, RM, and UNM) about 30 years ago. SIDA 22(2): 867 — 872. 2006 868 BRIT.ORG/SIDA 22(2) intend to make appropriate subspecific combinations in Ericameria nauseosa ina future paper so that variation within the wider ranging subspecies can then be recog- nized through quadrinomials. Some argue that quadrinomials are cumbersome, but at an applied level, some field workers may wonder why the populations in their regions have the same subspecific name that is applied to somewhat different-looking popula- tions in other regions, and quadrinomials would help them see the relationships. There- fore, in the discussion section of the present paper, I will use subspecies names as they occurred in Chrysothamnus nauseosus (Anderson 1986a) for comparative purposes. ve nauseosa (Pallas ex Pursh) Nesom & Baird subsp. cual ue LG. een: nov. (Figs. 1, 2). Type: UNITED STATES. COLORADO. Saguache Co: dsheet ae a National Wildlife Refuge, ca. 4.5 air mi SW of Crestone, elev. 2528 m (7600 ft), Lat. 37° 56 12" N, Long. 105° 44' 29" W, 2 Sep 2005, L.C. Anderson 21303 (HOLOTYPE: BRIT, IsoTyPEs: COLO, FSU, NY). Frutices usque ad 14 dm alti 2 I foliis {1 ibus; folia linearia, 20-45 mi longa, 0.6-1.2 mm f] iculati itula cylindrica, 9.3-14 mm alta, straminea bracteis exterioribus ovatis, comentalosis vel glabratis, neous oblongis, glabris, apicibus obtusis vel acuminatis; ura geas 5, pallide flavi, corollibus 10.5-13 mm pone, lobis pecuaus unuequen lam longis, 1] ] & I ‘= aliquot y ion cterieis plerimane longioribus; achaenia aa. Fastigiate shrubs, 3.5-7(-14) dm tall, usually broader than tall from considerable branch- ing; ultimate flowering branches (8-)10-20(-25) cm tall, shallowly grooved, yellowish- green with compact tomentum, somewhat viscidulous. Leaves alternate, entire, linear, stiffly ascending or slightly spreading, (20-)25-30(-45) mm long, only slightly shorter toward stem tips, (0.6-)0.9-1.0(-1.2) mm wide, subterete with adaxial groove, light green, pubescence of sparse, compact, tomentum and a few longer, villous hairs. Inflorescences tightly congested paniculate cymes; heads cylindrical, (9.3-)11-12.5(-14) mm tall, (2-)2.5(- 3)mm wide, stramineous; receptacle with central cusp (0.7-)1-2(-2.5) mm long. Phyllar- ies (15-)16-18(-20), in vertical rows, outermost (lowest) bracts ovate, 1.2-2 mm long, tomentulose to glabrate, inner bracts oblong, 8-10 mm long, glabrous with + ciliate mar- gins (few cilia to 0.4 mm long), weakly keeled, tips (rarely with tufted pappilae 0.1mm long) obtuse to acuminate. Disk flowers 5; corollas (10.5-)11.5-12.5(-13) mm long, nar- rowly cylindric gradually flaring to erect lobes (0.6-)0.7-LO(-L.3) mm long, yellow, sparse hairs on tube generally 0.2 mm long but a few 1-2(-3.5) mm long; styles (13-)14-16(-17.5) mm long, style branches 3.9-4.7 mm long, the stigmatic lines more or less equal to branch tips in length (40-56%), anthers 4 mm long with appendage 0.8-0.9 mm long; mature achenes cylindric, 5-nerved (the nerves when noticeable because of brownish material in associated secretory canals), +.5-6(-6.8) mm long, sericious with hairs (0.4-)0.5-0.7(- 1.0) mm long, pappus (9-)10-11(-12.5) mm long, finely barbellate, white. Phenology.—A few shrubs of the new subspecies start to bloom in late August; most are in full bloom in mid-late September, whereas E. nauseosa subsp. consimilis, or var. oreophila (A. Nelson) G.L. Nesom & G.L. Baird, found in nearby more mesic, saline sites, blooms earlier in August. Habitat.—The new rabbitbrush occurs (in descending frequency) on: sandsheet, dune fields, stabilized sand dunes, and sandy soil of open pinyon-juniper (old dunes?). The term sandsheet is variously defined by geologists; here it refers to flat expanses of stabilized coarse, poorly sorted sands that occur extensively along the margins of the high dunes of the Great Sand Dunes National Park (tallest in North America). The rabbitbrush is the dominant plant (hiding nearly equally frequent grasses) on the sandsheet, and sandsheet LC / L oO I 1 ANDERSON, ERICAMERIA NAUSEOSA SUBSP. AMMOPHILA, A NEW SUBSPECIES FROM COLORADO Fic. 1. Close-up of of flowering branches of Ericameria nausosa subsp.ammophila showing characteristic yellowish-green stems, nearly glabrous involucres, and short corolla lobes. 870 BRIT.ORG/SIDA 22(2) rabbitbrush is very appropriate as a common name for this new subspecies (whose epi- thet ammophila means ‘sand loving’). From a distance, the vegetation on much of the Baca National Wildlife Refuge appears as a more or less evenly spaced, ‘unbroken sea’ of sandsheet rabbitbrush. Dune fields are generally flat but have some undulating topogra- phy. The sandsheet rabbitbrush occurs at elevations from 2518 to 2735 m (7550 to 8200 ft). Associated species.—The grasses, Achnatherum (Oryzopsis ) hymenoides (Roem. & Schult.) Barkworth and Chondrosum (Bouteloua) gracile H.B.K.,are usually co-dominants with the sandsheet rabbitbrush. The following are found at one or more sites: Chrysothamnus greenei (Gray) Greene, Eriogonum cernuum Nuttall, Heterostipa (Stipa) comata (Trin. & Rupr.) Barkworth, Ipomopsis longiflora Torr.) V. Grant, Nuttallia (Mentzelia) rusbyi (Wooton) Rydb., Opuntia polyacantha Haworth, Psoralidium lanceolatum (Pursh) Rydb., Rhus trilobata Nuttall, Senecio multilobatus Torr. & Gray, Sporobolus cryptandrus (Torr.) Gray,and Yucca glauca Nuttall. The flora of the sandsheet- sandhill ecosystem of this region is sparse in numbers of taxa. Additional collections: COLORADO. Alamosa Co.: Great Sand Dunes National Park Visitor's Center, 2 Sep 2005, . Anderson 21,280 (FSU); ie sheet by road to Medano Ranch, 9.9 mi E of Hwy 17, 2 Sep 2005, L.C. Anderson 21,291 (FSU). Saguache Co.: ca. 3.7 mi SSE of Crestone, 21 Aug : 2001, J. Erdman O13 (FSU), 6 Sep 2001, J. Erdman O116 (FSU), 27 Sep 2001, J. Erdman 0121 (FSU); 1 Sep 2005, L.C. Anderson 21,276 (FSU); rd to San Isabel Cz ae trailhead, 21 Aug 2001, J. Erdman O1l4 (FSU); 7 Sep 2001 J. oud man O119 (FSU); 28 Sep 2001, J. Erdman 0123 (FS near Pinyon Flats, Great Sand Dunes National Park, 2 Sep 2 L.C. Anderson 21,286 (FSU); dune field near “Ox- bow Pond” between Spanish Creek and Cottonwood aie - 22 Aug 2001, J. Erdman O115 (FSU); 7 Sep 2005, J. Erdman O18 (FSU); 27 Sep 2001, J. Erdman 0122 (FSU), 2 Sep 2005, L.C. Anderson 21,301 (FSU). DISCUSSION The sandsheet rabbitbrush (subsp. ammophila) is perhaps most closely related to subsp. nitidus L.C. Anderson which occurs in northern Arizona and adjacent New Mexico. The two share the features of yellow-green foliage and stramineous, shiny involucres. They dif- fer in geography and habitat (subsp. nitidus prefers sandy gravels of dry stream beds) and in that subsp. ammophila fomrs shorter shrubs usually 35-70 cm tall (60-150 cm in subsp. nitidus), and has flowering stems fastigiate and more crowded than in subsp. nitidus, leaves usually 25-30 mm long and! mm or less wide (30-50 by 1-L5 mm in subsp. nitidus), corol- las mostly 11.5-12.5 mm long (9.5-11 mm in nitidus), unusually long trichomes on the corolla tubes and pubescent achenes (usually glabrous in nitidus). Subspecies turbinatus M_E. Jones is like the sandsheet rabbitbrush in height and branching, but it occurs gener- ally in sandy regions of the Great Basin and has mostly longer involucres with tomentulose phyllaries that are acute to acuminate (phyllaries in subsp. ammophila are mostly glabrous with obtuse to acute tips), and shorter corollas with lobes that are vil- lous. Subspecies consimilis (e., var. eae of Nesom & Baird 1993) rarely grows with subsp. ammophila, but subsp. consimilis usually grows in more saline or disturbed (mesic) habitats in this region, and it differs in forming generally taller, narrower shrubs with darker green foliage and smaller involucres and flowers. The unusually long hairs (up to m) on the corolla tubes of the sandsheet rabbitbrush are 10 times longer than any others found in the species or in all of the former genus Chrysothamnus (Anderson 1970). A portion of the Urbatsch et al. (2006) key to the varieties (listed here as subspecies) of Ericameria nauseosa is given below with the new subspecies included; subsp. ammophila shares a couplet with subsp. bernardinus, but the two are not closely related (ecologically, geographically, or taxonomically within the species). au AMARAQDUII A ANDERSON Fig.2.5 Ad aun 872 BRIT.ORG/SIDA 22(2) 1. Cypselae a (plants 60-150 cm; involucres 10-12.5 mm) subsp. nitidus (in part) 1. Cypselae hai 16. Corolla ices villous (sometimes sparsely). 19. Leaf blades 30-50 mm, faces glabrate subsp. nitidus (in part) 19. Leaf blades 10-20 oe tomentulose subsp. turbinatus 16. Corollas lobes glabro 23. Phyllaries a epbeus outer sometimes sparsely hairy. 24. Corollas 1 2[13] mm;involucres [9.3]10-14 mm. (24a. an lobes 1.7-2.3 mm subsp. bernardinus (in part) [24a. Corolla lobes 1 mm or less] subsp. ammophila 24. Corollas 6—9+ mm; involucres 6-10 mm ___ subsp. consimilis [i.e., var. oreophila] ACKNOWLEDGMENTS James Erdman provided camaraderie, accommodations, transportation, and valuable general knowledge of the flora and geology of the area and obtained access for collecting in restricted areas. Richard Sparks of the USDA and Fred Bunch and Phyllis Pineda Bovin of Great Sand Dunes National Park also gave excellent assistance. Mark Garland provided the Latin diagnosis; William A. Weber and an anonymous reviewer offered important suggestions for improving the manuscript. Funds from the Friends of the Robert K. Godfrey Herbarium (FSU) helped defray publication costs of this paper. REFERENCES ANDERSON, L.C. 1970. Floral anatomy of Chrysothamnus (Astereae, Compositae). Sida 3:466-503. ANDERSON, L.C. 1986a. An overview of the genus Chrysothamnus (Astereae), In: E.D.McArthur and B.L. Welch. Proc. Symp. Biology of Artemisia and Chrysothamnus. USDA, Forest Service, Intermountain Research Station, Ogden, Utah. Pp. 29-45. ANDERSON, L.C. 1986b. Sympatric subspecies in Chrysothamnus nauseosus. In: E.D. McArthur and BLL. Welch. Proc. Symp. Biology of Artemisia and Chrysothamnus. USDA, Forest Service, Intermountain Research Station, Ogden, Utah. Pp. 98-103. Hatt, H.M. 1928. The genus Haplopappus; a phylogenetic study in the Compositae. Carnegie Inst. Wash. Publ. 389, Washington, D.C Hait, H.M.and F.E. Clements. 1923. The phylogenetic method in taxonomy. The North American spe- cies of Artemisia, Chrysothamnus, and Atriplex. Carnegie Inst. Wash. Publ. 326, Washington, D.C Nesom, G.L. and G.I. Baird. 1993. Completion of Ericameria (Asteraceae: Astereae), diminution of Chrysothamnus. Phytologia 75:74—93. Urearscu, L.E., L.C. ANDERSON, R.P. Roberts, and K.M.Neusic. 2006. The genus Ericameria Nuttall (Asteraceae). Fl. North Amer. 26:50-77. A NEW SPECIES OF BOLTONIA (ASTERACEAE) FROM THE RIDGE AND VALLEY PHYSIOGRAPHIC PROVINCE, U.S.A. John F. Townsend Vesna Karaman-Castro oe of Conservation and Recreatio Louisiana State University Division of Natural Heritage Department of Biological Sciences a7 ee St, 3° Floor 107 Life Science Building Richmond, Virginia 23219, U.S.A. Baton Rouge, Louisiana 70803, U.S.A. Email: John. Townsend@dcr.virginia.gov ABSTRACT Boltonia montana, a new f 1 i ine habi in the Appalachian Mountains of as New elie and Pennsylvania is described. This taxon has most often been es under the name onia roides (L.) LHer. due to its relatively large heads which are usually few in number, and vegetative ] similarities. C of the new taxon, however, is dissimilar to that of B. oe and most closely resembles Boltonia ca relent na (Walt. ) Fern.,a species characteristic of coastal plain and Oia bottomlands of the Carolinas and Virginia. Boltonia montana differs from the latter taxon in terms of inflorescence structu head dimensions, ray color, phyllary morphology and size, plant stature, and leaf shape. Evidence of its eae tiveness is provided and its wetland habitat is discussed. RESUMEN omer | slA5 Ral . ¢ eee | “ J mak eS ci naae | ae i Vee Nee pas ui Pennsylvania ES taxon se ea eolecaas la mayor ce las veces con el usualmente en un numero pequeno, y hae vegetativas. La ene de las cipselas del nuevo Bon, sin embargo, es diferente de la de B. asteroides y se parecen mucho mas a Boltonia caroliniana (Walt.) Fern., una especie caracteristica de la llanura costera y base del pie de monte de las Carolinas y Virginia. Boltonia montana difiere e este Ultimo taxon en términos de estructura de la inflorescencia, dimensiones de la cabezuela, color de los radios, morfologia y tamano de los filarios, tamano de la planta, y forma de la hoja. Se aportan pruebas de su diferenciacion y se discute su habitat de zonas himedas INTRODUCTION The first author was alerted to the presence of a taxonomically problematic Boltonia in Augusta County, Virginia (Ridge and Valley physiographic province) by Gary P. Fleming, J. Christopher Ludwig (Virginia Department of Conservation and Recreation, Division of Natural Heritage), and Thomas F Wieboldt (Massey Herbarium, Virginia Polytechnic Institute) in late 2001. This plant had been identified as Boltonia asteroides by collectors, but certain details of the plants, particularly those of cypselae and phyllaries, indicated the Augusta County plants were not a good fit for that taxon. Subsequent investigations of the taxonomic literature, herbarium material, and living populations of Boltonia led to the realization that the plants from sinkhole ponds (dolines) in the Virginia mountains possessed a unique set of characters. During morphological and molecular anesrieaHens: of the nee plants, correspondence with - eos author led to the discovery of morphologically similar herbarium specimen dfrom the Ridge and Valley physiographic province of northwestern Nor iecee: and eastern Penn- sylvania. Field studies in the early 1980s and late 1990s by David Snyder and Kathleen Strakosch Walz of the New Jersey Natural Heritage Program uncovered or reinventoried the Boltonia populations in the northwestern part of that state and provided much data SIDA 22(2): 873 — 886. 2006 874 BRIT.ORG/SIDA 22(2) on the plants discussed here. Subsequent herbarium and field investigations of the Vir- ginia, New Jersey, and Pennsylvania plants confirmed that these plants belonged to the same taxon. It is described here as new. Boltonia montana J.- Townsend & V. Karaman-Castro, sp. nov. (Figs. 1, 2). Typr: U.S.A. New Jersey. Sussex Co.: Muckshaw pond complex, located 3.2 km SW of Newton sual 14 km W of US 206, nearly dry, herb-dominated sinkhole pond, 3 Oct 2005, J.k Townsend 3570 (HOLOTYPE: US; IsoTYPES: MO, NY, VPD. Boltonia asteroides (L.) VHer. a oo maxime simile ob capitulis magnus et paucus sed acheniis glabris, 7 ] R alis et pap] (Walt. Fern. similis. Phyllariis costa angustata, herbaceous et distalis viridis. Glabrous, perennial herbs 1.2-15 dm tall, ascending to erect (often prostrate in lower por- tion to ca. 1/2 stem length, sometimes rooting at nodes in this section). Stems glabrous, light to yellowish-green, ribbed with yellow or golden striations (more prominent in up- per 1/2 of plants), branching above into small, simple, corymbose panicles to diffusely paniculate in larger plants. Leaves basal and cauline, glabrous; basal leaves 0-18, alter- nate, in a rosette, oblanceolate to oblanceolate-spatulate, attenuated at the base (where often slightly widened), dark green or blue-green, often whitish or reddish colored at the base, with prominent, lighter-colored midrib and inconspicuous, arcuate secondary veins, apices bluntly rounded, mucronulate to entire, 1-17.5 cm long, 0.1-2 cm wide; cauline leaves oblanceolate to obovate or spatulate, bases sessile, often partially clasping, apices rounded to acute, mostly with mucronulate tips, appearing entire but minutely serrulate with narrow, wnlash, aoe margin, 2.7-11 cm long, 0.5-1.9 cm wide, sometimes with smaller leaves clustered at nodes. Capitulescences oes paniculate, few-headed (1-25) in average at many-headed (-100+) in large individuals. Peduncles bracte- ate, 0.5-11.8 cm long, 0.5-L1 mm thick, bracteal leaves oblanceolate to obovate or oblong, tips obtuse to acute, mucronulate, 0-5 per pedicel, 0.1-9.7 cm long, 0.02-1.4 cm wide. In- volucre 5.2-7.8 mm tall, 6.0-11.3 mm wide; phyllaries 30-00, lance-attenuate, serrulate to lacerate, in 2-4 series, 2.5-4.4 mm long, 0.4-0.9 mm wide, midrib 0.12-0.3 mm wide (1/10- 1/5 of phyllary width), proximally whitish, distally green; receptacles conic, naked, L.9- 3.0 mm tall, 1.9-3.8 mm wide. Ray flowers 27-406, pale lavender to pinkish (-white), corol- las glabrous, 4-5 nerved with golden or amber colored veins, 8.1-12 mm long, 0.8-1.9 mm wide, tubes 0.45-0.95 mm long; styles 1.3-2.45 mm long, style branches 0.25-0.4 mm long, cypselae glabrous, narrowly oblong, triquetrous, with | adaxial and 2 abaxial ribs, often slightly curved outward, 1.25-2.05 mm long, 0.5-0.9 mm wide, wings vestigial or absent, 0-0.05 mm wide; pappus awns reduced to a crown of minute scales, 0-0.1(-0.4) mm long, Disk flowers 220-350, pale yellowish green to pale yellow-white as cypselae mature, su- ture lines pale amber, disk appearing more yellowish during anthesis, corollas glabrous, L.1-2.3 mm long, tubes 1.0-1.85 mm long, lobes 0.15-0.36 mm long; stamens included to exceeding corolla length by 0.4 mm, anthers 0.65-1.1 mm long, appendages 0.15 mm long, styles 1.05-2.55 mm long, style branches 0.2-0.4 mm long, cypselae glabrous, narrowly oblong, strongly flattened laterally with two marginal ribs, 1.3-1.9 mm long, 0.55-1.25 mm wide, wings vestigial, O-0.09 mm wide, pappus awns usually reduced to a crown of minute scales 0-0.1 (-0.6) mm long. Distribution.—Distributed in sinkhole pond habitats and associated riversides and streamsides of the Ridge and Valley physiographic province in western Virginia, north- western New Jersey, and eastern Pennsylvania. TOWNSEND AND KARAMAN-CASTRO, A NEW SPECIES OF BOLTONIA FROM THE EASTERN U.S.A zk ‘ es Wh av I , } terial collected in $ Count y, New Jersey. 1 C T. IOouUW. factrn Wl ra Pe L J hyll y ( t ).6 Ray corolla. 7. Disk corolla. Scale pareatale iif Fic. Habit. 2. Ray cypselae. 3. Disk cypselae. 4. Head. 5. Out except in 1 (= 1 dm) and 4 (=1 cm). 876 BRIT.ORG/SIDA 22(2) Fic. 2. Low ing individual of Boltoni ith Eriocaul ticum, Augusta County, Virginia. Photo by Gary P. Fleming. I J 1 Etymology.—The name refers to the species’ montane habitats. Phenology and development.—Flowering primarily occurs from early August to mid- October with precocious plants flowering in July, especially during dry years or in dr microhabitats. Hydrologic fluctuations in the species’ sinkhole pond habitat can ey a dramatic effect on the timing of flowering in a population. Even when water levels re- main high enough to inundate these ponds during the growing season, seeds are capabl of germination and rosettes may develop while still under water (Lassen and nee 2003; David Snyder pers. comm. 2006) but the plants may not develop further, and if inundated for successive years the plants may die (David Snyder, pers comm. 2006). Au- tumnal plants produce basal offsets nearly identical in appearance to the rosettes pro- duced by young plants. In the case of reclining or ascending plants of Boltonia montana, adventitious roots may form at the nodes as autumn approaches. In many cases, these nodes also produce leafy offsets complete with root systems. During collection of flower- ing or fruiting plants, these vegetative propagules are often dislodged due to their weak attachment to the parent plant. A concentric gradient of environmental conditions is created in these sinkholes by fluctuating hydrology and sloping shorelines. It is therefore possible to find several co- occurring growth stages of Boltonia montana. Recent field work in New Jersey with Natu- ral Heritage Program Botanist David Snyder and Ecologist Kathleen Strakosch Walz (au- tumn 2005) was conducted following a long period of pond inundation. On the pond margins that were not recently inundated, relatively well-developed plants (with flow- ers, fruits, and basal offsets) were found. In more recently inundated portions of the ponds, large numbers of Boltonia rosettes were found. These plants were either newly germi- 7 TOWNSEND AND KARAMAN-CASTRO, A NEW SPECIES OF BOLTONIA FROM THE EASTERN U.S.A. 877 nated seedlings, rosettes persisting from a previous germination event, or vegetative off- sets now independent of the parent plant. Regardless of origin, these rosettes were unable to flower or proeuce fruit before the end of the growing season due to their ey ceccpuonally late Septem ber —October). In general, when f as ground- water levels drop after such periods of inundation, new ae develop fro rom the seed bank (David Snyder, pers. comm. 2006). Observations indicate that Smith and Mettler’s (2002) description of the life cycle of the Federally Endangered Boltonia decurrens (Torr. & A. Gray) Wood provides a frame- work for understanding the life cycle of Boltonia montana as well: “Although B. decurrens is a perennial under optimal conditions, various life stages may behave as winter annu- als (winter seedlings bolt, flower, and senesce the following summer), as summer annu- als (summer seedlings bolt, flower, and senesce the same summer), or as biennials (ca. 25 percent of spring seedlings overwinter as rosettes that bolt, flower, and senesce the fol- lowing summer).” Such a patchwork of potential strategies makes sense for a species adapted to the highly variable flood pulse of Midwestern rivers (B. decurrens) as well as the dramatic hydrological fluctuations of isolated wetland systems (B. montana). HABITAT AND DISTRIBUTION Boltonia montana is known from localized sections of the Ridge and Valley physiographic province in three U.S. states: Virginia (one county), New Jersey (two counties), and Penn- sylvania (one county). All occurrences are found within areas of gently sloping, rolling, or nearly level ground between 149 and 488 m (490-1600 ft) altitude (Walz et al. 2001, Va. Division of Natural Heritage files), but many of the New Jersey ponds are surrounded by rough terrain with extensive bedrock outcrops (Kathleen S. Walz, pers. comm. 2006). The sinkhole pond complexes which support this species span an area measuring roughly 14x 15 km near Newton, New Jersey and 5.25 x 4.5 km near Waynesboro, Vir- ginia. Within these areas, plants are known from a total of 36 sinkhole ponds and lakes (22 in New Jersey and 14 in Virginia; Va. Division of Natural Heritage files; David Snyder, pers comm. 2006). Although the vast majority of the records for this new taxon come from isolated wetlands, a very few have been found along rivers. Besides two collections made the same day in 1936 along the South River in Virginia, the only other such collec- tions were made in 1864 (1 specimen) and 1865 (1 specimen) along the Susquehanna River in Dauphin County, Pennsylvania—apparently the only observations of this species in Pennsylvania. The Virginia occurrence was taken from an area extremely close to, and just downslope from, a sinkhole pond complex known to contain Boltonia montana, while the disposition of the Pennsylvania material relative to potential sinkhole pond seed sources is unclear. It is assumed here that only occasional plants become established in such riverine habitats from nearby “parent” populations. The lack of additional collec- tions at least indicates that such riverine habitats are rarely occupied. Like Boltonia montana, many species seemingly defined by their isolated wetland habitat are occasionally found in other community types, indicating that their affinity for these ponds is not restrictive. Examples include the co-occurring Helenium virginicum Blake in Virginia, which is almost entirely restricted to sinkhole ponds but also occurs in a wet meadow of the nearby South River (Va. Division of Natural Heritage files). The diminutive sedge Fimbristylis ee eae ex Small and Britt.was also thought to bea strict pond endemic and func hin most localities, but discoveries in North and South Carolina (Leonard 1981. 1987), established the species as an occupant of river- 878 BRIT.ORG/SIDA 22(2) ine “draw-down” bars, banks, and sloughs as well. A final example involves Boltonia decurrens, a perennial species of floodplains in the Ilinois River valley (Stoecker et al. 1995) which was originally dependent on natural flood pulses for habitat maintenance and germination. Most of the habitats occupied by this species have been described as wet prairies or bottomland communities of various types, but McClain et al. (1997) also reported the species from “sand ponds” near the Illinois River in Cass and Mason coun- ties, Illinois. They described water levels in the ponds as fluctuating in response to the local aquifer rather than overbank flooding, but it is not known to what degree human manipulation of the river has lessened the importance of this water source in the ponds. The sinkhole pond habitats occupied by Boltonia montana are ultimately formed by the solution of underlying carbonate bedrock and subsequent volume loss or collapse of this foundation (Fleming @ Van Alstine 1999; Walz et al. 2001). The habitat types found within sinkhole ponds in Virginia and New Jersey are floristically and physically differ- ent, owing to their means of origin, Pleistocene history (glacial scouring and surficial geologic deposits), and distance of calcareous materials from the soil surface. In Virginia, weathering of limestones and dolomites of the Cambrian Shady and Elbrook Formations is ultimately responsible for the formation of these pond features, but a deep overburden of relatively acidic, infertile soils covers these carbonate rocks toa depth of 30 to 150m (Fleming & Van Alstine 1999). This phenomenon is similar to that described in Maryland and West Virginia by Bartgis (1992) who studied ponds support- ing the rare pond endemic Scirpus ancistrochaetus Schuyler. In Virginia, surficial depos- its are the result of alluvial fan formation along the western slopes of the Blue Ridge mountains. Materials range from quartzite cobbles and boulders to sand, silt, and clay and range from the late Pleistocene to Pliocene or Miocene in age (Whittacar et al. 1999). Due to the depth where the carbonate rocks reside, soil and water chemistry of the ponds is acidic (pH 3.9-4.0 at 0-30 cm). Soils are predominantly gray to blackish clays or clay loams with lighter colored mottling evident below 3 to 10 cm. This clay, weathered from metasiltstone and phyllite alluvium, impedes drainage but most ponds draw down as the growing season progresses and evapotranspiration increases. Although some of these sinkholes are relatively stable hydrologically, Boltonia montana is almost exclusively found in those ponds with highly variable and unpredictable water levels. The Boltonia montana “zone” in these ponds is typically doughnut-shaped and extends from the rim of the pond to the edge of the most deeply flooded portion of the basin (Fig. 3). The y types occupied by the species in Virginia are the Quercus palustris / Panicum rigidulum - Panicum verrucosum ~ Eleocharis acicularis Wooded Herbaceous Vegetation Association and the Cephalanthus occidentalis / Polygonum hyd ropiperoides - Glyceria acutiflora - Proserpinaca palustris Shrubland Association (Grossman et al. 1998; NatureServe 2005). The former community type is endemic to Virginia and assigned a rarity rank of G1G2 (NatureServe 2005). The latter type is somewhat more widespread (ranked G3?/S1) but is considered endemic to mid-Atlantic region. These communities are open-canopied except on pond edges, where the forest canopy can provide partial shading. Associated species in these communities include Quercus palustris, Nyssa sylvatica, and Cephalanthus occidentalis (mostly occurring on pond margins), Panicum rigidulum var. rigidulum, P. philadelphicum, P. verrucosum, Eleocharis acicularis, Eleocharis melanocarpa, Eriocaulon aquaticum, Lysimachia hybrida, Ludwigia palustris, Leersiaoryzoides, Hypericum boreale,and the federally Threatened Helenium virginicum. In addition to relatively undisturbed sinkhole ponds, Boltonia montana is also found with — TOWNSEND AND KARAMAN-CASTRO, A NEW SPECIES OF BOLTONIA FROM THE EASTERN U.S.A. 879 Fic. 3. Sinkhol 1 habi f Roltoni, i Augu ta County. Virdinia. Photo by G oc J y dry Fr. PIecninniy. Fig. 4.2 ied by Boltoni | in of “Piggyback Pond,” Warren County, New Jersey. Photo by John F. Townsend. Pr 7 J J 880 BRIT.ORG/SIDA 22(2) H. virginicum in several altered ponds, most of which are in pasture. As a result, these ponds include a number of species with broader habitat associations such as Juncus effusus, J. canadensis, J. tenuis, Bidens frondosa, B. polylepis, Ambrosia artemesiifolia, Phytolacca americana, Daucus carota, Vernonia noveboracensis, Digitaria ischaemum, Echinochloa crusgalli, Datura stramonium, Eclipta alba,and Erigeron annuus,among others. The New Jersey ponds are decidedly calcareous, occurring within a glaciated karst landscape on dolomite bedrock. These sinkholes are found within the Ordivician Allen- town Dolomite and Beekmantown Dolomite formations, with surficial deposits composed of a glacial till/ bedrock complex. In rare instances, Boltonia montana populations may be found on glacial outwash (kettles). In all cases, the amount of overburden present is significantly less than (and more alkaline than) that found in the Virginia pond com- plexes, and the carbonate rocks or marl deposits are usually at or near the soil surface, providing a slightly acidic to mildly alkaline soil environment (pH 6.2-7.8 at 0-30 cm; Walz et al. 2001). In New Jersey the presence of marl and Chara spp. are characteristic of the sinkhole pondshore habitat for B. montana. Although soil chemistry of these ponds differs from Virginia examples, dramatic fluctuations in the seasonal water table (as much as 5.5m per season in New Jersey) are characteristic of ponds occupied by the species in both states. Soils in New Jersey are shallow silt loams, silty clay loams, silty clay, or rarely silt marl overlaying clay or bedrock. Marly muck or peat soils are also present in rare in- stances and small dolomite rocks are often found on the surface (Walz et al. 2001). Deep marl deposits often occur in the center of ponds occupied by B. montana, but as with the Virginia populations, Boltonia montana usually occurs ina distinctive doughnut-shaped zone or “ring” between the pond edge and the more frequently and deeply flooded pond center (Fig. 4). The community type occupied by the new taxon in New Jersey is the Boltonia asteroides var. asteroides - Symphyotrichum racemosum - Mentha arvensis Herbaceous Vegetation Asociation (Grossman et al. 1998; Natureserve 2005; Walz et al., 2001), a com- munity type that is ranked G1G2 by Natureserve (the naming of B. montana will involve a concomitant change in the community name). Characteristic species in this commu- nity type include: Asclepias incarnata, Cyperus strigosus, Carex cryptolepis, C. lurida, C. typhina, C. viridula, Cuphea viscosissima, Eleocharis palustris, Eragrostis frankii, Eupa- torium perfoliatum, Iris versicolor, Dichantheliumacuminatum var. fasciculatum (=Pani- cum lanuginosum var. tennesseense), Panicum capillare, Panicum rigidulum var. pubescens (=Panicum longifolium), Polygonum amphibium, Sium suave, the green algae Chara spp., and the bryophytes Hypnum lindbergii, Campylium stellatum,and A serpens (Walz et al. 2001). Species in this community type which are rare in New Jersey include: Boltonia montana, Carex bebbii, C. cryptolepis, C. haydenii C. retrorsa, C. typhina, C. viridula, Eleocharis quadrangulata, and Eragrostis frankii (Snyder 2005). Many endemic wetland plant species are confined to habitats with widely fluctuat- ing hydroperiods, and it has been suggested that the ability of these species to tolerate such stressful conditions releases them from competition with other taxa (Knox 1997). This aspect of the life history of Helenium virginicum was studied by Knox (1997), who concluded that the ability of plants to tolerate the twin stressors of potentially toxic soils and fluctuating hydrology was rare and contributed to the species’ endemism. Boltonia montana and H. virginicum co-occur at several Virginia ponds where these adverse soil conditions (low pH and elevated levels of aluminum and arsenic) have been documented, with the remainder of Virginia ponds thought to have similar soil characteristics. Data TOWNSEND AND KARAMAN-CASTRO, A NEW SPECIES OF BOLTONIA FROM THE EASTERN U.S.A. 881 on soil relations in the New Jersey ponds are incomplete, but indications of high alumi- num, calcium, and magnesium levels have been found and erratic water level fluctua- tions are well-documented there (Walz et al. 2001). Additional specimens examined: NEW JERSEY: Sussex Co.: near Lake Grinnell, 17 Sep 1887 N.L. Britton, s:n(NY); Swartswood Lake, 6 Jul 1907, Philip Dowell 4951 (GH), low swale, Brighton, 4 Sep 1910, K.K. Mackenzie 4778 (GH, NY); borders of pond E of Swartswood Lake, Aug 1911, K.K. Mackenzie 4922 (NY, PH); border of Muckshaw Pond, Springdale, Aug 1917, K.K. Mackenzie 7986 (NY); Ea ae Pond,” 3.1 km E of Swartswood, 4.7 km NW of Newton, 3 Oct 2005, J.F Townsend 3556 (VPI, NCU, CLEMS); “Duck Pond,” Swartswood State Park, 1.2 km N of Paulinskill, 5 km WNW of Newton., 3 Oct 2005, J.F ay nsend 3557 (PH, PENN); “Frog Pond,” Swartswood State Park, 1.75 km NNE of Paulinskill, 5 km WNW of Newton., 3 Oct 2005, J.E Townsend 3558 (NY); “Little Frog Pond,” Swartswood State Park, 1.75 km NNE of Paulinskill, 5 km WNW of Newton, 3 Oct 2005, J.F Townsend 3559 (VPI); “Spring Lake,” Swartswood State Park, 3.1 km NW of Newton, 2.1 km SE of Swartswood, 4 Oct 2005, J.E Townsend 3562 (BRIT, GMUF GH); northernmost “Muckshaw Ponds,” 1.6 km SW of Newton and 0.4 km W of US 206, 4 Oct 2005, J.E Townsend 3565 (MARY, WYVA, VPI): “Muckshaw Ponds.” 2 km SW e Newton and 0.6 km W of US 206, 4 Oct 2005, J.F Townsend 3566 (CHRB, PH, NY); “Muckshaw Ponds,” 2.75 km SW of Newton and 0.6 km W of US 206, . Townsend coe NCU, GMUF). Warren Co.: Muddy calcareous ae and flats, Shyster Pond, 24 Jul 1920, K.K. Mackenzie Y, PH); muddy shore of limestone sink pond, eee 4 Oct 1981, DB. Snyder 551-6P - ee Pig ac ee located ca. 5 km NE of Blairstown and 1.6 km N of Squires Corner, J.E Townsend 3560 (GH, CLEMS, USCH); (Warren/Sussex Co. line) “Four-Angle Pond,” fae 16 km NW of Tranquility and 1.25 km a W om es, 3 Oct 2005, J.F Townsend 3561 (VPI). PENNSYLVANIA. Dauphin Co.: banks of the Susquehanna, near Dauphin, 2 Aug 1865, CE Parker, s.n. (NY): Harrisburg, Jul 1864, LC. Martindale, s.n.(PH). VIR- GINIA. Augusta Co.: Sandy, wet soil, bank of South River, vic. Lipscomb, 18 Jul 1936, L.L. Carr 47 (GH); growing on bank of South River vicinity of Lipscomb, abundant in Stuart's Draft area around ponds and along South River, 18 Jul 1936, E.S. Rawlinson 58A (VPP: Kennedy Mt. Meadow, (unintelligible) bog, 14 Sep 1937, L.L. Carr, s.n. (PH); moist edge of pond (open sit.), vic. Lipscomb, 5 Aug 1938, L.L. Carr 365 (GH); edge of dried up pond, Big Levels refuge, 8 Sep 1939, A.B. Massey, s.n. (VPI); Lipscomb Pond, in water, vicinity of Lipscomb, Shenandoah Valley, 8 Sep 1940, EP Killip, 36069 (US); adjacent to Spring Pond, near Stuarts Draft, bottom of dried up pond, 16 Sep 1964, R. S. Freer Sete (GH, ve ieee to Spring Pond, near Stuarts Draft, bottom of dried up pond, 23 Sep 1964, R.S. Freer 2869 (VPD); edge of pond near Spring Pond, 5 Aug 1967, A.M. Harvill 17240 (FARM, NCU); Twin Ponds, near Sherando, common in aca portions of pond, elev. 1550 ft, 19 Sep 1970, CE. Stevens 2781 (VPI); dried up pond, open area, Twin Ponds, Big Levels Game Management Area, 19 Sep 1970, LE Wieboldt 271 (WILLD; around the margins and drawdown areas of Spring Pond, elev. 1600 ft, 26 Sep 1970, G. Ramsey, R.S. Freer, R. Bruce, et al. 18260 (WILLD; common in shallow water of open edge of Twin P Pond, near Sherando, elev. 1540 ft, 16 Oct 1971, CE. Stevens 4772 (VPI, WILLD; Wood duck Pond, scattered in aod around edges of shallow es hole pond 1.2 mi SE of Lipscomb, elev. 1440 ft, 2 Oct 1984, C.E. Stevens 19078 (FARM); Campbell’s Pond, 1.7 mi N of Sherando, SW side of St. Rt. 634 1 mile N of jet. with St. Rt. 610, elev. 1470 ft, 6 Aug 1986, G. Fenwick, s.n. aD common with Eriocaulon septangulare in dried muddy bed of intermittent sinkhole pond (“Twin Pond”), 2.3 mi W of Sherando, elev. 1540 ft, 22 Sep 1991, G.P. Fleming 6126 w/ CE. Stevens (WILLD; Twin Ponds, located ca. 2.26 mi due W of Sherando, 8 Aug 2002, J.K Townsend 2869 (VPI, WILLI, Va. Div. Natural Heritage Herbarium). Relationship to Boltonia asteroides var. asteroides and B. caroliniana The gross morphology of Boltonia montana is nearly identical to that of B.asteroides var. asteroides, a fact which explains the use of this name in both Virginia and New Jersey for many years. Boltonia asteroides var. asteroides will be referred to as B. asteroides for the sake of brevity in the following discussion. The cypselae of Boltonia montana are found to lack several key characteristics found in B. asteroides, namely: cypselae wings, cypselae trichomes, and pappus awns. Occasional plants of B. montana will posess cypselae with slender pappus awns that are longer than the typically low crown of scales. In this respect, such plants may have awns approach- ing the length seen in B. asteroides, but the other key features, glabrous cypselae with wings absent or vestigial, readily separate them from B. asteroides. Fernald (1940) de- scribed B.asteroides as having obovate, thick rimmed cypselae with awns wanting or up to 0.7 mm long and apparently included the plants of northwestern New Jersey in his 882 BRIT.ORG/SIDA 22(2) concept of B.asteroides at that time, which probably accounts for his description of awn- less B. asteroides in Gray’s Manual (Fernald 1940, 1950). In many specimens of B. asteroides, phyllaries are mostly sclerified and whitish on either side of the midrib and almost to the phyllary tip, becoming greenish gradually in the distal portion, and have a relatively large, resinous midrib (1/5 to 1/4 of phyllary width) which continues to the phyllary tip. In contrast B. montana phyllaries are only sclerified and whitened in their lower half or less, becoming green and herbaceous in the distal portion. The midrib is relatively nar- row (1/10 to1/5of phyllary width) in the lower half and anastamoses in the distal half to produce a network of fine veins within the green portion of the phyllary. It was at first assumed that the phyllaries seen in B. montana were always unlike those found in B. asteroides, but several specimens of B. asteroides from the Susquehanna River in Pennsyl- vania have similar phyllary morphology to the former. Whether this represents past gene exchange with B. montana or character convergence is not known. Due to the strong resemblance between the cypselae of Boltonia montana and B. caroliniana, the latter species was investigated to discern other morphological similari- ties, if present. Other than cypselae morphology, these two taxa have few character states in common. Phyllaries of Boltonia caroliniana are quite dissimilar to B. montana, and most closely resemble B. asteroides in terms of their white color and relative midrib size, but are shorter and narrower than both B. asteroides and B. montana. Head size, ray corolla length, and pedicel diameter of B.caroliniana are smaller than in B. asteroidesand B. montana, giving a delicate appearance to the inflorescence of B. caroliniana. In addition, typical individu- als of B. caroliniana are profusely branched and have numerous heads. Most individuals of B. montana and B. asteroides have relatively few heads (often 1-25), although plants of both species have been noted to produce large plants with numerous heads (Karaman- Castro and Urbatsch 2006, see below). Such large reproductive outputs are seldom seen and may be a strategy to sidestep adverse meterological or edaphic conditions. Ray corol- las of B. caroliniana, in addition to being smaller than those of B. montana, are white, fading to pink or lilac in age (Anderson 1987, pers. obs.), while the rays of B. montana are usually lilac to pinkish throughout the flowering period, though they may also be extremely pale (see Fig. 2). Most plants of B. montana are characterized by widely oblanceolate leaves whereas those of B. caroliniana are linear to narrowly oblanceolate or linear-obovate. This difference is most evident when bracteal leaves of the two species are compared. Morphological similarity of Boltonia montana to B. asteroides and B.caroliniana sug- gest that B. montana may have arisen as a result of an ancient hybridization between the two species. Phylogenetic analyses of the internal and external transcribed spacers (ITS and ETS) from the nuclear ribosomal DNA suggested sister relationaship of B. montana and B. caroliniana, while only distant relationship with B. asteroides (Karaman-Castro et al. in prep.). Thus the treatment of the awnless and wingless specimens of B. asteroidesas a new species, B. montana was supported by molecular data. However, it remains unclear whether B. montana isa result of an ancient hybridization between B. caroliniana and B. asteroides. Some observations on the morphology of Boltonia montana The stature of Boltonia montana varies from ascending to erect, with most individuals exhibiting the former condition (Fig. 1). It seems probable that this growth pattern is a response to edaphic conditions in the ponds rather than genetics. For instance, observa- tions at “Twin Ponds” in Augusta Co., Virginia were generally of few-flowered, ascending TOWNSEND AND KARAMAN-CASTRO, A NEW SPECIES OF BOLTONIA FROM THE EASTERN U.S.A. 883 or erect plants from 2 to 6 dm in vertical extent (Fig. 2). This growth form is present in most populations, but was completely absent during the 2002 season at this site, where all plants were erect, .5-2.0 m tall, bore numerous heads, and had narrower bracteal leaves. Since this growth form was produced at the height of a multi-year drought, it seems pos- sible that soil moisture relations were involved in mediating this morphological shift. Herbarium specimens and observations of New Jersey populations also produced the general impression of decumbent or low growth, but plants in somewhat drier micro- habitats tended to be taller, more strictly erect, have more heads, and possess narrower bracteal leaves. In light of this variability, the particulars of inflorescence size and shape, head distribution, and plant stature should not be considered diagnostic of this taxon. Adventitious roots sometimes form at nodes along the decumbent portion of stems in Boltonia montana. These roots are either associated strictly with the stem or may be attached to leaves similar in appearance to basal rosettes. These “offsets” are a means of vegetative propagation and are often easily detached from the plant in the fall of the year. Such rooting along the stem was encountered more often in New Jersey populations than in Virginia localities and basal offsets were produced in all populations studied. These structures are hypothesized to be the primary means of perennation in the species. Smith and Keevin (1998) describe the adaptations for long-distance water dispersal in Boltonia decurrens cypselae and pointed to the evidence of such dispersal under field conditions. They found the wide, flat profile of the cypselae and wings, along with the trapping of air bubbles by cypselae trichomes, allows them to easily rest on the surface of the water. In one experiment, 20% of cypselae were still floating after + weeks of simu- lated wave action. No similar experiments have been done with B. montana, but cypselae morphology implies a reduced level of dispersability when compared to all other mem- bers of the genus except B. caroliniana. Given that nearly all B. montana populations are found in small, island-like wetlands within an upland matrix, these physical character- istics may have developed due to the improved survival of sedentary (wingless, awnless, hairless) cypselae. Schiffman (1997) noted wing width reductions in cypselae from is- and populations of Coreopsis gigantea (Kellogg) Hall when compared to conspecific mainland individuals. She theorized that this atrophy occurs because “dispersal-enhanc- ing morphologies increase probabilities of being transported beyond the bounds of these narrow habitats,” a classic argument applied to many specialized taxa of islands. In the case of Boltonia montana, the reasons for its restriction to such narrow “island-like” habi- tats are only speculative but it seems likely that a competitive advantage is enjoyed by the species in sinkhole ponds and enhanced dispersal structures would represent an un- necessary expenditure of energy. Occasional overland flows do provide occasional con- nections between neighboring sinkhole ponds in Virginia, (Christopher S. Hobson 2005, pers. comm.) and despite lacking obvious dispersal mechanisms, B. montana propagules could conceivably disperse to adjacent ponds during these relatively rare overflow events. A KEY TO BOLTONIA MONTANA AND RELATED SPECIES Given the morphological variability observed in Boltonia species, several morphological characters should always be used when identifying these taxa. Taken together, the fea- tures of phyllaries, and cypselae awns, wings, and surfaces are diagnostic of Boltonia montana. The following key distinguishes Boltonia montana from B. asteroides var. asteroides and B. caroliniana, the two congeners most likely to be confused with the new taxon. 884 BRIT.ORG/SIDA 22(2) 1. Cypselae with two distinct pappus awns in addition to a shorter ring of scales, the awn stly 0.8-1.1mm long; cypselae wings obvious, mostly 0.3-0.4mm wide; lateral faces of cypselae pubes Boltonia ast Jes var. asteroides ent ; seer with pappus reduced to a short ring of scales to 0.15mm long or with occasional slender awns to 0.6mm; cypselae wings lacking or up to 0.1mm wide; lateral faces of cypselae glabro —_ N ee 0.2-0.5mm wide, sclerified throughout or with herbaceous a midrib relatively prominent (1/5-1/3 phyllary width); heads 4.3-6.1mm wide, peduncles 0.25-0.5 mm in di- ameter;ray corollas white fading to pinkish in age; cauline and bracteal a narrowly oblan- ceolate to linear Boltonia caroliniana Phyllaries 0.4—0.9mm wide, sclerified in lower 1/3 to 1/2 distal portion dilaae mid- rib relatively narrow (1/10-1/5 phyllary width);heads 6-11.5mm wide, peduncles 0.5—1.1 mm in diameter; ray corollas lilac or pinkish (-white); upper leaves oblanceolate to oblong Boltonia montana N CONSERVATION Isolated wetlands of the eastern United States are known for harboring a large number of rare species and for being heavily impacted by human activity (Weakley & Schafale 1994; Edwards & Weakley 2001; Comer et al. 2005). Research has focused heavily on depres- sion wetlands of the coastal plain, but some studies have dealt with rare and disjunct species affiliated with depression wetlands of the interior (Steyermark 1963; Homoya 1983: Bartgis 1992; Fleming & Van Alstine 1999), including the rare pond endemics Helenium virginicum and Scirpus ancistrochaetus. Ponds containing Boltonia montana have been the focus of conservation efforts in both Virginia and New Jersey due to the large number of globally and regionally rare plants and animals already known to occur in them. A patchwork of private and public ownership exists for these ponds, and the level of threat to these wetlands is generally high due to their small size and location within upland areas. A recent Supreme Court ruling also called into question the jurisdiction of the Clean Water Act over isolated wet- lands (Comer et al. 2005), potentially relaxing protections for these natural ponds and their characteristic flora. Given their vulnerability and relative scarcity on the landscape, efforts to protect these unique wetlands should continue to be a high priority. ACKNOWLEDGMENTS We thank staff of the Virginia Department of Conservation and Recreation, Division of Natural Heritage for their insight and support during this project. Thomas F. Wieboldt of Virginia Polytechnic and State University provided taxonomic expertise and access to specimens. David B. Snyder of the New Jersey Natural Heritage Program and Kathleen Strakosch Walz of the New Jersey office of the Nature Conservancy provided detailed collections data, maps, field expertise, and interpretation of the ecological communities and karst landscape of northwestern New Jersey. Greg Plunkett of Virginia Common- wealth University coordinated loans from various herbaria and provided work space. The staff of the Gray Herbarium, Harvard University (GH), the University of North Carolina- Chapel Hill (NCU), Longwood University (FARM), and the College of William and Mary (WILLD are thanked for their hospitality while studying specimens. Karen Townsend is thanked for scanning and manipulating the illustrations comprising Figure 1. Alan Weakley and an anonymous reviewer provided valuable suggestions for improving the manuscript. Special thanks go to Adam and Zena Rudzki for access to, and permission to collect, the type voucher specimen from their private property in New Jersey. TOWNSEND AND KARAMAN-CASTRO, A NEW SPECIES OF BOLTONIA FROM THE EASTERN U.S.A. 885 REFERENCES ANDERSON, L.C. 1987. Boltonia apalachicolensis (Asteraceae): A new species from Florida. Syst. Bot. 12: 33-138 Bartais, R.L. 1992. The endangered sedge Scirpus ancistrochaetus and the flora of sinkhole ponds in Maryland and West Virginia. Castanea 57:46-51. Comer, P., K. Goopin, G. Hammerson, S. MENARD, M. Pyne, M. Reio, M. Rostes, M. Russo, L. SNEDDON, K. SNow, A. Tomaino, and M. Turrty. 2005. Biodiversity values of geographically isolated wetlands: an analysis of 20 U.S. states. Natureserve, Arlington, VA. Eowaarps, A.L.and A.S. Weakiey. 2001. Population biology and management of rare plants in depres- sion wetlands of the southeastern coastal plain, U.S.A. Nat. Areas J.21:12-35. Feanato, M.L. 1940. A synopsis of Boltonia. Rhodora 42:482-492. FerNaco, M.L. 1950. Gary’s manual of botany (ed.8).D.Van Nostrand Co., New York. Fiemine,G.P.and N.E.VaN Atstine. 1999. Plant communities and floristic features of sinkhole ponds and seepage wetlands in southeastern Augusta County, Virginia. Banisteria 13:67-94. Grossman, D.H., D. FaBeR-LANGENDOEN, A.S. WEAKLEY, M. ANDERSON, P. BourGERON, R. CRAWFORD, K. GOODING, S. LANDAAL, K. Metzter, K.D. Patterson, M. Pyne, M. Rein, and L. SNeooon. 1998. International classification of ecologi- cal communities; terrestrial vegetation of the United States. 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Identifica- tion and protection of reference wetland natural communities in New Jersey: calcareous sink- hole ponds of the Kittatinny Valley. New Jersey Department of Environmental Protection, Divi- sion of Parks and Forestry, Office of Natural Lands Management, Natural Heritage Program, Trenton, New Jersey. Weaxtey,A.S.and M.P. ScHarace. 1994. Non-alluvial wetlands of the southern Blue Ridge—diversity in a threatened ecosystem. Water, Air, Soil Pollution 77:359-383. Wuittacar, R.G., D.F. Durry, and A. Nowrooz!. 1999. Late Cenozoic history of alluvial fans near Stuarts Draft, Virginia. Banisteria 13:210. oy A NEW SPECIES OF WATERLILY (NYMPHAEA MINUTA: NYMPHAEACEAE) FROM MADAGASCAR Kenneth Landon Richard A. Edwards 2906 Waco Street 2417 Brass Lantern Road San Angelo, Texas 76901, U.S.A. Cedarcreek, Missouri 65627, U.S.A. P Ivan Nozaic & Veivers Drive Sneewah Kuranda pb Queensland, 4881 AUSTRALIA ABSTRACT A new species of waterlily, Nymphaea minuta, is described from Madagascar It is closely related to N. stellata Willd. but displays a characteristic rompnenon of Sanaa chatacters aud two ewes forms in its life re The undersurfaces of the | | to brownish-viol f the taxon. mphaea minuta in nature is a dwarf with small eee , cleistogamous flowers. In eulayaiein it exhibits two eosin growth forms: a submerged form and an emergent form with floating leaves and larger, emergent, chasmogamous flowers. Key Worps: Waterlily, Nymphaea, Madagascar RESUMEN especie de iitto de agua, Nymphaea minuta, de Madagascas, eae Ee emparentada con N. ellie Willd. Pero ti én peculiar d y ale a Els envés de sus ee es de color gris-pardo a marrén- violace, una caracteristica singular del taxon. i aes una planta enana, sumergida En cultivo, muestra dos f d imiento diferentes: una ane sumergida y una forma orient et con hojas flotantes y flores mas grandes, emergente y casmogaimas. The senior authors’ long-time interest and expertise in the cultivation, research, and iden- tification of waterlilies has occasioned the present paper. The junior author collected seeds from Madagascar that were subsequently grown at the International Waterlily Preserva- tion Repository in San Angelo, Texas. Upon examination of these garden-grown plants, it became apparent that the material concerned represented an undescribed taxon, the de- scription of which follows. i des minuta K. Landon, R.A. Edwards & PI. Nozaic, sp. nov. (Figs. 1-4). Type: MADA- ASCAR: near village of Tampolo, Coastal Forest in shaded rain pools (ca. 49°...26'E, 17°... 15'S), 1Jul 1999, PI. Nozaic s.n. (HOLOTYPE: TEX). Nymphaeae stellata Willd. Verd imilis sed differt floribus di libi celal bien 11 | et/vel ents portatis et paginis foliorum emergentium supra viridibus infra limoso-canis vel dilute brunneolo-violaceis Plants rhizomatous growing horizontally, of two distinct forms: submerged and emersed. Submerged (underwater) form: leaves very thin, flaccid, ovate, the margins entire and wavy, surfaces glabrous, pustulose to wrinkled, bright green above, the veins distinct, lower surfaces medium-green with a bronze to reddish cast, the veins bright green, obvious and distinct, having 5 primary veins on each side of the midvein, the surface glabrous, pustulose to wrinkled, the sinus slightly open with some overlap near attachment to peti- SIDA 22(2): 887 — 893. 2006 888 BRIT.ORG/SIDA 22(2) DRAWING by PITTMAN Fig. 1 Al n ° fA Dian#aith rT 4 1 £] +; | (fi Loa lae \ ole; blades mostly 10 cm = 14cm, the petioles usually less in length than the blade, keep- ing habit of plant compact and rosette-like in shape; petioles 2-3 mm wide, olive green, glabrous. Flowers produced on submerged plants rarely reaching surface, cleistogamous, rarely opening underwater; base of cleistogamous bud 0.5-2.0 cm wide, 1.5-2.0 cm long; ripe fruit to 3cm = 3cm, petioles slightly curving down as fruit matures, fruit ruptures to release seed, seeds brown, oval, smooth, ca. 0.5 mm wide, at release enclosed by aril. Emer- gent (above water) form: leaves round, 7-12 cm wide, margins entire to slightly sinuate near sinus, sinus open with 10° to 30° angle, the surface medium-green with no other pigmentation or marking at any stage of development, lower surface pale violet with brown tones, muddy-gray appearance, bright green veins distinctive, petioles olive-green, maximum length observed 24 cm, all surfaces free of hairs or scales, overall growth pat- tern a circle of ca. 60 cm wide. Flowers small, 2.5-4.0 cm wide; sepals 3 to 4, somewhat longer than the petals, tips blunt-pointed, twice the width of petals at the base, sepal olive green changing to magenta at base with faint lighter stripes on outside, 7 to 9 veins, pale pink to white on inside, with faint greenish stripes. Petals 6 or 7, pale pink to white, 1.5-2.0 cm long, 0.5 cm at base, tapering to a blunt-pointed tip, with 3 main veins and 2 smaller veins on the outer edges. Stamens 8-32, shorter than petals, yellow with append- ages following petal color. Anthers comprising most of the length, ripe pollen released on the Ist day, self-pollinating but also receptive to pollination by other waterlilies in the subgenus Brachyceras. Carpels 5 to 15, carpellary appendage a hook-like extension on outside wall of carpel, 2-3 mm long. Flowers open near noon and close by late afternoon, LANDON ET AL., A NEW SPECIES OF NYMPHAEA FROM MADAGASCAR 889 DRAWINGS by PITTMAN Fic. 2.N I h inuta. A Seedling B. Bud. C. Sut g {and ial fl D. Aerial flower. E. Fruit submerge on third day, open to ca. 45° on first day, and slightly more on 2nd day, often do not open on 3rd day, first day flowers usually produce stigmatic fluid, peduncles curve down into water as fruit matures. Fruit orbicular, ca. 3 cm x 3 cm, rupturing to release seeds; seeds brown, smooth, oval, ca. 0.5 mm wide, enclosed by aril at release. Distribution and Ecology.—This species is known in Madagascar only from the type locality, where it grows in jungle pools along slow-moving streams. The new species is adaptable to growing conditions other than those encountered at the type location and easily cultivated, especially in low-light environments. Etymology.—The type plants discovered in Madagascar were all dwarf. Under typi- cal waterlily cultivation, all plants grown remained dwarf. Only under intensive atypi- cal cultivation regimes have larger plants been produced. This species is by far the small- est known species of Nymphaea. The epithet ‘minuta’ was proposed by the author who discovered this species. After five years of cultivation and multiple generations for con- sideration, all authors believe ‘minuta’ to be accurate and appropriate. Life history of Nymphaea minuta Nymphaea minuta in nature is a dwarf with a submerged habit, including small, cleisto- gamous, submerged flowers. These flowers rarely reach the water surface and apparently are self-fertilized without the sepals opening or exposing the petals. Various species from all subgenera of Nymphaea are self-pollinating, but only N. minuta has been documented to do this underwater. Plants of N. minuta in cultivation display both submerged and BRIT.ORG/SIDA 22(2) s * Fic. 3. Top, submerged plants under artificial light in an aquarium, not submerged bud on center plant. Bottom, submerged plant grown in pond under nature conditions in a tropical climate. LANDON ET AL., A NEW SPECIES OF NYMPHAEA FROM MADAGASCAR 891 disk, | Il | 1 1 I 1 fl fully open. 892 BRIT.ORG/SIDA 22(2) emergent growth forms, the flowers are chasmogamous and vary from floating on the water to rising above the surface. Floral morphology and color remain constant in all growth forms. In response to changing environmental conditions, an individual plant can switch from one form to the other. During these transitions, intermediate phases with both floating and submerged leaves are produced, along with both emergent and sub- merged flowers (Fig. 2a, 2b). Light intensity and duration appear to be the controlling factor in these transitions. In temperate latitudes, mature plants of Nymphaea minuta exposed to full sun will develop the emergent phase throughout the summer but revert to the submerged phase as the daylight lessens in length and intensity. The same plant often reverts back to the emergent phase the following summer. Almost all cultivated specimens have been dwarf. However, under cultivation with fertilization and optimum conditions of light and climate, plants have been produced roughly four times larger (including flowers and all vegetative parts of the plants) than the dwarf plants first discovered. Seeds [of cleistogamous and chasmogamous flowers] remain the same size, although the number of seeds produced from the larger flowers and resulting larger fruits, increases proportionally to the rest of the plant. The number of carpels and stamen also increase at the same ratio as the size increase of vegetative parts. Seedlings produced in full sun in late spring through mid summer will immediately grow into an emergent plant, with floating leaves and flowers at or above the water sur- face, typical of brachyceras tropicals. Seeds germinated in the late summer or fall will often grow in the submerged form until the following summer or until artificial light of enough intensity and duration are provided, causing the plant to transform into the emer- gent form. Seedlings grown in shade remain in the submerged phase but continue to grow and produce flowers and fertile fruits. In chasmogamous flowers, stigmatic fluid is usually produced on the first day of anthesis, and ripe pollen is released. As the fruit ripens, the peduncles bend downward and fruits mature beneath the water surface. As the plants mature, a perennial rhizome is produced. Dormant tubers have been produced in cultivation and would be expected to occur in the wild populations in ephem- eral conditions. Relationship of Nymphaea minuta to other species Characters of the flowers place the new species in Nymphaea subgenus Brachyceras. The cary ee at the sides with separate walls for each ovary. Carpellary styles are present asa small hook-like extension at the back of each carpel. The stamens have distinct ap- pendages beyond the anthers. Nymphaea minuta at a glance ee more closely resembles Nymphaea stellata Willd. (not N. nouchali Burm. f., a different species, in our opinion) of tropical Asia than any other known species of aoe Both species display a coarsely dentate margin on floating leaves. Flowers display very low petal numbers in both species. The type habitat for N. minuta would also support N. stellata although ina stunted form. Such plants would in fact more closely mimic the dwarf nature of N. minuta rather than a typically proportioned N. stellata. Upon closer inspection several differences between the two species would become apparent. Nymphaea minuta has no contrasting pigmentation upon the leaves in either LANDON ET AL., A NEW SPECIES OF NYMPHAEA FROM MADAGASCAR 893 form. Nymphaea stellata often display slight mottling or flecks of purple pigmentation on floating leaves as they mature. The lower surface of the leaves of both species exhibit distinct and different venation and pigmentation patterns. Like most Nymphaea species, N. stellata produces submerged leaves as seedlings and emerging from dormancy, but they are diminutive, quickly deteriorate and do not represent a submerged growth form. Nymphaea minuta produces substantial and persistent submerged foliage, flowers and fruits, completing a full life cycle completely submerged. Nymphaea minuta apparently will hybridize with other species and cultivars of subgenus Brachyceras. With N. minuta as pollen donor, cross-pollination was attempted on N. micrantha Guill. & Perr. and N. colorata Peter. and seed development proceeded in normal fashion. Similar results were obtained using three additional cultivars as seed parents. Nymphaea minuta was also used as the seed parent with crosses to hybrids with distinctive features, such as pigmentation on leaves. Further tests must be performed to see if cross-pollination actually occurred (vs. asexual seed development), but both senior authors have progeny from crosses displaying obvious hybridization. Crosses within the subgenus Brachyceras, with resulting progeny displaying obvious characteristics, sucha pigmentation on the leaves similar to the pollen parent, provides further evidence this species should be placed within the subgenus Brachyceras. Dimorphism in growth form is a rare feature in the genus Nymphaea. No published material exists discussin g this feature with any other known subgenus of Nymphaea. At the International Waterlily Preservation Repository, we have taxon in the subge- nus Brachyceras that appears to exhibit some degree of persistent submerged growth, however, the research is incomplete. ACKNOWLEDGMENTS We are grateful to Guy Nesom for the Latin diagnosis and for a detailed review of the manuscript and to B.L. Turner for his encouragement and help with this project. We also thank Ed Schneider for a thorough review of the paper. Drawings for the new species were kindly provided by Katherine Pittman. REFERENCES Conard, H.S., 1905. The waterlilies: a monograph of the genus Nymphaea. Publ. Carnegie Inst. Wash- ington. No. 4. Carnegie Institution Washington, Washington, DC. BRIT.ORG/SIDA 22(2) Book REVIEW Donato G. Crossy. 2004. The Poisoned Weed, Plants Toxic to Skin (ISBN 0-19-515548-3, hbk.). Oxford University Press. (Orders: Oxford University Press, 2001 Evans Road, Cary, NC 27513, U.S.A., 800-451-7556, 919-677-1303 fax, wwwoup.com). $59.95, 214 pp. figures, graphs, 24 color figures, 6 1/2" x 91/2" about the plants that Dermatoxic plants are all around us. The book entitled helps readers learn much more a cause itchiness, rashes and other very painful symptoms. This book gives a detailed background with compre- chemicals involved in causing dermatoxic reactions. first gives a background history of a variety of plants that cause Informatior n the hensive, technical The book consists of ten chapters. The reactions such as allergic contact dermatitis (ACD) in humans. Much of this history centers on the naming, his- ived causes of the ACD reaction from poison ivy and its relatives. The second chapter focuses on ir relatives. The third chapter discusses other —_ tory and the taxonomy, habitat and distribution of the poison ivies and the plants that can cause ASP in humans. Plants covered by the “other allergens” include wild species such as onwood), Quinone en. plants such as the amano 20 trees or ex- id \ recian lau- fetopium otic bardaroads — eacel (Dalbe is gid sp.), | lactone containing § species such as Lat ts and others rel), weeds, food pl chapter four discusses preetone pes from the Rutaceae family, Apiaceae family, and Moraceae nae and Urticaceae families. Chapter five | Brassicaceae, Alliacea Fats 6a along with irritant plants from is a chapter detailing chemical and physical properties of the allergens related to urushiol. This nee ation meuniees aan ication ane anal has ondation, puoi. and bio-degradation, reactions with proteins and more. [ | properties of allergens such as quinines, lactones, acety- sand rubber latex. Chapter seven discusses the physical and chemi- (oni alcohols, essential oils, lic] b cal properties of phyioreac and irritant (non- a ae ee neuen Furocoumarins, Hypericins, Irri- xalates, Chapters eight and nine — tant esters, organosulfut ( ompounds — forms of exposure to various ACD substances and their adverse effects. hapter ten includes methodologies to prevent exposure to ACD substances as well as discussion on treat- ment options. The chapters are followed by multiple appendices, containing chemical structures and physical Calis for various substances discussed in the text and some key references. Indices are separated into plant a general index inder common names and The Poisoned Weed: Plants Toxic to Skinis a very aa mative and timely book. P rimarily designed for pro- elds, it looks to be the new authority on dermatoxic name fessionals in the biochemical, chemical, and medicina plants and substances which cause allergic contact pees Luckeydoo, Herbarium, Boe Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A SIDA 22(2): 894. 2006 TWO NEW COMBINATIONS IN PEYRITSCHIA (POACEAE: POOIDEAE: AVENINAE) Victor L. Finot Paul M. Peterson eae de Concepcion Department of Botany nomia eee ee Depart to de Produccién Animal mithsonian fe Casilla 537, Chilldn, CHILE ie DC 20013 ae U.S.A, vifinot@udec.cl ae, Fernando O, Zuloaga Instituto de Botanica Darwinion 3 0 San Isidro 1642, ARGENTINA fzuloaga@darwin.edu.ar ABSTRACT Trisetum howellii Hitchc., an endemic species from the Galapagos Islands, Ecuador, and Trisetum pinetorum wa Ae an aos Species pom Guatemala, are pineal into Peyritschia IP. howell fEitehe. ) Finot & PM. PM. Peterson]. P with new areas of distribution documented in South America. A key disti he species and able com- paring the salient morphological features of all species within be are erouded RESUMEN Tri howellii Hitchc., especie endémica de | la cuador, y Trisetum pinetorum Swallen, especie endémica de Guatemala se transfieren a Peyritschia [P. howellii ieee Finot & P.M. Peterson; P. pinetorum (Swallen) Finot @& P.M. Peterson]. Peyritschia, asi circunscrita, incluye siete especies con nuevas areas de distribucion documentades en Sudan ence) Pioporeona una epi peas distinguir las especies y un cuadro onde se compar g las especies de Peyritschia. The genus Peyritschia was described by Fournier (1886) and originally included only one species, P. koelerioides (Peyr.) E. Fourn., based on Aira koelerioides Peyr, from south- ern México and Guatemala (Finot et al. 2004). A second species from México, Deschampsia pringlei Scribn., was transferred to Peyritschia by S.D. Koch in 1979[= P. pringlei (Scribn.) S.D. Koch], and is also known from Guatemala, Costa Rica, Venezuela, and Ecuador (Hernandez-Torres & Koch 1988; Pohl & Davidse 1994; Finot et al. 2004). This small ge- nus (Peyritschia) was later included within Trisetum by Hernandez-Torres and Koch (1987). When placed in Trisetum, P. koelerioides and P. pringlei are named: Trisetum altijugum CE. Fourn.) Scribn. and T. kochianum Hern.-Torres., respectively. Peyritschia has isomorphic to sub-isomorphic, linear, 1-3-nerved glumes; bilobed lemmas that are awned near the base or the middle of the back or the awn reduced toa subapical mucro; paleas that are tightly enclosed by the margins of the lemma; linear lodicules; and an androecium composed of two stamens. In contrast, Trisetum has het- eromorphic, ovate-lanceolate or oblanceolate, 1-5-nerved glumes |first glume 1(-3)-nerved, second glume 3(-5)-nerved|]; lemmas with 2 to 4 short setae at the apex due to the prolon- gation of the nerves and a dorsal awn born on the upper third of the lemma, rarely lo- cated near the middle of the back; paleas that are free from the margins of the lemma; lodicules with 2- or 3-lobed apices; and an androecium with three stamens (Finot et al. SIDA 22(2): 895 — 903. 2006 896 BRIT.ORG/SIDA 22(2) 2004, 2005a, 2005b). Peyritschia includes perennial herbs with flat leaf blades, mem- branous ligules, narrow to contracted-spiciform or lax and somewhat open panicles, 2(- 3)-flowered spikelets, rachillas disarticulating above the glumes and between the florets, glabrous ovaries, and caryopses with liquid endosperm. Previous studies showed that the micromorphology of the lemmatal epidermis has good characters to distinguish Trisetum from Peyritschia. Most species of Peyritschia lack prickles hairs (present in all species of Trisetum), although, if present, they are restricted to the keel or the apex of the lemma in P. deyeuxioides and P pinetorum. All species of Peyritschia have bordered hooks alter- nating with epidermal long cells (hooks not bordered, nor alternating with long cells in Trisetum) and lack macrohairs (macrohairs are present in some species of Trisetum)[Finot et al. 2006]. These characteristics along with the salient features of the spikelet, were con- sidered important for resurrecting the genus by Finot (2003a) who transferred three ad- ditional species into Peyritschia: P. conferta (Pilg.) Finot (=Trisetum confertum Pilg.) from Bolivia, Ecuador and Venezuela; P deyeuxioides (Kunth) Finot (=T. deyeuxioides Kunth) widely distributed in México, and extending into Central and South America (Colombia, Costa Rica, Ecuador, El Salvador, Guatemala, Honduras, Nicaragua, Panama and Venezu- ela); and P. humilis (Louis-Marie) Finot (=T. humile Louis-Marie), an endemic in México (Finot et al. 2004). Trisetum confertum was originally described from an Ecuadorian specimen (Crescit in Prov. Imbabura, ad Loma de Canaballa et locis vicinis; alt. 2100-2300 m; A. Sttibel 152: holotype, B; isotypes, US-81771 ex B!, CONC fragm. and photo ex B!). Hitchcock (1927) cited this species as occurring in Cochabamba, Bolivia. Later, Valencia (1941) moved the species to Deschampsia[D. conferta (Pilg.) Valencia]. However, Parodi (1949) and Chiapella (2000) excluded it from Deschampsia. More recently, two species, Irisetum de yeuxioidesand T. humilis, were transferred in Peyritschia, based on spikelet and floret characteristics (Finot 2003a; Finot et al. 2004). Three of the five currently recognized aps o oe il Sot id are found in Guatemala: P. koelerioides (the type species of the g thern México; P. pringlei also present in México, Costa Rica, Venezuela, and Ecuador; and P. deyeuxioides ranging from southern México to Ecuador (Pohl & Davidse 1994; Espejo-Serna et al. 2000; Finot 2003a; Finot et al. 2004). The following seven species of Trisetum have been mentioned as occurring in Guate- mala: I’ angustum Swallen from San Marcos near Volcan Tajumulco, also extending into southern México; T. pinetorum Swallen here transferred to Peyritschia, from Quetzaltenango near Volcan Santo Tomas, and now known from México; T. viride (Kunth) Kunth (syn. T. altum Swallen) from El Progreso also widespread in México; T. irazuense (Kuntze) Hitche. from Huehuetenango near Tunima, mentioned for the first time by Finot et al. (2004), also from Costa Rica, Honduras, Panama, Colombia, Venezuela, Ecuador, and Peru; I. rosei Merr. from Huehuetenango from near Volcan Santa Maria, also extend- ing into México; T. pringlei (Scribn. ex Beal) Hitchc. from Huehuetenango and Totonicapan, also found in México, Costa Rica, and Panama; and T. spicatum (L.) K. Richt. with a wide distribution in the Americas (Swallen 1955; Pohl 1980; McVaugh 1983; Hernandez-Torres 1988; Pohl & Davidse 1994; Finot 2003b, Finot et al. 2004, 2005a, 2005b). In Ecuador, the following four species of Trisetum have been reported: T. andinum Benth., endemic to Ecuador; T. spicatum and T. oreophilum Louis-Marie var. oreophilum ranging from Ecuador to Pert and Bolivia (Finot et al. 2005b); and T. howellii here trans- FINOT ET AL., NEW COMBINATIONS IN PEYRITSCHIA 897 ferred to Peyritschia. Species of Peyritschia found in Ecuador are: P deyeuxioides and P pringlei (Finot 2003a). Preliminar Vi | using Mor phological characters within Trisetum s.l. (including Trisetum. s.s., Peyritschia, and Sphenopholis) [Finot 2004], depicts a well- supported clad (bootstrap of 95 %) that includes Peyritschia koelerioides, P pringlei, P deyeuxioides, P. conferta, and T. howellii Hitchc. This clade is supported by three syna- pomorphies: terete lemmas in lateral view, two stamens, and ornamentation of the lemma. In this paper we make two new combinations, Peyritschia pinetorum(Swallen) Finot & PM. Peterson, from Volcans Atitlan and Santo Tomas, Guatemala, and P howellii (Hitchce.) Finot & PM. Peterson, from Galapagos, Ecuador. In addition, we include a key to distinguish all seven species, a table comparing their salient morphological features, and a new illustration, description, and SEM photo of the lemmatal surface of P. pinetorum. Peyritschia howellii (Hitchc.) Finot & PM. Peterson, comb. nov. (Table 1). Trisetum howellii Hitchc., Proc. Calif. Acad. Sci. ser. 4, 21(24):296. 1935. Type: ECUADOR. GALAPAGOS ISLANDS: Indefatigable Island, Mt. Crocker, 9 May 1932, J.T. Howell 9208 (HOLOTYPE: CAS-211262!: ISOTYPE: US-1611545)). Perennials; culms 35-60 cm tall, weak; internodes glabrous, 5-6 nodes per culm, nodes glabrous. Leaf sheaths shorter than the internodes, glabrous, striate; ligules 0.5-1.5(-2) mim long, apex obtuse, not ciliate ti denticulate, glabrous; blades 8-15cm x 2mm, flat, glabrous, smooth, margins sometimes scabrous. Panicles 8-18 cm long, (0.5-)10-20 mm wide, contracted, narrow, lax, somewhat open, exserted or sub-included in the up- per sheath; rachis and pedicels glabrous; spikelets 4.5-5.5 mm long, (1-)3-flowered; rachillas 1.5 mm long, copiously pilose, the hairs ca. 1.5 mm long, as long as the rachilla: glumes almost covering the florets, isomorphic, linear to linear-lanceolate, narrow, some- what scabrous on the keel, the margins hyaline, apex acute, purplish; first glumes 4.3-4.7 x 0.3mm, l-nerved; second glumes 4.7-5.5 x 0.4 mm, 1-3-nerved; first lemmas 4-4.5 = 0.5 mm, linear-cylindrical to linear-lanceolate, glabrous, smooth, or scaberulous only towards the apex, purple towards the apex, rounded on the back, the margins enclosing the palea, 5-nerved, awned, the awn 5.5-7 mm long, twisted, geniculate, somewhat scabrous, borne on the upper third of the lemma, at 1-1.5 mm below the apex, the apex bidentate, toothed, acute with the intermediate and marginal nerves conspicuous towards the apex; callus obtuse, with stiff hairs, the hairs 0.5 mm long; paleas 3-4.3 mm long, shorter than the lemmas, 2-nerved, the nerves scabrous owas the apex, apex acute, ale ovary gla- brous; lodicules 0.6 mm long, linear, entire; stamens 2, anthers about 0.8 mm long. Cary- opses 2 mm long, glabrous; endosperm liquid. Phenology.—Flowering from September to January. Matures fruits are found from February to March. Distribution and habitat.—Peyritschia howellii is an endemic species from Ecuador. Most of the studied collections come from Isla Santa Cruz (Indefatigable), Galapagos, grow- ing between 480-1000 m; Jeppesen 76 (AAU) is the first collection of the species made from Isla San Cristobal (Chatham). Comments.—Peyritschia howellii was included by Valencia et al. (2000) in the Red Book of the endemic plants of Ecuador (as Trisetum howellii). Additional ined. ECUADOR. Galapagos Islands: Isla Santa Cruz, in area that includes W slope of Media Luna, E slope of Cerro de los Caminos and flat plain between, 4 Sep 1975, D.A. & D.B. Clark 447 (AAU); Puntudo, 17 Mar 1977, A. & H. Adsersen 1413 (QC - ca. uN m, 8 Nov 1966, U. & I. Eliasson 557 (S); sur de Mt. Crocker, 710 m,17 Oct 1966, U.& I. Eliasson 284(S) Bella Vista, 480 m, 19 Oct 1966, U. & I. Eliasson 356 (S); no date, Fagerlind & Wibon 2957 (S); no date, Paver lind & Wibon 3320(S); highest mountain top, 860 m, 23 May 868 Taste 1. Salient morphological characteristics to distinguish among the seven species of Peyritschia. P.conferta P. deyeuxioides P. howellii P. humilis P. pinetorum P. koelerioides P. pringlei Panicle shape narrow, contracted — lax, somewhat narrow, narrow, contracted narrow,contracted — spiciform narrow, open and nodding somewhat open somewhat open Number of florets 2 2073 (1-)3 2 2 2or3 per spikelet Spikelet, length (mm) 5.0-55 4.5-8.0 4.5-5.5 -7 - 5 Glumes, shape oval-lanceolate linear linear to linear- lanceolate lanceolate to lanceolate to lanceolate lanceolate ovate-lanceolate ovate-lanceolate Glumes, size verses longer equal, shorter or equal longer slightly shorter longer as long as the spikelet longer florets or longer Glumes, length (mm) 5.2-5.5 (3.5-)4-5.5 (-8) 4.3-5.5 4 45-6 3-5 Lemma, length (mm) 3.5-5.0 4-6 4-45 3-3.5 5-6 2.5-4 3-43 (lower floret) Lemma, apex bilobed, lobes bidentate, toothed, bilobed, lobes bifid, toothed, bilobed, lobes bilobed, lobes obtuse Rachilla,indument sparsely pilose Awn, position basal Awn, shape geniculate, twisted Awn, length (mm) 6-7 Callus indument short pubescent entire or bilobed to bidentate copiously pilose median geniculate, twisted 4.5-8(-12) short pubescent acute copiously pilose upper third geniculate, twisted short pilose obtuse sparsely pilose middle or lower third geniculate, twisted sparsely pilose acute copiously pilose pubescent median geniculate, twisted 7-12 pilose btuse glabrous to sparsely apical if present (as a short mucro) straight or absent glabrous to short pubescent obtuse sparsely pilose basal geniculate, twisted short pubescent (Z)% vals/OYO'LINA FINOT ET AL., NEW COMBINATIONS IN PEYRITSCHIA 899 1959, Harling 5764 (S), 1850 ft, 27 Jul 1974, van der Werff 1330 (S); Mt. Crocker, 840 m, 18 Feb 1967, Wiggins & Porter 652(S); fern-sedge zone along trail to Mt. Crocker, 500 m, 6 Feb 1964, Wiggins 18577 (S); Isla San Cristobal, El Junco, 650 m, 28 Feb 1976, Jeppesen 76 (AAU). aa L Septet (Swallen) Finot & P.M. Peterson, comb. nov. (Fig. 1, Table 1). Trisetum um Swallen, Phytologia 4:424. 1953. Type: GUATEMALA. QUEZALTENANGO: Volcan Santo Tom on pine- ines clad slope, 2500-3700 m, 22 Jan 1940, ].A. Steyermark 34824 (HOLOTYPE: F-1048257! ISOTYPES: MO fragm. ex FI, US-2240525 ex Fl, US-2236478 fragm. ex F)). Perennials; culms 33-70 cm tall, weak, internodes glabrous, 3-4 nodes per culm, nodes glabrous. Leaf sheaths shorter than the internodes, glabrous, striate; ligules 1.5-3 mm long, truncate, dentate-laciniate, glabrous; blades 5-10 cm long, 1-1.5(-2) mm wide, flat, gla- brous above and below. Panicles 5-15 cm long, 5-10 mm wide, contracted, narrow, some- what eS somewhat interrupted, exserted or sub-included in the upper leaf sheath; ra- chis g] Is scabrous or glabrous. Spikelets 5-7 mm long, 2-flowered; rachillas ca. 1 mm lene. copiously pilose, the Haus about 1-2 mm long; glumes slightly shorter than the florets, sub-isomor phic anceolate, the keel smooth to slightly scabrous towards the apex, apex acute; first glumes 4.5-6 x 0.7-0.8 mm, a little shorter and narrower than the second glumes, l-nerved; second glumes 5-6 x 0.7-0.8 mm, l-or 3- nerved; first lemma about 5-6 mm long, lemmas glabrous, awned, rounded on the back with involute margins on the lower 1/2 enclosing the palea, the awn 7-12 mm long, twisted, geniculate, and inserted on the median portion of the back, the apex deeply bifid with acute teeth, each tooth with two short awns derived from the prolongation of the mar- ginal and intermediate nerves; callus obtuse, with short hairs, the hairs ca. 1 mm long; paleas about 3.5-4 mm long, shorter than the lemmas, 2-nerved, the nerves scabrous to- wards the apex, apex acute; lodicules about 0.5 mm long, linear, sometimes with a little lateral lobe near the median portion, the apex subacute; stamens 2, anthers 0.9-1.5 mm long; eel glabrous. Caryopses not seen. mma micromorphology. Long cells rectangular, 3-12 times longer than wide; anti- ae ents parallel and highly undulate, the undulations U- or V-shaped; periclinal walls flat; silica cells about 10 um in diameter, short, elliptical; stomata absent; prickle hairs about +5 um long, ovate-elliptical, restricted to the keel and with a short apical barb; long cells about 10-12.5 um in diameter, alternating with bordered hooks; macrohairs absent (Fig. 2.). Phenology.—Flowering in January. Distribution and habitat.—Known only from southwesterm Guatemala in Departa- mentos Quetzaltenango and Solola. Apparently, P pinetorum is restricted to volcanic soils since the only two localities are from higher elevations (2500-3700 m) on slopes of Volcan Atitlan and Santo Tomas. Comments.—This new combination can be distinguished from all other known spe- cies of Peyritschia by having lemma apices that are deeply bifid, and these apices end in four hyaline setae derived from the apical extension of the marginal and intermediate nerves (Table 1). The presence of these setae suggests that P. pinetorum, ina phylogentic sense, lies very near Trisetum. Nevertheless, because of the shape of the lodicules (entire in Peyritschia and bilobed or toothed in Trisetum), the lemmas rounded on the back (ver- sus keeled in Trisetum), the awn inserted on the median portion of the lemma [inserted on upper 1/3 in Trisetum except in Trisetum subgen. Deschampsioideum (Louis-Marie) Finot], paleas that are tightly enclosed by the margins of the lemma (gaping in Trisetum — ite eee. oe KK: eX SSK ‘es a A. Habit. B. Sheath liaule y eee d t BRIT.ORG/SIDA 22(2) f the blade. C. Spikelet. D. First floret. E. Palea, dorsal view.F. Palea F 4 Y Fic. 1. Pe is, dorsal view. J. 0 Lf é T és é F is, ventral view J b ventral view. G. Rachilla. H. Lodicules.1.0 P FINOT ET AL., NEW COMBINATIONS IN PEYRITSCHIA 901 # Fic ») CEM nhat bh af tho | | ef ANN\ .£ D reps Pe) H rr {C+ L 2AQDVA\ ch 2 / \ nating with long cells. Scale bar = 55m. and not tightly enclosed by the margins), an androecium with two stamens (3 stamens in Trisetum), and lemmatal epidermis with bordered hooks alternating with long cells (verses epidermis without bordered hooks, the hooks not alternating with long cells in Trisetum) [Fig. 2.], we place the new species in the genus icibeciaee [Or i 5 o) Sy & a a pe am aN wr e; Gi oO Co Z, Oo ha 3 No) ron ee = => a wi io) Additional specimen examined. GUATEMALA. Solola: Volcan Atitl: (3540 m), 23 Jan 1907, W.A. Kellerman s.n. (US-2181368). KEY TO THE SPECIES OF PEYRITSCHIA . Lemma without a dorsal awn, apex muticous or with a subapical mucro P. koelerioides . Lemma with a dorsal awn, the awn geniculate and extended beyond the glumes 2. Rachilla copiously pilose, the hairs 1-3 mm long; awn borne on the middle or the upper third of the back of the lemma 3. Lemma with the apex deeply bifid; panicles 5-15 cm long, contracted, narrow P. pinetorum 902 BRIT.ORG/SIDA 22(2) 3, Lemma with the apex entire or shortly bidentate or bilobate; panicles 8-35 cm long, lax, somewhat open 4. Rachillas with hairs 2-3 mm long; glumes linear 5.2-5.5 mm long; awn borne on the middle portion of the lemma P. deyeuxioides 4. Rachillas with hairs about 1.5 mm long; glumes linear to linear-lanceolate 4.3-5.5 mm long; awn borne on the upper third of the lemma P. howellii 2. Rachilla pubescent, the hairs 0.2-0.8 mm long; awn borne near the base of the lemma. 5. Culms 5-12 cm tall; leaf blades 2.3-3.5 cm long; panicles about 2.5 cm long, 0.5 cm wide P. humilis 5. Culms 20-200 cm tall;leaf blades 5-15 cm long; panicles 5-20 cm long, 1-4(-4) cm wide. 6. Spikelets 2- or 3-flowered, 4-5.5 mm long; glumes 4-5.3 mm long; lemma 3-4.3 mm long P. pringlei 6. Spikelets 2-flowered, 5—5.5 mm long; glumes 5.2-5.5 mm long; lemma 3.5-5 mm long P.conferta ACKNOWLEDGMENTS We thank the Directors and Curators of the following herbaria: AAU, BA, BAA, BAF, C, CR, CONC, F LP, MERL, P PR, QCA, SGO, §S, SI, US. The first author gratefully acknowl- edges the Myndel Botanical Foundation for a fellowship to study types of Trisetum and allied genera at Paris (P) and Stockholm (S); a grant from MECESUP-Universidad de Concepcion UCO-9906/01 to study Trisetum at the Smithsonian Institution (US) in Wash- ington, DC, U.S.A and Instituto de Botanica, Darwinion (SI) in Buenos Aires, Argentina; Projects DIUC 204.121.009-L0 and 205.111.047-1.0 for financial support; Oscar Matthei and Clodomiro Marticorena for directing my Ph.D. studies, and Susan J. Pennington (US) for helping me during my stay at the Smithsonian. In addition we thank Vladimiro Dudas (SD for the beautiful illustration and Yolanda Herrera Arrieta and Ana Maria Planchuelo for reviewing the manuscript. This paper is part of the doctoral thesis of the first author at Departamento de Botanica, Universidad de Concepcion, Concepcion, Chile. REFERENCES Cuiapetta, J. 2000. Taxonomy, morphology, phylogeny and biogeography of Deschampsia (Poaceae- Aveneae). Universidad San Carlos de Bariloche, Doctoral thesis. Espeso-Serna, A., A.R. Lopez-Ferrari, and J. Vacpes-Reyna. 2000. Poaceae. In: A. Espejo-Serna and A.R.L6pez- Ferrari (eds.). Las Monocotiledéneas Mexicanas. Poaceae Barnhart. Una Sinopsis Floristica. Partes IX—XI. Pp. 8-236. Finor,V.L. 2003a. Peyritschia.P.478.In: Soreng, R.J.,PM. Peterson, G. Davidse, EJ. Judziewicz, F.O.Zuloaga TS. Filgueiras, and O. Morrone. 2003. Catalogue of New World grasses (Poaceae): IV. subfamily Pooideae. Contr. U.S. Natl. Herb. 48:1—730. Finor, V.L. 2003b. Trisetum. In: Soreng, R.J., PM. Peterson, G. Davidse, E.J. Judziewicz, F.O. Zuloaga, TS. Filqueiras,and O.Morrone. 2003.Catalogue of New World grasses (Poaceae): IV. subfamily Pooideae. Contr. U.S. Natl. Herb. 48:659-676. Finot, V.L. 2004. Sistematica del género Trisetum Pers. (Poaceae: Aveneae) en América. Una aproximacidn filogenética basada en datos morfoldgicos. Universidad de Concepcion, Doctoral thesis. Finor, V.L., PM. Peterson, R.J. Sorenc, and F. Zutoaca. 2004. A revision of Trisetum, Peyritschia, and Sphenopholis (Poaceae: Pooideae: Aveninae) in México and Central America. Ann. Missouri Bot. Gard. 91:1—30. Finor, V.L., PM. Peterson, R.J. Sorene, and F.O. ZuLoaca. 2005a. A Revision of Trisetum and Graphephorum (Poaceae: Pooideae: Aveninae) in North America. Sida 21:1419-1453. FINOT ET AL., NEW COMBINATIONS IN PEYRITSCHIA 903 Finot, V.L., RM. Peterson, F.O. ZULOAGA, R.J. SoreNG, and O. MattHel. 2005b. A Revision of Trisetum (Poaceae: Pooideae: Aveninae) in South America. Ann. Missouri Bot. Gard. 92:533-568. Fino, V.L., C.M. Baeza, and O. MattHel. 2006. Micromorfologia de la epidermis de la lemma de Trisetum y géneros afines (Poaceae: Pooideae). Darwiniana.|n press. Fournier, E. 1886. Mexicanas plantas Il. Gramineae 1-xix, 1-160, pl. 1-10. Paris. HerRNANDEZ-Torees, |. and S.D. Koch. 1987. The status of the genus Peyritschia (Gramineae: Pooideae). Phytologia 61:453-455, HerNANnbez- Torres, !.and S.D. Kock. 1988.Revisidn taxondmica del género Trisetum (Gramineae: Pooideae) en México. Agrociencia 71:71-102. HitcHcock, A.S. 1927. The grasses of Ecuador, Pert, and Bolivia. Contr. U.S. Natl. Herb. 24:291-556. HitcHcock, A.S. 1935.The Templeton Crocker expedition of the California Academy of Sciences, 1932, No. 24,new species of grasses from the Galapagos and the Revillagigedo Islands. Proc. Calif. Acad. Nat. Sci.,ser.4 21:25-300. Koch, S.D. 1979. The relationships of three Mexican Aveneae and some new characters for distin- guishing Deschampsia and Trisetum (Gramineae). Taxon 28: 225-235. McVauch, R. 1983. Gramineae. Flora Novo-Galiciana 14:1-436. Parooi, L.R. 1949. Las gramineas sudamericanas del género Deschampsia. Darwiniana 8:415-475. PoHt. RW. 1980. Gramineae. In:W. Burger, ed. Flora Costaricensis. Fieldiana Bot. n.s. 4:1-608. Pout, R.W.and G.Daviose. 1994. Trisetum Pers. ln:G. Davidse, M. Sousa, and A.O.Chater (eds). Fl. Mesoamer. G253=23 3. Swaiten, J.R. 1955. Gramineae. In: PC. Standley and J.A. Steyermark, eds. Flora of Guatemala, Part Il: Grasses of Guatemala. Fieldiana Bot. 24:1-390 VALENCIA, J.1. 1941. Especies criticas de Trisetum que deben pasar al género Deschampsia. Revista Ar- gent. Agron. 8:1 22-130. VALENCIA, R., N. PITMAN, S. LEON-YANEZ, and P.M. Jorcensen (eds.). 2000. Libro Rojo de las Plantas Endémicas del Ecuador 2000. Pontificia Universidad Catolica del Ecuador, Quito. 904 BRIT.ORG/SIDA 22(2) Book REVIEW Marr Waite. 2000. Prairie Time: A Blackland Portrait. (ISBN 1-58544-501-0, hbk.). Texas A&M University Press, 4354 TAMU, College Station, TX 77843-4354, U.S.A. (Orders: 979-458-3982, 979-847-8752 fax; http://www.tamu.edu/upress/). $19.95, 272 pp., b/w figures, 1 map, index, 51/2" x 81/2" Texas-prairie enthusiasts will relish author Matt White’s new memoir of the Blackland Prairie called Prairie Time: A Blackland Portrait. His new book intertwines the author's interests in the Blackland Prairie with per- fek sonal touches of family, natura, and land histories, with prairie ecology—all with the background of his senti- mental memories of the Blackland Prairie region. The book is a collection of Matt White's experiences in exploring prairie remnants; among these are his ay interactions with land stewards and also stories of unfortunate losses of prairie remnants to various causes. Throughout the book there is information on animals that have inhabited the prairie, such as the prairie chicken and wild turkeys, as wel as information on the importance of prairie as wildlife habitat. Prairie ecology is dotted through the text as well. For eXé ae, yowll find information on the importance of fire to the prairie system, the for t] | ch . t stics of prairie soils, flora indicative of prairie areas, and flora invasive to the prairie. A discussion of historical land uses and views of the prairie over time is also included. The author's deep interest and enjoyment of the prairie is delicately sprinkled throughout the text. Take a sentimental journey with author Matt White and experience his excitement in a aa ing I ee ad | Prairie remnants in Texas. White shares both joyous experiences of preserving remnants an feats in loss of Blackland Prairie areas. Readers with an interest in learning more about the Blackland Paine should seek out this very readable book—it includes many details on the natural history and ecology of the Black- land Prairie within the author's stories. es Luckeydoo, Herbarium, Berane al Research iia of Texas, 509 Pecan Street, Fort Worth, 1X 76102-4060, U.S. SIDA 22(2): 904. 2006 POA MATRI-OCCIDENTALIS (POACEAE: POOIDEAE: POEAE: POINAE), A NEW SPECIES FROM MEXICO Paul M. Peterson and Robert J. Soreng Yolanda Herrera Arrieta Department of Botany CHDIR- Dgo, ae ee Wen im elinee History Instituto Polit nstitution Sigma s.n. Fracc. 20 de Noviembre II ee ee DC 20013-7012, U.S.A. 34720D Durango, MEXICO peterson@si.edu; sorengr@si.edu yherrera@ipn.mx ABSTRACT Poa matri-occidentalis PM. Peterson & Soreng, sp nov, is described and illustrated. Poa matri-occidentalis subsp. matri-occidentalis is known from steep rocky slopes of Cerro Gordo, Durango, and Poa matri-occidentalis subsp. mohinorensis Soreng & P.M. Peterson, subsp. nov. is known from rocky cliffs of Cerro Mohinora, Chihuahua. The new species is morphologically similar to Poa kelloggii but differs by having leaf sheaths that are closed near the base 2/3 to 4/5 their length, entire ligules that are 3.5-6 mm long, nodding panicles, ovate lemmas with obtuse to acute apices, and non-visible rachilla internodes. A key distinguishing 11 species now reported from northern MAp 7 ] iq ] Is PlOVIGEC, RESUMEN s] Pay i bs Se Gescue e ilustra Poa matri- uceidentalis P.M. Peterson & Soreng, sp. nov. Poa matri -erro Gordo, Durango, y Poa matri- ‘cccldentalts subsp. mohinorensis Soreng & PM. Peterson, subsp. nov. se encuentra en la zona de laderas rocosos del Ce Mohinora, Chihuahua. La nueva especie es morfol6gicamente similar a Poa kelloggii pero difiere de ésta ce por tener las vainas de las hojas cerradas en la parte inferior de la base de 2/3 a 4/5 de su longitud, por tener igulas enteras de 3.5-6 mm de largo, paniculas recurvadas, lemas ovadas con apices agudos a obtusos y raquilla con entrenudos no-visibles. Se proporciona una clave para distinguir las 11 especies reportadas actualmente del — norte de México. Pod L. is the largest genus of grasses, including some 500-575 species that occur ina wide range of habitats from throughout the World (Soreng 1990; Gillespie & Soreng 2005). The genus is characterized by having rather small, multi-flowered spikelets, lemmas that are keeled, unawned, usually 5-nerved, commonly with web-like hairs emerging from the dorsal side of the callus, caryopses that are firm with lipid and a short hilum, lodi- cules that are broadly lanceolate with a lateral lobe, leaf sheaths closed above the base more than 1/20 the entire length, leaf blades that generally have two rows of bulliform cells (one on either side of the midnerve, these appearing as railroad tracks) and no addi- tional rows of bulliform cells, and blades commonly with naviculate (boat-shaped) api- ces (Soreng, in press). Poa has been divided into subgenera and sections and subsections for North America (Soreng 1998; Soreng in press), the New World (Soreng et al. 2003a), and the World (Gillespie et al. 2006; Zhu et al. 2006). Northern México (excluding Baja California and Baja California Sur) is not thought to be especially rich in Poa species. The following 11 species have been reported from the states of Aguascalientes (Ags), Chihuahua (Chih), Coahuila (Coah), Durango (Dur), Jalisco Jal), Nuevo Leon (NL), San Luis Potosi (SLP), Sonora (Son), Tamaulipas (Tamp), and Zacatecas (Zac): Pod annua L. (Ags, Chih, Coah, Dur, Jal, NL, SLP, Tamp, Zac), P. bigelovii Vasey & Scribn. (Chih, Coah, NL, Son), P. compressa L. (Coah), P. conglomerata Rupr. (= P. scaberula Hook. f.)[Coah is an error since a specimen (J.A. Garcia 49) cited in Beetle et al. SIDA 22(2): 905 — 914. 2006 906 BRIT.ORG/SIDA 22(2) (1999) is = P. pratensis LJ, P fendleriana (Steud.) Vasey lincludes P fendleriana subsp. albescens (Hitche.) Soreng](Chih, Coah, Son, Zac), P infirma Kunth (listed as synonym of P.annua and probably with similar distribution), P mulleri Swallen (NL), P orizabensis Hitche. (NL possibly an error for CH. & M.T. Mueller 1248 from Cerro Potosi as deter- mined by A.A. Beetle =P. mulleri!, SLP-not yet verified by us from there), P. pratensis (Chih, Coah, NL), P. ruprechtii Peyr. includes P. sharpii Swallen as a synonym] (Coah, NL,), and P. strictiramea Hitche. (Chih, Coah, Dur, NL, Zac) [Beetle et al. 1999; Espejo Serna et al. 2000; Hitchcock 1913]. Of the 10 species known from the region, P annua, P. compressa, P. infirma, and P. pratensis are introduced, and P. bigelovii, P fendleriana, P. mulleri, P. orizabensis, P ruprechtii, and P. strictiramea are native. The annuals, P annua and P infirma have been placed in Poa sect. Micrantherae Stapf, and P. bigelovii was placed in Poa sect. Homalopoa Dumort. (Soreng et al. 2003a). Of the remaining natives, P mulleri, P orizabensis, P. ruprechtii, and P. strictiramea have been placed in Poa sect. Homalopoa s.l, and P.fendleriana was placed in Poa sect. Madropoa Soreng (Soreng et al. 2003a). While making determinations of material in the field using Las Gramineas de Durango (Herrera Arrieta 2001) and then checking the herbarium at Centro Interdiciplinario de Investigacion para el Desarrollo Integral Regional (CIIDIR) in Durango, the first author recognized the unique features of a recent collection from Cerro Gordo. Upon further inspection of collections on loan from various herbaria for a com- plete revision of Poa in México, the second author discovered another, generally pubes- cent form, occurred near Cerro Mohinora, Chihuahua. The third author then contrib- uted the anatomical portion. We describe these two new forms as a new species of Poa with two subspecies. The new species with two subspecies is clearly aligned within sub- family Pooideae, tribe Poaeae, and subtribe Poinae (Soreng et al. 2003a, 2003b, 2005). Poa matri-occidentalis P.M. Peterson & Soreng, sp. nov. (Figs. la-c; 2¢e-1). Type: MEXICO. DURANGO: Sierra Madre Occidental, SW slope of Cerro Gordo just below twin rock outcrops (23°12'32.5"N- 104°56'54.1"W), 3130-3200 m, 26 Sep 2005, P.M. Peterson 19145 & F Sdnchez Alvarado (HOLOTYPE: US! ISOTYPES CHDIR!, MEXUN). A Poa kelloggii Vasey vaginibus connatus 2/3-4/5 longe lateque, ligulis 3.5-6 mm longis integris, paniculus nutans, lematibus ovatis apicibus obrusus ad acutus, rachillibus no visibilis, recedit. Loosely caespitose and rhizomatous perennials with intra and extravaginal basal branch- ing. Culms 45-80 cm tall, solitary to several, erect or bases slightly decumbent, terete or weakly compressed; nodes 2-4, terete, glabrous and smooth, 1-3 exserted. Leaf sheaths mostly 3-14 cm long, closed about 2/3 to 4/5 their length, compressed, glabrous and smooth, or sometimes the lower ones retrorsely scabrous to puberulent and with ciliate collars, most sheaths 0.4-L.1 times as long as their blades; collars smooth or with a few hooks, glabrous or ciliate; ligules 3.5-6 mm long, membranous to hyaline, glabrous and smooth, or sometimes puberulent, apex obtuse to acute, entire; blades 2-6 mm wide, flat, abaxial nerves and margins lightly scabrous, adaxially smooth, glabrous throughout, apices narrowly prow-shaped; flag blades 12-22 cm long. Panicles 12-26 cm long, nod- ding, pyramidal, open, sparse with 24-85 spikelets, longest internodes 2.5-5.5 cm long; branches (1-)2(-3) at the lower nodes, longest 5.5-10 cm long bearing 3-15 spikelets, as- cending to spreading, lax, angled, the angles sparingly scaberulous. Spikelets 4-8 mm long, 1.8-2.7 mm wide, laterally compressed, greenish to stramineous, florets 2 or 3, per- fect, occasionally the upper floret is much reduced and sterile; lower rachilla internodes 1-2 mm long, usually not visible, glabrous, smooth; glumes equal to subequal, lanceolate, PETERSON ET AL., POA MATRI-OCCIDENTALIS, A NEW SPECIES FROM MEXICO = ee EF, = ga i o TL BAG Oye 4 Le {Dd Al Fic. 1. Poa matri-occidentalis subsp. matri-occi (Peterson 19145 & Sd and blade. Poa tri identali } ie + rar AFJEOM-N lo). A. Habit. B. Inflorescence. C Sheath, ligule, Id). D. Sheath ligule and blade BRIT.ORG/SIDA 22(2) 908 Poa matri-occidentalis subsp. matri- glume. F. Upper glume. G. Lemma. H. Palea, dorsal K. lodicules. L. Young caryopsis Id). A. Spikelet. B. Floret AA-ND do). C. Spikelet. D. Floret. E. Lower ATCO Fic. 2. Poa matri Al hb occidentalis (Peterson 19145 & Sa at view. I, Palea, ventral view.J. Stamens, ovary 7 J 7 PETERSON ET AL., POA MATRI-OCCIDENTALIS, A NEW SPECIES FROM MEXICO 909 scabrous along the upper keel, distinctly keeled, 3-nerved, the lateral nerves often only evident near base, scaberulous along the nerves, apex acute to acuminate, of ten appear- ing mucronate because the margins are often involute, margins hyaline; lower glumes 3- 5 mm long, lanceolate to ovate usually shorter than the upper; upper glumes 3.7-5.6 mm long, ovate to obovate, wider that the lower with more prominent nerves; calluses with cob-webby hairs, the hairs about 1/2 to 1/3 the length of the lemma; lowest lemmas 4.6- 6.3 mm long, ovate, distinctly keeled, surfaces between nerves finely muriculate (with minutely pointed, whitish bumps on the short cells) or sometimes densely scabrous, 5- nerved, the lateral nerves moderately prominent, keel and lateral nerves scabrous, keel and marginal nerves glabrous or sometimes puberulent below, margins hyaline, apex obtuse to acute; lowest paleas 4.4-6 mm long, usually a little shorter than the lemmas, strongly laterally compressed with a deep adaxial furrow between the nerves, muriculate, scabrous along the nerves; anthers 2-2.2 mm long, yellow; lodicules 2, membranous; ovary glabrous. Caryopses 2.6-2.9 mm long, fusiform, light brownish. Phenology.—Flowering August through October. Comments.—The new species can be distinguished from Poa kelloggii by having leaf sheaths that are closed near the base 2/3 to 4/5 their length, entire ligules that are 3.5-6 mm long, nodding panicles, ovate lemmas with obtuse to acute apices, and non-visible rachilla internodes. Poa matri-occidentalis subsp. matri-occidentalis (Figs. la—c; 2c-I; 3a-c; 4). Lower leaf sheaths glabrous and smooth; collars smooth or with a few hooks, glabrous: ligules glabrous and smooth. Lowest lemmas 4.7-5.4 mm long, surfaces between nerves finely muriculate (with minutely pointed, whitish-bumps on the short cells), keel and marginal nerves scabrous, glabrous throughout. Distribution and habitat—Known only from a single locality at Cerro Gordo, Durango, between 3130-3200 m where the species was found growing on steep, rocky, and grassy slopes beneath open forests of Pseudotsuga menziesii (Mirb.) Franco, Quercus sp., Pinus spp., with other associates, such as: Arctostaphylos pungens Kunth, Bromus carinatus Hook. & Arn., B. richardsonii Link, Aegopogon cenchroides Humb. & Bonpl. ex Willd., Trisetum viride (Kunth) Kunth, Festuca sp., Carex sp.,and Agrostis sp. Leaf anatomy.—Cross-sections were done by hand on re-hydrated material then mounted on temporary slides and stained with concentrated safranin. A diagram of a cross-section and abaxial leaf scrape were made with the aid of a camera lucida (Figs. aaa: Cross-section view (Fig. 3a-c.).—The lamina are V-shaped with primary, secondary, and tertiary vascular bundles of decreasing size. The primary vascular bundles are well differentiated into xylem with metaxylem, phloem, anda double bundle sheath (mestome and parenchyma bundle sheath). The ribs are rounded and furrows are <1/5 as deep as the width of the blade adaxially and abaxially. Abaxial projection of the midrib caused by inflated parenchyma cells and sclerenchyma is conspicuous. In half-the-width of the blade there is a single primary vascular bundle comprising the midvein; 3-4 additional primary vascular bundles; 3-4 secondary vascular bundles placed between consecutive tertiary vascular bundles; 7-8 tertiary vascular bundles placed between each consecu- tive primary and secondary vascular bundles; and two tertiary vascular bundles at the margin (i-iii-ii-iii-i-iii-ii-iii-i-iii-ii-iii-i-iii-ii-iii-iii). All vascular bundles are placed in the median layer of the blade and are round in outline. The xylem of the primary vascular 910 BRIT.ORG/SIDA 22(2) n £hiad + £D, tr). id tpt oy - 1D. OVAL &. CAnrt Af, din\ 7 adaxial 7 Scale bar = ‘ jie surface uppermost in all drawings. A. Primary vascular bundle. B. S Jary vascular bundle. C. T 50um. Fic. 4. Abaxial leaf epidermis of Poa matri-occidentalis subsp. matri-occidentalis (Peterson 19145 & Sanchez Alvarado) showing long cells, silica bodies (dark), cork cells (clear), and th t ta. Scale bar = bundles contains two or occasionally three metaxylem vessels adjacent to the phloem, that are larger than the parenchyma bundle sheath cells, and sometimes one or two pro- toxylem vessels are located adaxially to the phloem. The mestome, or inner sheath, is always present in primary and secondary vascular bundles surrounding the xylem and phloem. The Mestome is composed of small cells with thick walls and interrupted abaxially by sclerenchyma fibers forming a girder in primary and secondary bundles, except in the central primary vascular bundle where it is continuous surrounding the bundle and the sclerenchyma is separated from the bundle by colorless cells and cen- tered at the very bottom of a conic abaxial base. The parenchyma bundle sheath, or outer layer, contains large, thin-walled cells; 10-12 cells are present on the continuous adaxial half circle of the primary vascular bundle sheath; 6-8 cells are present in the secondary vascular bundles; and the number of cells varies between three and four on each side of the interrupted tertiary vascular bundles. Chlorenchyma cells surround the bundles and are conspicuously continuous between the vascular bundles in a loose arrangement. Five PETERSON AND SORENG, POA MATRI-OCCIDENTALIS, A NEW SPECIES FROM MEXICO 911 to eight sclerenchyma fibers are present adaxially in all bundles. Five to eight rows of sclerenchyma fibers form the abaxial girders of the primary and secondary vascular bundles, which widen near the epidermis and narrow toward the vascular bundle. Deep fan-shaped rows of bulliform cell are located adaxially near the central part of the blade between the central primary vascular bundle and two lateral tertiary vascular bundles: these are continuous to adaxial epidermal cell walls. Epidermis in abaxial view (Fig. 4).-Costal and intercostals zones are clearly distin- guishable. Three to six rows of short cells are found in the costal zone, each separated by a row of long cells. Irregularly-shaped silica bodies are alternate and adjacent to short cells of the costal zone. Cork cells are less frequent and the same shape as the silica bod- ies. Fifteen to seventeen rows of long cells are common in the intercostal zone. Long cells are 8-10 times longer than wide with wavy cell walls. No papillae or hairs are present. Low dome-shaped stomata are infrequent and found in single row. Parallel subsidiary cells are rounded in outline. A few antrorse bristles are present at foliar margin. Poa matri- cece subsp. mohinorensis Soreng & PM. Peterson, subsp. nov. (Figs. 1d; 2a, ).T EXICO. CHIHUAHUA: Municipio de Guadalupe y Calvo, Sierra Nees ee east Nside of Cerro 1 tall, inal gynostegial corona o slightly vertically “lobed apieally (.e., each ridge Le rising vertically immediately before descending to the base); anther wings slightly shorter than the corpusculum; terminal appendage of stylar head conic (volcano-shaped; broader at the base than at the apex),0.9-1.2 mm tall M. pauciflora 2. Gynostegial corona (C(is)) lacking a hardening, lobe-like, translucent yellow-golden secre- tion in the interstaminal position; interstaminal gynostegial corona (Ci) lobed (or if not, then the stylar head with a knob-shaped terminal appendage); anther wings horizontally conver- gent at least at the apex or parallel; stylar head exhibiting a terminal appendage 4. Calyx colleters absent; corolla lobe bases each with two round, distinct, lateral thicken- ings, basal lobe surfaces concave ("scooped-out’); interstaminal gynostegial corona (Ci) unlobed, staminal gynostegial corona (Cs) apically ungrooved, unlobed laterally, scallop- shaped in broad outline, margins entire or somewhat undulating; anther wings parallel; terminal appendage of stylar head capitate (e.g., reminiscent of a round door knob), to 1.1 mm tall M. torulosa 4.Calyx colleters 1 per sinus;corolla lobe bases lacking round lateral thickenings, lobes plane; interstaminal gynostegial corona (Ci) shallowly to broadly lobed, staminal gynostegial corona (Cs) apically grooved or laterally lobed, not scallop-shaped; anther wings conver- gent at least apically; terminal appendage of stylar head clavate or narrowly elongate- conic (obelisk-shaped), > 1.8 mm tall. 5. Interstaminal gynostegial corona (Ci) a large lobe, staminal gynostegial corona (Cs) grooved apically, lacking two broad, lateral lobes; terminal appendage of stylar head clavate . corynephora 5. Interstaminal gynostegial corona (Ci) a shallow lobe, staminal agynostegial corona (Cs) apically not grooved, broadly lobed laterally to appear 3-lobed; terminal appendage of stylar head narrowly elongate-conic (obelisk-shaped) M. rhynchocephala Matelea corynephora Krings, sp. nov. (Fig. 1). Type: HAITE Ile la Tortue, la Vallée, top of Morne Bar- ranca, ca. 300 m, 21 Mar 1928 (£1), E.L. Ekman H9740 (HOLOTYPE: S!). Species nova Matelea Aubl. diftert a Matel ill ilol i i lol eee h Slender, woody vine. Latex unknown. Stems ca. 1.8 mm diam., pubescent, glandular-capi- tate trichomes throughout, ca. 0.09 mm long, eglandular sharp trichomes mostly in 2 lines, one on each side of the stem, retrorse, ca. 0.4 mm long. Leaves opposite, simple, blades oblong to linear-oblong, (7.2-)12-25 « (3.6-)5.2-8.6 mm, apex obtuse, apiculate, base broadly cuneate to rounded, margins entire, adaxial surface dark green, glossy on drying, glabrous or minutely pubescent along midvein with short glandular-capitate trichomes, abaxial surface pubescent along midvein and sometime secondary veins, trichomes mostly glandular-capitate, eglandular sharp trichomes few and scattered, colleters 2; petioles 3.8-7 = ca. 0.6 mm, glandular-capitate trichomes dense, ca. 0.1 mm long, eglandular sharp trichomes dense, antrorse, ca. 0.2 mm long, both trichome types found throughout, but the longer sharp trichomes tending to be slightly denser along the adaxial KRINGS, NOVELTIES IN MATELEA FROM HISPANIOLA 943 po Fic. 1. Motel } a A hah; 1 dotail nflaat-D ),aW = anther win gs; Haloh ! c Fee, eee tanial Wee: dane oh aa corolla lobes , CO = cOrpusculum,; © gy g rf pollinium, t PI g y head. chomes ca. 0.07 mm long, eglandular sharp trichomes ca. 0.2 mm long, predominantly antrorse, some retrorse, sparser than glandular-capitate trichomes; pedicels 4.5-6.1 x 0.4 mm, densely pubescent, glandular-capitate trichomes ca. 0.07 mm long, eglandular sharp trichomes ca. 0.18 mm long, predominantly antrorse or antrorse appressed. Calyx 5-lobed, lobes lanceolate or lance-ovate, ca. 1.8 x 0.8 mm, apices obtuse, margins entire, scattered pubescent, glandular-capitate trichomes to 0.07 mm long, eglandular sharp trichomes to 0.2 mm long; colleters lacking. Corolla subrotate, bases subcampanulate, tube 0.9 x | mm, 5-lobed, lobes imbricate in bud (not dextrorse), linear-lanceolate, 4.5-5 x 1.3-16mm, apices obtuse, margins entire, marginally thickened at the base on each side, scooped- out and forming a distinct pit near the base, otherwise plane, adaxial surface papillate, abaxial surface glabrous or witha few, eglandular sharp trichomes ca. 0.2 mm long. Faucal annulus (corolline corona or Ca) indistinct. Gynostegial corona of fused staminal (Cs) and interstaminal (Ci) parts, Cs rising to meet the lower portion of the anther, rising seg- KRINGS, NOVELTIES IN MATELEA FROM HISPANIOLA 953 ment scallop-shaped in outline, ca. 0.5 mm tall, not lobed on either side, yet exhibiting a smaller segment immediately in front, margins somewhat undulating to entire, Ci unlobed; anther wings parallel, nearly touching, well-developed toward the center of the stylar head, ca. 0.4-0.5 mm long, the vertical segment immediately subtending the corpusculum ca. 0.2 mm long, slightly longer than the corpusculum or as long: laminar dorsal anther appendages (Cd) absent. Pollinarium: corpuscula dark reddish-brown, ca. 1.8-1.9 mm long; pollinia borne horizontally, ovate, ca. 0.37 X 0.25 mm. Stylar head with a terminal appendage, appendage capitate (“door knob-shaped”), ca. 1-L1 mm tall, apex ca. | mm broad. Follicles unknown. Distribution.—Apparently endemic to the Dominican Republic [known only from the typel. Etymology—The epithet refers to the capitate, door knob-like terminal appendage of the stylar head. Discussion.—The appendage is among the shortest of the four species bearing them in Hispaniola. Only Matelea corynephora (Fig. )) exhibits an appendage similarly broader at the apex than at the base, but the appendage is much longer than in M. torulosa and clavate. The two species also differ in a number of additional floral characteristics, such as the morphology of the gynostegial corona. The other appendaged species—M. pauciflora (Fig. 3) and M. rhynchocephala (Fig. 5)—exhibit terminal appendages that are narrower at the apex than at the base. ACKNOWLEDGMENTS [thank the curators and staff of the following herbaria for searching, or providing access or loans of their collections: B, BG, BH, BKL, BM, BOLO, BR, BREM, BSC, BUE.C, CGE, COLO, CR, DUKE, E, F FI, FLAS, FR, FIG, G, GH, GOET, H, HAC, HAJB, HBG, IA, IJ, ISC, JBSD, JE, K, LD, M, MICH, MIN, MO, MSU, NEU, NSW, NY, O, OXE P, PH, RSA, S, U, UC, UPS, US, USE TUR, WILLI, WU, Z. l also thank Roxana Yockteng for her kind assistance during, and subsequent to, a visit to P. The North Carolina State University Libraries Inter-Library Loan Service was instrumental in obtaining necessary titles not held in the collections. Alain Liogier and an anonymous reviewer provided thoughtful reviews of a previous version of the manuscript. REFERENCES Enpress, M.E. and P.V. Bruyns. 2000. A revised classification of the Apocynaceae, s.|. Bot. Rev.66:1-56. Krincs, A. 2005. Notes on the Matelea bayatensis-correllii-tigrina complex (Apocynaceae - Asclepiadoideae - Gonolobinae) in the Greater Antilles and Bahamas. Sida 21:1525-1533. Kunze, H. 1995. Floral morphology of some Gonolobeae (Asclepiadaceae). Bot. Jahrb. Syst. 117:211-238. Liebe, S. and H. Kunze. 1993. A descriptive system for corona analysis in Asclepiadaceae and Periplocaceae. Pl. Syst. Evol. 185:275-284. Liocier, H.A. 1994. La flora de la Espanola, VI. Publ. Univ. Central Este. 70:1-517. LivsHuttz, T. 2003. Systematics of Dischidia (Apocynaceae, Asclepiadoideae). Ph.D. dissertation, Cor- nell University. SCHLECHTER, R. 1899. Asclepiadaceae. In: Symbolae Antillanae, |. Urban,ed.Gebriider Borntraeger, Berlin Pp. 236-290. Stevens, W.D. 2005. New and interesting milkweeds (Apocynaceae, Asclepiadoideae). Novon 15: 602-619, Woooson, R.E.1941.The North American Asclepiadaceae. Ann. Missouri Bot. Gard. 28:193-244. 954 BRIT.ORG/SIDA 22(2) Book REVIEW ANTHONY Huxtry. 2005. Green Inheritance: The WWF Book of Plants.SBN 0-520-24359- 5, pbk.). The University of California Press, Berkeley, CA 94704, U.S.A. (Orders: Cali- fornia Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A. Tel: 609-883-1759; Fax 609-883-7413, www.ucpress.edu). $29.95, 192 pp., 234 color, OE LO fe Why is environmental conservation important? If you have ever asked this question, this book will provide your answer. Focusing on the “green mantle” of plant life that surrounds us, ie ne lela ae reader how conser- vation of the environment is integral to human survival. Color pl inc uding botani- cal illustrations from the collection at Kew Gardens, complement os clear writir ] f ig style of the late Mr | Huxley, a prolific writer and an editor at Amateur Gardening. In this book, Mr. Huxley reminds us that “Without plants, animal lif Id ist.” From the very beginning, plants have acted as the “life support systems” of the planet, providing human and animal life with protection, nourishment, and even the air we panes It was the emer- gence of green plants that changed the poisonous primeval atmosphere into today’s | ygen-bearing atmosphere. In Chapter 2, “Guardians of the Environment”, the role of plants in protecting the air we breathe, preventing soil erosion, protecting us from harmful radiation, fixing nitrogen in the soil, stabilizing the climate, and producing rain is discussed. The role of cultivated plants in providing us with food and spices is celebrated, as well as the problems resulting from modern agricultural techniques. The 5,000-year history of “green medi- cine” is astanee from traditional pera! cures to modern drug aIScONeEy ao orts ane the resurgence of herbal | 1.Inachapter titled “Plants and Seeley" the sp aia of pants in our be: day lives and in the lives of other cultures is discussed. Mr. medicine f medicinal f t Huxley pter to the roles of in situ conservation, such as land reserves, and ex situ conserva- tion, such as modern bio- i pinnae and gene hanks, in protecting our resources and f feeding a nungty P planet. The author reminds us throughout the book that * tho ight- less impatience and greed, failing to appreciate the value of what we ae The last ciapeer is appropriately titled “Saving the Plants that Save Us” and summarizes the environmental problems that are depleting plant life on our planet, as well as solutions. The primary approaches to saving our “green inheritance” include halting environmental degradation, rehabilitating land, and conserving the pristine natural and semi-natural vegeta- tion that still remain. Mr. Huxley does not sugar-coat the issue, he acknowledges the difficulty of getting nations to work together, as well as the role that economic disparity plays in the battle. But, he paints a beautiful picture of the reward, “Besides ¢ thus providing for our io nee plants can also pone solace for the mind. They are endlessly fa Ascin AtiNg, we asia g word, and the ey are diversely beau- tiful too.”—Marissa Oppel, MS, C dR *h Assistant, Botanical Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A. SIDA 22(2): 954. 2006 NEW COMBINATIONS IN THE GENUS CYMOPTERUS (APIACEAE) OP THE SOUTHWESTERN UNITED STATES Ronald L. Hartman Rocky Mountain Herbarium Department of Botany University of Wyoming . University Ave. Laramie, Wyoming 82071-3165, U.S.A. Rhartman@uwyo.edu — ABSTRACT Preparation ol the pmiateas for Bolack San Juan Basin Flora has necessitated the following new combinations: C. davidsonii, C. glomeratus var. fendler i, and C. ses ili lorus eA 1 RESUMEN La preparacion de las cane ee la Flora de Bolack San Juan Basin | do] combinaciones nuevas: Cymopterus breviradiatus, C. davidsonii, C. glomeratus var. fendleri, y C. eciils Orus. In revising Cymopterus Raf. (Apiaceae) of the greater Four Corner’s region for the San Juan Flora (Heil et al.), it has become necessary to make the following nomenclatural innovations. The aforementioned treatment represents a continued reliance on charac- ters of the fruit and of vegetative morphology (Mathias & Constance 1944-45; Cronquist et al. 1997). Recently, several phylogenetic analyses of nuclear (ITS) and plastid (rpsl6 intron) sequences (Downie et al. 2002; Sun @ Downie 2004; Sun et al. 2004) have defined a group, likely monophyletic, of western North American apioids. Genera include Aletes J.M. Coult. & Rose, Cymopterus [here circumscribed to include Oreoxis Raf., Pseudocymopterus J.M. Coult. & Rose, and Pteryxia (Nutt. ex Torr. & A. Gray) J.M. Coult. & Rose], HarbouriaJ.M. Coult. & Rose, Lomatium Raf., Musineon Raf., Neoparrya Mathias, Oreogenia S. Wats., Oreonana Jeps., Podistera S. Wats.,and Shoshonea Evert & Constance. Thus far the molecular trees are poorly resolved with the basal branches containing sev- eral to many polytomies, thus obscuring relationships among terminal groups. A num- ber of terminal clusters make sense to one steeped in western umbels while others seem- ingly defy logic. It is hoped that meet pequcuce data will provide sufficient insight into the evolution of the western ay at a stable classification can be established. Cymopterus breviradiatus (W.L. Theob. &@ CC. Tseng) R.L. Hartm., comb. @ stat. nov. BASIONYM: Aletes macdougaliiJ.M. Coult. & Rose subsp. breviradiatus WL. Theob. & CC. Tseng, Brittonia 16:306, fig. 6.1964. The genus Aletes is defined by its laterally Hoses (versus terete to dorsally flattened) fruit. This lateral compression is prominent in the type species, A. acaulis (Torr.)J.M. Coult. & Rose and its presumed closest relatively, A. humilis J.M. Coult. @ Rose. None of the three taxa of Aletes for which nomenclatural innovations are here made have fruit that are flattened laterally. Although molecular studies may eventually show that this is a fickle character, it is one that has been relied upon traditionally. Based on gestalt and technical features of the fruits and leaves, Cymopterus breviradiatus is extremely similar to Cymopterus sessiliflorus (Theobald et al. 1964) despite the fact SIDA 22(2): 955-957. 2006 956 BRIT.ORG/SIDA 22(2) that the two taxa do not group near one another based on molecular data (Fig. 2, Sun & Downie 2004). There is ample justification for removing subsp. breviradiatus from sub- jugation to Cymopterus macdougalii J.M. Coult. & Rose) Tidestrom [Oreoxis m.].M. Coult. & Rose, 1913; Aletes m. J.M. Coult. & Rose, 1935]. The former has fruit with corky-thick- ened wings whereas C. macdougalii has weakly developed, thin wings. Cymopterus bechii S.L. Welsh & Goodrich appears virtually identical to C. macdougalii morphologically. This is reinforced thanks to the number of collections of each that have been amassed in recent years. Consequently the former is treated as a synonym of C. macdougalii. Fur- thermore, as sequence data indicate (Figure 2, Sun @ Downie 2004), C. macdougalii, C. (Pteryxia) davidsonii, and C. beckii group with C. lemmonii (Coult. & Rose) R.D. Dorn [Pseudocymopterus montanus J.M. Coult. & Rose]. C. davidsonii strongly resembles C. macdougalii morphologically and they may represent sister taxa. eS i davidsonii (J.M. Coult.& Rose) R.L. Hartm., comb. nov. Basiony: Aletes? davidsonii M. Coult. & Rose, Contr. U.S, Natl. Herb. 7:107. 1900. Peuaeconun we davidsonii J.M. Coult. & Rose) Mathias Ann. Missouri Bot. Gard. 17:282, 316. 1930. Pteryxia davidsonii .M. Coult. @ Rose) Mathias & Constance, Bull. Torrey Bot. Club 69:248. 1942. This species was most recently treated in Pteryxia (Mathias & Constance 1944-45) and tentatively attributed to Pseudocymopterus (Cronquist et al. 1997). The reader is referred to comments provided under Cymopterus breviradiatus. Cymopterus glomerattis (Nutt. ) DC. var. fendleri (A. Gray) R.L. Hartm., comb. & stat. nov. BASIONYM: Cy terus i A. Gray, Mem. Amer. Acad.,n 56.18 Specific epithet: C sitentars glomeratus (Nutt. ) DC,, Prodr. 4: nen 1830. BASIONYM: Thapsia glomerata Nutt., Gen. N. Amer. Pl. 1:184. 1818, for which C s Raf., Herb. Raf. 2:40. 1833, is illegimate and — surperfluous (nom. nov. for Selinum eee Push FL Annee Sept. 2:732, ne non Cav, 1799; Internationa Plant Names Index 2005). Cymopterus fendleri long has been treated as distinct from the related C. glomeratus (C. acaulis, Mathias & Constance 1944-45) or as a variety of it (Cronquist 1997 et al; Goo- drich 2003). A recent study (Sun et al. 2005) using principal component analyses failed to discriminate among the five varieties recognized by Goodrich (2003). Despite this lack of resolution, the above combination is made for botanists, including myself, wishing to recognize var. fendleri. Cymopterus sessiliflorus (W.L. Theob. & C.C. Tseng) R.L. Hartm., comb. nov. BasionyM: Aletes sessiliflorus WL. Theob. & C.C. Tseng, Brittonia 16:309. 1964. The reader is referred to comments provided under Cymopterus breviradiatus. ACKNOWLEDGMENTS I appreciate the comments of an anonymous reviewer. REFERENCES Cronauist, A., N.H. Houmaren, and P.K. Houmaren. 1997. Intermountain flora: Vascular plants of the Inter- mountain West, U.S.A. Vol. 3A: Subclass Rosidae (except Fabales). New York Bot. Garden, Bronx. P. 372. Downie, S.R., R.L. HARTMAN, F.-J. Sun, and D.S. Katz-Downie. 2002. Polyphyly of the spring-parsleys (Cymopterus): molecular and morphological evidence suggests complex relationships among the perennial endemic genera of western North American Apiaceae. Canad. J. Bot. 80:1 295-1324. HARTMAN, NEW COMBINATIONS IN CYMOPTERUS 957 GoooricH, S. 2003. In: S.L. Welsh, N.D. Atwood, S. Goodrich, and L.C. Higgins, eds. A Utah flora, ed. 3. Brigham Young Univ., Provo, Utah. P.654. Heit, K. et al. Bolack San Juan Basin flora. Missouri Botanical Garden Press, St. Louis, Missouri.In prep. INTERNATIONAL PLANT Name INoex, THE. 2005. (Attp://www.ipni.org) Maruias, M.E.and L. Constance. 1944-45. Umbelliferae. N.Amer.Fl.28B:187.New York Bot.Garden, Bronx Sun F-J. and S.R. Downie. 2004. A molecular systematic investigation of Cymopterus and its allies (Apiaceae) based on phylogenetic analyses of nuclear (ITS) and plastid (rps16 intron) DNA se- quences. 5S. African J. Bot. 70:407-416. Sun F.-J., S. R. Downie, and R.L. Hartman. 2004. An ITS-based phylogenetic analysis of the perennial, endemic Apiaceae subfamily Apioideae of western North America. Syst. Bot. 29:419-431. Sun F.-J.,G.A. Levin, and S.R. Downie. 2005. A multivariate analysis of Cymopterus glomeratus, formerly known as C. acaulis (Apiaceae). Rnodora 107:359-385. THEOBALD, W.L., C.C. Tsenc, and M.E. Matias. 1964. A revision of Aletes and Neoparrya (Umbelliferae). Brittonia 16:296-315. 958 BRIT.ORG/SIDA 22(2) Book REVIEW HAKAN Rypinand Joun K. JecLum. 2006. The Biology of Peatlands. Oxford University Press (ISBN 0-19-852872-8, pbk.). Oxford University Press, 2001 Evans Road Cary, NC 27513, U.S.A. (Orders: 800-451-7556; http://www.oup.com/us/). $49.50, b/w figures and photographs, tables, graphs, and glossary, 343 pp., 61/4" x 91/4" The Biology ae tua nds by ae Rydin and oe is part of Oxford's Biology of Habitats series and offers readers a Peatlands. This is a well-illustrated book with many text- summarizing and research- eed illustrations. This book would be very suitable as a textbook for classes on peatlands or as a supplemental reading for exploring wetland habitats. Additionally, this title could also be a beneficial reference text a classes such as agronomy, ee or forestry. a 1) ¢. } The book < well alo oi ae on previous cha cere intorma- = tion. Chapter topics include (1) Peatl I | habitats, (2) at eee peatland habitat, (4) Sphagnum - the builder of boreal Jaen (5) Peat and organics soil, on re peat archives, (7) Peatland succession and development, (8) Hydrol peatlands, (9) Nutrients, lig )) Peatland pat- terns and landforms, (11) Peatlands Sau the world, (12) ae oe carbon bs nee ie oS) Uses, func- tions, and cue oncas of peatlands CI he text is very readable and is accompanied | by many supportive illustrations, graphs and charts. The authors’ eae chapters provide an introduction to terminology associated with peatlands which will help the reader both tear a are — ter Tminology and determine the relationship of peatland to } t other wetlands types. These openin the reader an overview of common plants and animals associated in peatlands as well as special sdepenions of these organisms to living in various peatland types. Chapter four focuses specifically on the life cycle, morp hology, physical attributes, chemical attributes and the colonization ability of Sphagnum mosses. All chapters are steeped with references to pertinent scientific studies. These references allow readers to learn more about methodology, data and results from such studies. The middle chapters of the book focus on peatland organic soils, hydrology including peat physical and chemical properties, peat as historical, fossil “ar- chives,” detailed explanations of the processes involved in peatland succession and devel t, descriptions of water flows, balances in peatland types and the roles of nutrients, light and appears in peatlands. The hy- drology coe includes a biguoats discussion of the variations in water chemistry along bog-rich fen gradi- Thi ents. from recent research studies on the subject and includes a summary chart of the surface water chemistry of various peatland sites. Within chapter nine is a wonderful example of research regarding forestry uses of peatlands in a discussion of “nutrients after drainage for forestry”; again the authors have presented many supportive research references in the discuss e final chapters discuss peatland patterns and formations “peatlands aed the world, peatland produc- tion n and catho balance, pec a and uses, peatland function and peatland management. These final chapters use the eatland ecology and ee The peatland landforms secon has a help ful inn on mire ste oue and c serene at different scales including eee ta able and detailed description of mire features, sites and types. Many aoe ert accom- pany this of mire types. The authors have focused this text on eas peatlands, but they do include sections contributed from any auLners on ana in Argentina, New Zealand, and southeast Asia. Peatland Ss acuen and a tie together much of what has been | | W icmp) i oli: : 1 fields. The pena: pues is loaded ++ 1 a | } ) ] a ae ] W and various parts of the carbon cycle within peatlands, ] biomass and rn Pp | as wellasa presentation of information on neat accum ae os limitations in accumulation “The nee chapter has many applications of lating susta lity and uses c in fields such as forestry, fuel use, horticulture a asa de- ies substrate. The Biology of Peatlands by authors Rydin and Jeglum provides a comprehensive overview to peatlands biology, ecology and peatland types. This mo) is Srull of supportive black and white illustrations, charts, tables an ndr ele erences to research studies. T Bi ls would set l textbook for classes focus- ing on peatlands, wetland habitat types, ind elated classes where peatland products may be of interest.—Lee Luckeydoo, Herbarium, Botanical Research Institute of lexas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. SIDA 22(2): 958. 2006 SIX NEW SPECIES OF SPHAGNUM (BRYOPHYTA: SPHAGNACEAE) FROM NORTH AMERICA Richard E. Andrus Environmental Studies sh Binghamton Univ Binghamton, New York 1 ne Ln U.S.A. randrus@binghamton.edu ABSTRACT Sphagnum mcqueenii, S. kenaiense, 5. bergianum, S. beothuk, S. sitchense, and S. talbotianum, are described and illustrated as new species for North America. RESUMEN raat | a ] N América Sp } S. kenaiense. S bergianum, 1 S. beothuk, S. sitchense, y S. talbotianum. INTRODUCTION Extensive field work in North America in recent years has revealed a number of new species of Sphagnum, mostly in more remote and poorly collected areas. Six of those spe- cies are described in this paper. Four are from Alaska, one from Newfoundland, one from both Newfoundland and Alaska and a sixth species from several states and Canadian provinces in northeastern North America. Sphagnum mcqueenii Andrus, sp. nov. (Figs. 1-6). Tyr: U.S.A. NEW HAMPSHIRE: Monroe Co. Town of Nottingham, Pawtuckaway State Park, Dead Pond, extensive poor fen, 18 Sep 1994, Andrus 9070 (HOLC TYPE: DUKE, ISOTYPES: BING, NY). Sphagnum sect. Cuspidata. Planta staturam mediam attingens, ate vel Devens escens, capitul pi pl ] i,g inali praedito. Cortex caulinuse i ietil litari it Folia caulina aequilaterali- triangularia, 0.75- te mm lata, 08> Ll mm Longa, late marginata, in Posen apical efibrillosa ie aballe osa pl | I I I iB ] [Rey emecreed cee : | Tels li 5 a | 7] praedita lanceola, 0.5- coe mm lata, ee 21 mm longa, in superficie convexa a cellulis hyalinus poros apicales O- i et L Ree i 12. 4-8 i diametro praedita. Plants robust and weak-stemmed,; yellow to light brown; capitulum typically flat- topped and with a + conspicuous terminal bud. Stems light green; cortex in 1-2 layers of moder- ately differentiated thin-walled cells. Stem leaves equilateral- triangular, 0.75-1.0 mm wide by 0.8-11mm long, usually spreading, apex obtuse, hyaline cells usually septate in lower half of leaf and efibrillose (or fibrillose near apex). Branches unranked, + straight, branch leaves moderately elongated at distal end. Branch leaves < 2.1 mm long, ovate- lanceolate, undulate and sharply recurved when dry; hyaline cells on convex surface with 0-1 apical pores and often with pseudopores, concave surface with up to 12 round wall- thinnings in cell angles and sometimes along commissures; chlorophyllose cells trian- gular in transverse section and just enclosed on the concave surface. Sexuality unknown. Distribution.—Western Hemisphere Atlantic; known to date from Maine, New Hamp- SIDA 22(2): 959- 972. 2006 BRIT.ORG/SIDA 22(2) Fis. 1. Sphagnum mcqueenii. Habit. ANDRUS, NEW SPECIES OF SPHAGNUM FROM NORTH AMERICA 961 Fic. 2. Sphagnum mcqueenii. Spreading branch. shire, Pennsylvania, and Vermont from the U.S.A. and Nova Scotia and Newfoundland in Canada. Identification—Sphagnum mcqueenii can be separated from S. cuspidatum Hoffm. by its enclosed chlorophyll cells, obtuse stem leaves and the relatively short, recurved and broad ( width: length ratio > 0.3) branch leaves. Sphagnum viride Flatberg. will dif- fer in having branch leaf chlorophyll cells not reaching the concave surface and pointed stem leaves. Both S. cuspidatum and S. viride are also smaller and slenderer plants. S. pulchrum (Braith.) Warnst. has strongly 5-ranked branch leaves, more pointed stem leaves and more well-enclosed branch leaf chlorophyll cells. Wet-growing forms of S. pulchrum will also not have branch leaves elongated at the distal end of the branches, a character- istic of the more “aquatic” Cuspidata species. The other “aquatic” Cuspidata with which S. mcqueenii can occur are S. torreyanum Sulland S.atlanticum Andrus. Both of these are larger plants and have longer branch leaves (> 2.1mm). They also have stem leaves more appressed to the stem than S. mcqueenii, in which the stem leaves are often spreading. Ecology.—Habitat preferences not well known but it has so far been collected in poor fen habitats with floating mats, wet carpets and wet peaty depressions. It was quite abun- dant in the New Hampshire type locality, where it occurred in the moat at the bog mar- gin associated with S. torreyanum, S. atlanticum, S. cuspidatum and S. flexuosum Dozy.& Molk Etymology.—The namesake of this species is the late Cyrus McQueen, a good friend and fellow sphagnophile, who led me to the type locality. Additional collections examined: CANADA. Newfoundland: N of Pouch Cove, Andrus 10309 (BING): Winterland, Andrus 10422, 10423, 10439 (BING, DUKE): N of Harbour Breton, Andrus 10632 (BING): Lark Harbour, Andrus 10683 (BING, DUKE). NOVA SCOTIA. Guysborough Co.: Sc is & Maass 3315 (DUKE). U.S.A. MAINE: Acadia National Park, Little 16 (BING). Sagahadoc Co.: Allen 27060 (DUKE). PENNSYLVANIA. Monroe Co.: Andrus & Damman 6641 (BING). VERMONT. Windham Co.: McQueen s.n., i Nov 1998 (BING, DUKE). Sphagnum kenaiense Andrus, sp. NOV. (Figs. 7-12). Type: U.S.A. ALASKA. Kenai Peninsula Co. Kenai Peninsula, Soldotna, Headquarters Lake large poor fen complex surrounding lake, 17 Aug 2004 (HOLO- TYPE: DUKE; IsoTyPEs: BING, NY). Sphagnum sect. Cuspidata. Planta Denv caule debili, pallide brunnea vel aetate aurea, capitulo plano; cortex caulinus parietibus tenuibus, tantum leniter dilatatus. Folia caulina ad caulem appressa, lingulata, ovato-lingulata vel triangulata, aequa vel minora quam 0.9 mm longa, apice obtuso aus sata eroso ve ius see tantum leniter 5-ordinati vel 5- seriata, in apice distali rameali vix elongati. F 2-3 pendentibus compositus. Folia ramulina 1.1-1.3 mm longa, ovata, rigentia, in statu sicco ae antele atque subrecurva, 962 BRIT.ORG/SIDA 22(2) Fics. 3-6. Sphagnum mcqueenii. 3. Branch leaves. 4. Stem leaves. 5. St } +] Veesenten “| a4. t lat ] cellulis hyalinis in parte mediana } s usque longitudinis 0.3- 0.4) et in superficie convexa in tas folii intone saepe jaa oro magno atque/usque 6 pone ae et { - ~ ¢ parte apical isae epe g i) 5 pe udoporis instructa, uperticie concava ad t E atque angulos areis parietum tenuium magnis rotundis Hee ee inconspicuuis vel nullis praedita; cellulae chlorophylligerae in section transversali triangulares, typice in superficie bene inclusae. Plants small.and weak-stemmed; pale brown to golden brown; capitulum flat-topped and ANDRUS, NEW SPECIES OF SPHAGNUM FROM NORTH AMERICA 963 12. Branch leaf Fig. 7-13 5} f 9 k j 7. Habit. 8. Branch. 9. Stem leaves. 10. Branch leaves. 11.B h leaf concave surface. 13. Stem section. Scale bars: A= 2 mm, 7; B = 2 mm, 8; C= 0.5 mm, 9-10; D = 50 pm, 11-13. only weakly 5-radiate. Stems pale yellow; stem cortex moderately well-differentiated but not much enlarged. Stem leaves appressed to stem; lingulate, ovate,to triangular; equal to or less than 0.9 mm long; apex obtuse and often erose to lacerate. Branches with leaves unranked to 5-ranked, leaves not much elongated at distal branch tip. Branch fascicles with 2 spreading and 2-3 pendent branches. Branch leaves ovate, 1.1-1.3 mm long, stiff, 964 BRIT.ORG/SIDA 22(2) weakly undulate and slightly recurved when dry; hyaline cells in mid-region quite short and broad (width:length = 0.3-0.4), in lower 1/2 of leaf on convex surface often with 1 large pore apically and/or up to 6 free pores, in apical region often with pseudopores along the cell margins; on concave surface with large round wall- Tenens in the cell ends and angles (these sometimes faint or absent); chlorophyllose cells triangular in trans- verse section and typically well-enclosed on concave surface. Sexuality unknown. Distribution.—South-Central and Western Alaska. Identification —Within its range, S. kenaiense may be confused with S. angustifolium (Russow) C. Jens., S. balticum (Russow) C. Jens.) and S. brevifolium (Braithw.) Roell. Sph- agnum angustifolium in the field has narrower ovate-lanceolate branch leaves. Micro- scopically S. angustifolium has a poorly differentiated stem cortex, chlorophyll cells not well-enclosed and narrower branch leaf hyaline cells. Sphagnum balticum has narrower branch leaves and typically only one hanging branch, with stem leaves markedly spread- ing. Microscopically the branch leaf hyaline cells are also narrower. Sphagnum brevifolium has stem leaves that are apiculate to acute and ovate-lanceolate branch leaves. It also has narrower hyaline cells in its branch leaves than S. kenaiense. Ecology.—Sphagnum kendiense occurs in poor to medium fens, often of a sedge domi- nated nature. It typically forms sprawling patches in hollows, often over bare peat. Etymology.—The species epithet kendiense is after the Kenai Peninsula where the species was found. a. Additional coll ined: U.S.A. ALASKA: King Salmon, Andrus 9306 (BING, DUKE); Selawik, Schofield 121,733 (BING): Anc horage C ampbell Airstrip fen Andrus 8448 (BING, DUKE); Kenai Peninsula, Mil epost 100 on Seward Hwy., Andrus 8752 (BING, DUKE, NY). Sphagnum bergianum Andrus, Sp. NOV. (Figs. 13-19). Type: US.A. ALASKA. Kenai Peninsula Co: Kenai Peninsula, Soldotna, H leadquarters | ake, extensive poor fen con nplex at lake en 17 Aug 2004, Andrus 10020 (HOLOTYPE: DUKE; ISOTYPEs: BING, NY). apa section Acutifolic pl] | a ] yb ] ] ] Ne omnino atro-fusco, St | centro Cz cenit aurato, in statu vivo saturate € rubritinge nti, in statu sicco nitore roseolo- pur Durascenti 1: praedito.C aulis atro-fuscus Folia caulina late triangulari- lingul ata, 1.15-1.25 x 0.8, in ae subobrusa vel late aiaicn limbo ad hasem ates oe cellulis hyalinis pro parte maxima |-septatis, [ vel isin parte mediana [ ami plus minusve 5-ordinati. Fasciculus see e cael: 2 patentibus atque | aie Folia ramulina 1.2-L3 x 0. 65-0.75 mm, late ovata, apice involuta, cellulis h yalinis in merosis rotundatis i ellipticis poris commissuras se- cus instructis, in apic idatim usque ab yasem maioribus sa, POTIS paucis magnis in epost dicate ibe paivasiint exceptis. Status sexualis ce Sporae 26-30 —- Plants moderate-sized to robust, capitulum more or less flat-topped and large; dark brown overall with a golden center to the capitulum and a distinctive deep red tinge in the field which becomes a pinkish-purple sheen upon drying. Stems dark brown; superficial cor- tical cells aporose. Stem leaves broadly triangular-lingulate, 1.15-1.25 = 0.8, apex slightly to broadly obtuse, border moderately broadened at base; hyaline cells mostly l-septate with afew to many 2 septate in the mid-region, shape rhomboidal. Branches more or less 5 ranked. Branch fascicles with 2 spreading and | hanging branch. Branch leaves |.2-1.3 mm x 0.65-0.75, broadly ovate, apex involute; hyaline cells on convex surface with nu- merous round to elliptic pores along the commissures, these grading from small pores at the apex to large pores at the base; concave surlace apor xcept for a few large pores in the lower side regions. Sexual condition dioicous. Sas 26-30 um. Distribution.—Newfoundland and South-central Alaska. ANDRUS, NEW SPECIES OF SPHAGNUM FROM NORTH AMERICA 965 Fics. 14-20. Sphagnum bergianum. 14. Habit. 15. Branch fascicle. 16. Stem leaves. 17. Stem leaf hyaline cells. 18. Branch leaves. 19. Branch leaf, concave surface. 20. Branch | face.A = 2 mm, 14; B = 2 mm, 15;C=0.5 mm, 16, 18; D = 50 pm, 17, 19-20. Identification. Sphagnum bergianum seems most clearly related to S. subfulvum Sjoers sensu lato, as described by Flatberg (1985). Both S. subfulvum and S. subfulvum ssp purpureum Flatb. are a lighter golden-brown than the quite dark brown of S. bergianum. Furthermore, both of these taxa have narrower stem leaves that are more acute at their apex. And neither of these taxa have distinctly 5-ranked branch leaves like S. bergianum. Sphagnum bergianum is a quite unusual looking species in the field, with its very dark brown color tinged with a deep red color. When dry this deep red becomes more purplish 966 BRIT.ORG/SIDA 22(2) but it still shows some red. This distinctive field appearance made it quite recognizable when collected in Newfoundland in 2005 Table 1 highlights the differences between S. bergianum and other larger brown Acutifolia. Because of its similarity, S. subfulvum ssp purpureum is included for complete- ness even though the author has not seen any North American specimens of that taxon and believes it to be of doubtful occurrence there. Ecology.—In the few sites where it has so far been collected, S. bergianum has been a hummock forming species of medium fen habitats, associated with such species as S. warnstorfii Warnst., S. subfulvum, S. platyphyllum (Braithw.) Warnst. and S. fuscum (Schimp.) H. Klinggr. Etymology.—This species named after Ed Berg, ecologist with the US Fish and Wild- lite Service, who first found this species in Headquarters Lake next to his office. Additional collections examined: CANADA. Newfoundland: 9 km N of Badger, Andrus 10646 (BING, DUKE, NY); 15 km SW of Corner Brook, Andrus laa DUKE, NY). U.S.A. Alaska: Kenai Peninsula, Milepost 40 on Seward Hwy., Andrus 10036 (BING, DUKE a Sphagnum beothuk Andrus, sp. nov. (Figs. 20-25). Type: CANADA. NEWFOUNDLAND: Lark Harbour, ev, Andrus 10687 (HOLOTYPE: DUKE; ISOTYPE: Naas} ca. 50 km NW of Corner Brook, medium rich fen, 20 m e BING, NY) eee sect. Ac a id. Pl. anta parva | 1 | | fi usco nitore 1 li Caulis brunneus, eda corticalis supericlie eporosis. Folia oan ieee ll- 124 mm oe in apice subapiculata vel maximam partem lata, erosaque vel lacerata, me Ss ad co tantem parce dilarato, cellulis hyalinis rhombiformibus, 0-1-septatis. Rami plus minusve 5-ordin 2 patentibus atque 1 pendenti compositus. Folia ramulina 0.95-1.3 mm nae ovata vel ovato-lanceolata, concava, stricta vel parce en cecunee in pie volta, cellulis hyalinis in ie Hole convexa ee commissuras secus numerosis 4 lell; | rot undatls ad 5 \ L ie} s | titudinic Thal vel perpusillis on m, minoribus quam 025 par form ane superticie concava in partibus proximalibus ne poris parcis, rotundatis per sate instructa. Status sexualis ignotu. Plants small to moderate-sized; capitulum rounded and dense; dark brown with a pur- plish sheen. Stem brown, superficial cortical cells aporose. Stem leaves lingulate, 1.1-1.2 mm long, apex slightly apiculate to mostly broad and erose to lacerate, border only slightly broadened at base; hyaline cells rhomboidal and 0-1 septate. Branches more or less 5-ranked. Branch fascicles with 2 spreading and 1 pendent branch. Branch leaves 0.95-1.3 mm long, ovate to ovate-lanceolate, concave, straight to slightly subsecund, apex involute; hyaline cells on convex surface with numerous round to elliptic pores along the commissures, grading from large pores at the base to a mixture of small and tiny (2 pm, less than 0.25 cell width) at the apex, concave surfacewith a few large, round pores/cell in lower side regions. Sexual condition unknown. Distribution —Known only from Newfoundland, Canada. Identification.—In the field S. beothuk looks like a very dark form of S. fuscumexcept for its distinct purplish sheen. It’s rather more robust than Sfu scumand also has 5- Sans branch leaves, which S.fuscum usually lacks. Microscopically, however, S. beothuk differ clearly in the tiny pores seen mixed in with more typical pores on the convex euiiace of the branch leaf at the apex. Among the known Acutifolia species, only the red pigmented S. warnstorfii has been noted with this character before. Ecology.—Sphagnum beothuk forms dense hummocks similar to S. fuscum. It appears to be a minerotrophic species and has been collected in the same mire as Ssubfulvum, S. inundatum Russow, and S. warnstorfii. Taait 1. Comparison of Sphagnum bergianum and related species. Characteristic 5. bergianum S$. subfulvum $. subfulvum ssp. purpureum 5. subnitens 5. flavicomans Color Branch fascicles, spreading + hanging branches Branch leaf ranking Branch leaf shape Stem leaf shape Stem leaf apex Stem leaf length in mm Stem leaf hyaline cells Distribution Dark brown with a deep red tinge Often 5 -ranked Broadly ovate Triangular-lingulate More or less obtuse .15-1.25 |—2 septate Newfoundland & SC Alaska Golden-brown with a purplish gloss sometimes 2+ Unranked Ovate Triangular-lingulate to broadly lingulate More or less acute ee 0-1 septate NE and NW North America Golden-brown with red tinge + 1-2 Unranked vate Triangular-lingulate More or less acute 1.3-1.4 O-1 septate NW Europe Reddish-purple mixed with golden brown 2+ 1-2 Unranked Ovate to ovate-lanceolate Triangular to triangular- lingulate Acute to apiculate 1.3-2.7 0-1 septate Eurasia, W North America & New Zealand Brown 2+ 1-2 Often 5-ranked Ovate-lanceolate fail triangular- lingulat Acute to ice 0-1 septate NE North America VOIYIWY HLYON WOUd WANOWHdS 40 S31D3dS MIN ‘SNYGNY 196 968 BRIT.ORG/SIDA 22(2) ABCD 1D | AY Op Oe C) EO o / ae gua7cee. i isc qs WANS exe 7 rO) i | Fics. 21-26. 5; k. 21. Habit. 22. Branch. 23. Stem leaves. 24. St | Ils. 25. Branch leaves. 26. Branch leaf hyaline cells, convex surface. A = 2 mm, 21; B = 1 mm, 22; C= 0.5 mm, 23, 25; D = 30 pm, 24, 26. Etymology.—The Beothuk are the aboriginal people of Newfoundland. Comments.—Several things suggest a possible hybrid origin between S.fuscum and S. warnstorfii. The tiny pores at the branch leaf tips, the purplish sheen and the 5-ranked branch leaves suggest S. warnstorfii and the lingulate stem leaves are a character of S. fuscum. Furthermore the type locality is a minerotrophic site in which both S. fuscum and S. warnstorfi also occur. ANDRUS, NEW SPECIES OF SPHAGNUM FROM NORTH AMERICA 969 YG; my it pia (i dart LE pe A KA SSE " Vy we Fics. 27-34. Sph itch 27. Habit. 28. Branch. 29. Branch leaves (“b”) 30. Stem with leaves. 31. St leaf hvall IIs. 32 £ 24 RQ hl £ A=2mm,27;B=1mm,28, 30;C=0.5 mm, 29, 32; hl Stem leaves (“s”).33.B D=50 pm, 31, 33, 34. Additional collections examined: CANADA. Newfoundland: Lark Harbour, Andrus 10685, 10686, 10687 (1orotyPEs: BING, DUKE); ca. 16 km S$ of jet. of Hwy 210 and Hwy 214, Andrus 10408 (BING, DUKE). Sphagnum sitchense Andrus, sp. nov. (Figs. 26-33). Type: US.A. ALASKA: Sitka Co. Sitka, Harbor Mt, alpine area at end of road, 700 m elev, dense patch in tundra, 19 Aug 2004, Andrus 10097 (HOLOTYPE: DUKE, IsoTyPEs: BING, NY). Sphagnum sect. Acutifolia. 970 BRIT.ORG/SIDA 22(2) nee F eee F ] dence. loat lo ni lesti Caulis roseolus . | c Ct ediis coneaioue fF aialibie p — Folia caulina anguste vane aria vel lingulato-triangularis, 1.2- ven mm longs x 2 6- 2 8) mm oe ratione enihtieticg ongicncinls usque SAUCES ca. 2:1, apice apiculato, limbo ellulis lum 2-septatis, saepe in apice kenbillods Bata non 5-ordinati. Fasciculus ramulinus e scene? 2 patentibus atque | absaiae peepee aes ramulina 1 = Wo mm longa, 0.5- ve 55 mm lata, ovato-lanceolata, concava, stricta, in l vel ellipticis ad basem poris magnis jen Bree a vege ad ee rotunealls quoae staturam mediocribus, in superficie concava poris magnis rotundatis g Plants cael saints flat-topped and dense; pale yellow and pink, lacking sheen. Stem pink, superficial cortical cells aporose. Stem leaves narrowly triangular to lingulate- triangular, 1.2-1.6 mm long x 0.6-0.8 mm wide, length:width ratio ca 2:1 apex apiculate, border strong and moderately broadened basally; hyaline cell rowly rhom- boidal and | to occasionally 2 septate, often fibrillose apically. Branches not 5-ranked. Branch fascicles with 2 spreading and | pendent branch. Branch leaves 1.3-1.5 mm long and 0.5-0.55mm wide, ovate-lanceolate, concave, straight, apex involute; hyaline cells on convex surface with numerous round to elliptic pores along the commissures, grading from large elliptical pores at the base to moderate-sized round pores at the apex, concave surface with a few large round pores scattered throughout. Sexual condition unknown. Distribution—Known only from the type locality in pe neeee Alaska but since al- pine areas in this area are poorly collected it is likely more widespr Identification Sphagnum sitchense is somewhat similar to S. cari ilyeinen (Ehrh.) Hedw. But differs ina number of important ways. Its stem leaves are often narrowly tri- angular whereas those of S. capillifolium are lingulate- triangular. Branch fascicles on S. sitchense have 2 spreading and one hanging branch whereas those of S. capillifolium have 2 spreading and mostly 2 hanging branches. The stem leaf hyaline cells in S. sitchense are 1-2 septate while those on S. capillifolium are 0-1 septate. In the field S. sitchense has a pale red tinge and quite flat topped capitulum which contrast with the deeper red tinge and rounded capitulum of S. capillifoilum. Sphagnum sitchense also bears some resemblance to Sphagnum subnitens but differs in the following ways. Sphagnum sitchense hasa strongly bordered stem leaf with afew 2 septate hyaline cells while S. subnitens has a narrow border and hyaline cells 0-1 septate only. Sphagnum sitchense also lacks the metallic sheen and brown coloration that are typical of S. subnitens. And lastly, the branch fascicles in S. sitchense have only one hang- ing branch whereas S. subnitens has mostly two. The stem leaves of S. sitchense are quite similar to those of S. quinquefarium, with which it occurred in the type locality, but the latter species has 5-ranked branch leaves and typically 3 spreading branches per fascicle. Ecology.—In the type locality, S. sitchense was collected several times in dense patches in the tundra. Growing similarly in the same area were S. girgensohnii Russow, S. quinquefarium (Braithw.) Warnst., and S. compactum D.C. Etymology.—Sphagnum sitchense is after the type locality. — Additional collections examined: U.S.A. Alaska: Sitka, Harbor Mt,. Andrus 10095, 10098 (BING). Sphagnum talbotianum Andrus, sp. nov. (Figs. 34-39). Type: U.S.A. ALASKA: Bethel Co: Bethel, 0.9 mi — es Rd. ie turnoff for Chief Eddie Hoffman Hwy, just W of Larson Subdivision, medium fen, =; DUKE; ISOTYPES: BING, NY, TRH). Sphagnum sect. ea Planta rubri-pigmentifera, capitulo in apice complanato, subdenso. Cortex caulinus eporosus. Folia caulina ANDRUS, NEW SPECIES OF SPHAGNUM FROM NORTH AMERICA 971 ABCD Fics. 35-40. Sphag lboti 35. Habit. 36. Two branches. 37. Stem leaves. 38. Stem | Branch leaf, convex surface. A = 2 mm, 35; B = 2 mm, 36;C = 0.5 mm, 37, 39; D = 30 pm, 38, 40. ve obtusa, 0.8-1.05 mm longa, 0-0.55 mm lata, limbo ad 0-1 septatis,et fibrillis et poris carentibus. Folia ramulina = j; ] ; lselasss falas lingulato-triangularia vel lingulata, in apice plusminus basem valde distincto, cellulis hyalinis rl bif il stricta et ovato lanceolata, 1.0-1.2 mm longa, 0.3-0.35 mm lata t k = a, convexa poris quoad amplitudinem mediocribus, in apice 4-6 um usque in base minoribus quam 20 pm, in ee ; eee 7 ] 1st superficie concava ef ae praeter interdum in partibus I g I Status sexualis ignotis. Plants pink-colored; capitulum flat-topped and moderately dense. Stem red-tinged, cor- tex eporose. Stem leaves triangular-lingulate to lingulate with a more or less obtuse apex, 972 BRIT.ORG/SIDA 22(2) 0.8-1.05 mm. long = 0.5-0.55 mm. wide; border strongly developed at the base; hyaline cells rhomboidal and 0-1 septate, efibrillose and eporose. Branches with leaves 5-ranked and loosely spreading. Branch leaves small, 1,0-1.2 mm. long « 0.3-0.35 mm. wide; con- vex surface with moderate- sized pores, grading from 4-6 jim near apex to more than 20 jum at base, concave surface eporose except for occasional large round pores in proximal side regions. Sexual condition unknown. Distribution.—Western Alaska. Currently known from Bethel, King Salmon, and Koyukuk National Wildlife Refuge. Identification.—As far as other red Acutifolia within its general range, S. talbotianum is most similar to S. rubellum Wils. and S. warnstorfii. Sphagnum rubellum has a lingulate stem leaf with at least some hyaline cells 2-septate whereas S. talbotianum has stem leaves that are often triangular-lingulate and with hyaline cells that are 0-1 septate. The branch leaves on S. rubellum are also often subsecund while those of S. talbotianum are straight. Sphagnum warnstorfii looks quite similar macroscopically but has much smaller ringed pores on the convex surface of the branch leaf tips. Ecology.—Collections to date indicate this to be a quite widespread tundra species of minerotrophic peatlands. It typically forms low dense hummocks. Compared to S.rubellum Wils. and S. warnstorfii Russow, the other abundant red quinquefarious Acutifolia with which its range overlaps, S. talbotianum occurs in richer sites than S. rubellum and poorer sites than S. warnstorfii.. Common associated Sphagna include S. lenense Pohle. S. squarrosum Crome, S. teres (Schimp.) Aongs., S. brevifolium (Braithw.) Warnst., S. arcticum Flatberg & Frisvoll, S. subsecundum Nees, S. perfoliatum Savicz- Lubitskaya, S. obtusum Warnst., S.alaskense Andrus & Janssens, and S. steerei Andrus. Etymology.—Sphagnum talbotianum honors Stephen Talbot, who has collected more Sphagnum in more remote areas of Alaska than any other botanist. He deserves more credit than anyone else for our current knowledge of the distribution of Sphagnum in that state. Additional coll ined: ALASKA: Koyukuk National Wildlife Refuge, 21 Aug 1989, Tal bot Talbot 8-8- 16 (BING); Yukon Delta National Wildlife Refuge, Kuskokwim River Delta, (mixed with S. lenense) 15 Aug 2001, Andrus 9465 (BING, DUKE, NY); King Salmon, near village, 5 Jul 2001, Andrus 9522 (BING, DU “ NY); King Salmon area, ca. 39 km SW on maritime tundra, 6 Aug 2001, Andrus 9354 (BING, DUKE, NY). REPERENCE FLATBERG, KI. Studies in Sphagnum subfulvum Sjoers, and related morphotypes. Lindbergia 11:38—-54. REVIEW OF CRATAEGUS SERIES PULCHERRIMAE (ROSACEAE) J.B. Phipps RJ.O’Kennon Department of Biolog Botanical Research Institute of Texas The University of Western Ontario 509 Pecan St. London, Ontario, N6A 5B7, CANADA Fort Worth, Texas, 76102-4060, U.S.A. jphipps@uwo.ca okennon@brit.org KA. Dvorsky Dept. of Biology The University of Western Ontario London, Ontario, N6A 5B7, CANADA ABSTRACT Crataegus series Pulcherrimae (Rosaceae) is reviewed, Beadle’s group Sargentianae being incorporated. Ten spe- cies, C. sargentii, C. gilva,C. mendosa, C. pulcherrima, C. pinetorum, C. venusta, C. opima, C. tecta, C. pallens, and C. eximid are recognized. In addition, a possible eleventh species, C. ?vobur, is suggested but the evidence is not yet final on this. All species are lectotypified here except C. pinetorum, which is neotypified. Epitypes are proposed for two species, C.opima and C. venusta. Synonyms are also typified. Each species is keyed out and is represented by the most detailed description to date. Detailed line illustrations and county level range maps, in both cases generally the first for each species, are provided. Repesentative specimens are cited. Key Worps: Crataegus, series Pulcherrimae, group Sargentianae, typification RESUMEN Se revisa Crataegus serie Pulcherrimae (Rosaceae), se incorpora el grupo ma eke nae de Beadle. Se reconocen diez especies, C. sargentii, C. gilva,C. mendosa,C. pulcherrima,C. pinetorum,C. venusta C.opima, C. tect pallen y C eximia. demas se sugiere una posible undécima So oe cad ee no hay pruebas finale aun. Se lectotipifican todas las ee es C. pinetorum £ c c [ A ft Ms . ona venus iste los sinonimos. Se aport llada hasta la se S li il i detalladas y | L Lt > i ] 1d Aantiacad Lh ] 4 d ie S a) Lr I r r I if ] y of OLICCE la pera | ve The group Pulcherrimae, without formal rank or description, was first used by Beadle (1902) in the preparatory paper for his treatment in Small’s (1903) Flora of the southeast- ern United States, edition 2. In the two publications Beadle’s treatments are identical and itis notable that all but the Type species, C. pulcherrima, were described by Beadle. Beadle’s Pulcherrimae, which he treated as a group of nine species, was noticed by Sargent (1905) and was used in Palmer's (1925) nomenclator but it was not formally recognized until Robertson (1974) carefully validated the name. Beadle (1902) also recognized another group, Sargentianae, of 15 species, all described by him, that in my opinion cannot mean- ingfully be distinguished from Pulcherrimae and it is therefore here included with the latter group such that it now has 24 described species. Later, Tidestrom, in Small’s Manual of the Southeastern Flora (1933), dispensed with all the species of Pulcherrimae and Sargentianae, and henceforth floras of the region usually followed this example. A few, however, like Kurz and Godfrey (1962) did recog- nize the existence of the series and thus, a handful of its species continued to appear sporadically. This paper, based on the examination of 374 herbarium sheets from 25 herbaria and numerous field trips to the relevant region, reviews series Pulcherrimae as a whole for SIDA 22(2): 973-1007. 2006 974 BRIT.ORG/SIDA 22(2) the first time since Beadle inaugurated the groups Pulcherrimae and Sargentianae in 1902. Like other members of this series of papers, this one is a precursor to the author’s treatment in Flora of North America, vol. 9, due to be published next year. As such, it is not intended to make a deep analysis of serial boundaries or relationships but rather has the principal aim of assessing the individual species to be recognized. Nevertheless, a few obviously necessary adjustments to serial boundaries are provided in papers for this se- ries. Accordingly, lam characterizing series Pulcherrimae as being almost completely glabrous at maturity, having possession of 20 stamens, red or yellow fruit and a some- what elevated fruiting calyx. Though this diagnosis now includes Beadle’s group Sargentianae the expanded series does not appear to meaningfully widen the original diagnosis for ser. Pulcherrimae. The relationships of ser. Pulcherrimae are clearly to ser. Intricatae. Indeed, ser. Intricatae,a mainly 10-stamen group with a number of very hairy species, is not in other respects very different from ser. Pulcherrimae and the two may eventually be fused, a procedure which is, however, not appropriate to this paper. Series Pulcherrimae, whether considered in its narrow sense or in the extended sense (includ- ing Sargentianae), is restricted to the southeastern United States except for some over- flow into southeastern Texas. The approach used for this work, after the series boundaries had been settled on, was to study the maximum number of specimens possible over an approximate 15-year time frame. Accordingly, loans were requested from over 40 herbaria in the southeast plus several national herbaria with significant Crataegus collections. Only 24 institutions, however, were able to produce specimens of this series. Therefore the first author, assisted often by O’Kennon, made numerous field trips to the region to search for further speci- mens of ser. Pulcherrimae as well as other Crataegus species. The results of this field- work proved very helpful and review of the appendix of cited specimens shows that UWO now has more Pulcherrimae exemplars than the totality of specimens received on loan from all other herbaria. The specimens received on loan plus the UWO collections were then grouped into the more distinctive and repeatedly encountered morphotypes with few if any intermediates between them. These were then described in detail, where pos- sible, by matching with type material, which is well represented for this series. The names used were typified, a key to taxa recognized made and then line illustrations and distri- bution maps were produced, nearly always for the first time, for nearly all of the species recognized. The taxonomic conclusions are those of the first author while the county distribution maps were produced by K.A. Dvorsky, working from the UWO database of specimens. Difficulties working with series Pulcherrimae stem from certain problems in work- ing with Beadle’s text, particularly his imprecise description of leaf shapes as exempli- fied by his use of the phrase “leaf margins lobed and incised, leaves ovate” (or very simi- lar) for nearly all taxa, although inspection nevertheless demonstrates some clear and striking differences in leaf shape. Indeed, this shape characteristic proves to be the main taxonomic discriminator between species as recognized in this paper, rather than Beadle’s emphasis on fruit colour (red and yellow being usual in the series), fresh anther color, or flower size which do not seem to correlate sufficiently well with other characters to be generally useful. Furthermore, most collectors, including Beadle, have not recorded color characters with their label data, making them especially difficult to use with herbarium material. UWO collections are among the few that routinely record these characters. Another problem is that series Pulcherrimae does not seem to be particularly common — PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 975 except in sporadic favored locations. It is difficult to tell how much more abundant mem- bers of the series were in Beadle’s time, but, although a number of label annotations indi- cate very high abundances, ie., ‘thousands’ (very unusual today), the main collectors of this period, Beadle himself and Harbison, Biltmore’s collector, collected few specimens that survive other than the early collections associated with the protologues. Thus it is essen- tially impossible to infer the level of change in frequency over the last 100 years or so. Apomixis, polyploidy and perhaps hybridization (Phipps, 2005) contribute to taxo- nomic difficulty in Crataegus but so far no one has studied breeding systems in ser. Pulcherrimae, and only two chromosome counts or ploidy level estimates exist. Talent and Dickinson (2005) report a triploid ploidy level from an unidentified species and Longley (1924) a triploid count from C. pinetorum. One cannot generalize from so little information. Hybridization is first suggested by morphological intermediacy and as such there is no definite suggestion of it occurring in ser. Pulcherrimae. This is, therefore, a somewhat provisional treatment of ser, Pulcherrimae and be- cause of the problems noted it is necessarily grounded in the work of Beadle, the only previous worker who has paid serious attention to the group. In attempting to place all of Beadle’s 1902 and 1903 names in his groups Pulcherrimae and Sargentianae into the syn- onymy for this paper I have occasionally come up with only a possible, rather than a definite match. This itself speaks to difficulties of species level synonymy in series Pulcherrimae. Further, whereas most specimens can be more or less straightforwardly accomodated in the taxa ‘a that Irecognize here, there is a general residue of about 3% (see Appendix of Cited S ) of specimens of adequate quality where this is less straight- forward. Because of ‘this somewhat provisional state of the taxonomy, we also provide a map to the collective distribution of the series (Fig. 1). Thus, this will not be the last word on ser. Pulcherrimae and even more collecting as well as some combination of morpho- metric work, biosystematic investigation and molecular studies will be needed to stabi- lize the understanding of all its species. It is therefore to be hoped that a competent sys- tematist, preferably conveniently based in the southeast, will be encouraged by the platform presented here to carry out such work. This paper is a continuation of the first author’s studies of Crataegus in the south- eastern United States, examples of which are Phipps (1988) and Phipps and Dvorsky (2006). Because these papers were originally intended to provide the detailed backup for the first author's treatment of Crataegus in the southestern United States for Cronquist’s now defunct flora of that area, our meaning of “southeastern United States” here is iden- tical to Cronquist’s, ie. Louisiana and Arkansas are the western limits for that flora. Texan records are, however, added but only because a few species are of sporadic occurence in the extreme southeast of that state. The text continues with the formal taxonomic presentation. TAXONOMY Series Pulcherrimae Beadle ex K.R. Robertson, J. Arnold Arbor. 35: 628. ate TYPE spect oe sae oneal Ashe, Pulcherrimae Beadle, group without rank or des Bot 1:66. 1 Pulcherrimae Beadle ex cnet in clavem, Man. Trees North Amer 1:400. 1905; group ens EJ. Palmer, in clavem, J. Arnold Arbor. 5:78. 1925. Note.—Robertson attributed sectional rank for this group by Palmer (1925), but I argue that Palmer was not formally using the rank section there (Phipps, Harvard Pap. Bot., submitted) Sargentianae Beadle, group without rank or description, Biltmore Bot. Stud. 1:59.1902. 976 BRIT.ORG/SIDA 22(2) oN Sak. SE e ra are yA a oe yt x LY rar — as gs ese PT | eS asiar ay Tree RES ioasemmmen <8 ry ny, im pe a. gel = ‘ie ie a a, Shrubs or small trees; l-year old twigs shiny golden-tan to shiny deep reddish-brown; 2- yr old twigs deep gray; thorns mostly 2-3 cm long, fine, very dark at 2 yr. Leaves decidu- ous, long-petiolate; petiole glabrous, bearing a few to several, someti stipitate, glands; blades 4-7 cm long, ovate to ovate-rhombic or narrow-ovate in general outline, unlobed or more frequently with 4-6 shallow lobes per side, sometimes all of the lobes or only the upper, obscure, lobe tips blunt to acute; venation craspedodromous, 4-1] veins per side; + glabrous. Inflorescences 5-10 flowered slightly domed panicles; branches glabrous, bear- ing often plentiful, caducous, membranous, linear, gland-margined bracteoles. Flowers 14-18 mm diam.; hypanthium externally glabrous; calyx-lobes narrow-triangular, glan- dular-serrate to nearly entire; petals + circular, white; stamens 20, anthers usually pink to purple, sometimes cream (e.g. C. venusta); styles 2-5. Fruit 5(Beadle)-13 mm thick, py- riform to + orbicular, glabrous, yellow, orange, ruddy, red, red-purple or green, calyx lobes usually reflexed on a short collar; pyrenes 2-5. Mature bark deeply corrugated at least in some. Series Pulcherrimae occurs from the Gulf Coast of eastern Texas to northern Florida. Its species are collectively fairly common over much of Mississippi and Alabama as well as the western part of Georgia. There are also sporadic records east to South Carolina, one record in southern Tennessee and one locality for North Carolina in Buncombe County. Ten definite and one possible species are recognized here. PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 977 For the whole series, flowering is in April through much of Mississippi, Alabama and Georgia but, in panhandle Florida, it may start late March and in North Carolina or Tennessee in May. According to the season, typical timing may be retarded or speeded up by cooler or warmer weather than normal. The south-north progression is typical for spring flowering plants in the region (e.g. Reader et al. 1974). Fruit is ripe anywhere from late August to early October and its retention past the early stages depends on the mesicity of the site and seasonal weather until full ripeness makes the fruit attractive to frugivores. Related species of Crataegus usually show little or no habitat variation in their physi- cal habitat such that niche differentiation, if it exists, must be based on other character- istics. Series Pulcherrimae is normative for the genus in this respect as far as present ob- servation indicates and the plants are woodland edge or woodland species of mesic localities, if the latter, usually under oak or oak-mixed canopy but not in the denser shade. Soil type preference for the mesomorphic members of the genus is also not normally pro- nounced with the exception of avoiding coarse-grained or more acid soils. Species of ser Pulcherrimae are mainly recorded on the finer-grained soils. A reviewer makes the sug- gestion that the distribution of C. venusta, however, may be restricted to blackland soils,a soil type that certainly harbors hawthorns, but this cannot be confirmed from data avail- able to me. Three of the mechanisms for niche differentiation in Crataegus are: differen- tial anthesis timing, different pollinator preference and fruit differentiation (color, time of ripening, size, perhaps taste) and several of these differences do occur with ser. Pulcherrimae but have not been investigated further. Differences in anthesis time in sym- patric Crataegus may be very large, up to six weeks, but usually correlate to taxononomic distance (for more on this see Phipps 1995, on hybridization), so they likely do not ini- tiate niche differentiation. In the case of ser. Pulcherrimae different taxa that occur at the same site do sometimes have slightly different, but overlapping, anthesis time but there is no consistent recording of this. There is also an untested suggestion that differences in fresh anther color (basically, anthocyanic or not) may affect pollinator faithfulness, thus beginning niche dif iation. In this respect it is interesting that such color differences quite often appear in very closely related taxa, though they are not particularly frequent within single species. There is a huge field of study here awaiting attention by an enter- prising biologist but it clearly needs preceeding by a plausible taxonomy against which it can be tested. The deeply corrugated bark of several species, found in few other series, glabrous or nearly glabrous plant parts (except in one aberrant specimen), about 20 stamens, rather glandular-bracteolate inflorescences, usually spreading, somewhat elevated calyx-lobes in fruit, and sometimes large numbers of lateral veins in the leaf make this series distinc- tive. It is thus a quite easily recognized series, except for its differentiation from ser. Intricatae, which latter normally possesses 10, not 20, stamens as in ser. Pulcherrimae and has some very hairy species. As in series Intricatae, yellowish-fruited forms are at least as numerous as red-fruited ones. Notes on terminology: 1. This paper uses the term ‘leaf incision index, abbreviated to LII, for the relative depth of sinuses. This is measured by joining the lobe tips and projecting a line from there through the deepest part of the sinus, parallel to the adjacent veins, to the mid-vein. A one hundred percent value indicates cut through to the mid-vein 50% cut half-way, cut 25% of the way to the mid-vein 2. “Leaf? always refers to short-shoot leaves unless indicated otherwise. 978 BRIT.ORG/SIDA 22(2) KEY TO SERIES PULCHERRIMAE A He tara branches glabrous [series Pulcherrimae, sens. str.]. . Well-developed short-shoot leaf blades fairly narrow, 1.5-2 * as long as broad. 2. Well-developed short-shoot leaf-blades narrow-elliptic to narrow-ovate in general out- line;lobes absent or very shallow, perh ne notches with Ll 5% or even 10%, but if so | regularly occurring in all or most eves ee veins 6-10/side. . Leaf blades broad-elliptic to narrow-ovate in general shape; apices subacute to blunt; margins with crenate or crenate-serrate teeth; fruit red 3.C. mendosa 3. Leaf blades narrow ovate to lanceolate in general shape; apices acute to acuminate; arginal teeth sharply acute; fruit yellowish 2.C.gilva 2. Well-developed short-shoot leaf blades broad-elliptic or broad-oblong to narrow-ovate in general outline; regularly and distinctly lobed; max. LIl 10-40%; main lateral veins 4—6/ side. 4. Lobes 2-3 per side, blunt, max. LI <15% 4.C. pulcherrima 4. Lobes 2-5 per side, blunt or acute, max. LIl <15—25% or more. 5. Leaf blades narrowly elliptic to very narrow-ovate in general outline; 1-2 (—3) shal- low, acute lobes per side, max. LI usually <15% 5.C. pinetorum 5. Leaf blades narrowly ovate in general outline; 4-5 deep, acute to subacute, usually narrow-cuspidate lobes per side, max. LI] often 25% or more 9.C. incilis 1. Well-developed short-shoot leaf blades broader in proportion, generally < 1.5% as long as wide 6. Many leaves of a distinctive type, + trullate-ovate in general shape, the terminal half + straight-sided across lobe-tips; 4-6 cm long; lobes 1—3/side, sharp, but very small to small (max. LI] 2-10%(-15%)); 5-7 lateral veins/side; anthers pink; fruit yellowish 1.C. sargentii 6. Leaves of quite different construction; if with few, short, very acute lobes then blades more ovate or oblong in form and the terminal half not + straight-sided across the lobe tips; length various; lobes nearly always much more pronounced, at least at ae ve- nation various; anthers ivory to cream or pink to purple; fruit yellowish or reddis 7. nae with well-marked lobes, max. LI] usually 15-25%, ee asia more. . Blades usually smaller, 3—5(—8) cm long, ovate to broad g angular or not; anthers and fruit various colors. 9. Majority of leaves broad-ovate in general on with 4—5 lateral veins per side and 3-4 lobes;lobes short or longer (max. LIl 10-40%). 10. Lobes + blunt, generally fairly short (m3max. LI] 20-40%) 8.C. tecta 9. Majority of leaves ovate in general shape with 5-8 lateral veins per side and 4—- 6(—7) lobes; lobes fairly short, OOIUEP to acute (max. LII 10-40%) 7a.C. cf.opima 8. Blades Be 5-8 cm long, rhombovate in general shape; lobes very angular;anthers 10. C. eximia cream; fruit red 7. Leaf pede: Hn alot lobes, max LI] 5— 10%. 11. | quite shar eaf blades ee oblong to ei ovate; lobes and teeth acuminate iim? evelopment, sometimes reflex 6.C. venusta 12. Leaf blades narrow ovate to ovate-oblong; tips of lobes and marginal teeth less pronounced. 13. Leaf blades ovate-oblong in general shape; angle at leaf- — Aas ca. 90° shee — form 13, ou blades narrow-ovate in general shape; angle at leaf- ‘ne ae 60° 2.C.gilva 11. Lobes cn to subacute. 14. Leaf blades generally large at maturity, mostly 5-8 cm long; lateral veins 5-7 per side; blades nearly unlobed to shallowly lobed; bases cuneate 3a.C.cf. C. mendosa 14. Leaf blades small to medium-sized, rarely many > 5 cm long; lateral veins 4—7 per side; blades regularly lobed, but sometimes shallowly; bases cuneate to rounded. PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 979 15. Leaf blades cuneate at the base, most blades approximately 1.5:1 4.C. pulcherrima Leaf blades broad-cuneate to rounded at the base, blades <1.5:1 .C.opima B. Inflorescence branches pubescent C. craytonii ser. Intricatae] 1. Crataegus sargentii Beadle, Bot. Gaz. 28:407. 1899. (Fig. 2.) Typr: U.S.A. ALABAMA. De Kalb Co: ial Head, 20 — 1899, C.D. Beadle 1289(LECTOTYPE selected here: US 969465; ISOLECTOTYPE: US 969466). 2=C ili {le, Biltmore Bot. Stud., 1:68. 1902. Type: U.S.A. FLORIDA. Jackson Co:Chattahoochee, 8 Apr 1901, TG. Harbison H4096 (LECTOTYPE selected here: US 969401). Shrubs or small trees 3-5 m tall; extending twigs glabrous, reddish; at 1 yr. shiny deep reddish-brown, 2 yrs grayish-red, older gray; thorns 2-5 cm long, few to numerous, slen- der, straight to slightly recurved, black at 2 yrs. Leaves: petioles 25-40% length of blade, slender, glabrous, with a few sessile glands; blades 4-6(-7) cm long, ovate-trullate in gen- eral outline; tip acute to acuminate, base rounded to broad cuneate; sides barely or shal- lowly 1-3 lobed; lobes acute, max. LII usually 0-10%; upper third of leaf often unlobed; margins finely and sharply serrate with numerous very small teeth usually <1 mm long: venation craspedodromous, with 5-7 main lateral veins/side; leaves glabrous except for scattered hairs on the veins above when young. Inflorescences 4-10(-11) flowered: branches glabrous, bearing numerous, caducous, linear, often somewhat curved, mem- branous to subherbaceous (green), gland-margined bracteoles; anthesis late March (Gulf Coast) to mid-April (Alabama canyon lands). Flowers 15-18 mm diam.; hypanthium ex- ternally glabrous; calyx-lobes 3 mm long, narrow-triangular, flaring to a broad base, tip somewhat obtuse, margins entire to glandular-denticulate, abaxially glabrous; petals + circular, white; stamens 20, anthers pale pink to pale purple; styles 3-4. Fruit 8-10(-12) mm diam., subglobose, yellow or flushed pink (occ. reddish); calyx lobes reflexed, on an elevated collar; nutlets 3-4, dorsally grooved, sides plane. Common name.—Sargent’s hawthorn Habitat and Distribution.—Our records are mainly from Alabama, extending west to central Mississippi, south to panhandle Florida and eastwards with scattered records from Georgia (Fig. 3). It is a locally common species of woodland edges, cut-over wood- land and other open woodland. Comment.—Crataegus sargentii was the first species to be described in the series and this account appears to be the first to characterize it fully. It is most similar to C. gilva but that species has paseo leaves and smaller fruit. Occasional more deeply lobed forms (LU ca. 15%) may ble oth ies tosome extent, but they will normally retain the characteristic overall leaf-form of the species. 2. Crataegus gilva Beadle. Biltmore Bot. Stud. 1:60. 1902. (Fig. 4). Typr: U.S.A. ALABAMA. Marshall Co.: rocky woods, Apr 1901, T.G. Harbison 4374 (LECTOTYPE selected here: A). Shrubs or small trees 2-5 m tall; extending shoots glabrous, reddish when young: at 1 yr deep reddish-brown, older dark dull gray; thorns fine, 2-4 cm long, straight or slightly recurved, dull purplish-brown at 2 yr. Leaves: petioles slender, ca. 20-30% length of blade, adaxially grooved, + glandular; blades 3.5-4.5 cm long, broad-lanceolate to narrow-el- liptic or very narrow ovate to narrow ovate or narrow ovate-rhombic; tip acute to acumi- nate and with 0-3 acute lobes per side these with max. LII usually < 15%: margins sharply fine-serrate or occasionally with quite large teeth; venation craspedodromous, lateral veins ca. 5-7(-9) per side on average or larger short-shoot leaves; glabrous above and be- 2) — BRIT.ORG/SIDA 22 ar, 5 SOONG aig A NS 980 TIVAINY h\UWY), 1cm.S. Laurie-Bourque del. single leaf at left from extension shoot. Scale bars PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 981 yee Ee ses ecetemeta! roe & | oie eemayeqwes’ 2 Shed @hancacey Her ee eh OOS TE m4 av eteree py SRO low except for some hairs on the veins above especially when young. Inflorescences usu- ally 3-7 flowered; branches glabrous, densely beset with caducous, narrow-oblong to very narrow obovate, membranous to subherbaceous, densely gland-margined bracteoles. Flowers (13-)18-20 mm diam. hypanthium externally glabrous; calyx-lobes 6-7 mm long, narrow triangular, margins glandular-serrate; petals + circular, white; stamens 20, an- thers pale pink; styles 3-4. Fruit ca. 5-8 mm wide at maturity, + orbicular, greenish yel- low to yellow; sepals + reflexed; nutlets 3-4. Habitat and Distribution.—Crataegus gilva occurs from Mississippi to Georgia and northern Florida (Fig. 5), in woodland edges and gaps, clear-cuts and pine-oak woodland. Comment.—Crataegus gilva leaves are rather like a sharp-tipped C. mendosa and and somewhat resemble a much less clearly lobed C. pinetorum. The leaves are also usually somewhat like a proportionately narrower C. sargentii and in the flowering Type they are narrow-elliptic, acuminate at the tip, cuneate at the base, virtually devoid of lobes and have particularly large marginal teeth, this being the most extreme specimen of the species seen. The suspicion that C. gilva may represent an extreme form of C. sargentii is heightened by the almost complete lack of known fruiting specimens of the species, the fruiting co-type excepted (Harbison 4374/2), which specimen, therefore, does suggest the possibility that the leaves often widen after anthesis. 3. Crataegus mendosa Beadle, Biltmore Bot. Stud. 1:65. 1902. (Fig. 6). Tyre: U.S.A. ALABAMA. Mar- shall Co.: near Albertville, 18 Oct 1901, [.G. Harbison 5219 (= 4336/2) (ectotype selected here: US 969431). 982 BRIT.ORG/SIDA 22(2) C. gilva C, pinetorum Fic. 4 g £1 ry I + Fe a! +. g 9 fT, {¢. | \£ J JTIMAINY ial! ipp In0907 (UWO) Ty pinetorum (bot- tom row) from, left to right, Lance 2174 (UWO), Phipps 8173 (UWO) Phipps & 0 8042 (UWO). Scale bars = 1 cm. S. Laurie- Bourque del. Shrubs or small trees 3-6 m tall; bark of trunk not recorded; l-yr old twigs shiny reddish or purplish brown, older dark or dull gray; thorns 2-5 cm long, fine, straight or slightly recurved, reddish black at 1 yr, frequent to absent. Leaves: petioles 30-50% length of blades, dorsally furrowed, bearing a few small glands; blades +-7 cm long, 2-3 cm wide, broad- oblong to elliptic or narrowly ovate; unlobed or with a few obscure lobes; apex subacute to blunt on some larger leaves, cuneate at the base; margins with numerous, small, sharp teeth; lateral veins 8-9 per side (fewer on smaller leaves), venation craspedodromous; gla- brous except for a few hairs on the midvein abaxially when young, dark green. Inflores- cences 4-7 flowered; branches glabrous, densely beset with caducous, narrow, semi-her- baceous and green to membranous, gland-margined bracteoles. Flowers 14-17 mm wide; hypanthium externally glabrous; calyx lobes tr lang ular, abaxially glabrous, margins glan- dular-denticulate; petals + circular, white; stamens 20, anthers pink or pink-purple; styles 3-5. Fruit 8-10 mm diam., + orbicular to broad ellipsoid, yellow-green with pink or salmon blush, or red (Beadle); calyx lobes reflexed; nutlets 3-5, dorsally furrowed, laterally smooth. Habitat and Distribution.—Scattered from extreme southeastern Texas to west- ern Georgia and northern Florida with a disjunct record in South Carolina (Fig. 7), Crataegus mendosa is a plant of woodland edges, cut-over woodland and brushy places. Comment.—Crataegus mendosa is one of the more distinctive members of the series although it is approached in form by C. gilva, which, however, has an acuminate leaf-tip and much sharper lobes (if present) and marginal teeth. The extension shoot leaves of C. mendosa are ovate to broad-ovate in outline with 3-4 sub-acute to acute lobes that are usu- ally quite broad and this can confuse if there are only few typical short-shoot leaves present. PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 983 Number of Records 0 Ty 4 - Crataegus gilva LUS Z Y a + ! | ee pare ae + rf; +. LS £. eo | Pl Fic. 5 y 3a. Crataegus cf. mendosa Beadle Shrubs or small trees 3 m tall; mature bark not recorded; extending twigs glabrous, red- dish; at l-yr shiny reddish-brown; older duller dark gray; thorns often on material seen. Leaves: petioles ca. 33% length of blade, glabrous, glandular, adaxially grooved; blades 5- 7 cm long, mostly broad-elliptic in general shape; apex subacute, base cuneate to broad cuneate; 0-3 lobed per side, lobes obtuse to subacute, max. LII usually 5-10%; margins serrate; venation craspedodromous, ca. 6 main lateral veins per side; both surfaces gla- brous. Inflorescences not recorded. Young infructescences with 2-5 fruit, branches gla- brous. Young fruit 7-10 mm diam., subglobose to + pyriform, green, glabrous; filament bases ca. 20; sepals mainly erose; nutlets ca. 4. Habitat and Distribution.—Only known from one locality in Alabama, where it is recorded from ‘bottoms’ from where three fine duplicates at DUKE, GA and NO of Demaree 50656 (MS: Alabama: Dallas Co., White Oak Creek, PO. Sardis, 29 June 1964) show a plant intermediate between C. mendosa, C. opima and C. pulcherrima but which cannot be confidently placed with any. More material is needed. 4. Crataegus pulcherrima Ashe, J. Elisha Mitchell Sci. Soc. 16:77. 1900. (Fig. 8). Typr; US.A. FLoripa. Gadsden Co: River Junction, 10-11 Aug 1895, G.V. Nash 2377 (LECOTYPE selected here: DOV 4634; ISOLECTOTYPE: US 250142). have selected from Ashe’s co-types the specimens with better foliage and fewer fruit rather than more fruit and poorer foliage. BRIT.ORG/SIDA 22(2) al, 8088, Smith 129 and Lance 2176 (UWO). Scale bars = 1 cm. S. Laurie-Bourque del. PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 985 FEE Sag ils foes pe TE 2 Spl egy 2 ane ae Man ar sn Sait eee tae Se aaa lgee i at BOA ?=Crataegus robur Beadle, Biltmore Bot. Stud., 169.1902. Type: U.S.A. FLORIDA. Leon Co.: Tallahasee, 18 Sep 1901, C.D. Beadle 2051 (LECTOTYPE selected here: A). Shrubs, 2-4 m tall; extending twigs glabrous, green or reddish-tinged; at 1 yr. reddish- black; at 2 yr. dark gray; thorns at 2 yr. 2-3.5 cm long, blackish, fine, + straight. Leaves: petioles 1-2 cm long, slender, glabrous, glandular, glands sessile or stipitate; blades 2.5- 5.0 cm long in our material, narrow ovate or + oblong through narrow obovate or nar- row rhombic in general shape, with 2-4 shallow obtuse to subacute lobes per side; tips acute, bases cuneate; margins serrate with small teeth, somewhat distant for their size; venation craspedodromous with 4-5(-6) main lateral veins per side; glabrous above ex- cept for along the main veins adaxially young, glabrous below; thin. Inflorescences 4—7- flowered with a very short central axis, branches glabrous, bearing plentiful, small, ca- ducous, oblong, membranous, stipitate-gland-bordered bracteoles; anthesis mid-April. Flowers 15-20 mm diam.; hypanthium externally glabrous; calyx lobes 3-4 mm long, narrow-triangular, margins glandular-serrate, abaxially glabrous, tip obtuse; petals + circular, white; stamens 20, anthers small, pink; styles ca. 3. Fruit in clusters of 2-5, 6-7 mm diam., glabrous, deep yellow sometimes flushed red, occasionally purplish-red; ca- lyx lobes reflexed, on a short collar; nutlets ca. 3, dorsally grooved, sides plane. Common name.—Beautiful hawthorn. Habitat and Distribution.—Mississippi to Georgia and northern Florida (Fig. 9). Open, rocky woodlands JBP) and swamp borders (Beadle), scarce. BRIT.ORG/SIDA 22(2) Nitti h /— A Se i a ee \ tu” J, 7 NY, Ps ape tie Mm Na He ae Lt: ( a % oe if 4 hipps et al. 8091 (UWO) Scale bars = 1 cm. S. Laurie- Bourque del. PHIPPS ET AL., CRATAEGUS SERIES PULCHERRIMAE 987 Vitae por ee om ET Ph EP KL AA OT i S HE RR ee ee ee gitqeSniomim meer “te ermaueieace esiere 3 nue BRaSISanay Cee Geecor eee an oa %G hae che me nee = ai Bere = be int oy = st) is Pilg rn ib Beale parat ee email ane ae & nee hen aa Vere yA ee Ss ce bee ee i fa 41, 1] I dictrihit? £ frat, I-fh i f, ll d qd Fic. 9. County p | t Comment.—The type species of its series was described by Ashe as having 10 sta- mens but neither of Ashe’s co-types (Nash 2377 and Nash 2568) from River Junction, Florida confirm this. This species is usually quite easy to distinguish though occasion- ally it resembles Crataegus cf. opima. 5. Crataegus pinetorum Beadle, Bot. Gaz. 28: 343. 1900. (Fig. 4). Type: U.S.A. ALABAMA. De Kalb Co.: De Soto State Park, 25 Sep 2001, R. Lance 2174 (NEOTYPE selected here: UWO; ISONEOTYPES: Anniston Nat. Hist. Mus., UNC, JSU). Shrubs or small trees 2-6 m tall; extending twigs glabrous, with a strong reddish cast; at lyr shiny reddish-brown; dull gray-brown at 2 yr; older dull gray; thorns usually present, mainly 2-35 cm long, fine, + straight, shiny black or reddish-black at 1 yr, becoming black. Leaves: petioles 25-30% of leaf blades, glabrous, sparsely to densely glandular or even stipitate-glandular, abaxially grooved; blades 4-6 cm long, nearly full-grown at an- thesis, narrowly ovate to elliptic in general shape; tip acute, base cuneate; lobes usually 2-3 per side, fairly shallow (max. LI] 10-20%), acute; marginal teeth extremely small, acute; venation craspedodromous, main lateral veins 5-6 per side; glabrous adaxially and abaxially. Inflorescences 2-4-flowered; branches glabrous, bearing plentiful, + persis- tent, very narrowly obovate, chartaceous to subherbaceous, densely gland-margined bracteoles. Flowers at anthesis not known; hypanthium externally glabrous; calyx lobes ca.6 mm long, narrow triangular, margins finely glandular-serrate, abaxially glabrous; petals not known; stamens 20, anther color not known; styles porly persistent. Fruit 12- 988 BRIT.ORG/SIDA 22(2) sae ee The role of Root Exudates in Rhizosp Genetics of Meiotic Prophase | in Plants Biology and Biochemistry of Glucosinolates Bioinformatics and its Applications in Plant Biology Leaf Hydraulics Plant Uncoupling Mitochondrial Proteins Genetics and Biochemistry of Seed Flavonoids “ytokinins: Activity, Biosynthesis, and Translocation Global Studies Cell Type-Specific Gene Expression in Plants Mechanism of Leaf-Shape Determination Mosses as vee Systems for the Study of Metabolism and Development Structure and Function of Photosystems | and II Glycosyltransferases of Lipophilic Small Molecules Protein Degradation Machineries in Plastids Molybdenum Cofactor oo and Molybdenum Enzymes Peptide Hormones in Plant Sugar sensing and ee in Plants: Conserved and Novel Mech Vitamin Synthesis in Plants: i and Carotenoids lanisms c Plastid-to-Nucleus R r etrograde Signalin Phe Genetics and Biochemistry of Floral Pigments Transcriptional Regulatory Networks in Cellular Responses and Tolerance to Dehydration and Cold Stresses Pyrimidine and Purine Biosynthesis and Degradation in Plants Phytochrome Structure and Signaling Mechanisms Microtubule Dynamics and Organization in the Plant Cortical Array SIDA 22(2): 1026. 2006 MORPHOMETRIC ANALYSIS OF AN AMELANCHIER (ROSACEAE: MALOIDEAE) COMPLEX ON THE DELMARVA PENINSULA (DELAWARE, MARYLAND AND EASTERN VIRGINIA) RESOLVES THE TAXONOMIC IDENTITIES OF AMELANCHIER OBOVALIS AND A. CANADENSIS Christopher T. Frye Maryl iat Depaminent of Natural Resources fe and Heritage Service ‘Netaral Heritage Program 90 e Mills Road Wye Mills, Maryland 21679, U.S.A. cfrye@dnrstate.md.us ABSTRACT Amelanchier obovalis and A. canadensis (Rosaceae: Maloideae) are similar taxa with overlapping geographical ranges on the Atlantic Coastal Plain of the United States. Conflicting circumscriptions, aces olen tion, and agamosperm y have contributed to the taxonomic confusion in these species. In this study I per- form a morphometric examination of nese two species onthe C eastal igh oe Deewale eee and the east- ern shore of Virginia—coll Peninsula. 1d tof eleven flowering, fruiting and vegetative characters cee from marked individuals. A second dataset contains six minimally correlated aes that were obtainable from the type specimen of Amelanchier obovalis. | use Principal Components Analysis (PCA) to summarize phenetic similarities and dissimilarities between individuals and agglomerative, ierarchical Cluster Analysis to aid in visualization of morphological groups. Selection of groups (putative taxa) PCA and to inform a vtestabie Sroup, model was Peon? aoe tvery using concordance in phenetic methods cl four-group model for which I found a sig- iff rall multivariate eff 1 (P Pillai’s Trace, F = 7.29, p> ee < 0.0001) using Multivariate Analysis of Vari- ance (MANOVA). Two well- oe groups (1 & 4) corresponded to Amelanchier obovalis and Amelanchier canadensis. The remainders of the sample are intermediate between Amelanchier obovalis and Amelanchier canadensis and may be the product of recurrent hybridization between these taxa. Lack of clearly delimited morphological character space and apparent ecological and genetic were in these taxa suggests the taxonomic rank of variety under the earliest combination in Amelanchier, e.g, Amelanchier canadensis var. canadensis and Amelanchier canadensis var. obovalis as originally ee by Michaux (1803). I present a taxo- nomic treatment and historical analysis of the species taxonomy and a dichotomous key to the varieties. RESUMEN Amelanchier obovalis y A. ca spiaetiala (Rosaceae: Maloiceae) son taxa psunllatcse con areas eee solapadas nlall dos Unidos. I polip oidia, hibridacion y ; ae psn nape copula: ala confusién taxonémica en estas especies. En este estudio realizo pecies en la parte costera as Delaware, Narang y costa este de Vir- 1] J a] ct Dp, < g Pee |e oa et omen ginia de flores, frutos y vase latipes pomaces de ane» pecseeie Un segundo onjinte de datos contiene seis écimen tipo de ace CG eae is. es -] Caer (c | el Analisis Componentes Principales (PC y un Analisis de Clusters jerarquizado aes ayudar en la visualiz zai ws grupos eis ase La seleccion de jelecde grupos (taxa putativos) para informar de un m usando conecraanci en los métodos renences (PCA y on ter mg) ye tadistica eecenpie inferencial. analisis da un e,F=7.29, P > F < 0.0001) usando Andlisis de Varianza Multivariante (MANOVA). Dos grupos muy ek (1 & 4) correspondieron a Amelanchier obovalis y Amelanchier canadensis. Los restos de la muestra son intermedios SIDA 22(2): 1027- 1048. 2006 1028 BRIT.ORG/SIDA 22(2) — entre ‘cious rier apovalisy mmelane/aer Eanes nsis y pueden ser is Progucto de hi bridacion recurrente entre estos taxa. | } y genético en estos taxa en el rango de vate ad bajo la combini acion mas antigua en Amel lamchier, e.g., Amelanchier canadensis var. canadensis y Amelanchier canadensi: a oe aux aves Presento un tratamiento taxonémico y amalisis historico de la taxonomia de ge especies y ra las variedades. INTRODUCTION Amelanchier Medic. (Rosaceae: Maloideae) is comprised of deciduous small trees and shrubs of the North Temperate Zone (Phipps et al. 1990). Amelanchier flowers are pen- tamerous, radially symmetrical and protogynous (Robinson 1982; Campbell et al. 1991) arranged as inflorescences of simple racemes (except in A. bartraminana). Amelanchier fruits are globose, berry-like pomes that attract a wide variety of frugivorous dispersers (Robinson 1986). The leaves of Amelanchier, and of the subfamily Maloideae in general, vary greatly in size, shape and dentation, and mature foliage on short shoots is often very different from those on juvenile (e.g., new sucker growth) or long shoots (Robertson et al. 1992). The total enumeration of taxa varies among authors. For example, in the north- eastern United States, Landry (1975) recognized six species; Fernald (1950a) includes 19 species but Gleason and Cronquist (1991) only nine species. Synonymy within Amelanchier is formidable, reflecting uncertainty surrounding taxonomic concepts of species and infraspecific taxa (see Fernald 1941; 1946). Reproductive biology of Amelanchier is characterized by polyploidy, agamospermy and extensive hybridization between most eastern North American species (Wiegand 1912; 1935; Jones 1946; Cruise 1964; Campbell et al. 1987; Weber and Campbell 1989; Campbell et al. 1991; Dickinson and Campbell 1991; Campbell and Wright 1996; Campbell et al. 1997b; Campbell 1999). Recent research has revealed several complex taxonomic issues arising from mating sys- tem versatility in the genus including the presence of persistent, morphologically dis- tinctive hybrids, e.g. Amelanchier x neglecta (Weber and Campbell 1989; Campbell and Wright 1996), microspecies derived from intraspecific hybridization followed by agamo- spermy,e.g., Amelanchier “rubra” (Dibble et al. 1998) as well as wide morphological varia- tion among populations of single species (Cruise 1964, Campbell et al. 1997a). Reproduc- tive versatility in Amelanchier, ie., the combination of vegetative spread, interspecific hybridization and formation of agamic complexes has contributed to their phenotypic diversity and ecological success, particularly in early successional environments (Wiegand 1912, 1935; Cruise 1964; Robertson 1974; Campbell et al 1991; Campbell 1999). Despite the abundance and ecological importance of Amelanchier in the southeast- ern United States this genus remains poorly studied. This may be due, in part, to the lim- ited field work that the primary authors of the genus (e.g., K.M. Wiegand, G.N. Jones, ML. Fernald) performed in the southern states but probably more persistently by the diffi- culty in gathering adequate material for study, and the limited usefulness of herbarium material, including some type specimens, (Wiegand 1912; Nielsen 1939: Fernald 1946; Jones 1946; Cruise 1964). In a long-term study (1999-present), mainly of the stoloniferous members of Amelanchier, the most problematic species for confident identification in the field and her- barium have been Amelanchier canadensis (L.) Medic. and Amelanchier obovalis (Michx.) Ashe. Amelanchier canadensis and A. obovalis occur along the east coast of North America with overlapping ranges in the southeastern United States. Herbarium sheets labeled as these species often have multiple annotations and the opinion expressed of ten seems cor- FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX 1029 related with the general habitat types written on the label and notes on plant habit or height noted by the collector and not their morphological characteristics (C. Frye, pers. obs.). Three species in addition to Amelanchier obovalis and A. canadensis are listed in flo- ras (Tatnall 1946, Brown and Brown 1980) specifically covering the geographic region known as the Delmarva Peninsula (Fig. 1), which is comprised of eastern Maryland, Dela- ware and eastern Virginia (hereafter referred to as the study area). Amelanchier arborea (Michx. f.) Fernald is a species of dry, upland forests that is ex- cluded in this analysis. Amelanchier intermedia Spach, is restricted to the Piedmont and Mountain physiographic provinces (Wiegand 1920) and was not considered here. Amelanchier stolonifera Wieg. is cited by Tatnall (1946) from the Piedmont physiographic province in Delaware (not included in the study area). Amelanchier canadensis (L.) Medic. var. subintegra Fernald (Fernald 1950b) has been observed by the author in the study area but no specimen corresponding to this taxon occurred in the sample. Shrubs and small trees that key to Amelanchier canadensis and A. obovalis are wide- spread and common in the study area but many characters purported to separate the species in taxonomic keys appeared highly variable, temporal, or were the subject of dis- agreements among authors. Characters such as plant habit (stoloniferous and colony form- ing versus clumped and multi-stemmed), plant height, and ovary pubescence that are often used in major couplets to distinguish these species I found were difficult to objec- tively determine. For example, nearly all individuals in the study vigorously sucker but under certain conditions such as mowing and burning form low, shrubby colonies with apparent vegetative spread. Long observation (1999-present) of Amelanchier in the study area suggested the presence of at least two coherent taxa; however, several morphologies could not be identified with existing taxonomic keys (Wiegand 1912; Fernald 1950a; Rad- ford et al. 1968; Gleason & Cronquist 1991). Taxonomic complexity in the study area may reflect the presence of numerous distinctive microspecies produced by hybridization events followed by asexual seed production and/or cryptic hybridization involving a number of agamosperms that would be difficult to detect using morphological charac- ters present at a single phenological stage. This study aims to determine whether Amelanchier occurring on the Delmarva Pen- insula exists as a complex of intergrading, minimally separable phenotypes or contains two or more well-defined taxa that are significantly different from each other in mor- phology, ecology or both. In order to examine morphological complexity and evaluate the various treatments of Amelanchier canadensis and A. obovalis | developed two datasets of morphometric characters. The first dataset comprised a “full” set of flowering, fruiting and vegetative characters taken from marked individuals (Full dataset). The Full dataset was used to explore multivariate relationships between individuals, reduce dimension- ality and select taxonomically informative characters. The second dataset comprises a reduced set of characters that includes only those characters present on the type speci- men of Amelanchier obovalis (Type dataset). The Type dataset is analyzed to determine the taxonomic identity of Amelanchier obovalis. A similar analysis using the type of ible given the few taxonomically informative char- Amelanchier canadensis was not p acters present. Due to the complex classification history of Amelanchier canadensis and A. obovalis I defined groups for analysis a posteriori, without taxonomic classification so as to avoid confounding a priori taxonomic identity with the partitioning of morphological varia- tion. | use a combination of Cluster Analysis and Principal Components Analysis (PCA) BRIT.ORG/SIDA 22(2) Fall Line 1030 - Western Delaware Pee ae Line Delaware N \ . Be a ‘gia Maryland |7} _ $ y we hath rd ae « Prey 6s B { Sussex as, Fee ts Cath if ca nt . \ a Dorchester Atlantic Le en Ocean KE, ry tf Wicomico Ob rh a — ee he Le Ff te tr NS oa Re oor Seenee me Worcester ms PB a r » " Fy > ¢ a sen -Accomack ‘. Virginia 2 Northampton Eastem US. ~N } J Fin 1G g phi VW re Let Creal TL Nal D 1 fel W Ic n.l TRI r \ - RA | A A rs VV JA I Jal LL ia 7 y g I FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX 1031 to delimit groups (putative taxa) and Multivariate Analysis of Variance (MANOVA) to test the best-supported group model. Multivariate analysis of morphological data has been used to resolve species complexes in Amelanchier (Dibble et al. 1998), classify taxa within groups defined a priori (Saarela et al. 2003; Leonard et al. 2005) and to resolve difficulties caused by morphological overlap in infraspecific taxa (Whang et al. 2002). The taxonomic treatment and arguments developed herein are presented in a his- torical context beginning with André Michaux’s Flora Boreali-Americana (1803) and relying heavily on the landmark treatment of Amelanchier by Karl Wiegand (1912). METHODS Study Site—The Delmarva Peninsula comprises the eastern shore of Maryland, all of Delaware (New Castle, Kent and Sussex Counties) and the eastern shore of Virginia (Accomack and Northhampton Counties) bounded on the west by the Chesapeake Bay and the Susquehanna River and on the east by the Atlantic Ocean (Fig. 1). The Delmarva Peninsula lies south of the fall-line, a term used to describe the boundary between the Piedmont Province and the Atlantic Coastal Plain and falls entirely within the Outer Coastal Plain Mixed Forest Province of Bailey (1995). Elevations on the peninsula are mostly less than 20 m ASL and are underlain by relatively young Tertiary and Quater- nary sediments of uniformly low relief (Schmidt 1993). It is mostly within the Oak-Pine forest region, with the northernmost counties transitional to the Oak-Chestnut region (Braun 1950). Significant habitat features include extensive fresh and salt tidal marshes, blackwater river swamps, barrier islands and Delmarva Bays (seasonal ponds). The cli- mate of Delmarva is moderated by the Delaware Bay, Chesapeake Bay and the Atlantic Ocean and is characterized as having cool winters and warm, humid summers. Character Selection and Sample Size Determination.— used data from two charac- ters: (1) length of the flowering inflorescence [hereafter abbreviated as Flinfllen] and (2) length of the lowest pedicel on the flowering raceme [hereafter Flpedlen] as a model for determining sample size requirements in the field study. These two characters have been used by many authors (Ashe 1903; Jones 1946; Fernald 1941, 1950; Robertson 1974; Wiegand 1912, 1920) to describe Amelanchier obovalis and A. canadensis and were readily obtainable from herbarium sheets. I took measurements of Flinfllen and Flpedlen from 54 accessions of both from herbarium specimens borrowed from CM, DOV, FLAS, GA, MARY, NA, NCU, PH, US, USEH Vv PL (acronyms follow Holmgren et al. 1990) and of personal collections from Delaware, Maryland, North Carolina and Virginia. Flpedlen had the lowest dispersion of scores (G = 11.88 mm, o = 5.46 mm, N = 119) and the ratio of the standard deviation to the mean, expressed as a percentage, (Coefficient of Variation) was 46%. Iterative sample size analysis (Sokal and Rohlf 1968, p. 247) converged on a sample size > 53 that would be sufficient for an 80% certainty of determining a 25% dif- ference between means at the 5% significance level by Analysis of Variance. For the field study I chose an additional thirteen characters that I determined might be informative in describing variation in the complex: (1) the proportion of the length of the flowering inf] mprised by the lowest flowering pedicel [Flped_flinfl]l, (2) ovary pubes- cence during flowering [Flovpubl, (3) flowering sepal position [Flsepos], (4) length of the fruiting inflorescence [Frinfllen], (5) length of the lowest pedicel on the fruiting raceme [Frpedlen], (6) the proportion of the length of the fruiting inflorescence comprised by the lowest fruiting pedicel [Frped_frinfll], (7) ovary pubescence on the developing fruit [Frovpub], (8) sepal position on the developing fruit [Frsepos], (9) petal length [Petlen], 1032 BRIT.ORG/SIDA 22(2) (10) petal width [Petwid], (11) the ratio of petal length/petal width [Petl_wratio], (12) the leaf length/width ratio[Lfl_wratio]and (13) the ratio of the widest point of the leaf rela- tive to total leaf length [Lfwidptl. Table | provides a complete list of characters, their de- scriptions and measurements. The means of five replicates per character on each indi- vidual were calculated and used in subsequent analyses for all quantitative characters. Of the total 15 characters, four were multi-state, categorical variables: flowering and fruit- ing sepal position [Flsepos, Frsepos] and flowering and fruiting ovary pubescence [Flovpub, Frovpub}| these characters are not included in analysis of variance but were collected for descriptive purposes and as a preliminary sample for future analysis. Sampling.—During the late winter and early spring | marked 75 individuals of Amelanchier with permanent tags on a north-south transect from Cecil County, Mary- land (39 32' 38" N) to Northhampton County, Virginia (37 08' 45" N). Additionally, I in- cluded a sample from within the core range of Amelanchier obovalis (Bladen County, North Carolina) matching the description of Amelanchier obovalis by Fernald (1950a) and Radford et al. (1968). I did not attempt to stratify the sample among disturbance lev- els, although in order to examine any correlations with disturbance and to avoid ecologi- cal bias in selecting the sample I marked individuals in both natural habitats (interior of swamp forests, hummocks along fresh-tidal rivers, pine flatwoods) and disturbed habi- tats (roadside verges, clearcuts, and powerlines). I returned to each marked individual during peak flower and mid-fruiting stages. I defined the peak flowering period as the stage when all flowers on the raceme had expanded and mature petals could be easily removed from the upper flowers. I defined the early fruiting stage as the period when the ovary on the developing fruit had swollen but the fruit was still green or just coloring and mature leaves were available. Additional effort was made to examine variation in elongation of the lower pedicels over time, a character noted by several authors (Ashe 1903; Wiegand 1912; Fernald 1941; Jones 1946) and one that I thought might be informative for understanding the various taxonomic concepts of Amelanchier obovalis and A. canadensis. This additional charac- ter isnot used in the multivariate analysis. lexamined the difference between mean length of the lowest pedicel at peak flower and at mid-fruit by analysis of variance. Inflores- cence and pedicel measurements were taken in the field, all other measurements were taken in the lab. All measurements utilized a digital caliper and an ocular micrometer scaled to tenths of a millimeter. Vouchers of flowering and fruiting material were col- lected from marked individuals at each site; these are deposited at the Maryland Depart- ment of Natural Resources Herbarium (Tawes), Annapolis, MD. Sampling began on 18 April 2005 and continued through 26 May 2005. Flowering, fruiting and leaf measure- ments were successfully gathered for 61 individuals. Quantitative Morphological Analyses.—My overall objective for the analysis of quan- titative morphological characters is to test the null hypothesis of no significantly distin- guishable groups of individuals. To test the null hypothesis I looked for discrete clusters using Principal Components Analysis (PCA) based on the correlation matrix of eleven (Full dataset) and six (Type dataset) quantitative characters standardized by log trans- formation. | also used the Full dataset to reduce the dimensions of the dataset by elimi- nating highly correlated characters (Pearson's r >0.7). Proportion of variance explained by each axis was compared with that expected under a broken-stick model Jackson 1993). I plotted the first principal component against the second to graphically portray any relationships among individuals. Additionally, I created a second matrix from an FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX Taste 1.List of the 15 character acronyms, their descriptions and measurement. All measurements taken to the nearest tenth of a millimeter. Character acronym Description and Measurement Flinfllen Length of the flowering inflorescence at peak flower. Measured from the base of the distal-most flower on the raceme to point of insertion on the stem Flpedlen Length of the lowest pedicel on the flowering raceme at peak flower. Measured from the base of the ovary to point of insertion on the raceme. Flped_flinfl Proportion of the length the flowering raceme comprised by the lowest pedicel. Flovpub vary pubescence on the terminal flower of the flowering raceme scored from 1 (glabrous) to 4 (densely tomentose) Flsepos Sepal position on the terminal flower of the flowering raceme scored along a scale from 1 (erect) to 4 (tightly reflexed) in comparison to an imaginary transverse plane through the ovary. Frinfllen Length of the fruiting inflorescence. Measured from the base of the distal-most fruit on the raceme to point of insertion on the stem Frpedlen Length of the lowest pedicel on the fruiting raceme at mid-fruit. Measured from the Frped_frinfl Frovpub base of the ovary to point of insertion on the raceme Asem of the length of the fruiting raceme comprised by the lowest pedicel. vary pubescence on the terminal fruit of the fruiting raceme scored along a scale an 1 (glabrous) to 4 (densely tome Nn cy Frsepos Sepal position on the terminal fruit of the fang raceme scored along a scale from 1 (erect) to 4 (tightly reflexed) in comparison to an imaginary transverse plane through the ovar Petlen Length of mature, expanded petals measured from distal-most point to insertion 1033 on the hypanthiu Width of mature smpahued petals measured at the widest point. Ratio of petal length/petal width. Lfl_wratio Ratio of leaf length/leaf width. Lfwidpt Ratio of the widest point along the leaf axis (measured from base)/total leaf length. agglomerative, hierarchical Cluster Analysis using Euclidean distance and Ward’s method of linkage (McCune & Grace 2002). This second matrix was used as an overlay on the PCA scatterplot to assist in visualizing groups. PCA and Clustering were performed on Pcord version 4.14 (McCune & Mefford 1999). A separate objective was to determine the a (group membership) of the type specimen of Amelanchier obovalis in multivari space. 1 performed both PCA and Clustering based on the correlation matrix of six quantitative characters derived from analysis of the Full dataset and that were also obtainable from the type specimen: Petlen, Flinfllen, Flpedlen, Flped_flinfl, Lfl_wratio, and Lfwidpt. The Type dataset was organized into a group by character matrix and exported to SAS version 9.1 (SAS Insti- tute, Cary, NC, USA) for Multivariate Analysis of Variance (MANOVA, GLM procedure). MANOVA tests for an overall multivariate effect of the group model and performs pair- wise comparisons between character means (mean vectors) for all groups for each char- acter. Approximate univariate normality of morphometric characters within each group was assessed by inspection of frequency histograms, accepting normality with skewness < 1 (McCune & Grace 2002). MANOVA is robust to multivariate non-normality and Levene’s test was used to ascertain homogeneity of variances within each group. MANOVA is used to test the null hypothesis of no significant differences between the mean vectors of each group. 1034 BRIT.ORG/SIDA 22(2) Assessment of Taxonomic Rank.—| utilize the concepts of genetic and ecological exchangeability introduced by Crandall et al. (2000) to assess population distinctive- ness and extend that method to assessing taxonomic rank. Species populations are ge- netically exchangeable if there is evidence of ample gene flow between them. Ecological exchangeability is demonstrated if individuals can be moved between populations and can occupy the same ecological niche. Exchangeability is viewed from both recent and historical perspectives, e.g., two taxa may currently experience gene flow due to break- down of intrinsic or extrinsic barriers via ant} causes but the populations may have historically experienced (e.g., presettlement) little gene flow. RESULTS Full Dataset.—Eigenvalues for the first two axes, representing 56% of the total variation, exceeded their broken-stick values and were chosen for interpretation (Table 2). PCA-1 comprises > 37% of the variance in the dataset. Five morphological characters ordinate individuals along PCA-1. These are, in descending order in the loading matrix (Table 3): Flpedlen, Frpedlen, Frinfllen, Flinfllen and Petlen. PCA-1 is thus a size axis. PCA-2 com- prises > 19% of the variance in the dataset. Five morphological characters ordinate indi- viduals along PCA-2. These are, in descending order in the loading matrix (Table 3): Flped_flinfl, Frped_frinfl, Lfl_wratio, Flinfllen and Frinfllen. PCA-2 is chiefly a shape axis. The characters most highly correlated with both PCA axes are also very highly cor- related with each other, in particular, flowering and fruiting stages of the inflorescence are redundant (Table 4). Flowering characters were selected for the final matrix. Type Dataset.—Six characters were used in the analysis of the Type dataset: Petlen, Flinfllen, Flpedlen, Flped_flinfl, Lf{l_ Wratio and Lf widpt. These six quantitative char- acters were obtainable from the type specimen of Amelanchier obovalis and serendipitously mirrored the set of minimally correlated characters derived from analy- sis of the Full dataset with the exception of Flpedlen (r = 0.708), which I maintained because of its use as an informative character in most treatments. The dendrogram pro- duced by Cluster Analysis (Fig. 2) is scaled by an objective function that measures the loss of information at each step; as groups are fused the amount of information decreases until all groups are fused and no information remains (McCune & Grace 2002). In the iterative process, five and higher order groups were composed of single (outlier) indi- viduals and selection of three groups combined groups 2 and 3 into a single membership cluster. I chose to overlay a four-group cluster on the PCA as this appeared to be the maxi- mum number of well-supported groups (e.g., long branches of the dendrogram) and cor- responded to my field observations of the number of putative morphological forms prior to sampling and roughly to the different habitat types that were sampled (e.g., tidal and nontidal swamps, low Pine woodlands, and sand ridges). The PCA scatterplot with Clus- ter overlay (Fig. 3) shows a single distinct cluster at the far left of PCA-1 and central to PCA-2. The eigenvalues for the first two principal components, representing >65% of the total variation, were selected for interpretation; however, only the second axis exceeded its broken-stick value (Table 5). PCA-1 comprises > 39% of the variance in this dataset. Three morphological characters ordinate individuals along PCA-L. These are, in descend- ing order in the loading matrix (Table 6): Flpedlen, Flinfllen and Petlen. PCA-1 is a size axis clearly separating group | (positive affinities) and group 4 (negative affinities) but with overlap between groups 2 and 3 at the origin. PCA-2 comprises > 26% of the vari- ance in the dataset. Three characters ordinate individuals along PCA-2. These are, in FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX 1035 Tas.e 2. Full dataset. Eigenvalues, percent variance extracted and broken-stick comparison of four Principal Component axes with eigenvalues > 1. Axis Eigenvalue Percent of variance Cumulative Broken-stick Percent Eigenvalue 1 4.106 37.325 37.325 3.020 2 2.112 19.204 56.529 2.020 3 1.482 13.470 69.999 1.520 4 1.199 10.902 80.901 1.187 Tasie 3. Full dataset. Principal components loading matrix of the correlations between eleven morphological characters and | t scores (eigenvectors). The five highest correlations with the eigenvector are in bold type. Character Eigenvector 1 2 Petlen -0.3267 0.0632 Petwid -0.2567 0.0249 Petl_wratio -0.0921 0.0471 Flinfllen -0.3826 -0.3641 Flpedlen -0,.4620 0.0131 Flpedl_flinfl -0.2524 0.4800 LfL_wratio 0.0045 -0.4525 Lfwidpt 0.0434 0.3073 Frinfllen -0.3906 -0.3242 Frpedlen -0.4433 0.0426 Frpedl_frinfl -0.2107 0.4732 descending order in the loading matrix (Table 6): Lfl_wratio, Flped_flinfl and Lf widpt. PCA-2 isa shape axis that most clearly separates group 3 (positive affinities) from group 2 (negative affinities). The type specimen of Amelanchier obovalis is nested near the center of group 1 out- side of the range of variation (along PCA-1) in other groups (see Fig. 3). Descriptive statis- tical analysis of the four-group model, calculating the means, standard deviations and 90% confidence intervals of morphological characters most highly correlated with PCA- l and PCA-2 provide some morphological coherence of the groups with strong positive or negative affinities along the principal component axes (Table 7). Group 1 (triangles) consists of individuals (on PCA-1) having short petals (6.8 + 0.9mm), short flowering inflorescences (28.7 + 0.9mm), and short lower pedicels (7.9 + 1.3 mm). On the second PCA axis group | consists of individuals with leaves averaging 1.8 times as long as wide (1.79 + 0.22) that are widest above the midpoint (0.548 + 0.037) and have more or less cylindrical inf] (0.283 + 0.053). Group 4 (stars) consists of individuals (on PCA- 1) having long petals (8.8 + 1.2mm), long flowering inflorescences (47.8 + 3.9mm) and long lower pedicels (15.8 + 1.7mm). On the second PCA axis group 4 consists of individu- als with leaves averaging 1.8 times as long as wide (1.80 + 0.18), that are widest near the midpoint (0.519 + 0.034), and have spreading inflorescences (0.335 + 0.042). Group 2 Taste 4. Full dataset. Cross-products matrix of Pearson's Correlation Coefficients (r) for eleven morphological characters. Values of r 0.7 = r ounded) are in bold type. Petl_wratio Flinfllen Flpedlen Fiped|_flinfl Lfl_wratio Lfwidpt Frinfllen Froedlen Froedi|_frinfl Petlen Petwid Petl_wratio Flinfllen Flpedlen Fipedl_flinfl Lfl_wratio Lfwidpt Frinfllen Frpedlen Frpedl_frinfl 1 0.669 0.396 0.368 0.497 0.327 0.005 0.137 0.385 0.398 0.156 1 -0.410 0.328 0.356 0.163 -0.049 0.164 0.314 0.326 0.118 - 1 0.062 0.185 0.202 0.043 -0.057 0.104 0.107 0.057 : 1 0.778 -0.018 0.229 -0.271 0.839 0.652 -0.021 - : = 1 0.612 -0.024 -0.117 0.708 0.794 0.372 = : : 1 0.329 0.146 0.058 0.434 0.616 = : : - - 1 -0.168 0.192 -0.039 -0.311 - ; - 1 -0.179 -0.094 0.073 - - : - - - 1 0.759 -0.042 : : - - - - 1 0.602 9E0L (2)@2 VaIs/DYO'LINS Q¢ 74 5.5E-04 6.4E-01 Distance (Objective Function) 1.3E+00 Cluster Analysis Information Remaining (%) 50 Group 1 Group 3 Q20n See za® Group 4 kings-d 3 jonmt-b f forestin X31dWO) YFIHDNVTIWY NV 4O SISATVNY DINLIWOHdYOW ‘JAYS ZE0L 1038 BRIT.ORG/SIDA 22(2) Principal Components Analysis 4 4 Group & 1 ¢ 3 x 4 J ray O02 e e 25 ¢ Sa . * * o K 7 & 7k ° * my “ A ' * & od xe Go a A O * ¢ = Oo *R O a aK oO A © & ray ray , Re TYPE~ A&A A @ & A 2 “7 oO oO A JJ © A -4 -3 -1 1 3 Raced Fic, 3. PCA scatterplot with Cluster overlay. Symbols d bership: triangl group 1, circles = group 2, diamonds = group 3, stars = group 4. Type specimen of Amefanchier sbolle indicated. (circles) consists of individuals (on PCA-1) having intermediate petal lengths (7.7 + 0.8 mm), flowering inflorescences (35.1 + 4.6) and lower pedicels (12.6 + 1.7mm). On the second PCA axis group 2 consists of individuals with leaves averaging 1.7 times as long as wide (1.67 + 0.14) that are widest above the midpoint (0.547 + 0.057) and have widely spreading flowering inflorescences (0.363 + 0.043). Group three (diamonds) consists of individuals (on PCA-l) having intermediate length petals (7.3 + 0.9mm), long flowering inflorescences (39.6 + 3.0mm) with short lower pedicels (10.8 + 1.1mm). On the second PCA axis group 3 consists of individuals with leaves averaging 1.9 times as long as wide (1.87 + 0.17), that are widest at the midpoint (0.503 + 0.029), and have cylindrical flower- ing inflorescences (0.272 + 0.026). FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX 1039 Taste 5. Type dataset. Eigenvalues, percent variance extracted and broken-stick comparison of two Principal Component axes with eigenvalues > 1. Axis Eigenvalue Percent of variance Cumulative Broken-stick Percent Eigenvalue 1 2.345 39.082 39.082 2450 2 1.575 26.243 65.325 1.450 Tape 6. Type dataset. Principal Components loading matrix of the correlations between six morphological characters and component scores (eigenvectors). The three highest correlations with the eigenvector are in bold type. Character Eigenvector 1 2 Petlen -0.4524 -0.1036 Flinfllen -0.4994 0.4079 Flpedlen -0.6326 0.0032 Flpedl_flinfl -0.3720 -0.5164 LfL_wratio 0.0144 0.5610 Lfwidpt 0.0842 -0.4914 Multivariate Analysis of Variance.—There is a significant overall multivariate effect of the four-group model (Table 8) and significant F ratios were found for five of the six quantitative characters (Table 9). Pair-wise comparisons between all groups (Table 9) de- termined significant differences between all pairwise group mean vectors for Flinfllen and five of six comparisons for Flpedlen. Petlen was significant for 4 of 6 group compari- sons, Flped_flinfl was significant in 4 of 6 group comparisons, and Lf widpt was signifi- cant for 2 of 6 group comparisons. Differences between group mean vectors for LfL__Wratio were not significant. Group 4 contains the most distinctive individuals, as expressed by all pairwise comparisons of mean vectors for Petlen, Flinfllen, and Flpedlen and two of three pairwise comparisons for Flped_flinfl. Group 1 is also distinctive as expressed by all group parisons for Flinfllen and Flpedlen, 2 of 3 pairwise comparisons for Petlen, 2 of 3 pairwise comparisons for Flped_flinfl,and 1 of 3 pairwise comparisons for Lf widpt. Group 2 is significant in all pairwise comparisons for Flinfllen, 2 of 3 comparisons of Flpedlen, 2 of 3 pairwise comparisons for Petlen, 2 of 3 pairwise comparisons for Flped_flinfl, and 1 of 3 pairwise comparisons for Lf widpt. Group 3 is significant in all pairwise comparisons for Flinfllen, 2 of 3 comparisons of Flpedlen, but only 1 of 3 pairwise comparisons for Petlen, 2 of 3 comparisons for Flped_flinfl and 2 of 3 comparisons for Lfwidpt. The presence of a large sample of individuals with intermediate phenotypes sug- gests ample gene flow (genetic exchangeability) between the groups. Ecological exchange- ability is demonstrated by the occurrence of morphologically distinct individuals oc- curring within the same habitat type. Taxonomic Usefulness of the Elongation of the Lowest Pedicel.—There is significant variation among groups in the elongation of the lowest pedicel from peak flowering to mid-fruiting as determined by Analysis of Variance. The difference between the mean 1040 BRIT.ORG/SIDA 22 = 2) Taste 7. Descriptive statistics of six characters most highly correlated with the principal components (PCA-1 and PCA-2) under a four-group model.Columns under Group labels N = number of individuals in g + standard deviations, (ranges), [minimum, maximum], {90% Conndence ees All mencurements ae raw (untransformed) scores. Character Group 1 Group 2 Group 3 Group 4 N=24 N= 13 N=14 N=11 Petlen 6.8 + 0.9 7.7+08 73+09 8.8 + 1.2 (3.0) (3.5) (3.0) (4.0) [5.6; 8.6] [5.9;9.4] [5.8; 8.8] (6.5; 10.5] {6.5—7.1} {7.3—8.1} {6.9—7.7} {8.1—9.5} Flinfllen 28.7 +09 35.1446 39.6 + 3.0 478 + 3.9 (17.3) (14.4) (8.8) (10.6) [18.7; 36.0] (28.4; 42.8] (35.0; 43.8] (42.2; 52.8] {27.1—30.3} {32.8—37.4} {38.2—41.0} {45.7—49,9} Flpedlen 79413 1236617 10.84 1.1 15.8+ 1.7 (5.7) (5.6) (3.4) (5.2) [5.6: 11.3] (9.6; 15.2] (9.3; 12.7] [13.2; 18.4] {7.4—8.4} {11.8—13.4} {10.3—11.3} {14.9—16.7} Flpedl_flinfl 0.283 + 0.053 0. oe + 0.043 Hees + 0.026 0. ee + 0.042 (0.238) (0. (0.108) (0.123) (0.167; 0.405] (0. ae 0.456] [0. ‘ 2; 0.320] (0. 0.396] {0.264—0.302} {0.342—0.384} {0.260—0.284} {0.332—0.378} LfL_wratio 1.79 + 0.22 1.67 + 0.14 1.88 + 0.17 1.80+0.18 (0.70) (0.45) (0.58) (0.59) (1.47; 2.17] [1.41; 1.86] (1.64; 2.22] [1.56; 2.15] {1.71—1.87} {1.60—1.74} {1.80—1.96} {1.70—1.90} Lfwidpt 0.548 + 0.037 0.547 + 0.057 0. ai + 0,029 0.519 + 0.034 (0.145) (0.171) (0.114) 097) (0.471; 0.616] (0.441; 0.612] (0. Hs 0.582] (0.473; 0.570] {0.535—0,.561} {0.522—0,572} {0.490—0.5 16} {0.500-0.538} flowering pedicel length and mean fruiting length are not significant in group 4 (P =0.15, F & 05 [1,20] = 2.23, F critical = 4.35) indicating that the inflorescence elongates very little from peak flower to mid-fruit. Significant differences between mean flowering and mean fruiting pedicel length were found in the other three groups. Group | has the largest F ratio (P < .OOOL, F «& .05 [1,44] = 20.39, F critical = 4.06) indicating that the lowest pedicel elongates significantly from peak flower to fruit. Group 2 and group 3 also show signifi- cant differences in this character (Group 2: P = .0O1, F « 95 [1.24] = 13.53, F eritical = 4.26; Group 3: P = .0006, F ».05[1.26)= 15.17, F critical = 4.23). DISCUSSION Multivariate analysis of morphological characters distinguishes four groups of specimens within the Amelanchier complex on the Delmarva Peninsula. Group | contains shrubby or multi-stemmed Amelanchier with short and rather fat petals, short, cylindrical flowering and fruiting inflorescences with short lower pedicels. This group of specimens is refer- able to Amelanchier obovalis; the type specimen nests within this group outside of the range of variation of other groups. This group may be more clearly visually defined by FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX 1041 Taste 8. MANOVA test criteria and F approximations for the hypothesis of no overall effect of the four-group model. Statistic Value F Value Num DF Den DF Pr.F Wilk’s Lambda 0.04823940 14.12 18 150.39 ,_ 0001 Pillai's Trace 147957567 7.29 18 165 , 0001 Hotelling-Lawley 9.83905376 27.18 18 100.15 , 0001 Taste 9. Univariate F values (GLM procedure) and P > F for six informative characters and pairwise group com- parisons of character means using Tukey’s Studentized Range (HSD). Critical value of Studentized Range = 3.74. *= Comparisons significant at a =.05, df =58; ns = not significant. Grp! = A. obovalis; Gro4 =A. canadensis. Character F ratio for P.F Group Difference Confidence Limits Character Comparison Ww Means 95% (Tukey) Flinfllen 61.59 0.0001 grp4-grp2 13.19 8.82—17.56 . grp4-grp3 8.25 3.95—12.55 . grp4-qrp1 19.10 15.21—22.98 * grp2-qrp3 4.98 0.827—9.05 grp2-qrpl 5.9] 2.32—9.58 * grp3-grp| 10.85 7.26—14.43 * Flpedlen 68.86 ,0.0001 = grp4-grp2 3.70 2.01—5.39 * grp4-grp3 5.05 3.39—6.72 . grp4-grp1 7.93 42—9 grp2-grp3 1.35 -0.24— 2.94 ns grp2-grp1 4.22 2.80—5.64 . grp3-grp1 2.87 1 48—4.26 . Flped_flinfl 10.16 ,0.0001 = grp4-qrp2 0.019 -0.032—0.070 ns grp4-grp3 0.063 0.012—0.113 . grp4-qrp] 0.053 0.007—0.098 - grp2-grp3 0.081 0.033—0.129 " grp2-grp| 0.071 0.028—0.114 i grp3-grp| 0.010 -0.032—0.052 ns Lfl_wratio 2.01 0.1228 grp4-grp2 0.104 -0.106—0.314 ns grp4-grp3 0.079 -0.127—0.285 ns grp4-grp| 0.016 -0.171—0.203 ns grp2-grp3 0.183 -0.015—0.380 ns grp2-grp | 0.088 -0.089—0.264 ns grp3-grp| 0.094 -0.078—0.267 ns Lfwidpt 5.99 0.0013 grp4-grp2 0.031 -0.009—0.072 ns grp4-grp-3 0.017 -0.023—0.057 ns grp4-grpl 0.030 -0.006—0.066 ns grp2-grp3 0.048 0.010—0.086 * grp2-grpl 0.002 -0.033—0.036 ns grp3-grp1 0.046 0.013—0.080 : Petlen 12.13 0.0001 grp4-grp2 1.11 0.079—2.15 ms grp4-grp3 1.53 0.510—2.55 [ grp4-grpl 2.06 1.14—2.98 : grp2-grp3 0.41 -0.559—1.39 ns grp2-grp1 0.94 0.073—1.81 * grp3-grp1 0.53 -0.320—1.38 ns 1042 BRIT.ORG/SIDA 22(2) eliminating outliers; however, eliminating these individuals in the PCA does not change group membership and these specimens do appear to represent natural variation within the group. One individual classified in group 1 (upper right corner of Fig. 3) deviates from the morphological group defined here by an extremely narrow cylindrical inflorescence. This individual may best be recognized as a separate entity; however, no other similar individual or specimen was located during sampling. Individuals forming group | (A. obovalis) were collected chiefly from sandy woodlands and barrens but did occur on hummocks along tidal creeks and thicket margins around non-tidal wetlands in disturbed and in nearly pristine habitats. Individuals are typically low and shrubby but may grow to several meters in absence of disturbance; this was ideally illustrated by single, apparently clonal individuals with stems that were inside and outside of a roadside-mowing regime. Group 4 corresponds to a group of tall, arborescent or multi-stemmed Amelanchier with large petals and long lower pedicels forming long, widely spreading flowering and fruiting inflorescences. This group of specimens is referable to Amelanchier canadensis, matching as well as can be expected the type specimen and at least part of the historical circumscriptions. Individuals in group 4 (A. canadensis) were collected from both tidal and non-tidal swamp forests and were easily distinguished in the field. This species oc- curs sympatrically with Amelanchier obovalis in some habitats. Group land group 4 correspond to known taxonomic entities (Amelanchier obovalis and A. canadensis respectively) but group 2 and group 3, despite the statistical signifi- cance of several pairwise comparisons of character mean vectors in the MANOVA could not be assigned to distinctive forms such as might arise as microspecies. | hypothesize that group 2 and group 3 are members of an agamic complex derived from hybridization between Amelanchier canadensis and A. obovalis. Evidence of apomixis has been dem- onstrated in Amelanchier canadensis (Campbell et al. 1987) but not in A. obovalis. How- ever, evidence to support this hypothesis may be drawn from my statistical analysis and observation. First, both Amelanchier canadensis and A. obovalis (as clear entities) were observed to occur within the same habitat patch in the study area and were observed to have overlapping phenologies, which increase the probabilities that natural hybridiza- tion occurs. Second, the means of important classification characters in group 2 and group 3 (with 90% confidence intervals) consistently occupy ranges intermediate between Amelanchier canadensisand A. obovalis (Table 7). Finally, the members of groups 2 and 3 appear morphologically heterogeneous and do not correspond to the four putative mor- phological forms that I noted in the field prior to sampling or toa particular habitat type. My field observations were that one form was referable to Amelanchier obovalis(group 1),a second to Amelanchier canadensis (group 4),a third group comprised the pubescent-ovary forms within group | and the fourth, an early flowering Amelanchier with small petals on ummocks along tidal creeks | now believe are consistent with Amelanchier obovalis. Additionally, the discrimination of multivariate character space held by group 2 versus group 3 is uncertain. Group 2 and 3 separate nearly at the origin of PCA-2 (Fig. 3) and the character most highly correlated with this second axis (r = .704) is Lfl_Wratio, a character that fails statistical significance tests between groups in Analysis of Variance (see Table 9). Further, comparison of mean vectors of two important characters (Flpedlen and Petlen) between group 2 and group 3 are not significant (Table 9). Finally, the itera- tions of Cluster Analysis join group 2 and 3 in a three-group model and then collapse into group 4 (A. canadensis) in a two-group model. If | accept more variation in Amelanchier canadensis and consider a two-group model composed of Amelanchier FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX 1043 obovalis and a new group containing all the members of groups 2, 3 and 4 then my per- formance in keying vouchers would certainly improve but accepting this variation comes with a heuristic cost in that lobscure the only spatially distinctive cluster resulting from PCA. Given the well-documented propensity for hybridization and the formation of ag- amic complexes in Amelanchier, the most reasonable assumption is that individuals within group 2 and group 3 do constitute intermediate forms due to incomplete repro- ductive isolation between Amelanchier canadensis and A. obovalis. I conclude that the multivariate character space occupied by Amelanchier obovalis exhibits significant phenetic dissimilarities with the other groups as expressed by its distinctiveness in Cluster Analysis (e.g., the long branch before joining) and by statistical significance of the majorities of group comparisons of character mean vectors for Flinfllen, Flpedlen, Petlen, and Flped_flinfl. The discreteness of multivariate character space occupied by Amelanchier canadensis can be determined both visually (occurrence of a distinct cluster in the PCA) but also by the statistical significance of the majorities of group comparisons of character mean vectors for Flinfllen, Flpedlen, Petlen and Flped_flinfl. However, the presence of numerous intermediate specimens violates most commonly accepted taxonomic conceptions of species status. Subspecies as a taxonomic rank is poorly defined in the literature of taxonomic botany and is used more or less interchangeably with variety (McDade 1995; Nesom and Lipscomb 2005). Wiegand (1912), Fernald (1950a) and Jones (1946) assigned the rank of variety to those distinctive forms of Amelanchier that occupied a particular geographical area and/or had a habitat pref- erence slightly different than that typical of the species. Campbell and Dickinson (1990) concluded that nonconcordance between morphological and breeding units in Maloid apomicts rendered the biological speci ept inappropriate and that a morphospecies concept may be necessary to deal with evolutionary complexity in the Maloideae. In this study I found no well-defined geographical or ecological separation between the taxa. How- ever, I must also consider that the high frequency of intermediate forms found in the study area may reflect the long disturbance history of the Delmarva Peninsula and this distur- bance history is responsible for recent, but perhaps not historical, genetic and ecological exchangeability (Crandall et al. 2000). That the frequency of disturbance often correlates with the number of hybrids and ‘perplexing forms’ has been the subject of repeated obser- vation and much discussion among authors since Wiegand (1912). The study populations of Amelanchier fall under Crandall’s et al. (2000) case of “anthropogenic convergence on demographic exchangeability” where we continue to treat the populations as distinct units but not as distinct species. Both taxa are distinctive enough morphologically and interme- diate forms may not occur with aia frequency in other regions. In the taxonomic treat- ment below, I utilize the taxon ncept of “variety” defined as the primary taxonomic level below species ae Code of Botanical Nomenclature, 4.1). os TAXONOMIC TREATMENT Amelanchier canadensis (L.) Medic. var obovalis Michx. Typ spilu idensis var. obovalis Michx., Fl. Bor. Amer. 1:29]. 1803. Amelanchier obovalis (Michx.) pas Bor Gaz. 35:434. 1903, in part. Amelanchier obovalis (Michx.) Ashe, M.L. Fern ald in Rhodora 43:566. 1941, in part. Amelanchier obl bengvl ia (Torr. & A. Gray) ne K.M. Wiegand in yeaa 14:147-150. 1912, in part. Amelanchier obovalis, P Landry, Bull. Soc. Bot. Fr. 122:248. 1 Amelanchier obovalis is a rather enigmatic species with a aie history of circumscrip- tion problems. André Michaux (1803) described Mespilus canadensis var obovalis from 1044 BRIT.ORG/SIDA 22(2) “carolina inferiore” (lower Carolina) as “humilior; folius oblongiscule obovalibus” (low with oblong obovate leaves) and “arbriss de deux pieds haut’ (a shrub about two feet tall). The Michaux specimen (P) is very well preserved having several stems with inflorescences and petals intact and a single stem with (apparently mature) ded leaves. A century later Ashe (1903) created a new combination, Amelanchier obovalis (Michx.) Ashe de- scribing a shrub of “swamps and loose soils” 9-15 dm tall or a small tree to four meters tall with flowering inflorescences 3-5 cm long, the lowest pedicels 3-9 mm but becom- ing elongate (2-3 cm) in fruit. Wiegand (1912), probably the most influential of Amelanchier taxonomists, includes Amelanchier obovalis (Michx.) Ashe under his “fasti- giate not stoloniferous” Amelanchier oblongifolia Torr. & A. Gray) Roemer. stating that it “seemed to be this species” but it is not clear that Wiegand ever saw any of Ashe’s collec- tions from the southeastern United States. Wiegand (1912) described his A. oblongifolia as a shrub with slender upright stems in rather dense fastigiate clumps with inflores- cences 2.5-6 cm long, the lowest pedicels 8-18 mm long but scarcely longer in fruit. M.L. Fernald (1941), after several field seasons exploring the Pine Barrens of southeastern Vir- ginia, disagreed with Ashe’s description of Amelanchier obovalis, particularly regarding plant height, and subsequently ignored his treatment, describing Amelanchier obovalis (Michx.) Ashe as a “dwarf and stoloniferous” shrub with flowering inflorescences 1-2.5 cm long with lower pedicels lengthening only to 3-8 mm. Fernald (1950a) distinguished the species in his key chiefly by plant habit in a major couplet: “stoloniferous or surculose loosely colonial shrubs” 0.2-2 M. high (A. obovalis); versus “fastigiate coarse and tall shrubs or arborescent” (A. canadensis). Jones (1946) closely followed Fernald’s (1941) treatment of Amelanchier obovalis as “low shrubs, surculose, forming loose colonies” but with flow- ering inflorescences 1-3 cm long with lower pedicels 1-3 mm, becoming 3-8(11-14 mm) long in fruit. Landry (1975) describes Amelanchier obovalisas a subspecies of A.canadensis with the major difference between the two subspecies being the general size of floral inflorescences and (very vaguely) the length of the lowest pedicels with Amelanchier canadensis ssp. canadensis having “rather large” flower clusters and lower pedicels at least 6 mm long versus clusters of flowers small and lower pedicels more than 6 mm long (A. canadensis ssp. obovalis). The members of Group 1 (Amelanchier canadensis var. obovalis) are both low shrubs with apparent vegetative spread and tall, slender, multi-stemmed shrubs that occurred ina variety of habitats including hummocks on margins of tidal creeks and rivers, sandy pine forests and barrens, roadsides and under powerlines. In both flowering and fruiting stages the lowest pedicels were the shortest in the dataset and although they did elongate in fruit the extremes that Ashe (1903) suggested (10-20 mm) were not observed. I suspect that Ashe, perhaps due to finding very similar phenotypes in a variety of habitats or by witnessing the same complex as analyzed herein conflated the characteristics of both taxa. M.L. Fernald (1941), on the other hand, accepted only the extreme shrubby forms of the species and somewhat idiosyncratically noted that the taller species was probably Amelanchier dustromontana, a species that Ashe (1918) described from the Southern Appalachian Mountains and not the southern Coastal Plain. Amelanchier canadensis (L.) Medic.var. canadensis. Typr: Mespilus canadensis L., Sp. Pl. 1498. 1753. Amelanchier oblongifolia (Torr. & A. Gray) Roemer, K. Wiegand in Rhodora 14:147-150. 1912, in part. Amelanchier obovalis (Michx.) Ashe, Bot. Gaz. 35:434. 1903, in part. Amelanchier canadensis (L.) Medic. is the earliest combination in Amelanchier, the type FRYE, MORPHOMETRIC ANALYSIS OF AN AMELANCHIER COMPLEX 1045 specimen (LINN) from “habitat in Virginia, Canada” containsa single stem with five leaves and the remnants of a fruiting inflorescence with three fruit. The type is of limited use- fulness as the characters used by major authors of the genus to separate Amelanchier canadensis from A. obovalis, e.g. plant habit, petal length, flowering inflorescence length and lowest pedicel length are absent. Wiegand’s (1912) circumscription of Amelanchier oblongifolia appears to include both Amelanchier canadensis and A. obovalis as the ranges of character measurements are inclusive of both taxa. Similarly, Ashe’s (1903) circum- scription of Amelanchier obovalis appears to include cl istics of both species. It is very difficult to generalize but overall most authors treat Amelanchier canadensis asa fastigate shrub forming alder-like clumps having larger petals, and larger inflorescences than the low, shrubby and stoloniferous Amelanchier canadensis var. obovalis. The members of Group 4 (Amelanchier canadensis var. canadensis ) comprise a group of tall, fastigiate shrubs and small trees having the largest petal dimensions, very long flowering and fruiting inflorescences with long lower pedicels forming widely spread- ing inflorescences. This variety is easily distinguished in the field however numerous individuals with characters intermediate between the two varieties are present in the study area. Individuals of Amelanchier canadensis var. canadensis were located, mostly, in natural habitats associated with tidal and non-tidal wetlands, however, there appears to be little habitat specificity and these individuals occurred often in close proximity to var. obovalis. Wiegand’s notion that the pedicels of his Amelanchier oblongifolia are “scarcely longer in fruit” appears consistent with the results of Analysis of Variance com- paring lowest pedicel lengths in flowering versus fruiting material; although, it is appar- ent that Wiegand also conflated the cl teristics of both taxa, perhaps because he was unfamiliar with the more southern forms constituting Amelanchier obovalis. KEY The following key to Amelanchier canadensis includes the other common species occur ring in the study area, Amelanchier arborea (Michx. f.) Fernald. In keying Amelanchier one should realize that many specimens would not key confidently without observa- tions (collections) made throughout the growing season. Those with interme- diate characters are likely part of the species complex and so should be simply identified as ‘comparing favorably’ with one variety or the other. The twenty-four members of group 1(A.c. obovalis) provide the mean and range (one standard deviation) of characters that should be used to identify this species and the eleven members of group 4 (A. c. canadensis) provide the mean and range of the same morphological characters. Specimens with entire leaves from the southeastern Coastal Plain may constitute var. subintegra Fernald but this putative taxon is not included in the key as its evaluation was not within the scope of this manuscript. Sepals in flower and in fruit tightly recurved and appressed to the ovary, leaves subcordate 7 base, acute to acuminate at the apex and coarsely serrate. Arborescent species of ia renee nes arborea Sepals in flower and in fruit not tightly appressed to ovary, leaves rounded at base, ae to broadly rounded at apex and finely serrate. Tall shrubs with slender stems or dwarf shrubs with evident vegetative spread, chiefly of swamps and barrens at low elevations __ Amelanchier canadensis ioe (5.9-)6.8(-7.7) mm long and (2.6-)3.1(-3.6) mm wide, inflorescences at peak flower (27.8-) 8.7(-29.6) mm, elongating in fruit to (27.2-)32.1(-37.1) mm, lowest pedicels (flowering) (6.6-) ah 9.2) mm, elongating in fruit to (8.2-)11.0(—13.8) mm. Inflorescences usually cylindrical, the lowest pedicel averaging 28% of the total inflorescence length at flowering, and 35% percent pee 1046 BRIT.ORG/SIDA 22(2) in fruit. Leaves on older branches chiefly obovate, the widest point occurring at 51-58 percent of the total leaf length. Sepals usually erect in flower, but often spreading during maturation of the ovary.Ovary summit usually glabrous but a variable amount of pubescence occurs in some forms (typically in dry, well-drained habitats). Plants are typically short and shrubby but may w to several meters in the absence of disturbance var. obovalis ea (7.6-)8.8(-11) mm long and (3.1-)3.7(-4.3) mm ae inflorescences at peak flower (43.9-) 47.8(-51.7) mm, elongating in fruit to (43.2-)48.2 (-53.2) mm, lowest pedicels (flowering) 15.8(14.1)-(17.5) mm, elongating in fruit to (15.1-)17.5 (-19.9) mm. inflorescences using widely spreading, the lowest pedicel averaging 36% of the total inflorescence length at flowering, and 38% in fruit. Leaves on older branches chiefly broadly elliptical or occasionally oblong, the wid- est point at 49-55% of the total leaf length. Sepals usually spreading to reflexed in flower and in fruit. Ovary summit glabrous. Plants are typically multi-stemmed with tall slender branches var. canadensis ACKNOWLEGMENTS [am grateful to Art Tucker, director, Robert Naczi, curator, and Keith Clancy, collections manager, of the Claude E. Phillips Herbarium (DOV), Delaware State Univ, Dover, DE for managing loans of herbarium material and providing access to a beautiful facility. Dur- ing the term of this research [have borrowed material from a number of institutions and I would like to thank the curators and staff at: CM, FLAS, GA, MARY, NA, NCU, PH, US, USCH, VPI, and WVA, who so efficiently handled the distribution of material to Dover. | would like to offer special thanks to Alain Changy (P) and Mark Spencer (LINN) who provided images of type specimens held in Paris and London, respectively. Gwen Brewer, Wesley Knapp and William McAvoy provided helpful comments on early drafts of the manuscript. Jason Harrison and Wesley Knapp assisted with fieldwork. Dan Sieh], with an artists care, mounted and processed hundreds of Amelanchier vouchers at the Tawes Herbarium, Maryland Department of Natural Resources, Annapolis, MD. The Maryland Department of Natural Resources, Natural Heritage Program, the Claude E. 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GAREX REZNICERILA NEW WIDESPREAD SPECIESOF CAREX SECTION ACROCYSTIS (CYPERACEAE) FROM EASTERN NORTH AMERICA David A.Werier 30 Banks Road Brooktondale, New York 14817, U.S.A. nakita@lightlink.com ABSTRACT Carex reznicekii is described as new from mesic to dry-mesic forests of Alabama, Arkansas, Connecticut, Dela- ware, District of Columbia, Georgia, Kentucky, Maryland, Mississippi, Missouri, New Jersey, New York, North Carolina, Pennsylvania, Rhode Island, South Carolina, Tennessee, and Went pees C. reznicekii had mostly been misidentified as either C. umbellata or C. nigromarginata. It is distingui d from these species as well as other members of section Acrocystis by a combination of lack of pasa) spikes, once culms, narrow leaves, and only ce red pistillate scales. A key to the 19 taxa of section Ac t rth America is provided. RESUMEN Se d ibe como nuevo Carex reznicekii de los bosques mésicos a xérico-mésicos de Alabama, Arkansas, Caro- lina del Norte, Carolina del Sur, Connecticut, Delaware, Distrito de ae Georgia, Kentucky, Maryland, Mis- sissippi, Missouri, New Jersey, New York, epee ibis Rhode Island, Tennessee, y Virginia. Previamente, C. bee habia ace en su mayor parte mal interpretada 3G, aan lataoC. eee a. Se distingue o de otros miembros de ie seccion eee ae por la combin cin de carencia de espiga basal, nee cor tos, ee Soa cn2s) jas. Se proporciona una clave para los 19 taxa de la seccion Acr s que ocurren en el este de Norteamérica. INTRODUCTION Carex section Acrocystis Dumort. contains approximately 35 species worldwide with 20 in North America (Crins & Rettig 2002), although the most recent comprehensive treat- ment is Kikenthal’s (1909) now outdated revision. Species distributions lie mostly in Eurasia and North American with one taxon known from South America (Ktkenthal 1909). It appears that Carex section Acrocystis as currently circumscribed is polyphyl- etic, but North American species and some Eurasian species form a clade (Roalson et al. 2001, Roalson & Friar 2004). Twenty-eight taxa, including the one described here, are currently known from North America, with 19 of these occurring in eastern North America (Crins & Rettig 2002). At least two additional undescribed species occur in east- ern North America (Crins & Rettig 2002; Roalson & Friar 2004; Anton Reznicek, pers. comm). Eastern North American taxa of section Acrocystis, share the following characters: perigynia pubescent (except C. tonsa (Fernald) Bicknell var. tonsa ), two veined, and abruptly narrowed toa distinct beak; stigmas three (except two and three in C.floridana Schwein.); non-basal pistillate spikes (when present) subtended by sub-sheathing to sheathless bracts, approximate, the distal ones short pedunculate to sessile; and habitats dry to mesic forests (tending to favor drier) or more open for a few taxa. During survey work in the Hudson Highlands of southeastern New York, I found a SIDA 22(2): 1049- 1070. 2006 1050 BRIT.ORG/SIDA 22(2) population of the New York State endangered C. nigromarginata. While delineating and describing this population I encountered a population of a strikingly different plant, which at first appeared to be C. umbellata because all of its culms were very short and hidden in the bases of the plants. Upon closer inspection, | observed that these short- culmed plants lacked the basal spikes characteristic of C. umbellata. These short-culmed plants were uniform throughout its population and appeared immediately adjacent to and clearly did not resemble C. nigromarginata. In contrast to the C. nigromarginata popu- ation, these short-culmed plants had much narrower leaves, uniformly very short culms, and pale pistillate scales. Wider sampling showed that the short-culmed plants formed discreet uniform populations, although sometimes occurring mixed with, but seemingly not intergrading with, C. nigromarginata. A cursory examination of herbarium material also suggested that the distributions of the two entities were different. This new species of Carex is described here as Carex reznicekii. MATERIALS AND METHODS Specimens of C. reznicekii were sought by examining all specimens labeled as C. nigromarginata Schwein. and C. umbellata Schkuhr ex Willd. (two superficially similar species) and in some cases other members of section Acrocystis from BH, BRIT, GH, MO, NCU, NY, NYS, PH, and US. In addition, putative C. reznicekii specimens were sent to me from collections at DOV, MICH, and VPI as well as by Philip E. Hyatt. Additional speci- mens were gathered from field work. Type material was examined of all species similar to or confusable with C. reznicekii, including their synonyms. Thirty-one populations were visited in the field, covering a large part of the species’ range. Site visits were conducted to assess habitat affinities, morphological variability, and habitat differences with other members of Carex section Acrocystis. Soil samples were collected at six distant sites throughout a large part of the range of the species (Fig. 1). Equal amounts of soil from four places at each of the six sites (adjacent to individual C. reznicekii plants) that were at least 10 meters apart were mixed together. These were sent to Cornell Nutrient Analysis Laboratories at Cornell University, Ithaca, NY for analysis. Soils were analyzed for particle size distribution, pH, and the minerals Al, Ca, Fe, K, Mg, Mn, P and Zn. To clarify the differences between C. nigromarginata, the most similar species to C. reznicekii,and C. reznicekii, specimens were compared using eight continuous charac- ters (see Table 1). Specimens were selected so as to have ten specimens per species from each of three geographic regions: 1) northeast (Virginia/Kentucky north), 2) southeast (North Carolina/Tennessee south), and 3) Arkansas/Missouri. Only six adequate speci- mens were available for C. reznicekii from the latter region. Within each geographic re- gion (excluding Arkansas/Missouri), only two specimens per state and one per county were selected to help cover the range of the species. For each species, specimens were selected randomly given the r ioned above and excluding specimens where all eight characters could not be measured. Analysis of variance (two-way ANOVA) and Pearson Correlation Coefficients were calculated using SPSS Version 13. Additionally, ranges, means, and standard deviations were calculated for both species for each region. Nonmetric multidimensional scaling (NMDS) ordination was chosen to compare the relationship of C. nigromarginata and C. reznicekii. In comparison to other ordina- tion methods such as canonical correspondence analysis (CCA) or principle components analysis (PCA), NMDS does not assume an underlying distribution for measured vari- WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1051 pares | ee ay: = ATE BLT racae es Tay ines Hh apaue zine en oy Serene cee ieee ro Z = La TS a a" on aeness Lt —+ iT Cail I Hartad in hla-l 1 rs Fic. 1.G hic distributi f C. reznicekii Gla} ables. A NMDS ordination was created using the slow thorough mode on PC-ORD (McCune & Grace 2002). Specimens measured for use in the ordination are delineated by a single asterisk in the isotypes listed for C. reznicekii, in the representative specimens of C. reznicekii, and in the specimens cited in Appendix A. A scatter plot graph using a larger sample size was created for the two most explana- tory characters. As additional evidence that C. reznicekii and C. nigromarginata are dis- creet entities, distribution curves were created and compared for the two most explana- tory characters for C. nigromarginata and C. reznicekii, as well as C. nigromarginata and C. reznicekii combined, and C. nigromarginata specimens from outside the geographic range of C. reznicehii. For the latter distribution curves all sp of C.nigromarginata that occur outside of the geographic range of C. reznicekii from BH, BRIT, MICH, and MO, excluding duplicates, were measured. Specimens measured for the scatter plot graph and distribution curves are delineated by either a single or double asterisk in the isotypes listed for C. reznicehii, in the representative specimen of C. reznicehii, and in the speci- mens cited in Appendix A. description of C. reznicekii and a key to all members of Carex section Acrocystis were created from original measurements except, for the key, measurements for C. com- munis Bailey varieties and C. inops Bailey ssp. heliophila (Mackenzie) Crins are adapted from Crins and Rettig (2002). Character states that were on an extreme edge or disjunct from most other measurementare placed in parentheses. The following notes clarify three characters listed in the description and the key. 1. Culm length is measured from the base of the culm to the top of the inflorescence as interpreted or implied by many other authors (LeBlond et al. 1994; Reznicek & Camelbeke 1996; Naczi et al. 2001; and others). 2. Terminal staminate spikes exceeding or exceeded by the rest of the inflorescence is 1052 BRIT.ORG/SIDA 22(2) Taste 1. Characters measured for ordination, ANOVA, and Pearson Correlations giving the codes used through- out the paper and any specifications for choosing which part to measure. Character Code Specifications Tallest culm height (cm) TaCuHe Tallest per specimen Widest leaf width (mm) ea Widest per specimen Anther length (mm) ean of two per specimen Largest length that a staminate setae Greatest length per specimen spike exceeds the dista most pistillate spike (mm) Perigynia length (mm) PeLe Mean of two per specimen Perigynia width (mm) PeWi Mean of two per specimen Taller staminate spike length (mm) StSpLe Mean of the length of two taller staminate spikes per specimen Staminate spike width (mm) StSpWi Mean of the width of the two staminate spikes measured for spike length measured by subtracting the difference of the length from the base of the bract subtend- ing the proximal-most non-basal spike to the apex of the terminal staminate spike, and the ae from the base of this same bract to the apex of the distal most lateral spike. 3. Perigynium body refers to the perigynium excluding the beak and stipe or stipe- like base. The beak and stipe begin at the deepest point in the concavity formed at the summit and base of the perigynium. Perigynium body shapes vary considerably and al- though their shape aids in distinguishing taxa (and is used in the key), they should be used cautiously. RESULTS Carex reznicekii Werier, sp. nov. (Fig. 2). Typr: U.S.A. VirGiNiA. Caroline Co: near North Anna River, Route 207 Jericho Road) W from Carmel Church to Oxtord Road (Route 689), 3.6 mi S and SE on Oxford Road (Route 689) to small dirt road on S side of road; SW along dirt road towards North Anna River, UTM coordinates in NAD-83 Zone 18 4196701N 280996E, 10 Apr 2004, Werier 1951 (HOLOTYPE: BH; ISOTYPES: GH*, MICH, MO, NCU, NY, US). Carici nigromarginatae similis, sed characteribus sequentibus differt: laminis foliorum latissimis 1.2-2.2(-2.5) mm latis; culmis singulae plantae saepe in longitudine similibus longissimis 1.9-9.9(-13.7) cm pe spicis staminalibus 3.3-8.3(-9.2) mm longis et 0.4-1.6(-1.8) mm latis; et colore rubenti squamarum pistillatarum ad regiones marginales restricto. Densely caespitose, herbaceous, evergreen perennial. Rhizomes horizontal, ascending, or erect, 0.1-3.8 mm long between shoots or branches of the rhizomes, 0.6-1.2 mm wide with leaf sheaths absent, enclosed by leaf sheaths and/or cataphylls, which disintegrate into long fibers. Vegetative shoots erect to ascending, arising from buds at the tip or side of the rhizomes, or directly from the apical meristem of rhizomes or previous year's vegetative shoots; bases consisting of cataphylls or leaf sheathes which disintegrate into long fi- bers; pseudoculms (represented by leaf sheaths) 1.2-7.6 cm long. Reproductive shoots erect, mostly produced directly from the apical meristem of vegetative shoots or rhizomes, or less frequently produced from buds at tip or side of rhizomes, bases mostly surrounded only by leaf sheaths which disintegrate into long fibers, or less frequently surrounded by cataphylls; culms 1.3-9.9(-13.7) cm long, the tallest 1.9-9.9(-13.7) cm long, 0.3-0.5 mm wide just below the inflorescence, trigonous, with 1 elevated vein on each angle and 2-5 elevated veins on each side, smooth to minutely antrorsely scabrous on angles and veins, 1053 WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1mm).Middle Fig. 2. Carex reznicekii. Bottom right, plant (scale (I to r), culm, inflorescence, and sheath (scale = 1 cm). Top left, staminate scale and anther (scale = 1 mm). , perigynia, and achenes (scale ft, Pistillate scales a 5 cm).Bottom le 1054 BRIT.ORG/SIDA 22(2) more prominently scabrous on angles and distally. Cataphylls at base of reproductive shoots 0-2, 1.6-10.8 mm long. Leaves of reproductive shoots 3-9. Leaf blades to 52.0 cm long, 0.7-2.2(-2.5) mm wide, the widest 1.2-2.2(-2.5) mm wide, flat to V-shaped, to occa- sionally M-shaped; adaxial surface dark green and smooth, papillose, or antrorsely sca- brous, more textured distally; abaxial surface smooth to occasionally scabrous, more tex- tured on mid-vein and distally; margins smooth to antrorsely scabrous, more scabrous distally; leaf sheaths 0.7-5.1 cm long, lower portion especially on outer sheaths have some red coloration; abaxial face with green to white or proximally red veins and whitish and translucent to proximally reddish and more opaque intervein regions; adaxial face with whitish or proximally red veins and whitish, thin, and translucent intervein regions, thin- ner and more translucent than abaxial intervein regions; vein edges with ascending, spreading, or reflexed minute, stiff, broad-based deltoid hairs; sheaths disintegrate into long fibers consisting of veins, which retain pubescence; adaxial sheath face apex con- cave to a depth of 0.3-0.8(-1.0) mm and sometimes slightly thickened; ligules typically wider than long, 0.3-1.0(-1.5) mm long (including the free portion), 0.6-1.4 mm wide, free portion of ligule 0.1-0.2 mm long, ciliate with abundant minute, stiff, broad-based del- toid hairs. Infructescences 5.2-12.9(-16.7) mm long, consisting of 3-5 approximate spikes at the summit of the culm; proximal-most internode 0.8-3.7(-5.2) mm long; bracts sheathless; proximal-most bract with blade 3.6-23.0(-38.5) mm long, 0.6-1.8 mm wide, 3.9(-4.3) mm shorter than to 10.9(-23.5) mm taller than infructescences; distal bracts re- duced. Spikes arising singly from nodes; terminal spike staminate, 3.3-8.3(-9.2) mm long, 0.4-1.6(-1.8) mm wide, on peduncle (0.2-)0.3-0.7 mm long, exceeding distal-most lateral spike by 3.7 mm to exceeded by distal-most lateral spike by 1.1(-1.6) mm; lateral spikes 2- 4, pistillate, short-pedunculate, with a cladoprophyll towards base of peduncle; proxi- mal-most spikes 3.4-7.2 mm long, (1.8-)2.2-4.4 mm wide, (3-)5-10(-12) flowered, on pe- duncles (0.3-)0.4-0.9(-1.4) mm long, with cladoprophyll (0.7-)1.7-2.8(-3.0) mm long. Staminate scales (2.5-)2.7-4.0 mm long, 0.7-1.8 mm wide, ovate to elliptic, apex acute to obtuse, with a green or yellowish longitudinal mid-stripe which includes the mid-vein, margins translucent, thin, whitish to occasionally yellowish-brown, often red sub-mar- ginally forming a longitudinal stripe, more so toward apex of scales and on distal scales, mid-vein present, extending further toward the apex of the scale in more distal scales; distal scales with mid-vein raised, slightly antrorsely scabrous, and ending close to apex of scale or sometimes with mid-vein projecting as a short awn attached just proximal to the scale apex and projecting up to just beyond the apex of the scale; Pistillate scales 2.6-4.3 mm long, 1.0-1.9 mm wide, 0.9(-1.) mm shorter than to 0.5 mm longer than associated perigynium, ovate to lanceolate to elliptic, apex acute to obtuse, with a green (to brownish) longitudinal mid-stripe which includes the mid-vein; margins translucent, thin, whitish to occasionally yellow-brown, often some reddish color sub-marginally forming a sub- marginal longitudinal stripe, more so distally on the scale; sub-marginal longitudinal red stripe Gif present) does not extend laterally to immediately adjacent to mid-stripe except sometimes at apex of scale; mid-vein antrorsely scabrous and ending just before apex of scale or mid-vein, projecting as a short awn attached just proximal to the scale apex and projecting up to just beyond the apex of the scale; bases of scales often with a narrow red horizontal stripe just above attachment to spike axis. Anthers 3, (1.2-)1.3-1L9(-2.3) mm long. Stigmas 3, withering a age. Perigynia (2.5-)2.7-3.9 mm long, (0.8-)0.9-1.3(-1.5) mm wide, light green sparsely red-punctate, obtusely trigonous to plano-convex in cross- section, with two prominent nerves that extend the length of the perigynia and occasion- WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1055 ally up to8 less nerves at base, papillose with papillae short-cylindric up to 0.01 mm. long, as well as pubescent with small (less than 0.1 mm), predominately antrorsely directed, stiff, deltoid hairs, which are denser on the prominent nerves that extend into the beak teeth; body 1.4-1.9(-2.0) mm long, ellipsoid, gradually tapering to beak and stipe: stipe 0.5-1L0(-LD mm long; beak (0.4-)0.6-1.0 mm long, bidentate; beak teeth 0.1-0.3 mm long. Achenes (1.4-)1.5-18(-19) mm long, 0.9-1.2(-1.3) mm wide, ovoid, acutely to obtusely trigonous to plano-convex in cross-section, minutely papillose, yellow-green to green when immature, light to chestnut brown when mature. Etymology.—I selected reznicekii as the epithet to honor Dr. Anton Reznicek, Univer- sity of Michigan, who has inspired me and many others interested in carices. He has cata- lyzed a revival in the study of Carex which has revealed a tremendous amount of new information. In addition, he has contributed directly toa greater understanding of these plants through numerous articles and classes. A total of 242 specimens (143 records) of C. reznicekii were examined. Of these, 125 specimens (88 records) were collections made prior to recognition of C. reznicekii. Sev- enty-one of these 125 specimens had at one point been labeled C. nigromarginata, 78 C. umbellata (including C. abdita Bicknell and C. umbellata var. brevirostris Boott), 6 C. floridana [including C. nigromarginata var. floridana (Schwein.) Ktikenthal], and 9 C. albicans Willd. ex Spreng. var. emmonsii (Dewey ex Torr.) Rettig [including C. emmonsii Dewey ex Torr.and C. nigromarginata var. minor (Boott) Gleason]. The sum totals to more than 125 because some specimens had been annotated numerous times. Type material examined comprised the following: C. umbellata, scan of holotype from B; C. umbellata var. vicina Dewey, holotype at GH, isotype at PH; C. umbellata var. brevirostris, isotype at US (holotype not examined but specimen is from Saskatchewan): C. abdita, holotype at NY; C. microrhyncha Mackenzie, holotype at NY: C. umbellata var. tonsa Fernald, lectotype and 2 isolectotypes at GH; C. rugosperma Mackenzie, holotype at NY, C. nigromarginata, holotype at PH; and C.floridana, holotype at PH. None of the types examined proved to be C. reznicekii. A detailed comparison of C. reznicekii to C. umbellata and C. nigromarginata is made below. In the past, C. reznicekii has been mistaken most often for these two species. Carex nigromarginata is most similar. The key compares all members of section Acrocystis from eastern North America. Carex umbellata superticially resembles C. reznicekii, but is actually quite distinct. The two species can easily be fully separated by noting the presence (C. umbellata) or absence (C. reznicekii) of basal spikes. Mackenzie (1913), clearly stated this when he ex- plained how to distinguish C. nigromarginata and C_floridana from C. umbellata and its allies. He stated, “these [C. nigromarginata and C.floridanal]are to be distinguished by the fact that while the spikes are on very short culms and may appear basal they are not on basal peduncles.” Fernald (1902) apparently did not understand the concept of basal spikes when he stated that “the best means of distinction between C. umbellata [misap- plied to C. tonsa var. rugosperma (Mackenzie) Crins] and C. nigro-marginata [sic] is of- fered by the thickness of the perigynia.” In his 8th edition of Gray’s Manual, Fernald (1950) placed C. nigromarginata with the basal spike members of section Acrocystis (as Montanae Fries) and then used perigynium characters and geographic region to distinguish C. nigromarginata from C. umbellata and its allies. This means of distinction appears to have been followed by at least Cusick (1992) and Tucker (1995), and may have in part resulted in C. reznicekii often being misidentified as C. umbellata. 1056 BRIT.ORG/SIDA 22(2) Other characters that separate C. umbellata from C. reznicekii include the following: C. umbellata has staminate spikes with 0-2 approximate pistillate spikes vs. 2-4 for reznicekii; some pistillate spikes on elongate peduncles vs. none on elongate peduncles for C. reznicekii; staminate spikes extending up to 9.0 mm beyond the approximate (if present) pistillate spikes vs. staminate spikes at most extending 3.7 mm beyond the pis- tillate spikes for C. reznicekii; and perigynia (2.2-)2.3-3.2(-3.3) mm long vs. perigynia (2.5-)2.7-3.9 mm long for C. reznicekii. In addition, leaf blades of C. umbellata are gener- ally lighter green and slightly wider than C. reznicehii. A full understanding of basal spikes seems particularly important to understand- ing . difference between these two species, and may enrich the understanding of sec- tion Acrocystis as a whole. Basal spikes are individual pistillate spikes which arise al- most directly from the base of the plant [subradical of Mackenzie (1913)]. These pistillate spikes have elongated peduncles, and as with all pistillate spikes in section Acrocystis, they are subtended by bracts. These bracts have sheaths and blades [contrary to Macken- zie (1935) and Roalson and Friar (2004) that state that section Acrocystis (Montanae of Mackenzie) has bracts which are sheathless or subsheathing]. These authors probably meant that bracts of non-basal spikes are sheathless or subsheathing. In addition, the culms of C. umbellata (defined as from the bases of the culms to the apices of the inflo- rescences) are often much taller than the apices of the basal pistillate spikes. Roalson and Friar (2004) consider basal pistillate spikes to be better described as a continuous character with somewhat distinct states, as opposed to a discrete character. While in es- sence this could be true, it appears that this character functions in two completely dis- tinct states with associated other characters (elongated peduncles and sheathing bracts). Even more interestingly, while neither C. nigromarginata nor C. reznicekii has basal spikes, both of ten have “grouped culms.” These “grouped culms” have one culm which is taller and one to three which are shorter (much less pronounced in C. reznicekii), mim- icking the tall and short aspect of C. umbellata, with its often tall culms and shorter basal pistillate spike peduncles. In addition, the shorter culms in a “group” of culms in C. nigromarginata and C. reznicekii have relatively shorter and less projecting terminal staminate spikes. Carex reznicekii and C. nigromarginata share a few characters, including no basal spikes, no elongated rhizomes, at least some culms short and hidden in leaf bases, perigynia of similar length, stigmas 3, and inflorescences composed of 3-5 approximate spikes. Likely because of these similarities, C. reznicekii has often been misidentified as C. nigromarginata. Several characters separate these two species. First, C. reznicekii has narrower leaves with the widest per plant 1.2-2.2(-2.5) mm wide vs. (1.9-)2.3-4.5 mm wide for C. nigromarginata. Second, the tallest culms per plant are shorter. For C. reznicekii the tall- est culms are 1.9-9.9(-13.7)cm long vs. (4.5-)6.6-38.0(-51.0) cm long for C. nigromarginata. Third, individual plants of C. reznicekii have culms often about the same length com- pared to widely different lengths among individuals of C. nigromarginata. Fourth, C. reznicekii culms remain erect even at maturity while some culms droop at maturity for C. nigromarginata. Fifth, C. reznicekii has margins of pistillate scales without red or red only submarginally. The red coloration (if present) can form a submarginal longitudinal stripe, but does not extend laterally to the green (or brown) longitudinal mid-stripe of the scale except at the apex. Carex nigromarginata has pistillate scales either similar in color to C. reznicekii or more often with dark reddish to dark purplish/black coloration WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1057 that extends laterally from the submargin to the green (or brown) mid-stripe of the scale. In addition, the red to purple color often extends all the way to the base of the scale in C. nigromarginata, while in C. reznicekii the reddish color (if present) does not extend to the base of the scale. Sixth, the apex of the staminate spike in C. reznicekii exceeds the apex of the distal-most lateral spike by at most 0.0-3.7 mm. In C. nigromarginata, the apex of the staminate spike which exceeds the apex of the distal-most lateral spike the most, exceeds it by (0.0-)0.5-5.\(-7.3) mm. While this character exhibits considerable overlap in these two species, it can sometimes be useful for distinguishing difficult speci- mens. Seventh, C. reznicekii has staminate spikes on average slightly shorter and nar- rower, measuring 3.3-8.3(-9.2) x 0.4-L.6(-18) mm vs. 4.2-10.9(-12.0) x 0.5-2.4(-2.8) mm for C. nigromarginata. Again this character shows significant overlap, but it occasionally provides useful help in distinguishing between these two species. Carex reznicekii and C. nigromarginata were compared for eight continuous charac- ters (see Table 1). Ranges, means, and standard deviations for all eight continuous charac- ters measured for the NMDS ordination are presented in Table 2. Comparing C. nigromarginataand C. reznicekii, PC-ORD recommended a two-dimensional solution for the NMDS ordination. This resulted in a final ordination (see Fig. 3) with low instability (<0.001) indicating convergence of the iterations, and low stress (6.61) indicating that the data was far from random and there was “no real risk of drawing false inferences” (McCune S& Grace 2002). Axis 1 of the ordination represents decreasing TaCuHe (-0.961) and WiLe Wi (-0.747) with the numbers in parentheses representing the Pearson Correlation Coefficients with Axis 1 (Fig. 3). Axis 1 explained 81% of the variability in the dataset. Axis 2 of the ordination represents increasing TaCuHe (0.772), WiLeWi (0.707), StSpExPiSp (0.883), StSpHe (0.899), and StSpWi (0.702) with the numbers in parentheses representing the Pearson Correlation Coefficients with Axis 2 (Fig. 3). Axis 2 explained an additional 17% of the variability for a total of 98% explained variability. Overall, the ordination summarizes a very Se dataset with several highly correlated variables and in addition shows that while the two species are distinct, some overlap of individual specimens exists. Pearson-correlations illustrated that all of the characters measured except width and length of the perigynia were strongly correlated with one another (p< 0.01), indicating a high level of redundancy of the characters used. The ordination is pre- sented mainly as a visual tool to show how C. reznicekii and C. nigromarginata compare with numerous characters. All of the characters measured except width and length of the perigynia differed significantly between C. reznicekii and C. nigromarginata (Table 3). The scatter plot graph, using a larger sample size for the two strongest characters separating the two species (tall- est culm height and widest leaf width), shows almost all individuals can be determined using these two characters alone (Fig. 4). Ranges, means, and standard deviations for this larger sample size are presented in Table 4. Mainly depauperate or aberrant individuals were not separable with these two characters. These specimens were determined by the additional characters as discussed previously with scale color being the most useful. Regional differences in the characters were significant only for width and length of perigynia (Table 2). Perigynia of C. reznicekii varied by geographical region with slightly narrower and shorter perigynia on plants from the northern region compared to the south- ern and Missouri/Arkansas regions. These statistically significant differences represent only minor variations and have questionable biological significance. For the distribution curves for widest leaf width and tallest culm height a total of 1058 BRIT.ORG/SIDA 22(2) oe. o a —) ~ ps —) Sd is aif Sd 52 o ° Se $ joe e @ ae ° : om af mt Qo) + . = ‘ a —} aa =F a o . a 2 oe o ¢ = oO ca : Se + oA ? Je Be > Q 2 & -0.5 ‘ eo KS 0 C. reznicekii ; 2m * C. nigromarginata s 2° < rs a a a Nn a - Nn -1.5) iE 3 + T 7 T -2.5 -1.5 -0.5 ie 1.5 Axis 1: TaCuHe (-0.961), WiLeWi (-0.747) Fic. 3. Non ic multidi ional scaling ordination showi tial relationshir ie ean C. nigromarginata based on Apidild J y y based On wider WIUCDSL IC dt VIidUe . Main characters correlated to each axis Taste 2.Ranges, means, and standard deviations for characters ee for the NMDS ordination (Fig 3).Char- acter codes are defined in Table 1.Regions listed below are ¢ NE = Virginia/Kentucky north; SE = North Carolina/Tennessee south; and AR/MO = Arkansas/Missouri, Carex reznicekii Carex nigromarginata regions egions Characters NE SE AR/MO Allregions NE SE AR/MO All regions TaCuLe 2.2-12.2 2.5-13.7 3.7-13.7 2.2-13.7 9.0-27.7 6.6-18.7 7.2-38.0 6.6-38.0 4+3.1 5.4+3.3 (3235 5:62:33 15.2452 13.3438 175+88 15.3+6.3 WiLeWi 1.3-2.2 1.3-2.0 1.7-2.2 1.3-2.2 24-40 2.4-3.5 2.5-3.5 24-40 1.6+£0.3 1.7£03 2.00.2 1.7+0.3 2.9+0.5 2/7404 3.0+0.3 2.9+0.4 Anle 1.30-1.80 1.30-1.95 1.50-1.75 1.30-1.95 1.50-2.20 1.45-2.25 1.60-2.50 1.45-2.50 1.50+0.15 1.52+0.17 1600.10 1.5340.15 1.82+0.23 1.84+£0.25 1.98+0.30 1.88+0.26 StSpExPiSp 0.2-2.5 0.2-2.2 ]-3.7 0.2-3.7 0.7-7.3 1.1-6.0 0.5-5.5 -73 1.4+0.7 1.2+£0.5 25 [eZ 1.6+0.9 3.4+2.0 2.7+1.6 3.0+1.7 S017 PeLe 2.75-3.45 2.80-3.70 3.35-3.55 275-3.70 2.70-3.30 2.90-3.60 3.00-3.75 2.70-3.75 3.16+#0.21 3.23+0.27 3.40+0.10 3.244023 3.1440.18 3.2740.25 3.3740.25 3.264£0.24 PeWi 0.95-1.10 05-1.20 Henan 0.95-1.20 0.95-1.35 0.90-1.35 1.05-1.25 0.90-1.35 1.02+0.06 1.13+0.05 1.1340.07 1.09+0.08 1.1140.11 1.14#0.12 1.1240.07 1.12+0.10 StSpHe 4.60-7.55 4.60-7.85 a ae ie na 5.85-12.10 6.15-11.15 5.35-10.70 5.35-12.10 6.31+0.89 6.05+1.05 733+ 1.141 841#2.13 7.79+1.58 8.02+1.74 8.07+1.79 StSpWi eee 1.55 Q.70-1.65 ees ae 1.65 1.05-2.25 1.00-2.00 1.35-1.75 1.00-2.25 27+0.20 1.20+0.28 1.3140.22 1.2540.23 1.62+0.36 1.504£0.31 1.5240.15 15440.28 WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1059 Tasie 3. Summary of F statistics and probability values from one-way ANOVA between C. reznicekii and C. nigromarginata for characters measured for ordination. Characters Fi 50 p TaCuLe 43.8 <0.001 WiLeWi 136.2 <0.001 AnLe 32.6 <0.001 StSpExPiSp 123 0.001 PeLe 0.0 0.914 PeWi 1.6 0.208 StSpHe 13.3 0.001 StSpWi 15.7 <0.000 Taste 4. Ranges, means, and standard deviations for characters measured for the scatter plot graph and charac- ter distribution curves (Fig.4,5, and 6). C. nigromarginata outside of C. reznicekii 150 specimens C. nigromarginata (includes specimens from inside and outside the range of C. reznicekii) 152 specimens range of C. reznicekii 81 specimens TaCuLe 1.9=13.7 4.5-51.0 4.5-51.0 Ss 216 165 2725 19.1+8.0 WiLeWi 1.2-2.5 1.9-4.5 2.1-4.4 AZO 2905 3.00.5 Taste 5.Ranges, means, and standard deviations of particle sizes, pH, and mineral contents of six soil samples. Measurements Measurements Sand 31-88% Mg = 33.67-221.09 ppm 57+20 136.10+71.84 Silt 9-54% Ca 220.28-5509.98 ppm #15 1732.66+ 1931.26 Clay 2-27% Fe 0.64-45.38 ppm 14+£10 16.00+ 20.97 pH 4.35-6.51 Al 15.51-317.91 ppm 549+1.02 111.964117.45 P 0.89-6.40 ppm Mn 20.43-93.99 ppm 2.35+2.06 69.17+27.38 K 24.76-268.13 ppm Zn 0.87-9.21 ppm 125.79+82.20 3.5443.02 150 specimens of C. reznicekii and 152 specimens of C. nigromarginata were measured. Eighty-one of the C. nigromarginata specimens were from outside of the geographic range of C. reznicekii. The distribution curves for widest leaf width and tallest culm height for C.nigromarginataand C. reznicekii combined shows a bimodal distribution of these char- acters (Fig. 5 and 6). The distribution curves for C. nigromarginata and C. reznicekii as separate taxa are overlapping normal or skewed normal curves (Fig. 5 and 6). The distri- bution curves for specimens of C. nigromarginata from outside the range of C. reznicekii closely mimic the distribution curve for C. nigromarginata throughout its range and also clearly contrasts with the distribution curves of C. reznicekii (Fig. 5 and 6). These distri- 1060 BRIT.ORG/SIDA 22(2) 60.07 a a0) °C. reznicekii *C. nigromarginata — 40.0 2 y , ~~ . | ee o B® 30.0 . oe ° : ee . = P ter 4, . — Sa 5 20.0 *4, ee Sere .? Sd ~ *$ woe ee o #3 % 2 * ¢ —| O ry $$+ 33 ¢ aa 10.0 2 o* $f*, , ° . 5 s Heteese + o = iH ol lUf iE 0.0 T T T rT T T T T T T r T T T rT T —_—_—— 1.0 1.2 1.4 1.6 1.8 2.0 2.2 2.4 2.6 2.8 3.0 3.2 3.4 36 3.8 4.0 4.2 44 4.6 4.8 Widest Leaf Width (mm) widest leaf width by t C. nigromarginata and C. reznicekii. Cc + lnat f£.4l1 Fic. 4. Scatter f f g 80.0 - 70.0 - : ie : ‘ —— C. reznicekii and C. nigromarginata -- °C, nigromarginata 60.0 5 . as — --C. reznicekii ea — — C. nigromarginata beyond range of C. reznicekii Number of individuals od vA * 7 N 0.0 t T 7 T T LJ v T T T 7 a a a a an Dh a D a an 7 a 4 . . “0 na e bi + YH eo < ~ S ma >: = rs = es _ ro] fo) ro] er) oe ‘© a + + . ° ye ioe) Widest leaf width (mm) Fic. 5. Distributi f idest leaf width for C. reznicekii, C. nigromarginata, C. reznicekii and C. ig gi bined, and C. f C reznicekii. J J r J bal | J WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1061 100) 907 807 C. nigromarginata and C. reznicekii 2. ~* "°C. nigromarginata = 70 = —""C. reznicekii 607 ee es ieee i C. nigromarginata outside of range of C. reznicekii Z 507 — S 407 Sam 5 307 & 3 207 Zz. 107 “ a n — ————" () ee a SSegRgeege eRRRERRAR RA ZR GB 4 Foto RA SAREE TS SG 1 1 a oO 2 FR aR HH RH GE ESSE Tallest culm height (cm) Fic. 6. Distributi f ll Imt ig fe reznicekii, C. nigromarginata, C. reznicekii and C. ig gi I 1,and C. ig gil I tside the geographic range of C. reznicekii. bution curves help to demonstrate that although there is some overlap in the two most explanatory characters that separate C. nigromarginata and C. reznicekii, the distinction between these two species is not a result of a sampling bias. The two species simply have slightly overlapping normal or skewed normal distribution curves. Geography Carex reznicekii occurs from New England (southern Rhode Island and southern Con- necticut) south to southeastern New York, eastern Pennsylvania, South Carolina, and Georgia west to southern Missouri and northern Arkansas (Fig. 1). It is quite common in the southeastern US. The range of C. reznicekii appears to be totally within the range of C. nigromarginata and C. umbellata (Crins & Rettig 2002). The distribution of C. nigromarginata extends beyond the range of C. reznicekii into southern Ontario, central and western Pennsylva- nia, southern Ohio, Indiana, Illinois, central Missouri, central and southern Arkansas, Oklahoma, Texas, Louisiana, southern Mississippi, southwestern Alabama, and the pan- handle of Florida (specimens examined, Reznicek & Catling 1982; Cusick 1992; Crins & Rettig 2002). The distribution of C. umbellata extends well beyond the range of C. reznicekii into Greenland, eastern and western Canada, and from Nebraska south to Texas (Crins & Rettig 2002). Ecology Carex reznicekii occupies mesic to dry-mesic mostly deciduous or less commonly de- ciduous-pine and/or Ilex opaca Aiton forests. Overstory tree canopy cover often exceeds ca. 60% but does not create dense shade. The shrub layer is usually not very dense. The herb layer varies from quite diverse to not diverse, and quite dense to not dense. Occa- 1062 BRIT.ORG/SIDA 22(2) sionally C. reznicekii grows in clearings in forests. Even at these sites, it is mostly found in the adjacent full-canopy forests. It also occurs along road edges immediately adjacent to forests as well as in the forest interior. Overall, it grows predominately in full-canopy forests, and unlike some other members of section Acrocystis (e.g., C. nigromarginata and C.umbellata), it usually does not do equally as well in more open environments. Carex reznicekii often occurs on slopes above drainages from the bases of the slopes to mid-slopes and occasionally on the upper slopes or crests. It usually does not occur in the actual floodplain of the adjacent drainage unless the drainage is relatively small. Additionally, C. reznicekii usually does not occur on upper slopes, especially if erica- ceous shrubs and Quercus montana Willd. are dominant. Many members of Carex section Acrocystis in eastern North America often grow together. Perhaps this is because they grow ina “generalized” habitat (Cusick 1992). Other members of section Acrocystis observed growing syntopically with C. reznicekii, often within centimeters of it, include C. albicans var. albicans, C. albicans var. emmonsii, C. nigromarginata, C. pensylvanica Lamarck, C. tonsa var. tonsa, C. tonsa var. rugosperma, and C. umbellata. Carex nigromarginata occurred as the most frequent associate, being found with C. reznicekii at 19 out of the 31 sites visited. Despite C. nigromarginata fre- quently growing syntopically with C. reznicekii, some habitat differences between the two were observed. Carex reznicekii usually grew without or with low densities of C. nigromarginata in more calcareous or richer sites as well as lower on slopes. Carex reznicekii usually did not occur or occurred in lower densities with C. nigromarginata at more acidic sites, higher up on slopes, or in more open disturbed habitats. Soils are deep or less frequently shallow over bedrock. Bedrock present at sites in- cludes diabase, limestone, shale, and sandstone. Soil samples showed a wide variety of soil textures ranging from sandy to sandy loam to silty loam to sandy clay loam to clay loam. Table 5 summarizes the ranges, means, and standard deviations of values for par- ticle size, pH, and mineral content of the six soil samples. IDENTIFICATION KEY FOR MEMBERS OF CAREX SECTION ACROCYSTIS EAST OF THE ROCKY MOUNTAINS OF NORTH AMERICA This key works best with ample specimens in mature fruit. Depauperate or especially robust specimens may not key correctly. Rhizomes help in identification, but collectors often fail to collect them. The key attempts, where possible, to get around relying on rhi- zome characters. Parts of the key dealing with C. communis varieties and C. inops ssp. i aii are adapted from Crins and — _ Ir fare fea ‘ lee . Cul pedt near base lige eae times s difficult ie assess pecsuee ne ere can hidden . sheathes of bracts and leaves until they emerge adjacent to distal half of culms); pistillate spikes arising from tee half of culms mostly 0- 2. Pistillate scales om slightly shorter to longer than the perigynia; terminal staminate spikes with or without approximate pistillate spikes; bracts at base of proximal-most non-basal ou spikes (if present) mostly shorter than to sometimes slightly longer i. inflores- ences; bases of old leaves often very fibrous : Perigynia (2.2—)2.3-3.2(-3.3) mm long, beaks a )0.5-1.0 mm long Carex umbellata 3. Perigynia (3.0-)3.1-4.7 mm long, beaks 0.9-2.1 mm long. 4. Perigynia often mostly glabrous; young — short, thick, often smooth adaxially, and ri Carex tonsa var. tonsa 4. Perigynia usually pubescent; young leaves long, thin, scabrous adaxially, and not rigid rex tonsa var. rugosperma WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1063 2. Many pistillate scales shorter than mature perigynia; terminal staminate spikes almost al- wae ae at ce one sabe inane Piauliete spike; Piags a nase of shee most y longer s notor only slightly fibrou > ne (2.2—-)2.6-3.1(—3.2) mm long, beaks 0.4-0.8 mm long, beak teeth up to 0.2 mm long; pistillate scales often shorter than the body of mature perigynia; staminate spike 3.5-5.9(-6.7) mm long; rhizomes slender; plants loosely caespitose Carex deflexa Hornem. var. deflexa 5, Perigynia 3.1-4.2 mm long, beaks 0.9-1.7 mm long, beak teeth 0.2-0.5 mm long; ae aia scales ond longer than the body of mature perigynia; staminate spikes 4.2-11.6 m lon j zomes ice) ut; plants densely Ca ng;r spitose pss rossii Boott . Culms with ectie pies: short- seauneulaie to sessile (proximal-most non-basal spikes oc- casionally elongate-pedunculate) and arising from distal half of culms [occasional culms will have one pistillate spike with an elongate peduncle arising from base of culms ( represent hybrids)]; pistillate spikes arising from distal half of culms mostly 2-4 6. Longest staminate spike at least 12.3 mm long (use ample specimens); rhizomes elongated (except C. albicans var. albicans, C. communis, and C. novae-angliae: these ane out in either lead of 6): perigynia bodies globose to sree as long as or longer than 7. Widest ae equal to or less than 1.2 mm wide; widest leaf ee wide; perigynia bodies ellipsoid, longer than wide. 8. Widest ae anes 1.8 mm wide; proximal internode of inflorescence 5.2—29(—40) m long; staminate ae peduncles 1.2-8.0(-8.6) mm long; proximal-most pistillate ae peduncles 0.8-9.8 mm long; spike above the proximal-most spike subtended by twh ftenh longat pistillat | tred, i with s these may 1.1-2.7 mm me ed-brown Carex novae-angliae Schwein. 8. Widest leaf blades 1.3-2.7 mm wide; proximal internode of inflorescence 1.0-13.2( 0.1) mm long; staminate spike peduncles 0.3-4.5(-8.8) mm long; proximal-most pis- tillate spike peduncles 0.3-2.2(-3.3) mm long; spike above the proximal-most spike subtended 2 a bract which usually has at most a short awn; pistillate scales some- times partly r 9. Rhizomes Seeaes and spreading 9. Rhizomes short and ascending to erect iS Widest poy ae cag anes 2mm wide; wi widest feat blades 1.5-6.1 out Carex albicans var. australis (Bailey) Rettig x albicans var. albicans mm wide; wide to slightly longer = than W 10. ene leaves (2.3-)3.0-6.1 mm wide; plants with large caespitose clumps; not colo- nial; rhizomes short and ascending to erect; bracts subtending the spike above the proximal-most with an elongate awn tip that arises from scarious margins that are truncate to convex at summit; staminate spikes 0.9-2.2(-2.5 wide 1. Perigynia beak teeth 0.1-0.2(—0.3) mm long; pistillate scales usually 1.6 mm wide or less; widespread 11. Perigynia beak teeth usually more than 0.2 mm long; pistillate scales usually more than 1.6 mm wide; Georgia and South Carolina Caren communis var. communis Carex communis var. amplisquama i Rettig 10. Widest leaves less than 3.0(-3.5) mm wide; plants with small caespitose clumps; colo- nial; rhizomes spreading and long; bracts subtending the spike above the proximal- most with a short or no awn tip that arises from lee margins that mostly taper into the distal part of the bract; staminate spikes (1.1-)1.5-3.2(—5.2) mm wide. 12. Perigynia 1.1-1.6(-1.8) mm wide. 13. Perigynia beaks 0.3-0.8(-0.9) mm long, beak/body ratio 0.13-0.50(-0.56) Carex pensylvanica 13. Perigynia beaks 0.9-1.6 mm long, beak/body ratio 0.50-0.92 Widest leaf blades greater than (1.7-)2.0 mm wide; northeastern North Am Carex lucorum Willd. ex Link var. lucorum 14. Woes leaf blades less than 2.0 mm wide; southern Appalachian Mountains Carex lucorum var. austrolucorum Rettig 1064 6. BRIT.ORG/SIDA 22(2) 12. Perigynia (1.6-)1.7-2.2 mm wide Carex inops ssp. heliophila Longest staminate spike less than 12.3 mm long (use ample specimens); rhizomes not elon- gated (except C. floridana and C. albicans var. australis: the latter keys out in either lead of 6) perigynia bodies ellipsoid, longer than wide (except C. deflexa and C. communis: the latter keys out in either lead o 15. Pistillate scales shorter than body of mature perigynia they subtend. 16. P roximal internode of inflorescence 5.2-29(-40) mm long; staminate spikes (4.6— 5.1-12.1 mm long Carex novae-angliae 16. Proximal internode of inflorescence 1.3-8.5(—14.5) mm long; staminate spikes 3.5 8.3(-9.2) mm lon 7. Culms mostly surpassed by leaves; perigynia (2.2-)2.6-3.1(-3.2) mm long Carex deflexa var. deflexa Culms surpassing leaves; perigynia (3.0-)3.1-4.2 mm long Carex peckii Howe 15. ee oan longer than body of mature perigynia they subtend. 18. Widest leaf blades (2.3-)3.0-6.1 mm wide; proximal internode of the inflorescence (5.9-)7.5-36.0 mm long; bract subtending the spike above the proximal-most w an oe awn tip that arises from s | j ia hod) re arious margins that are truncate to convex at tol lly ellipsoid g to slightly longer to broadly ely about than wide. 9. Perigynia beak teeth 0.1—0.2(—0.3) mm long; pistillate scales usually 1.6 mm wide or less; widespread communis var. communis 9. Perigynia beak teeth usually more than 0.2 mm long; pistillate scales usually more t 1.6 mm wide; Georgia and South Carolina arex communis var.amplisquama 18. Widest leaf blades 1.1-4.5 mm wide (if widest leaves are over 2.7 mm wide then proximal internode of the inflorescence 0.8—5.4 (8.0) mm long); bract subtending the spike above the proximal-most with or without an elongate awn tip that arises from scarious margins that mostly taper into the distal part of the bract (occasion- ally truncate to convex at summit); perigynia bodies ellipsoid, longer than wide 20. oe leaf blades 1.1-1.8 mm wide; proximal internode of inflorescence 5.2— -40) mm long; staminate ae oooada at most pistillate spike peduncles 0. the proximal-most spike subtended by a bract waied often has an elongate awn; pistillate scales without true red, sometimes with some red-brown Car . Widest leaf blades 1.2-4.5 mm wide; proximal eae of inflorescence 0.8- 13.2(-20.1) mm long; staminate spike peduncles 0.1—4.5(-8.8) mm long; proxi- mal-most pistillate spike peduncles 0.0-2.2(—3.3) mm long;spike above the proxi- mal-most spike subtended by a bract which usually has at most a short awn pistillate scales sometimes with s 1.2- ool 8.6) mm long; ‘Proximal: NO (Se) arex novae-angliae 21. Longest leaf blades up to 2.2(- oe cme longas th median culm length; see 2.2—3.2(-3.3) mM long; proximal internode of inflorescence 1.0- 13.2(-20.1) mm long; staminate spike peduncles 0.3-4.5(-8.8) mm lo 22. Rhizomes elongate and spreading 22. Rhizomes short and ascending to erect 23. Staminate spikes (5.5-)6.6-1 Carex hes var. australis 2.2(-14.5) mm long; middle and upper staminate scales mostly with midrib faint, not raised, not scabrous towards tip of scale, and mostly without a minute awn arising just below tip of scale ex albicans var. albicans . Staminate spikes 4.0-8.7(—9.3) mm long; middle and er stami- nate scales mostly with midrib more prominent, raised, antrorsely scabrous towards ae of scale, and with a minute awn ary from just below tip of sc C NO WW No mart var. emmonsii ieee Hee ee )2.5 times as long as the median culm length; perigynia (2.5—)2.7-4.0 mm long; proximal internode of inflorescence 0.8 5.4(-8.0) mm long; staminate spike peduncles 0.1—0.8(—1.6) mm lon 24. Rhizomes horizontally spreading to ascending and elongate; mostly at WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1065 least some stigmas 2; mostly at least some achenes with only 2 pale nerves; sheathes of old leaves at base of plant usually weakly ote Carex floridana 24. Rhizomes ascending to erect and short; stigmas 3; achenes with 3 pale nerves; sheathes of old leaves at base of plant often strongly fibrous. 25. Widest leaf blades 1.2-2.2(-2.5) mm wide; culms often about equal length, the longest 1.9-9.9(-13.7) cm long; any reddish color on pis- tillate scales below the distal tip not extending laterally Helin near the margin to the green or brown longitudinal mid-strip side of mid-vein; the longest staminate spike up to 3.7 mm oiler oS the distal most lateral cass am staminate spikes 3.3-8.3( m long X 0.4-1.6(-1.8) m e Carex reznicekii : fe leaf blades (1.9 cee mm sere culms often of quite vari- able length, the longest (4.5-)6.6-38.0(-51.0) cm long;at least some pistillate scales often with reddish to purplish/black color below the distal tip extending laterally from near the margin to the green or brown longitudinal mid-stripe on either side of lea long- N [oa] est staminate spike (0.0-)0.5-5.1(-7.3) mm taller t! stal most lateral spike apex; staminate spikes 4.2-10.9(-12.0) mm x 0 2.4(-2.8) mm wide Carex Taremerenat Representative specimens of Carex reznicekii: poemce sabes with a single artes were measured for the ordination (Fig, °) ale ones marked witha single or d tk 1€ sCallel plot graph and character ibution curv Fig 4 5a nd .A, ALABAMA. Dekalb Co.: near top of Red Mountain ae NW of Fort Payne, 21 Apr 1943, Harper 3918 (MO, ust ), Harper 3919 (BH**, GH, US). Houston Co.: vicinity of Columbia, Thomas Creek up to bluff along Ommussee Creek, 12 Mar 1998, MacDonald 10970 (BRIT**). Jackson Co.: ca. 3 mi W of Carns, along E side of county Route 33, 22 Apr 2002, Naczi 9034 (DOV™*); just E of Route 35, 0.1 mi S of junction with Route 40, 21 Apr 2004, Werier 2075 (BH*), county Route 33, 8.05 mi by road SE of junction with Route 79, 21 Apr 2004, Werier 2081 (BH**, MICH, NCU). Limestone Co.: by 1-65, ca. 5 mi N of US 31 turnoff to Athens, 26 Mar 1986, Kral 73127 (BRIT **). Morgan Co.: by US 31,1 mi S of Lacon, 12 Apr 1970, Kral 38342 (BRIT*). tascalovss Co.: by AL 69,9 mi N of Moundville, 1 Apr 1984, Kral 71138 (BRIT**). ARKANSAS. Baxter Co.: Cole F | pple Turkey Road, 26 Apr 1993, Hyatt 5243 (MICH*). Independence Co.: 6.95 mi S of Sharp Co. Line (in Cave ae on US Hwy. 167,9 Apr 1994, Hyatt 5838 (MICH*), Hyatt 5839 (MICH*). Izard Co.: ca. 0.5 mi SE of Baxter Co. on AR Hwy. 5, 18 Mar 1998, Hyatt 7879 (MICH**). Johnson Co.: [Ozark National Hore st], Horsehead Lake Recreation Area, western most picnic area, 12 Apr 1996, Hyatt 6896 (MICH*). Marion Co.: Buffalo N | River, 100-200 m NE of Buffalo River Bridge of state Hwy. 14, 10 Apr 1994, Hyatt 5844 ne Newton Co.: ‘= of Harrison, 0.8 mi N of Harp Creek, along Hwy. 7,14 Apr 1984, Castaner 7992 (DOV**). Stone Co.: Mountain View vicinity, about 10 mi SSE on AR Hwy. 5,8.5 mi N of Cleburne Co. line, 18 Mar 1998, Hyatt ae a), CONNECTICUT. New Haven Co.: ee a New Haven, West Rock Ridge, a little N of “Judges Cave,” 20 May 2005, Werier 2533 & Haines (BH** Kent Co.: above Choptank River floodplain at Sandtown landfill, N of Sandtown, 30 Apr 2003, pee = (DOV*). [New Castle Co.]: Mount Cuba, | May 1898, Bicknell 3255 (NY*); near Centreville, 28 Apr 1876, Commons sn. (PH**). Sussex Co.: 3 mi E of Milford, along S side of Beaverdam Branch, ca. 0.2 mi W of its crossing by road 200, 21 Apr 2003, Naczi 9521 (BRIT, DOV™). [Sussex Co.]: Laurel, 25 Apr 1882, Smith 717 (US*™). [DISTRICT OF COLUMBIA]. [Washington Co.]: Washington, 25 Apr 1897, Steele s.n.(GH**); District of Columbia and vicinity, 7 ia 1899, Williams s.n.(NY**), Anacostia to Twining City, 7 Apr 1918, Leonard 139a (US**). GEORGIA. Clarke Co.: of Athens, ce of Rock and Shoals Road off Barnett Shoals Road, 24 Feb 1981, Manhart 202 (BRIT**, MICH**, a NYS,). Clay Co.: 1 mi E of Fort Gaines, 27 Mar 1949, Thorne G& Muenscher 9312 (GH**). Dade Co.: Sitton’s Gulch, Lookout Mountain, 2 mi SE, Trenton, 17 Mar 1939, Hermann 10192 (GH*, NY*™). Hall Co.: Chipowee Woods (Elachee Preserve), 19 Mar 2005, Hyatt 11802 (BH**), 9 Apr 2005, Hyatt 11807 (BH**). Randolph Co.: near Griers Cave, N of Cuthbert, 3 Apr 1948, Thorne & Muenscher 7910 (BH**). Walker Co.: W side of GA Route 136, 1.6 mi by road W of GA Route 193, 0.4 mi W of Cooper Heights Community, 8 Apr 2000, Hill 32220 ot Ww a 136, 1.6 mi by road W of junction with Route 193, 20 Apr 2004, Werier 2069 (BH*, MICH, NCU). KE Butler Co.: Roundabout Swamp, at end of Biggerstaff Road, off Hwy. 1153, 22 Apr 1993, ees 5360 es Edmonson Co.: Mammoth Cave National Park, Ugly Creek S eae, with Little Jordon Cemetery Road, 23 Apr 1991, rane aa 4622 (BRIT**). Rowan Co.: Lic ek ork, 4 1938, Braun 1758 (US**). MARYLAND. Caroline Co.: Tuckahoe State Park, off Cherry Lane l mi qorniae with Mason Branch Road, J wl 1066 BRIT.ORG/SIDA 22(2) 14 Apr 1997, Frye 779 (MICH**), 2 May 2004, Werier 2176 (BH*), Werier 2178 (BH**, MICH, US); Tuckahoe State Park, entrance to C Lane picnic area., 2 eo reall Werier 218] (BH**, MICH, US). Cecil Co.: along Bohemia Creek, Middle Neck, 7 May 1932, Long 37288 (PH**). Dorchester Co.: N side of Puckum Branch, W of Puckum Road, N of Eldorado, 15 oe 2003, McAvoy 5797, ae GE Biot (DOV*). Kent Co.: above the “Big Marsh,” W of Betterton, 27 se 2003, McAvoy 5739 (DOV**). Montgomery Co.: Plummer’s Island in the Potomac River, n Cabin John, 10 May 1915, Van Eseltine 325 (BH**). Wicomico Co.: S side of Bear Swamp Road, SW of See ‘ Apr 1999, a 4268 (DOV**). MISSISSIPPI. Lafayette Co.: NE of Oxford, 5.4 mi NE of junction of MS 7 and 30 on MS 30 then ca. | mi S on Hopewell Road, 12 Apr 1997, Bryson 15732 (BRIT, DOV**). Lowndes Co.: 2.5 mi N of Steens, N side of Vaughn Robertson Road, 20 Apr. 1986, Bryson 4217 & Curtis (MICH, MO), 12 Apr 1987, Bryson 5341 (MICH*™*). MISSOURL. Shannon Co.: above intermittent tributary of E branch of Dennis Hollow, 13 May 1997, Brant 3696 (MO**): Rocky Creek Falls, 24 Apr 1983, Merriott 61 (MICH*). NEW JERSEY. Burlington Co.: Bordentown, along Crosswicks Creek, 27 Apr 1910, oe 3103 (NY**). [Gloucester Co.]: Swedesboro, 25 Apr 1917 Mackenzie 7559 (NY**). Sussex Co.: Andover Township, SW of Lake Mohawk, just N of Route 613, 0.5 mi by road W of junction with Route 617, 4 May 2004, Werier 2191 (BH*™, iene NY); Stokes State Park, S of county Route 636, 0.25 mi E of US Route 206, 4 May 2004, Werier 2194 (BH*, MICH, NY). [Warren Co.]: between Pike and Pequest River ca. 1/4 to 3/8 mi E by slightly NE of Buttsville, 18 May 1919, Pretz 9602 (PH**). NEW YORK. New York Co.: Manhattan Island, Inwood, 27 May 1893, Bicknell 1237 (NY**). Orange Co.: Town of aa SE end of sor es aan N of the village of Greenwood Lake, SW of Route 17A, 7 May 2003, Werier 1507 & Barbour (BH*). Ulster Co.: Town of Saugerties, 1.0 km E of Old Dutch Road, 1.1 mi S$ eg road from Route 32, 22 May 2006, Werier se Barbour & Barbour (BH). NORTH CAROLINA, Davidson Co.: 1 mi E of Yadkin River near NC 4 Apr 1956, Radford 9476 (NCU*™*); S of Route 49, 0.6 mi E of bridge over Yadkin River, 14 vy 2004, Werier 1995 (BH*, MICH), Werier 2000 (BH**, MICH, NCU), Werier 2001 (BH**, MICH). Macon Co.: Nantahala National For- est,ca.5 mi E of Franklin, then 5 mi N of Ellijay Road, 16 Apr 2000, Hyatt, ae a Me Millan 9112 (MICH**); W of Ellijay Road and Ellijay Creek, 3.7 mi by road NE from Route 64, 19 Apr 2004, Werier 2048 (BH*, MICH, NCU); E of Route 2, 5.4 mi by road NE from Route 64, 19 Apr 2004, Werier 2052 (BH**, DOV, MICH). Madison Co.: 0.5 mi WNW of Marshall on NE side of ave n Broad River, 26 Apr 1958, Ahles 38966 & Duke (NCU**). Mecklenberg Co.: 2.4 mi SE of Davidson on NC 73, 22 Apr 1958, Ahles 38691 & Britt (NC — Co.: Phills Creek, N side, 3 mi W of Chapel Hill, 18 Apr 1940, Radford & Stewart 469a (NCU**). Sw : Nantahala National Forest, Wessen vicinity, S side of US Hwy. 19, 10 Apr 2000, Hyatt 9071 (MICH**). PENNSYLVANIA. Berks Co.: 5/8 mi ENE of Walnut & 14th Street, Reading, 20 May 1943, Berkheimer 3563 (NYS*). [Lancaster Co.]: Lancaster, aaa [no collector] s.n. (MICH**); [no location information], [no date], [Muhlenberg ?]s.n. (PH**), [no location information], [no datel, Mu ae: s.n.(PH**). RHODE ISLAND. [Washington Co.]: South Kingston, 13 May 187 pee ie 1776 H CAROLINA. Chester Co.: near Catawba River at Great Falls, 15 Apr 1961, Radford 43511 (NCU™). ame Co.: W of Catarrh, just E of Lynches River, and just S of Route 903, 17 Apr 2004, Werier 2013 eee MICH). Fairfield Co.: 2 mi NE of Altston, 15 Apr 1961, Radford 43473 (N NCU**). Greenville Co.: E of Marietta, small road, 20 m SE from Route 414, 2.1 mi by road NE from Route 276, 18 Apr aoa - erier 2029 (BH**, MICH). Horry Co.: 2 mi S of Myrtle Beach, 18 Apr 1932, Weatherby & Griscom 16461 (PH **) Lancaster Co.: near Catawba River, 2 mi NE of Great Falls, 15 Apr 1961, Radford 43524 (NCU): E of Great 7. anda little bit E of the ees ba River, 17 Apr 2004, Werier 2023 (BH**, MICH), Werier 2027 (BH**); E of US 601, S side of Flat Creek, 21 Apr 19 Williamson F138 & Ahles (NCU*™), 16 Apr 2004, Werier 2010 (BH*, MICH). Pickens Co.: SC Department of ee ral Res stern tract of Lake Jocassee, S of Mill Creek entrance, 19 May 1998, Waldrop W-274 & Townsend (MICI > just E of tributary to Oolenoy River and SSR 69, 0.1 mi by road N from Route 288, 2.5 mi by road E from Route 178, 18 Apr 2004, Werier 2038 (BH*™*, MICH), Werier 2041 (BH**, DOV NCU). TENNESSEE. Franklin Co.: x 6 road mi S of Huntland, along E of oo 97, 2.6 road mi S of junction of Routes 97 and 122, 24 tel 1998, zi 7249 & Ford (VPI*™). Grundy W of state Route 50 (Elk Head Rd.), 5.7 mi by road SW from junc- tion an Route 108, 22 Apr 2004, ara oe (BH*, MICH); E of Route 56 at hair pin turn, 6.6 mi by ee S from Warren Co, line, 22 Apr 2004, Werier 2098 (BH™, ea Co.: W side of the Cumberland River at its junction with Cub Creek, 19 Apr 1998, Phillippe 29579 (MICH**); NE of Cub Creek just before junction with Cumberland River, 23 Apr 2004, Werier 2102 (BH**, MICH). Jefferson Co.}: French Broad River, near eee aa 1844, Rugel s.n. oe NY**). [Knox Co.]: Knoxville, Cherokee Bluffs, 9 Apr 1930, Underwood 5 (GH*™*). Marion Co.: Send of Little Cedar Mountain, W side of Nickajack Lake, and 2.8 km N of Interstate 24 bridge W i Haletown, 8 Apr 2000, wea 10418 (MICH**, VPD; E slope of Little Cedar Mountain, ca. 3.5 km N of interstate 24 bridge over Nickajack Lake, NNW of Haletown, 8 Apr 2000, Wieboldt 10423 (MICH™*, VPI). Maury Co.: bluffs of Duck River ca. 2.5 mi SE of Columbia and 1.2 mi E on Sowell Pike from TN 50, 13 Apr 1974, Kral 42463 (BRIT**, MO). Polk Co.: N of Ocoee Lake and Route 64, 1.7 mi W of junction with Route 314, 20 Apr 2004, Werier 2062 (BH*), 20 Apr 2004, Werier 2063 (BH**, MICH). Putnam Co.: bluffs along Caney Fork River by I-40, 10 Apr 1992, Kral 80112 WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1067 & Moore (BRIT**). Rutherford Co.: just N of Luvergne, 9 Apr 1976, Kral 57547 (BRIT**, MO). Sevier Co.: N of Little Cove a Road, 4.8 mi by road E of Wear Valley Road, 27 Apr 2004, Werier 2132 (BH**, MICH). VIRGINIA. Accom .: Sside of road to Sinnickson, 18 Apr 1999, McAvoy 4266 (DOV**). elbow arle Co.: Mount Alto, near nae 21 Apr 1973, Stevens 6523 (NCU*), 21 Apr 1973, Stevens 6525 (NCU*™*). [Albemarle Co.]: Monticello Mountain, the Grove, 21 Apr 1980, Wieboldt M-26 (PH**). Alleghany Co.: N of Route 18, N of small creek which is a tributary of Potts Creek, 0.6 mi by road S from Route 657, 30 Apr 2004, Werier 2153 (BH**, DOV, MICH, VPD, 30 Apr 2004, Werier 2155 (BH**); 30 Apr 2004, Werier 2156 (BH**, MICH, VPD. Botetourt Co.: NW of Route 615, 0.7 mi by road N of junction with Craig Co. line, 29 Apr 2004, Werier 2143 (BH**, MICH, MO, NCU, VPD, 29 Apr 2004, Werier 2146 (BH**). Brunswick Co.: along Hwy. 670, 2.3 mi NE of its junction with Hwy. 46, 2 May 1986, Rettig 1523 (NYS**); Route 670, 2.2 to 2.3 mi by road N of Route 46, 11 Apr 2004, Werier 1983 (BH**, MICH, VPI); along Hwy. 712, 7.6 mi N of its junction with Hwy. 58 in Edgerton, 2 May 1986, Rettig 1525 (MO, NYS*™*); 7.7 to 7.8 mi by road N of Route 712 from Route 58 in Edgerton, 1] Apr 2004, Werier 1972 (BH**, MICH, VPD, Werier 1979 (BH**). Buckingham Co.: <— oe of David Creek, ca. 1 mi NE of Bent Creek, 9 May 1983, Wieboldt 4635 & Britten (MO**, VPI. C e Con rth Anna River, 10 Apr 2004, Werier 1954 (BH**), Werier 1958 (BH**), Werier 1960 (BH*™*, en ‘Werier 1961 oe MICH). Chesterfield Co.: 7 mi WNW of Winterpock, along S side of Route 360, 0.2 mi E of Appomattox River, 30 Apr 2005, Naczi 10792 (BH, DOV**, MICH). Culpepper Co.: 2nd peak, Buz- zard Mountain, 11 May 1941, Allard 8589 (VPI*). Fauquier Co.: C.F Phelps Wildlife se ie Area, ca. 1.9 mi SW ss Kelly Ford, E of Rappahannock River, | May 2004, Werier 2168 (BH**, MICH, NCU, VPI). Henry Co.: along Smith River about 0.9 mi below state Route 636 bridge and ca.5 mi NNE of Ridgeway, 28 Apr 1987, lee see Wiebol idt oe Isle of Wright Co.: near Walters, 6 Apr 1938, Fernald & Long 7775 (GH, PH**). King e Co.: 2 mi SSW of Owens, along E side of Route 301, 1.5 mi SW of its junction with Route 218, 29 hs 2005, ae 10778 ca ee Montgomery Co.: N of state Route 637 at NW end of Coffee Valley, ca. 7 km E of Ellett, 1l May 1993, Wieboldt 8509, Ludwig, Rawinski, et al. (DOV**, MICH*™, VPD; N of Coffee Hollow at NW end of hollow, NE of Seneca Hollow Road, 28 Apr 2004, Werier 2138 & Wield boldt (BH**, MICH). New Kent Co.: 4m of Seu along E side of Route 33, 0.5 mi SW of its junction with Route 273, 1 May 2005, Naczi 10822 on DOV**, MICH). Newport News City: behind baseball field on Monroe, Fort Eustis, 22 Apr 1974, Appler 1064 & ee) (NCU). Patrick Co.: along Spoon Creek ca. 10 mi SE of Stuart, 27 Apr 1983, Wieboldt 4619 (MO**, VPD. Pittsylvania Co.: along N-facing bluff of the Staunton River ca. 4 mi N of Straightstone, 23 Apr 1985, Me 5472 (MICH*™, VPI} aoe Mountain, ca. ; km NE of Leaksville Junction on Southern Railroad, 28 Apr 1 Wieboldt 8454 (DOV™, MICH**, VPD. P . Powhatan hee Wildlife Management Area, NW of i con, 11 Apr 2004, Werier 1964(BH* ‘ -MICHD, ele eon MICH, VPD. Prince Edward Co.: 7 mi SSW of Farmville, along tributary of Briery Creek, ca.0.3 mi N of dam olf Briery eet. Lake, S of road 790, 30 Apr 2005, Naczi 10811A (DOV**). Prince George Co.: Coggins Point, 4 Apr 1939, Fernald & Long 9699 (GH, PH**). Shenandoah Co.: S end of Short Mountain, Massanuttens, ca. 3 mi SE of Mount Jackson, | May 1982, Wieboldt 4284 & Davenport (VPI*™). Southampton Co.: by the Nottoway River near Davis School, Indian Road, NW of Courtland, 4 Apr 1940, Fernald, Long, & Pease 11654 (GH*™, PH**). Surry Co.: W of Claremont, 19 Apr 1942, Fernald, Long, & Abbe 14117 (GH, PH**), ca. 1/2 mi NE of reactors at VEPCO Surry Nuclear Power Station, 13 Apr 1974, Ware 5391 & Wieboldt (NCU). Sussex Co.: 4 mi S of Stony Creek, 5 Apr 1937, Fernald & Long 6952 (GH, PH**). Wythe Co.: Rock Creek 2.6 km E of community of Cripple Creek, 6 May 2004, Wieboldt 11390 (VPI*™*) oe APPENDIX A Specimens of Carex nigromarginata measured for the ordination (Fig. 3) are marked with a single asterisk; the ones marked with a single or double asterisk were measured for the scatter plot graph and character distribution curves (Fig. 4, 5, an CANADA. ONTARIO. Norfolk Co.: Long Point, Squires Ridge, 10 1/2 to 13 mi from base of point, 25 May 1980, ae icek 5496 (MICH**); Long Point, middle of a Creek Ridges to mee Bay, 15 to 19 mi from base of point, 5 Jun 1980, Reznicek 5636 (MICH**). U.S.A. ALABAMA. Autauga Co.: Autauga County 9, W of Ivy Creek, ca.0.5 mi E are with Autauga County 15, 16 ee 1980, Wiersema 1537 (BRIT**). Butler Co.: by county ei ae ol Pigeon Creek, 11 Mar 1991, Kral 78407 (BRIT**); by I-65,4 mi N of exit to AL 106, 26 Mar 2002, 14 (BRIT Dallas Co.: ca. 1 mi WNW of Cahaba, 31 Mar 1972, Kral 45340 (BRIT**). Escambia Co.: N side a Rolle 55,ca.05 mi W of Route 41,19 Mar [19]82, Wilhelm 10013 (BRIT**). Jackson Co.: just E of Route 35,0.1 mi S of junction with Route 40,21 Apr 2004, Werier 2072 (BH”*); off of county Route 33,8.05 mi by road SE of junction with Route 79, 21 Apr 2004, Werier 2079 (BH**). Monroe Co.: E bank of Alabama River above landing at Haines Island, 15 Apr 1980, Wiersema 1482 (BRIT**). Morgan Co.: by US 31,1 mi.S of Lacon, 12 Apr 1970, Kral 38342 (BRIT**). Perry Co.: Hwy. 219,3.0 mi S of Bibb County line, 9 Apr 1986, Rettig 1443 (BRIT**). ARKANSAS. Bradley Co.: Warren Prairie Natu- 1068 BRIT.ORG/SIDA 22(2) ral Area, SE of Warren, 17 Apr 1986, Rettig 1494 (MO**); Warren Prairie, ca. 15 mi W of Monticello, 3 Apr 1983, arc 2962 (BRIT*™); about 1 mi W of pe 18 Mar 1995, ean 6161 (MICH**). Calhoun Co.: Hwy. 160 mi W of Bradley County, 18 Mar 1995, Hyatt 6166 (MICH**). Clark Co.: Caddo Valley vicinity, 1 May 1995, yt oe (MICH**). Columbia Co.: US Hwy. 79 at Ouachita ees Te 18 Mar 1995, Hyatt 6173 (MICH**), Crawford Co.: just S of Rudy, 20 Apr 1985, ae 8218 (MO*); Road 220 just S of Lee’s Creek, 20 Apr 1985, Castaner 8224 (MO**). Drew Co.: Monticello, A.& M. College Farm, 21 Apr 1937, Demaree 14558 (MICH**); 2.8 mi S of Lincoln County on US Hwy.425,29 Mar 1997, Hyatt 7287 (MICH**). [Garland] Co.: Hot Springs National Park, Mar 1930, Connell s.n. (MO*). Howard Co.: Saline River at its junction with Hwy. 84, 15 Apr 1986, Rettig 1474 (MO**). Miller Co.: along Hwy. 237,0.7 mi N of its junction with Hwy. 160, 14 Apr 1986, Rettig 1470 (BH**); Hwy. 237,0.3 mi S of junction with Hwy. 134, 14 Apr 1986, Rettig 1472 (BRIT**). Nevada Co.: AR Hwy. 4 at Ouachita County line, 18 Mar 1995, Hyatt 6172 (MICH**):; by I-30, 3.3 mi SW of junction with AR 26, 16 Apr 1999, Kral 88225 BRIT**). Quachita Co.: NW of Camden, AR Hwy.4 loop, 1.6 mi E of AR Hwy. 376 junction, 18 Mar 1995, Hyatt 6169 (MICH**). Pike Co.: 0.3 mi S of Montgomery County on AR Hwy. 369, 15 Apr 1996, Hyatt 6938 (MICH**). Polk Co.: Ouachita National Forest, Along Forest Service Road 38 to Shady Lake, 3.2 mi N of its junction with Hwy. 246, 15 Apr 1986, Rettig 1477 (MO**); Ouachita National Forest, along Forest Service a 38, 1.0 mi N of entrance to Shady Lake at j EOL with pores Service Road 508, 15 Apr 1986, Rettig 1480 (MO**). Pope Co.: Long Pool Recreation Area,O rest, above Big Piney Creek, 16 Apr 1986 cee 1491 (MO*). Sevier Co.: Saratoga So 5 Apr Reese (MICH*™); DeQueen, 1.9 mi N on US Hwy. 71,23 Apr 1995, Hyatt 6949 (MICH). ott Co.: Ouachita National Forest, approximately 5.5 mi S, 1.5 mi E of junction of Hwy. 80 and business loop pee 71 in Waldron, 9 Apr 1999, Morse 2843 (BRIT**); Ouachita National Forest, approximately 2.5 mi S, 3.0 miW of Boles, 10 Apr 1999, Morse 2904 (BRIT**). Sharp Co.: N of Cave City on 167,2 mi S of junction with 167/56,6 May 1979, Castaner 5667 (MO*), Stone Co.: Ozark National Forest, =) aici DISHES ETON, R11W,S15,NW4 of NW4, 27 Apr 1993, Hyatt 5251 (MO*).Union Co.: S of El Dorado, at a roadside lcome center 5.7 mi N of Louisiana on US Hwy. 167,18 Mar 1995, Hyatt 6159 (MICH**). DELAWARE. INewCastle Co.]: Newcastle, [no date], Canby n. (BH*). DISTRICT OF COLUMBIA. [Washington Co.]: Brookland, 14 May 1899, Holm s.n.(MO*);Washington, ne date], Vasey 187 (BH**). FLORIDA. Gadsden Co.: near Flat Creek, general vicinity of Sycamore, 6 Mar 1977 Godfrey 75723 (BRIT**); N of FL Route 20 between Ochlockonee River and Hosford, 15 Mar 1975, Godfrey 74186 (BRIT**). Liberty Co.: ca.6 mi NE of Bristol, N of Route 12 just W of its junction with Route 271, 14 Mar 1994, Naczi 3620 (BRIT**). GEORGIA. Clarke Co.: near Winterville, 2 Mar 1929, Miller & Maguire 297 (BH**); GA State College Agricultural Farm, 9 Mar 1929, Miller & Maguire 298 (BH**); Princeton Woods, § side of Oconee River, 6 Apr 1929, Miller & Maguire 300 (BH). Dade Co.: E side of Route 136, 3.55 mi by road NW of junction with Route 189, 20 Apr 2004, Werier 2067 (BH**). Lincoln Co.: ner sum Gio Mountain, 5 Apr 1939, Clausen & Trapido 3794 (BH*). Rabun Co.: SE of Glade Mountain, 17 Apr | 33 (BH**). Upson Co.: Dripping Rocks, 30 Mar 1948, Smith et al. 3667 (BH**). ILLINOIS. Al ee Shawnee National Forest, Jonesboro Ranger District, Compartment J-104, 5 May 1994, Phillippe 23977 ee Union Co.: Ozark Hill Prairie cane Natural Area, Shawnee National Forest, 8 Apr 1992, Phillippe 19778 (MICH**). KENTUCKY. Reale Co.: along Hwy.411,0.4 mi E from Hwy. 85, 27 Apr 1987, McKinney 2473 (BRIT*). Lawrence Co.: above state Route | ae ae 0.8 mi NE of eek of Benbow Road, 12 Apr 1990, Cusick 28744 (BRIT*). LOUISIANA. te Parish: State Line Road 14.8 mi of junction with Hwy. 171,13 Apr 1986, Rettig 1457 (MO**). Claiborne Parish: Kisatchie National Forest Caney ee district, 10 Apr 1996, Hyatt 6876 (MICH**). Grant Parish: Kisatchie National Forest Catahoula Ranger district, 6 Mar 1995, Hyatt 6157 (MICH**): Kisatchie National Forest, about 3 mi NE of Williana, 3 Mar 1997, Hyatt 7865 (MO**); Dry Prong community vicinity, 22 Apr 1996, Hyatt 6857 (MICH**). Morehouse Parish: S of LA 835 by Bryant Main Line Road then E to end of Kitpatrick Road, 30 Apr 1992, Bryson 11441 (MICH**). Natchitoches Parish: Kisachie National Forest,ca. 1 mi SE of Goldonna at Saline Bayou on LA Hwy. 156,24 Feb 1998, Hyatt 7860 (MO**); Kisatchie National Forest, Kisatchie Ranger district, compartment 4, 10 Mar 1998, Hyatt 7872 (MICH**). Ouachita Parish: 3 mi SW Cheniere, 11 Apr 1959, Kral 8510 (BRIT**). Rapides Parish: Brushy Creek Riparian Site, 23 Mar 1998, a ee eee Sabine Parish: LA 175 at Pleasant Hill, 3 Oct 1967, Thomas 5635 (BRIT**). Union Parish: y. 2, mi E of H 33/15 in Farmerville, 11 Apr 1986, Rettig 1456 (BRIT**). MARYLAND. Baltimore Co.: ieee 7 mi NE ie Baltimore, 5 May 1910, Churchill s.n. (BH**), Caroline Co.: Tuckahoe State Park, W of Cherry Lane, 1.2 mi S from junction with Mason Branch Road, 2 May 2004, Werier 2177 (BH*), Werier 2179 (BH**).Montgomery Co.: Great Falls, 30 May 1917, ee 1302 (BH**). Prince George Co.: near Congress Heights, 25 May 1915, VanEseltine 385 (BH**). MISSISSIP tala Co.: 5.5 mi SSE of McCool, 12 Apr 1986, Bryson 4152 (MO**); 2.6 mi SSE of Attala-Carroll County line fa MS Hwy. 35,12 Apr 1986, Bryson 4138 (MO**).Holmes Co.: Holmes County State Park, 16 Mar 1968, McDaniel 10376 (BRIT**). Kemper Co.: White Horse Mountain, ca. 10.5 air mi N of Dekalb, 6 Apr 1972,McDaniel 15865 (BRIT**). Lafayette Co.: 4.2 mi S of intersection of MS Hwys. 7 and 6 bypass,s of Oxford,9 Apr 1987, Bryson 5327 (MO*). Leake Co.: near Yockanooka ny Creek, ca. WERIER, A NEW SPECIES OF CAREX FROM EASTERN NORTH AMERICA 1069 8 mi WSW Carthage, 2 Apr 1972, McDaniel 15819 (BRIT**). Oktibbeha Co.: 0.5 mi NW of Sturgis, Hannah Hill, 20 r 1989, Bryson 8463 (MO*).Winston Co.: W Betheden Lutheran Church, 6 Mar 1974, Bryson 318 (BRIT**); 2.5 mi NE Louisville,6 Mar 1974, Bryson 317 (BRIT**). MISSOURI. Butler Co.: Poplar Bluff Bottomland Hardwood Natural Area, 24 Apr 1982, Castaner 6749 (MO*). Callaway Co.: 0.25 mi E of Williamsville, 25 Apr 1997, Summers & Vander- bilt 8062 (MO**). Carter Co.: Mark Twain National Forest, 3 mi S of Van Buren off Hwy. 103 in Spring Valley, 6 May 1995, Summers & Wallace 7334 (MO*). Laclede Co.: Mark Twain National Forest, Gasconade River, 2 mi W of Adam Ford, 24 Jun 1995, Summers 7460 (MO**).Madison Co.: 8 mi S of Twelve Mile Creek, along Hwy.67, 26 Apr 1987, Castaner 9602 (MO*). Shannon Co.: Middle Hollow, 7 May 1997, Brant 3687 (MO*). Ste. Genevieve Co.: Hickory Canyon Natural Area, ca. 2 mi N of Sprott on Hwy. EE, 23 May 1996, Summers et al. 7800 (MO**); Pickle Springs Natural Area, ca.0.25 mi E of Hwy. AA, ca. 1.5 mi SE of Hwy. 32, 20 Apr 1996, Summers et al. 7695 (MO**). Wayne Co.: Mark Twain National Forest, 2 Apr 1998, Brant 3882 (MO*). NEW JERSEY. Somerset Co.: Chimney Rock, Bound Brook, May 1935,Edwards s.n. (BH**). [Gloucester Co.]: Newfield, 20 May 1873, Ellis s.n. (BH**). NEW YORK. Orange Co.: Town of Warwick, SE end of Bellvale Mountain, N of village of Greenwood Lake, SW of Route 17A, 7 May 2003, Werier 1506 and Barbour (BH*). NORTH CAROLINA. Davidson Co.: a little E of Tuckertown Road, 1.5 mi by road S of Route 49, 14 Apr 2004, Werier 1992 (BH**), Werier 1993 (BH*), Werier 1994 (BH**); S of Route 49,0.6 mi E of bridge over Yadkin River, 14 Apr 2004, Werier 1996 (BH**). Montgomery Co.: Flint Hill Road, ca. 2.5 mi by road W of Lovejoy Road, 13 Apr 2004, Werier 1986 (BH**). Swain Co.: 100 yards inside Graham-Swain County line on US 129, 11 Apr 1983, Reznicek 7132 (MO*); SE of Route 19 at junction of Route 28, 26 Apr 2004, Werier 2128 (BH**). Transylvania Co.: N side of Route 64 just E of Lake Toxaway, 18 Apr 2004, Werier 2044 (BH™*). OHIO. Gallia Co.: Symmes Creek Road, 1.25 mi S of Gallia-Centerpoint Road, 9 Apr 1986, Cusick 25054 (MICH*™). Jackson Co.: Camp Canter's Cave, N of Caves Road, 16 Apr 1990, Cusick 28752 (MICH**). Scioto Co.: N side of Dark Lick Road ce mi Ne oi Perici Road, 19 Apr 1990, McCormack 2094 (MICH**). OKLAHOMA, nate Co.: 3miWonSH63 klal li Apr 1992 Magrath 1 18592 ee “ side OK Sees Imi SE of Little ie eee at Honobia, 19 py 1993, Rezn oe 0353 mer oe ‘ord Co.: 0.75 mi WSW of Breezewood, 16 May 1946,Berkheimer7110(BRIT*™).B eee ne = 3 May 2004, ney 185 (BH**). [Carbon Co.]: ca. 1/8 mi W of Lehigh Gap LV.R R station, 22 May 1927, Pretz 12925 (BH™), Centre Co.: Ingleby, 2 mi E of Coburn, 5 May 1936, [Wahl] 2486 (BH*™*); ee 6 a) ae eaten & Wale 2528 (BH**); abandoned railroad at Ingleby, 7 May 1977, Keener 3401 (MO**). Huntingdo 29 May 1979, Lahham & Keener 15 (BRIT*). Potter Co.: Town of Os N of US Route 6,ca.0.5 mi NW of Galeton, May 2005, Werier 2546 (BH*). SOUTH CAROLINA. Anderson Co.: 2 mi SE of S Springs, 6 Apr 1939 ee - (BH**). Chesterfield Co.: N side of Route 9,W of Thompson Creek,W of Zoar Road, 15 Apr 2004, Werier 2003 (BH**). Darlington Co.: Society Hill, [no date], Curtis 106(BH**).[Darlington Co.]: W of Hartsville,9 Apr 1921,Norton C21 (BH), Norton s.n.(BH**).Greenville Co.: 20 meters SE from Route 414, 2.1 mi by road NE from Route 276, 18 Apr 2004, Werier 2028 (BH*). Lancaster Co.: E of Route 601, S of Flat Creek, 16 Apr 2004, Werier 2011 (BH**);E of Great Falls and a little bit E of the Catawba River, 17 Apr 2004, Werier 2024 (BH**). Oconee Co.: SE side of Stump House Mountain, 7 Apr 1939, Clausen & Trapido 3811 (BH**). Pickens Co.: just N Route 288, 1.2 mi by road E of Pumpkintown and Route 8, 18 Apr 2004, Werier 2031 (BH**), Werier 2034 (BH**), Werier fe (BH**); just E of SSR 69,0.1 mi by fea N from Route 288, 2.5 mi by road E from Route 178, 18 Apr 2004, Werier 2039 (BH*). TENNESSEE. Davidson Co.: Dyer Observatory, 30 Mar 1976, Kral 57466 (MO*).Grundy Co.: NW of state Route 50, 5.7 mi by road SW from junction with Route 108,22 eee Werier 2093 (BH**).Polk Co.:N of Route 64,11.1 mi by road E of junction a Route 314,20 Apr 2004, Werier 2059 (BH**). Sevier Co.: N of Little Cove Creek Road,4.8 mi by road E of Wear Valley Road, Apr 2004, Werier 2134 (BH**). TEXAS. Cass Co.: Hwy. 155, 1.5 mi SW of its junction with Hwy. 59 in Linden, 14 Apr 1 At 1469 (MO**);3 mi SE Hughes Springs, 2 Apr 1979,Kral 63243 and Carter (BRIT**). Houston Co.: FR 227,0.1 mi NW on junction with Davy Crockett Road (NF 526), 10 Apr 1990, 1309 and Wipff (MO**).Smith Co.: Tyler State Park N of Tyler, 13 Apr 1986, Rettig 1465 (BRIT**). Upshur Co.: Hwy.155,9.5 mi “SW of its junction with Hwy.259 in Cedar Springs, 14 Apr 1986, Rettig 1467 (MO**). VIRGINIA. Botetourt Co.: NW of Route 615,0.7 mi by road N of junction with Craig Co.line, 29 Apr 2004 Werier 2148 (BH**). Brunswick Co.: 7.7 to 7.8 mi by road N of Route 712 from Route 58 in Edgerton, 11 Apr 2004, Werier 1969 BH**), Werier 1971 (BH**), Werier 1975 (BH*); Route 670, 2.2 to 2.3 mi by road N of Route 46, 11 Apr 2004, Werier 1981 (BH*™*). Caroline Co.: near North Anna River, 10 Apr 2004, Werier 1952 (BH), Werier 1957 (BH**), Werier 1959 (BH**).Craig Co.: Potts Mountain ca.6 mi NW Castle, 18 May 1983, Wieboldt 4657 (MO*). Powhatan Co.: Powhatan State Wildlife Manage- t Area, NW of M 11 Apr 2004, Werier 1963 (BH**). WEST VIRGINIA. Binge Go: county Bette 2, ] TSS — Route 2 at Sevan 17 Apr] 991, Cusick 29399 (MICH**). Pleasants Co.: a! f Micki land Creeks, 10 1984, Cusick 23333 and Ortt Ce aie oo county Route 8, 1 1/2 mi S of US Route 50,8 May 1985, chee aes (BRIT*). Tyler Co.: j y Fairview Church, 3 May 1989,Cusick 28021 and Ortt (BRIT**). = = ACKNOWLEDGMENTS I thank Anton Reznicek for advice, thoughts, comments, and help on various aspects of 1070 BRIT.ORG/SIDA 22(2) my research project; Susan Reznicek for drawing the illustration; Anita Barbour, James (Spider) Barbour, Arthur Haines, and Thomas Wieboldt for assistance with field work: Nat Cleavitt for help with analysis of data; Neil Golder for help translating my abstract into Spanish; Guy Nesom and Thomas Vining for assistance with the Latin diagnosis; Peter Ball, Nat Cleavitt, Bob Dirig, Philip E. Hyatt, Richard LeBlond, Robert Naczi, Anton Reznicek, and Troy Weldy for reviewing my manuscript and providing valuable sugges- tions; the Bailey Hortorium and its staff for providing space to do my research; and Bob Dirig of BH for processing all of my loan requests and helping with other details related to my work at BH. Lalso thank the curators from B, BRIT, DOV, GH, MICH, MO, NCU, NY, NYS, PH, US, and VPI for loans of specimens and/or assistance during my visits and with my inquiries; and Philip E. Hyatt and Robert Naczi for sending recently collected speci- mens. The New York State Museum Biological Survey provided funding for survey work in the Hudson Highlands of New York which led to the initial discovery of C. reznicekii. REFERENCES Crins, W.J.and J.H. Rettic. 2002. Carex Linnaeus sect. Acrocystis Dumortier. In: Flora of North America Editorial Committee, eds., Flora of North America North of Mexico, Vol. 23. Magnoliophyta: Commelinidae (in part): Cyperaceae. Oxford Univ. Press, New York. Pp. 532-545, Cusick, A.W. 1992. Carex section Acrocystis (Cyperaceae) in Ohio. Michigan Bot. 31:99-108. Fernaco, M.L. 1902. Contributions from the Gray herbarium of Harvard University. The variation of some boreal carices. Proc. Amer. Acad. Arts 37:495-513. Fernaco, M.L. 1950. Gray's manual of botany. 8th ed. American Book Co., New York. KUKENTHAL, G. 1909. Cyperaceae-Caricoideae. In: A. Engler, ed., Das Pflanzenreich. IV. Vol. 20, Heft 38: Leipzig: Wilhelm Engelmann. Pp. 1-824. LEBLOND, R.J.,A.S. Weakey, A.A. Reznices, and W.J.Crins. 1994. Carex lutea (Cyperaceae), a rare new coastal plain endemic from North Carolina. Sida 16:153-161. Mackenzie, K.K. 1913. Notes on Carex VII. Carex umbellata and its allies. Bull. Torrey Bot. Club 40:529- 554. Mackenzie, K.K. 1935. Montanae. N.Amer.Fl. 18:185—206. McCune, B.and J.B. Gract. 2002. Analysis of ecological communities. MJM Software Design, Gleneden Beach, OR. Naczi,R.F.C.,R. Krat, and C.T. Bryson. 2001. Carex cumberlandensis,a new species of section Careyanae (Cyperaceae) from the eastern United States of America. Sida 19:993-1014. Reznicek, A.A. and K. Cameteeke. 1996. Carex porrecta (Cyperaceae), a distinctive new species for north- ern South America and Costa Rica. Novon 6:423-425, Reznicek, A.A. and P.M. Catuna. 1982. Cyperaceae new to Canada from Long Point, Norfolk County, Ontario. Canad. Field-Naturalist 96:184-188. Roatson, E.H., J.T. CoLumpus, and E.A. Friar. 2001. Phylogenetic relationships in Cariceae (Cyperaceae) based on ITS (nrDNA) and trnT-L-F (cpDNA) region sequences: Assessment of subgeneric and sectional relationships in Carex with emphasis on section Acrocystis. Syst. Bot. 26:318-341. Roatson, E.H. and E.A. Friar. 2004. Phylogenetic relationships and biogeographic patterns in North American members of Carex section Acrocystis (Cyperaceae) using nrDNA ITS and ETS sequence data. Pl. Syst. Evol. 243:175-187. Tucker, G.C. 1995. Preliminary keys to Carex (Cyperaceae) in New York State. New York Fl. Assoc. Newsl. 6(2):3-18. A NEW SPECIES OF EUGENIA (MYRTACEAE) FROM THE MONTEVERDE REGION, COSTA RICA Fred R. Barrie Missourl Botanical Garden PO. Box 299, St. Louis, Missouri 63166, U.S.A. ang fe; IdM artment rela 1400 5, lake Shore Drive, cee lin inois 60605 fbarrie@fieldmuseum.org ABSTRACT on oe | -R eT | \ 1c gee VSR co eer | ] Puntarenas Anew species, E provinces of Costa Rica. It may be distinguished from E. rhombea Dy chatacters of the leaves and flowers, espe- cially the calyx. The species is apparently restricted to cliff edg 1100 to 1350 meters. ABSTRACT to (AA y Se describe una nueva especie, Eugenia haberi B de las provincias de Guanacaste y Puntarenas en el lado pacifico de Costa Rica. Puede diferenciarse de E. rhombea por caracteres de las hojas y flores, especialmente el caliz. La especie esta restringida aparentemente a bordes de acantilados a elevaciones de 1100 a 1350 metros. INTRODUCTION With about 700 species, Eugenia is the largest of the New World genera of Myrtaceae. Forty-nine Eugenia are currently known to occur in Costa Rica, including 21 recently described (Barrie 2005) and the species described here. Eugenia is characterized by a race- mose or fasciculate inflorescence, flowers with an open, four-lobed calyx and an embryo that is an undifferentiated mass of tissue. reco haberi Barrie, sp. nov. (Fig. 1). Type: COSTA RICA: PUNTARENAS. Monteverde, Eston’s Cliff Edge, 350 m, | Jan 1979 (£1, fr), W. Haber 263 (HOLOTYPE: MO}; ISOTYPES: MEXU!, MO!, N lint? oO r , pedicellis brevioribus et lobis Eugenia rhombea (O. Berg) Krug et Urb. aemulans, differt foli calyce minoribus. Small tree; young growth glabrous or with a few hairs on the margins of the bracts, bracteoles and calyx. Branchlets compressed at the outer nodes; bark reddish brown. Leaves narrowly elliptic, the blades 3-8 x 1-2.5 cm, 2-5 times as long as wide, coriaceous, drying a concolorous green or somewhat darker above, the midvein and margins tan; midvein convex on both surfaces, often with a narrow ridge above: lateral veins 15-20 per side, straight, often obscure above; marginal veins similar to the laterals, straight, ca. 1mm from the margins; glands numerous on both surfaces, sometimes punctuate above: base narrowly cuneate to weakly concave; margins cartilaginous, decurrent along the petiole; apex acuminate to caudate-acuminate, the tip ultimately rounded; petioles 2-5 mm long, flattened or weakly concave dorsally. Inflorescences axillary, racemose, soli- tary; axis 1-5 x 1 mm long; flowers 4-10; bracts 0.5 mm in diameter, persistent, circular, the margins scarious, with or without a few hairs; buds 2-3 mm long, ovoid. Flowers pedi- cellate, the pedicels 4-12 mm long, with prominent pellucid glands; bracteoles 0.7-1 0.5 mm, persistent at least through anthesis, ovate, prominently glandular, the base free SIDA 22(2): 1071- 1073. 2006 1072 BRIT.ORG/SIDA 22(2) MISSOURI BOTANICAL GARDEN HERBARIUM N° 2740686 HOLOTYPE of Eugenia haberi Barrie F.R, Barrie 2006 Mi COSTA RICA Puntarenas Family: MYRTACEAE Eugena ct, COStariCensit kery Monteverde, Eston's cliffedge, 1350 m. Small tree. W. Haber 263 2 January 1979. MISCANID! BATAMIFAL ‘RA OMeM UeRR anne cans Fic. 1. Holotype of Eugeni ( , MO) BARRIE, A NEW SPECIES OF EUGENIA FROM COSTA RICA 1073 or slightly connate, the margins entire or scarious, with or without a few hairs; hy- panthium 1.5-2 mm long, campanulate; calyx lobes in subequal pairs, 0.7-1 x 0.5 mm, elliptic, prominently glandular, the margins entire or witha few hairs, the apex rounded: petals ca. + x 3.5 mm, ovate, the margins with or without a few hairs, the apex rounded; disk 1.2-2 mm in diameter, glabrous; stamens ca. 50, 3-5 mm long; style 5-6 mm long, glabrous. Fruits 6-11 x 6-10 mm, globose or obovoid, mature color black; pericarp thin- walled, glabrous; calyx persistent but reduced. Eugenia haberi is a small tree found along cliff edges on the Pacific slope in Guanacaste and Puntarenas provinces at elevations of 1100-1350 meters. In many respect it resembles E. rhombea (O. Berg) Krug & Urb., a species of coastal areas with a circum- Caribbean distribution. Although E. rhombea occurs in both Panama and Nicaragua, it has yet to be found in Costa Rica. The similarities between the two species include gen- eral habit, the overall absence of vestiture, with hairs restricted to the margins of the bracts, bracteoles and perianth, and the coriaceous leaves with cartilaginous margins. The two may be differentiated by the shape of the leaves, which in E. haberi is narrowly elliptic with a tapering, narrowly cuneate base and an acuminate to caudate-acuminate apex. In E. rhombea, the leaves are ovate to elliptic with a rounded or broadly cuneate base and an acute apex. The inflorescences in E. rhombed are often superficially fasciculate, and the pedicels of the flowers and fruits are up to 30 mm long. In E. haberi the inflorescence rachis may be short but it is apparent and the pedicels do not exceed 12 mm. The calyx lobes of E. rhombea are much larger and more prominent, up to 4 mm long and 2 mm wide, and are prominent in flower and fruit, retaining a pale color that contrasts with the dark body of the mature fruit. The calyx persists but is not conspicuous on the fruits of E. haberi. Etymology.—The species is na med for Bill Haber, the collector of the type, whose deep and extensive knowledge of the flora and fauna of the Monteverde region is exceeded ae by his Best in sharing the information with others. Addi Ico] |. COSTA RICA. Guanacaste: Canton Liberia, Parque Nacional Guanacaste, ee Cacao, 1100 m, 10°55'45'N, 85°28'15"W, 22 Nov 1990 (v?) C. Chavez 391 (NB); Estacion Cacao, 7 May 1 Chavez 553 (CR, E INB, MO); Estacion Cacao, 11 Apr 1991 (fr), C. Chavez 541(CR, INB, MO). Sania ner Hoge Uppan, 10 Feb 1979 (fl) W.A. Haber 263 (CR); Monteverde, lower community on Pacific slope, 1350 m, 10°18'N, 84°48'W, 5 Jul 1991 (fr) W. Haber & C. Ivey 10720 (CR, F, INB). ACKNOWLEDGMENTS This research was conducted while in residence in the Botany Department of the Field Museum. I thank the department and the chair, Michael O. Dillon, for their generous, con- tinuing support. Thanks to Bill Haber for sharing his knowledge of the field characteris- tics of this species, and to Bruce Holst, Lucia Kawasaki, and Les Landrum for reviewing this manuscript. REFERENCES Barrie, FR. 2005. Thirty-five new species of Eugenia (Myrtaceae) from Mesoamerica. Novon 15:4-49. 1074 BRIT.ORG/SIDA 22(2) Book NOTICE Neat K. VAN ALFREN, GEORGE BRUENING, and WILLIAM O. Dawson (eds). 2006. Annual Review of Phytopathology: Volume 44, 2006. (ISBN 0-8243-1344-5, hbk; ISSN 0066-4286). Annual Reviews Inc., 4139 El Camino Way, P.O. Box 10139, Palo Alto, CA 94303-0139, US.A. (Orders: ane awe org, 800-523-8635, 650-493-4400, 650-424-0910 fax: onlineactivati rg). $188.00 (USA), 538 pp., 7 3/8" x 91/4" Annual Review of Phytopathology: mH Contents of Volume 44 of A Retrospective of an Unconventionally Trained Plant Pathologist: Plant Diseases to Molecular Plant Pathology The Current and Future Dynamics of Disease in Plant Communities A Catalogue of rhe Effector Secretome of Plant Pathogenic Oomycetes Genome Packaging by Spherical Plant RNA Viruses Quantification ie Modeling of Crop Losses: A Review of Purposes Nonsystemic Bunt Fungi—Tilletia indica and T. horrida: A Review of H Significance of Inducible Defense-related Proteins in Infected Plants Coexistence of Related Pathogen Species on Arable Crops in Space and Time ng Nonpersistent and Semipersistent Transmission of Plant Viruses istory, Systematics, and Biology Virus-Vector Interactions Mediatir Breeding for Disease Resistance in the Major Cool-Season Turf grasses Molecular Ecology and Emergence of Tropical Plant Viruses Biology of Flower-Infecting Fungi A Model Plant Pathogen from the Kingdom Animalia: Heterodera glycines, the Soybean Cyst Nematode Reveals What a An Enterobacterial Plant Pathogen Comparative yeno The Dawn of Fungal Pathogen Gen Fitness of Human Enteric = ee on nPl ants and Implications for Food Safety The Role of Ethylene in Host-Pathogen Interactions mae Cc ae ae in Fluorescent Pseudomonas spp. Biosynthesis and Regulation Long-Distance RNA-RNA Interactions in Plant Virus Gene Expression and Replication Evolution of Plant ca ie in Streptomyces Climate Change Effects on Plant Disease: Genomes to Ecosystems SIDA 22(2): 1074. 2006 TAXONOMIC REVIEW OF SYMPHYOTRICHUM PATENS (ASTERACEAE: ASTEREAE) Guy L.Nesom Botanical Research Institute of Texas 509 Pecan Street Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT The eastern North American Symp} ichum patens has been treated by Jones (1983, 1992) and other olan to include var. patens, var. patentissimum, and var. gracile. These taxa h ily by habit, involucral size and vestiture, and phyllary orientation. aa cisHine Hons between v var. patens and var. gracile have been sedaiee as or APE ne and nS two ie a have b wide region; the present study h them inan Sn or evolutionarily mean- ingful way, and var. gracile is not fecconiecd Variety patens sensu lato appr eney Comprises two open on three or four tetraploid population systems. Variety } tetraploid, and it ene ae ae eae var. paren: ai their oe are mera or peaks A county-level map shows th the two infraspecific taxa a S ee as interpreted in the present | Loi previous studies. Ana) Alt +1 “ cr BE aaron ‘ sak ack study. A similar RESUMEN eal patens del este de Norte América ha sido tratado por Jones (1983, 1992) y otros botanicos para incluir las var. patens, var. patentissimum, y var. gracile. Bae taxa han sido esters panels mente por el ae tamano involucral y pubescencia, y la orientacion de los filarios. I g var. patens y var. gracile ha sido vista como solapada y los dos ta ido i d impa u region ae el presente estudio concluye que no es ae distinguirlos ae un moso que no sea Oranere oen una linea evolutiva significativa, y la var. gracile no se reconoce. La vaneudG ae sensu lato comprende aparentemente dos sistemas de poblaciones diploides y tres 0 cuatro tetraploides. La variedad patentissimum es ead lanes diferente ys constantemente tetraploide, y se intergrada con la var. ci es es ToT IOIGS, alli do ua 1 Un mapa a nivel de condado muestra la dist de los dos taxa eee de S. ee tal como se secre en a Diesenis estudio. un mapa similar muestra la distribucion ploidal de la especie, The eastern North American Symphyotrichum patens (Aiton) G.L.Nesom has been treated (especially in detail by R.L. Jones 1983, 1992, as Aster patens) to include var. patens, var. gracile (Hook.) G.L.Nesom, and var. patentissimum (Lindl.) G.L.Nesom. Infraspecific taxa have been distinguished primarily by habit, involucral size and vestiture, and phyllary orientation. Morphological differences, however, especially between var. patens and var. gracile, have been construed as overlapping and those taxa have been viewed as broadly sympatric. Differences among the varieties have been characterized as in the key below, which is amalgamated mostly from contrasts and comments by Cronquist (1980), A.G. Jones (1992), Jones (1983), and Brouillet et al. (2006). 1. Involucres broadly turbinate, 8-12 mm oa phyllaries often in 5-7 series, appressed, obtu mid phyllaries ovate-lanceolate, 1.2-1.7 mm wide, densely strigillose to sericeous- pene eglandular to sparsely and minutely a aeanae var. patentissimum Involucres campanulate or slenderly turbinate, 5.0-7.5 mm long; phyllaries in 4-5(-6 ee often at least somewhat squarrose, acute to acuminate, mid phyllaries linear-lanceolate, 0.7—1 mm wide, sparsely to densely strigillose, sparsely 2 oo sessile- or stipitate- glandular. 2. Plants slender branches long conspicuous! 1d minutely bracteate; princ | cauline leaves —_— aii SIDA 22(2): 1075- 1080. 2006 1076 BRIT.ORG/SIDA 22(2) 1-3(-4) cm long, 0.5-1(-1.5) cm wide; involucres 5.5-6.5(-7.5) mm long; phyllaries sparsely glandular and densely strigillose, mid 0.7—1 mm wide var. gracile 2. Plants more robust, branches shorter, with fewer and larger bracts; principal cauline leaves 3-6(-8) cm long, 1-2 cm wide; involucres 6-7.5 mm long; phyllaries densely glandular and sparsely strigillose, mid 1—1.2 mm wide var. patens Variety patentissimum occupies a geographic range in the Interior Uplands west of the Mississippi River, except for populations in southern Illinois. Jones (1983) mapped scat- tered collections of var. patentissimum from Texas and Louisiana; Cronquist (1980) included Mississippi in the range. Variety patens has been recorded from Maine (Magee & Ahles 1999), New Hampshire, and Masschusetts south to Florida, thence westward to Texas, Okla- homa, and Kansas (Jones 1983). Variety gracile was recorded by Jones (1983) in Texas, Okla- homa, Kansas, Missouri, Arkansas, Louisiana, Mississippi, and Alabama — sympatric in each state with var. patens and sympatric in Missouri, Arkansas, and Oklahoma with var. patentissimus. A.G. Jones (1992) viewed all three taxa as present in Oklahoma. Brouillet et al. (2006, Flora of North America treatment) presented an expanded geographic concept of var. gracile, following Cronquist (1980), both of whom recognized its occurrence in Florida, Georgia, South Carolina, North Carolina, Virginia, Tennessee, and Maryland, far into the range of var. patens. In the view of both R.L. Jones and of Brouillet et al, the range of var. gracile is completely sympatric with that of var. patens. Variety patens in the concept of Brouillet et al. occurs alone from New Jersey and Delaware northward to Maine and west- ward from Pennsylvania and New York to Indiana and Illinois. All chromosome number reports of var. patentissimum have been tetraploid (2n = 20). Many populations of var. patens (sensu Jones 1983) have been reported as tetraploid (counts from Massachusetts, Illinois, Indiana, New Jersey, Kentucky, Tennessee, Virginia, North Carolina, South Carolina, Georgia, Alabama, and Mississippi); diploids have been reported from western Alabama (Jones 1983; Semple 1984). Variety gracile has been re- corded at diploid level (2n = 10) in Alabama, Mississippi, Louisiana, Texas,and Oklahoma. Tetraploid counts of var. gracile have been reported from Louisiana and Texas Jones 1983; Semple 1984). Artificial intervarietal crosses between tetraploids (var. patens x var. patentissimum) “produced good seed sets, and the hybrid offspring were intermediate in morphology” (Jones 1983, p. 60). Crosses between var. patens and Symphyotrichum phlogifolium (both tetraploid) were as successful as intervarietal crosses. In contrast, “only a few crosses were attempted between diploid and tetraploid S. patens, and none of these were successful” OO); — The concept of Symphyotrichum patens in earlier literature has included S. georgianum (Alexander) Nesom and S. phlogifolium (Muhl. ex Willd.) Nesom, but Jones (1983) elevated these taxa to specific rank, and most later authors, including Brouillet et al. (2006), have followed these concepts, with Cronquist (1980) and Gleason and Cronquist (1991) being notable exceptions. Neither S. georgianum nor S. phlogifolium intergrades morphologically with other taxa; S. georgianum is a decaploid (2n = 50) while S. phlogifolium isa tetraploid (2n = 20), and both occur mostly within the geographic range of tetraploid S. patens var. patens. Another distinct but closely related species, S. grandifolium (L.) Nesom, is known only as a dodecaploid (2n = 60). The chromosome counts for these three taxa were first reported by R.L. Jones and A.G. Jones (1979). The present study reexamines the taxonomic distinctions and geographic distribu- tions of the infraspecific taxa of Symphyotrichum patens. The study is primarily based NESOM, TAXONOMIC REVIEW OF SYMPHYOTRICHUM PATENS 1077 on specimens examined from BRITSMU, MISS, MO, NCU, TEX-LL, and VDB (including many of the collections made by R.L. Jones in his Vanderbilt dissertation study). The col- lection localities of these are mapped in Figure 1. Semple (1984) mapped chromosome number reports for Symphyotrichum patens, as similarly done here, but the present map (Fig. 2) is more precisely represented and the modified taxonomy gives it a different perspective. Chromosome counts have been made by various botanists but mostly by Semple and Brouillet (1980), Jones (1983), and Semple (1984), as summarized by Semple (1984), with an addition by Jones and Smogor (1984). Several chromosome counts, vouchered by collections by John Semple in 1991 and depos- ited at TEX and MO, apparently have not been reported in literature: Kentucky, Whitley Co., 2n = 20 (MO); North Carolina: Wilkes Co., 2n = 20 (MO); Ohio: Scioto Co., 2n = 20 (MO); Texas: Harrison Co., 2n = 10 (TEX) and Fort Bend Co., 2n = 20 (TEX), West Virginia: Pendleton Co., 2n = 20 (MO). RESULTS AND DISCUSSION Observations of Jones (1983) and of Brouillet et al. (2006) are confirmed here: plants re- ferable to var. patens (using criteria as in the key above) occur essentially throughout the range of var. gracile, and plants referable to var. gracile occur through part of the range of var. patens. Map 14A of Jones (1983) indicates that the eastern extension of var. gracile is abruptly demarcated along the Mississippi-Alabama border, but the present study has not confirmed this. Typical var. gracile-like plants occur northeastward at least into North Carolina. Similarly, collections from Texas and Louisiana resemble plants identified as var. patens in Atlantic states. There appears to be a weak east-to-west trend in increase of nonglandular involucral vestiture and, beginning in Alabama westward, development of longer and more bracteate branches, but the present study has not found any geo- graphic zone of morphological discontinuity that would enable or justify the recognition of two taxa. Continued formal recognition of var. gracile perpetuates a typological defi- nition. In the present view, recognition of broadly sympatric varieties (as var. gracile with var. patens) would imply either that the two were reproductively isolated (thus each bet- ter treated at specific rank) or else that they represent populational variants (thus not justifiably treated at varietal rank). Variety patentissimum isa distinctive entity. It overlaps in distribution (Fig. 1) with tetraploid var. patens, and putative hybrids and introgressants apparently occur in the region of sympatry: Arkansas, Oklahoma, the southwest corner of Kansas, and (east of the Mississippi River) southern Illinois and extreme western Kentucky (intermediates not represented on map). Some measure of reproductive isolation may be active, as typi- cal plants of both taxa have been collected in relatively close proximity in their region of sympatry. Typical var. patentissimum occurs as far south as Cleburne and Dallas coun- ties in south-central Arkansas and Sevier and Little River counties in the southwest cor- ner of the state; var. patent issimum and var. patens and inte Li occur in Craighead and Poinsett counties near the northeast corner of the state. Plants and populations of var. patens sensu lato of different ploidy level do not ap- pear to be intermixed. More counts, of course, might reveal that diploids and tetraploids occur in sympatry, but based on current evidence, diploid populations within var. patens apparently occur in two geographic groups, as hypothesized in Fig. 2. Tetraploids appar- ently occur in four groups (or three groups, if tetraploidy has been derived only once on the east and west sides of the Mississippi River) (Fig. 2). More counts might reveal that 1078 BRIT.ORG/SIDA 22(2 A var. patentissimum @ var. patens Be 1. eDistabuten i Symphyoricnum patens var. patens and var. patentissimum. Symbols indicate counties from which specimens y. The record for var. patens in tay oe Kansas, is s fide R.L. Jones (1992); that for var. patens in Portage Co., Ohio, is fide Fisher (1988) Th g | New York, Rhode Island, Connecticut, New Hampshire, Massachusetts, and Maine. diploids are continously distributed along the outer coastal plain The nomenclatural type (lectotype) of var. gracile was collected in Washington Par- ish, Louisiana, a region where tetraploids, but not diploids, have been found. Morphologi- cal or ecological features that might be diagnostic of these population systems of differ- ent ploidy level are not apparent. Diploid and tetraploid plants of var. patens in Mississippi, for example, as well as in Texas, show similar ranges of morphological variability. Even so, it may be assumed that the ploidal systems within var. patens are isolated from adja- cent ones by production of triploid hybrids or some other incompatibility Jones noted that experimental 2x-4x crosses were not “successful”. ACKNOWLEDGMENTS A loan of specimens from MISS was critical for studying the pattern of variability in the Gulf Coast region. | also am grateful for help from staff at MO, NCU, and TEX-LL during visits there and for helpful reviews by Ron Jones and Luc Brouillet. NESOM, TAXONOMIC REVIEW OF SYMPHYOTRICHUM PATENS 1079 ge es eel ge os a ried Decree Tes oe 1 Let ie 4 age: He FR EP STN ci re Or eke eons MELEE Rameas cammMite MCPUIR EE Eor nes Varelit CC@p7 CN & Zauec eeeeeeneee — r tt Ape en sepeaeey \ © var. patens, tetraploid @ var. patens, diploid Fic? Nictrihuti linc L * oh nw [n* a occur in two groups. Tetraploids of var. p I k 1 population sy REFERENCES Broulttet, L., J.C. Sempte, G.A. ALLEN, and S.D. Sunpeer. 2006. Symphyotrichum (Astereae).In: Flora of North America Editorial Committee, eds. Flora of North America north of Mexico. Volume 20. Oxford University Press, New York and Oxford. Pp.465—539, Cronauist, A. 1980. Vascular flora of the southeastern United States, Vol.|. Asteraceae. Univ. of North Carolina Press, Chapel Hill. Fisuer, T.R. 1988. The Dicotyledonae of Ohio. Part 3. Asteraceae. Ohio State Univ. Press, Columbus. Gteason, H.A. and A. Cronquist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada (ed. 2). The New York Botanical Garden, New York. Jones, A.G. 1992. Aster and Brachyactis in Oklahoma. Sida, Bot. Misc. 8:1-46. Jones, A.G. and R.A. Smocor. 1984. In:IOPB chromosome number reports LXXXII. Taxon 33:1 26-134. Jones, R.L.and A.G. Jones. 1979. In: IOPB chromosome reports LXIV. Taxon 28:391-408. Jones, R.L. 1983. A systematic study of Aster section Patentes (Asteraceae). Sida 10:41-81. Jones, R.L. 1992. Additional studies of Aster georgianus,A. patens, and A. phlogifolius (Asteraceae). Sida 55305 5.15. 1080 BRIT.ORG/SIDA 22(2) Macee, D.W. and H.E. Atits. 1999. Flora of the Northeast: A manual of the vascular flora of New En- gland and adjacent New York. Univ. of Massachusetts Press, Amherst. Sempit, J.C. 1984. Cytogeographic studies on North American asters. |. Range surveys of Virgulus adnatus, V. concolor, V. georgianus, V. grandiflorus, V.novae-angliae, V. oblongifolius, V. patens, and V. walteri. Amer. J. Bot. 71:522-531. Semece, J.C. and L. Brouiter. 1980. Chromosome numbers and satellite chromosome morphology in Aster and Lasallea. Amer. J. Bot. 67:1027-1039. == DREPANOSTACHYUM FALCATUM VAR. SENGTEEANUM: IDENTITY AND ORIGINS (POACEAE: BAMBUSOIDEAE) C.M.A. Stapleton Royal Botanic Gardens, Kew Richmond, Surrey TW9 3AB, UK chris_stapleton@onetel.com ABSTRACT } } } } Leet 5 mies Drepanostachyum falcatum var. sengteeanum Stapleton, a variety of | g g I Himalayas and cultivated in the west for over a century is discussed. The application and the typification of the _ name under which it was once grown, Arundinaria falcata var. glomerata Gamble, are considered, along with the application of the name Arundinaria falcata Nees. Arundinaria falcata var. glomerata is lectotypilied in accor- dance with the protologue and current use of the name in India, placing it in synonymy of Drepanostachyum falcatum (Nees) Keng f,, for which an epitype is designated to support its own incomplete lectotype. RESUMEN Se discute Drepanostachyum falcatum var. sengteeanum Stapleton, una variedad de bambu que se piensa que se origino en el Himalaya y se ha cultivado en el oeste durante mas de un siglo. Se consideran la aplicacion y la ape ficacion ose pole palo el ane He cultivado, Ar undinaria aay var. glomerata Gamble, junto con la ifica A | ta var. slomenas de sewerdoc conel ae Pn . t ej pro 4] 1 de] } la India. colocandolo en | 8 A LULU Keng 7 ee el que se designa un epitipo para apoyar su lectotipo i plet A bamboo with a prominent and distinctive ring of hairs around the culm sheath base and on the young culm nodes was in cultivation at Kew around 1900, without any record of its provenance. It was misidentified (Bean 1907) as Arundinaria falcata Nees var. glomerata Gamble, a little-known variety from Uttar Pradesh in the NW Indian Himalaya, described on the basis of a very dubious floral characteristic. Arundinaria falcata is curently placed in the genus Drepanostachyum Keng f. A bamboo with very similar char- acteristics started to flower in California in 1994, and its seed has been widely distrib- uted, under different names. Ithough initially identified as Himalayacalamus falconeri, this graceful Califor- nian bamboo was later distributed as Drepanostachyum glomeratum Hort. based on D. falcatum var. glomeratum, and then as Drepanostachyum sengteeanum Hort It is proving somewhat hardier than all other bamboos in the genus Drepanostachyum, and is thus of considerable horticultural merit. It has recently been described as D. falcatum var. sengteeanum (Stapleton 2006), buta detailed nomenclatural treatment is required for the application of names, D.falcatum and D. falcatum var. glomeratum, to both wild and cul- tivated plants. Drepanostachyum falcatum When describing Arundinaria falcata, Nees (1834) cited ‘Nepalia’ collections made by Royle. Chao and Renvoize (1989) designated a Royle collection labelled ‘NW India’ from Hooker’s Herbarium (K) as lectotype of Arundinaria falcata. Although this appears con Becton no Royle collection from Nepal is extant, and it is actually quite unlikely that Royle or his collectors ever entered Nepal, as they concentrated their collecting activities SIDA 22(2): 1081— 1086. 2006 1082 BRIT.ORG/SIDA 22(2) to the north and west of their base in Saharanpur, Uttar Pradesh (Edmondson, pers. comm.). The Himalayan passes further to the east were little-known, except to some 17th Century Jesuits, because of the ‘jealous policies’ of Chinese and Nepalese powers (Royle 1839). For identification of species of Drepanostachyum, culm sheath characters are criti- cal, but the culm sheaths of D.falcatum are not well represented in the original material at K, nor in the Roylean Herbarium (LIV). One small, glabrous culm sheath is present in the lectotype. Munro (1868), Gamble (1896), Tewari (1993) and Negi and Naithani (1994) all record the culm sheath of D. falcatum as glabrous, reflecting the characteristics of a large number of later collections, made from Himachal Pradesh to Nepal, with which the lectotype collections agree in other characters. Drepanostachyum falcatum in the wild is clearly interpreted asa bamboo with glabrous culm sheaths (which actually have sparse, inconspicuous, white hairs when young). Plants of Drepanostachyum falcatum introduced into western cultivation on differ- ent occasions also have essentially glabrous culm sheaths. Stapf (1904) was the first to identify these plants correctly. They were initially cultivated in S Europe as Bambusa gracilis A.& C. Riviére, while the name A. falcata was misapplied to the hardier species now knownas Himalayacalamus falconeri(Munro) Keng f., known then as Arundinaria nobilis Mitford. Stapf (1904) concluded that early introductions probably came from Naini Tal or Mussoori in Uttar Pradesh around 1840. The consensus is clear that the name A. falcata applies to a species found in the Himalayas from Himachal Pradesh to Kathmandu, and also found in cultivation in Eu- rope, where it was known as Bambusa gracilis. This species has almost completely gla- brous culm sheaths, although the culm nodes may initially have sparse, short hairs be- low. A specimen from Uttar Pradesh is selected here as epitype, to support the incomplete Royle lectotype by indicating culm sheath characteristics. Arundinaria falcata var. glomerata Gamble Arundinaria falcata var. glomerata Gamble was initially distinguished solely on the ba- sis of agglomeration of 3-4 spikelets in close racemes or panicles. This followed an ear- lier, identical distinction (Munro 1868) between two unnamed developmental forms of A. falcata, listed as var.a and var. b. Munro clearly explained that these forms merely rep- resented different stages of inflorescence development. He observed that as the season advances branch proliferation leads to compound rather than paniculate forms, with shorter racemes of 3-4 spikelets (var. b), totally unlike t earlier on younger shoots (var. a). He cited a December collection, Wallich 5035 (K-W), from Chisapong, near Kathmandu, Nepal as representative of the later developmental form, var. b. It is not clear whether Gamble (1896) followed Munro’ interpretation of the devel- opmental nature of this distinction. Gamble’s brief description repeated that of Munro for var. b. He indicated in a plate caption that illustrations of var. glomerata were from collections made by Bagshawe in 1879. This action was interpreted by Chao and Renvoize (1989) as designation of a type for A. falcata var. glomerata Gamble, and they cited a Bagshawe collection at K as holotype, albeit a collection made in 1878 not 1879. They listed var. glomerata in synonymy of Sinarundinaria falcata. Tewari (1993) subsequently gave the variety full recognition, providing a detailed description attributed to Pandey ined. This, however, differed little from that of the type variety. Culm sheath hairs were not mentioned at all. The description of the culm sheaths STAPLETON CENCTEEANIIN 4083 as shining below probably meant having a glossy adaxial, internal surface, rather than being basally glabrous, but there was certainly no mention of any prominent basal ring of hairs. Naithani and Chandra (1998) and Seethalakshmi and Kumar (1998) merely in- cluded it in synonymy of Drepanostachyum falcatum. The name A. falcata var. glomerata has thus been applied to two taxa, one a bamboo of unknown origin cultivated at Kew, the other an Indian bamboo from the borders of Himachal and Uttar Pradesh. The principal diagnostic character of the cultivated plants is a prominent ring of hairs around the base of the culm sheath and the culm node. This is not seen convincingly in any of the collections of wild material. Other characteristics of the cultivated plants include long sparse hairs on only one side of the base of the leaf midrib (costa) rather than both sides, densely scabrous lemmas and paleas, and pubes- cence only at the tips of rhachilla segments. These characteristics are also not present in the wild material, which therefore cannot be considered the same taxon as the cultivated plants. Typification of var. glomerata Original material consists of three differently labelled collections at K from Gamble’s herbarium, all annotated by him as Arundinaria falcata var. glomerata. To complicate matters, as is usually the case with older bamboo collections including both fertile and sterile material, these collections represent at least two, and possibly three different spe- cies, with mixed collections mounted together on the same sheets. Only one collection, from Jaunsar Bawa, Bagshawe s.n. in 1879, was literally cited in the protologue. The culm nodes and persistent portions of the culm sheath bases are al- most completely glabrous. A second, different collection includes the specimen actually illustrated by Gamble, and this, therefore, can be construed as part of the protologue. This sheet, annotated as type by Chao, is simply labelled ‘Comm. Brandis, Jaunsar’. As ‘specimens’ collected by Bagshawe’ was the citation this could be a further Bagshawe 1879 collection, sent by Brandis. It is clearly mixed, with old, unidentifiable flowers of a Drepanostachyum spe- cies as well as leaves of a different species, probably Himalayacalamus falconeri, with smooth culms, entire leaf sheath ligules and distinguishable tessellation of veins on some of the leaves, which are broader, glabrous, and less cuneate than those of D. falcatum. Gamble dissected spikelets from these two collections. His drawings remain attached to the Bagshawe 1879 collection, with the recognisable spikelets he illustrated remaining in capsules on the two sheets. These drawings were reproduced when describing his var. glomerata, but interestingly were used for the plate of the type variety, not that of var. glomerata, suggesting that Gamble himself actually had scant regard for any difference between the two varieties. The third collection from Gamble’s herbarium annotated as var. glomerata, also from Jaunsar Bawa, but in 1878, is Bagshawe 6608. This was cited as holotype of A. falcata var. glomerata by Chao and Renvoize (1989), and is annotated as type by Renvoize. It again has no culm sheaths, but like the second collection, has some short hairs below the culm node. To summarise ‘specimens collected by C. Bagshawe.,.... in 1879’ was the citation in the protologue. A collection labelled thus is at K. A second, mixed collection, sent by Brandis, was actually illustrated in the protologue, and annotated as type by Chao, while a third collection, Bagshawe 6608 in 1878, a date that conflicts with the protologue, was cited as 1084 BRIT.ORG/SIDA 22(2) ‘holotype’ by Chao and Renvoize (1989). The collection cited in the protologue has com- pletely glabrous culm nodes, while the other two collections have distinct, though very short, hairs below the nodes. It seems more appropriate for the collection bearing the details cited in the protologue to be taken as holotype, rather than the collection stated as ‘holotype’ by Chao and Renvoize (1989). Although that collection was part of the original material, as it was nei- ther cited nor illustrated in the protologue, their citation of it as holotype cannot be con- sidered an acceptable lectotypification. The designation here as lectotype of the only sheet actually labelled Bagshawe, Jaunsar Bawa, 1879, is in accordance with both the protologue, and with current applica- tion of the name in India (Tewari 1993). As the collection actually illustrated in the protologue is recognisable, and citation of Bagshawe, 1879 was merely indicating (possi- bly incorrectly) which collection was illustrated, there would appear to be two elements to consider as syntypes. The name D. falcatum var. glomeratum Gamble is misleading, and seems merely to representa later stage of inflorescence development, as Munro originally intended (1868). There is indeed no separate variety of D.falcatum with agglomerated spikelets. It might be considered more appropriate for Gamble to have followed the typification of Munro’s ear- lier unnamed variety. However, Wallich 5035 is one of the worst bamboo specimens ever preserved in a herbarium, being so decrepit that it has practically no spikelets left at al Drepanostachyum falcatum var. glomeratum, as interpreted and described in Tewari (1993) and lectotypified here according to the protologue, differs little if at all from the type variety. The precise origin of this type is not known, but it is probably from the same district as the epitype of D. falcatum designated here. Culm sheaths are not known, but basal portions remaining attached to the culm nodes of the lectotype are completely gla- brous. — ee falcatum (Nees) Keng f., J. Bamboo Res. 2:16. 1983. Types: INDIA:‘N.W India, Royle (LECTOTYPE: K, anne in Chao and Renvoize 1989); INDIA. UTTAR PRADESH: Chakrata, Sep 1898, Gamble 27256 (epityy gnated here: kK) Sides Nees, Linnaea 9:478. 1834. Chimonobam busa falcata (Nees) Nakai in, J. ce Arbor. 6:151. 1925. Fargesia falcata (Nees) TP. Yi, Fl. Xizangica 5:33. 19% Sinarundinaria falcata (Nees) CS. Chao & ent €, ieee Bull. 44:357. 1989, head one men ele To ae Bot. Zhurn. 75:225. 1990. . Riviére, Bull. Soc. Acclim. sér. 3, hae 1878. Drenavactas hyum a atum (Nees) Keng fC var. ¢ ramble, Ann. Bot. Gard. Calcutta 7:13. 1896; emend. Pandey, D.N. Tewari, Monogr. Bamboo 84. | 1903 TYPE: INDIA ul FAR PRADESH: Jaunsar Bawa, 1879, Bagshawe s.n. (LECTOTYPE designated here, K, superseding previous citation of type by Chao & Renvoize, Kew Bull. 44:358. 1989, Further Sensseiey INDIA. ae Pradesh: Jaunsar, Aug 1878, Bagshawe 6608(K): Jaunsar, Comm. Brandis s.n. a Ss) u K). (flowers) only, t Drepanostachyum falcatum var. sengteeanum and other similar bamboos Having excluded the name Drepanostachyum falcatum var. glomeratum from the plants once cultivated at Kew and currently in horticultural use in the USA, it is neces- sary to consider other possible names. Arundinaria interrupta Trin. from Nepal has variable development of light, white hairs on the culm nodes, noted in the description. Culm sheaths are not present in the —_— STAPLETON CNCTECCANIINA ogee type, but recent collections from Nepal from a similar location have, in addition to simi- lar white hairs on the culm nodes, the distinctive sparse white hairs on most of the culm sheath and the mainly glabrous culm sheath bases of other collections of D. falcatum. Even the lectotype of A. falcata has some white hairs on the persistent culm sheath base, and this character was also given in Tewari (1993) for the type variety. The possibility of Arundinaria interrupta Trin. representing a separate species was raised (Stapleton 1994), but the new collections suggest that it should remain a synonym of D. falcatum. Other bamboos with hairs at the base of the culm sheath have been collected else- where in the Himalayas. Himalayacalamus fimbriatus Stapleton also has culm sheaths with a basal ring of hairs and a fimbriate ligule, but the larger stature, the much larger leaves, as well as the asymmetry of the culm sheath with its broader ligule and less sca- brous interior distinguish it clearly. Drepanostachyum annulatum Stapleton from Bhutan also has a ring of hairs at the base of the culm sheath, but the hairs are darker, longer, more upright and ina narrower ring. That species also differs in having dark brown culm sheath hairs, especially on the distal third of the culm sheath, which also bears erect, basally scabrous oral setae when young, as well as darker brown, longer cilia of up to 1.5mm on the edges. It also has well-developed leaf sheath auricles, and thicker culm wax. In addition the ligules are neither as delicate nor as laciniate, and the culm sheath is less pubescent on the inside at the base of the ligule and below it. Although it shares the pos- session of a basal ring of culm sheath hairs with H. fimbriatus and D. annulatum, the bamboo currently cultivated in the US, recently described as D. falcatum var. sengteeanum (Stapleton 2006) is clearly much closer to the type of D.falcatum. The 2-3 flowered florets and the densely pubescent culm sheath interior apex of var. sengteeanum are sufficient to place this cultivated bamboo indisputably in Drepanostachyum. The absence of oral setae and the distribution of hairs on the culm sheaths are sufficient to separate it clearly from all previously described taxa. Following current taxonomic concepts in bamboos, culm sheath pubescence and presence of oral setae are important and consistent specific characters. They are usually supported by minor differences in spikelet or floret detail, while leaf sheath and blade characters are inherently more variable. However, in Drepanostachyum more variety is seen within spe- cies than in most other bamboo genera, and it was felt inappropriate to describe this taxon as a new species, especially as its origin remains unknown. The florets and spikelets of D. falcatum var. sengteeanumare more scabrous than those of the type variety and the palea is less bifid. The rhachilla segments are slightly longer and have rather different apical pubescence: the beard is mainly restricted to the distal 0.25mm of the rhachilla, while in the type of D. falcatum the distal 1mm becomes progressively more puberulent. The lemma and palea are more densely scabrous than any other material of D.falcatum from the Western Himalayas, and approach the state seen in other species from the Eastern Himalayas, suggesting a Nepalese or Sikkimese origin. ACKNOWLEDGMENTS The author would like to express his deep gratitude to Seng-Tee Lee, in whose honor the variety D.falcatum var. sengteeanum was named, for funding the Sino- Himalayan Bamboo HSO post at Kew for six years from 1994 to 2000. Staff at LIV are thanked for loan of Royle specimens. Gib Cooper is thanked for initially sending material from Oregon. The Ameri- can Bamboo Society is thanked for supporting travel to lecture and collect cultivated mate- rial in California. Reviewers are thanked for helpful comments on the manuscript. 1086 BRIT.ORG/SIDA 22(2 REFERENCES Bean, W.J. 1907. The flowering of cultivated bamboos. Bull. Misc. Inform. Kew 1907:228-233. Cao, CS. and S.A. Renvoize. 1989. A revision of the species described under Arundinaria (Gramineae) in southeast Asia and Africa. Kew Bull. 44:349-367, Game, J.S. 1896. The Bambuseae of British India. Ann. Roy. Bot. Gard. (Calcutta) 7:1—133. Munro, W. 1868.A monograph of the Bambusaceae. Trans. Linn. Soc. London 26:1-157. NaiTHANI, H.B. and S. CHanpra. 1998. Flowering of a bamboo (Drepanostachyum falcatum). Indian For- ester 124:663-666. Nees Ab Esenseck, C.G, 1834. Bambuseae Brasilienses. Linnaea 9:461-494, Neal, S.S. and H.B. NarrHani. 1994, Handbook of Indian bamboos. Oriental Enterprises, Dehra Dun, India. Rovie, J.F. 1839. Illustrations of the botany of the Himalayan Mountains. Allen, London SEETHALAKSHMI, K.K. and M.S. MuktesH Kumar. 1998. Bamboos of India: A compendium. Kerala Forest Re- search Institute & INBAR. Star, O. 1904. Himalayan bamboos. Gardeners’ Chronicle May 14:305-306; May 21:325-326; June 4:356. STAPLETON, C.M.A.1994.The bamboos of Nepal and cs Part Ill: Drepanostachyum, Himalayacalamus, (Gramineae: Poaceae, Bambusoideae). } Ampelocalamus, Neomicrocalamus, and Chi Edinburgh J. Bot. 51:301-330 STAPLETON, C.M.A. 2006. New taxa and combinations in cultivated bamboos (Poaceae-Bambusoideae). Sida 22:331-332. Tewari, D.N. 1993.A monograph on bamboo. International Book Distributors, Dehra Dun, India. REJECTION OF LECTOIYEIRIGATION OF ASTER AMELLUS (ASTERACEAE: ASTEREAE) AND SELECTION OF A NEW LECTOLYPE John C.Semple Department of Biology University of Waterloo 00, Ontario, CANADA N2L 3G] jcsemple@sciborg.uwaterloo.ca Water, ABSTRACT Selection of 997.18 (LINN) as the lectotype of Aster amellus L. is rejected because the specimen is in conflict with the peat aes A new Oy, pe is designated f rom 1 authentic sian seen by Linnaeus. oe the original or conservation or redefining the eee of the genus Aster L. Herb, is inn. 997, 16 (1 -INN) is designated the new lectotype o Aster cls RESUMEN Se rechaza la seleccién del pliego 997.18 (LINN) como lectotipo de Aster amellus L. porque el espécimen esta en conflicto con i Protloeo, Se eclen un nuevo lectotipo a pare de malete auténtico visto ES Linnaeus. oo rechazo de la ipi la seleccion de vacion o redefinicion del cone del ae Aster L. El Biesollieit Linn, 907. 16 (LINN) se eee nuevo lectotipo de Aster amellus. Semple (1998) designated Herb. Linn. 997.18 (LINN) as the lectotype of Aster amellus L. (Astereaceae: Astereae). Recently, this lectotypification was found to be flawed and would result in a change in the application of the name Aster of which A. amellus is the generitype. The simplest solution to this situation is to reject the original lectotypification because the selection was in conflict with the protologue. An alternative solution would be to propose conservation of A. amellus with a conserved type, which does not seem necessary in this case. In choosing 997.18 (LINN) as the lectotype for Aster amellus for the Linnaean Typifi- cation Project (Semple 1998), I worked from a set of small photographs I had taken in 1984 during visits to the Linnaean Herbarium (LINN) and the British Museum of Natu- ral History (BM) and from additional notes made in 1994 on numerous authentic speci- mens of asters and goldenrods. Other authentic specimens were rejected as possible types then, and additional specimens and photographs have subsequently come to my atten- tion. 997.18 (LINN) is here rejected as the lectotype for Aster amellus on the basis of being in conflict with the protologue. Article 9.17(b) of the ICBN (McNeill et al. 2006) permits the supersession of a lectotype, if it is in serious conflict with the protologue, and an- other element that is not in conflict is available. At least one specimen that does not con- flict with the protologue, Herb. Linn. 997.16 (LINN), is available. In the protologue of Aster amellus, Linnaeus (1753, p.873) indicated the occurrence of some variation in leaf and phyllary traits, and he noted the range as “Habitat in Europae australis asperis collibus,” that is, native to rugged hills in southern Europe. Leaves were described as “lanceolatis obtusis scabris trinervis integris” and phyllaries as “calycinis obtusis” in the diagnosis. LINN 997.18 has linear, uninervate rather than lanceolate trinervate leaves and linear acute rather than obtuse phyllaries. The small black and white SIDA 22(2): 1087— 1089. 2006 1088 BRIT.ORG/SIDA 22(2) photograph of 997.18 (LINN) that | worked from was cropped and lacked critical annota- tions that are obvious in the colored image of the specimen currently posted on the Lin- naean Typification Project web site http://interntnhm.ac.uk/jdsml/research-curation/ projects/linnaean-typification /index.dsml. Upon seeing this image, | knew immediately that |had made a mistake in my choice of lectotype for Aster amellus. The specimen was annotated “non A. Amellus” by James Edward Smith (Charlie Jarvis, pers. comm.), who bought Linnaeus’ herbarium and moved it to England, and as “Calimeris A.G” by Asa Gray. Clearly, these annotations are critical because they indicate the practical conse- quences of retaining 997.18 (LINN) as the type of Aster amellus, the generitype; the mod- ern strict application of the name Aster (see Nesom 1994; Semple et al. 2002) would fun- damentally have to be changed. 997.18 (LINN) is likely a member the Galatella Cass. / Crinitaria Cass. complex rather than Aster The second paragraph of the protologue deals with the first synonym Linnaeus cited from Hortus Cliffortianus. The voucher of the synonym cited as “Hort. Cliff. 407.” is Clif- ford Herb. No. 407 (BM!) and was diagnosed by Linnaeus as having leaves “lanceolatis scabris semiamplexicaulisbus serratus” and phyllaries as “lanceolatis.” Linnaeus noted that the “Hort. Cliff.” reference was previously cited in “Royen” and “Gmelin.” Clifford Herb. No. 407 (BM) is a possible choice for lectotype for Aster amellus. The specimen is a fragment, and in leaf features it is similar to Herb. Linn. 997.17 (LINN), a specimen of A. pyrenaeus Desf. ex DC. in Lam. & DC. The Clifford herbarium specimen is rejected as a choice for the new lectotype of A.amellus. —y Retypification Herb. Linn. 997.16 (LINN) is also an authentic specimen and is annotated “5 Amellus” likely by Linnaeus himself (Charlie Jarvis, pers. comm.). | have no data indicating I saw this specimen in 1984 or 1994. A photograph of this specimen was given to me after 1998 by Paul Van Faasen, who took the picture. The plant illustrated is very similar to plants in my research collection of Aster amellus collected for me in Switzerland by Dr. Erich Weber. 997.16 (LINN) and the Weber collections match the description of Aster amellus in the diagnosis of the protologue in both leaf and phyllary traits. These correspond well with the description and figures of A. amellus in Wagenitz (1979) and the description of the species in Merxmiuller, Schreiber and Yeo (1976). Therefore, Herb. Linn. 997.16 (LINN) is here designated the new lectotype of Aster amellus. Aster amellus L., Sp. Pl. 2:873. 1753. Type: EUROPE: habitat in Europae australis asperis co oe [native to rugged hills in southern Europel, Herb. Linn. 997.16 (LECTOTYPE here designated as new: LINN ACKNOWLEDGMENTS This work was supported by a Natural Sciences and Engineering Research Council of Canada Discovery Grant to the author. Charlie Jarvis of the British Museum of Natural History is gratefully acknowledged for his assistance with Linnaean collections at BM and LINN. Fred R. Barrie and Charlie Jarvis are thanked for their useful and timely re- views of the submitted manuscript. lam grateful to Paul Van Faasen for having chosen to give me his extensive collection of photographs of type species of asters and copies of species protologues. SEMPLE, RE-LECTOTYPIFICATION OF ASTER AMELLUS 1089 REFERENCES Linnaeus, C.1753. Species Plantarum. Stockholm. McNett, J., FR. Barrie, HM. Buroet, V. DEmMouLIN, D.L. HAwKSworTH, K. MARHOLD, D.H. Nicotson, J. PRADO, PC. Sitva, J.E. Skoa, J.H. Wiersema, and NJ. Turcanp. (eds.) 2006. International Code of Botanical Nomenclature (Vienna Code)...Adopted by the Seventeenth International Botanical Congress Vienna Austria, July 2005. (Regnum Veg. 146). Koeltz Scientific Books, KOnigstein, Germany Merxmutter, H., A. ScHreiber and PF. Yeo. 1976. Aster. In: Tutin, T.G.,V.H. Heywood, N.A. Burges, D.M. Moore, D.H.Valentine, S.M.Walters and D.A.Webb, eds.Flora Europaea.Vol.4. Plantaginaceae to Compositae (and Rubiaceae). Cambridge University Press, Cambridge, U.K. Nesom, G.L. 1994. Review of the taxonomy of Aster sensu lato (Asteraceae: Astereae), emphasizing the New World species. Phytologia 77:141-297. Sempte, J.C. 1998. Typification of Aster amellus, A.dumosus, A.novae-angliae, A. puniceus, A. sibiricus, Sol- idago flexicaulis, S. lanceolata and S. latifolia. |n: Jarvis, C.E. and N.Turland, eds. Typification of Lin- naean names in the Asteraceae (Compositae). Taxon 47:347-370. Semee, J.C., S.B. Heard and L. Brouiter. 2002. Cultivated and native asters of Ontario (Compositae: Astereae):Aster L. (including Asteromoea Blume, Diplactis Raf.and Kalimeris (Cass.) Cass.), Callistephus Cass., Galatella Cass., Doellingeria Nees, Oclemena E.L. Greene, Eurybia (Cass.) S.F. Gray, Canadanthus Nesom, and Symphyotrichum Nees (including Virgulus Raf.). Univ. Waterloo Biol. Ser. 41:1-134. Wacenitz, G. 1979. Aster. In: Hegi, G., ed. Illustrierte Flora von Mittel-Europa. Band 6(4), Compositae. 1:35-71. Hehman Verlag, Munich. 1090 BRIT.ORG/SIDA 22(2) Book NOTICE S. InpbeRIt (ed). 2005. Invasive Plants: Ecological and Agricultural Aspects. (ISBN 3-7643- 7137-4; 978-3-7643-7137-1, hbk.). Birkhauser Verlag, PO. Box 133, CH-4010 Basel, SWITZERLAND. (Orders: Springer, 333 Meadowlands Parkway, Secaucus, NJ 07094, U.S.A.; 800-SPRINGER:; fax 201-348-4505: WW w.spt inger.com). $159.00, 283 pp., illustrations, 6 1/2" x 91/4". A set of studies toward understanding the biology of invasive species, comprising 17 papers written by an inter- national group of invasions biologists. The chapter titles give a good view of the content. * In search of an operational lexicon for biological invasions. * The ecology of biological invasions: past, present, and future. * Ecological niche models and the geography of biological invasions: a review and a novel applica- tion. * Importance of species replication in understanding plant invasions into North American grass- lands * Residence time determines the distribution of alien plants * The relationship between community diversity and exotic plants: cause or consequence of inva- sion? * Invasive plants: the process within wetland ecosystems. * Understanding invasions:the rise and fall of diffuse knapweed (Centaurea diffusa) in North America. * Defining a regional approach for invasive plant research and management. * Allelopathy as a mechanism for resisting invasion: the case of Polygonella myriophylla. * Ecology and management of an exotic weed Phalaris minor. * Reducing agroecosystem vulnerability to weed invasion. * Tillage intensity affects weed communities in agroecosystems. * Distribution and biological control of Chromolaena odorata. * Biology and anthropology of plant invasions. * Characters, significance, and human dimension of global invasive weeds. * Environmental consequences and economic costs of alien species. —Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, UX 76102-4068, U.S.A. SIDA 22(2): 1090. 2006 TRICHOME MORPHOLOGY IN SELECTED MEXICAN RED OAK SPECIES (QUERCUS SECTION LOBATAE) M. Lucia Vazquez! Department of Plant Biology Cornell University Ithaca, New York 14850, U.S.A. vazquez.lucia@uis.edu ABSTRACT Mature leaves and twigs noe oy red oak Species, meaty nas to oes were examinee with SEM to assess trichome eee ee i g g glandular. Described for the first ti fi icul df icul d d ltipl 1] trichomes. Furthermore, tl ick i I 1 for the fi ime i 1 oal llate, fused stellate and glandular branched. Tl | 1 f l | | lti li vhile f. ery | cf ese eee ea ay see aoe 4 1p; the most frequent types on the eee llea trichomes. In contrast to the significant anes ees variation found in oak leaf a ‘cichomle ee within each species are Eeacanye invaria ble ane t De useful in the iain canes of most species. In summary, together with veg i gy | ional information for -_ the identification and chactenause of Mexican oak sp RESUMEN Hojas maduras y ramillas de treinta especies de encinos rojos, principalmente endémicos a Mexico, fueron mas con pena ae Secu onica de BarEdo pana evaluar la variacion morfolégica de tricomas. En este glandulares. pe estos, se ‘ii iben por tr abajo of primera vez para encinos jesus tricomas s fasciculadoc contorto, lagen ado Gsiado y estr \demas se reportan por panes para j p llado fusi y glandu- larramifieado.| r ic] j ill | fasciculado sésil y el multiradiado, mientras S| ape | al : ] . ] : Ane es ‘ leael Beil £. G la- tin} ] f ic : r y el fies A ] ‘ fologia fol L i=) Oo .o L en cada especie son te por | utiles en la identificacion de la mayoria de las especies. En resumen, los caracteres vegetativos y reproductivos junto con la morfologia de trichomes proporcionan informacion valiosa para la identificacion y caracterizacién de las especies Mexicanas de encino. INTRODUCTION The genus Quercus (Fagaceae) consists of approximately +00 species of trees and shrubs distributed in temperate and subtropical regions of the Northern Hemisphere. In the Americas, Quercus is found from southern Canada to Colombia with representatives of three sections within the subgenus Quercus: Quercus (white oaks), Protobalanus (inter- mediate oaks), and Lobatae (red oaks) (Nixon 1993). Due to the large diversity of white and red oaks, Mexico is considered the center of diversity for the genus in the New World (Rzedowski 1965; Nixon 1993). he number of red oak species has been estimated around 195 (Jensen 1997) with about 55 species endemic to Mexico (Nixon 1993). In his treatment on the American oaks, Trelease (1924) segregated the red oak species into 73 series and also proposed a classifi- cation based on leaf and fruit morphology. However, his hypothesis on the relationships of red oak species has not been tested with phylogenetic methods. Although the mono- ‘Current address: Biology Department, University of Illinois Springfield at Springfield, Illinois, 62794, U.S.A. SIDA 22(2): 1091- 1110. 2006 1092 BRIT.ORG/SIDA 22(2) phyly of the red oak section is well supported (Nixon 1984; Manos 1999), an intrasectional phylogeny for the red oaks is yet to be conducted. As in other oak groups, Mexican oak species are difficult to study because most of them show significant amounts of foliar polymorphism. Species identity becomes even more complex when either hybridization or introgression has taken place. Therefore, taxonomic problems in many oak species persist, particularly in those species that have similar or overlapping leaf morphology. The utility of trichome morphology for species identification has been demonstrated in several plant families such as Brassicaceae, Geraniaceae, and Sterculiaceae (Inamdar & Rao 1983; Metcalfe & Chalk 1950; Oosthuizen 1983; Shanmuka Rao 1987). In oaks, tri- chome characterization has been carried out for several eastern North American, Asian, and European species. The first study on trichome morphology of eastern North Ameri- can oak species was reported by Dyal in 1936, who classified them, based on secretory function, into glandular and nonglandular. Camus (1934-1953) described seven trichome types that fall into the glandular and nonglandular categories previously described by Dyal (1936). Hardin (1976) published the first comprehensive work on oak trichome mor- phology for 58 eastern United States (US) oak species and recognized 10 trichome types. Thomson and Mohlenbrock (1979), Jones (1986), and Manos (1993b) conducted similar studies on other U.S. oak species. Trichome morphology of European and Mediterranean oak species has also been extensively documented by several authors (Olsson 1976; Kissling 1977, 1993; Safou & Saint-Martin 1989; Gellini et al. 1992; Llamas et al. 1995; Bussotti & Grossoni 1997). Trichome morphology, together with other vegetative and re- productive characters, has been used to discriminate several USS. oak species, as well as a few Mexican species distributed in the northwest of the country (Tucker 1952; Tucker & Muller 1957; Nixon & Steele 1981; Spellenberg 1992, 1998; Nixon & Muller 1993; Bacon & Spellenberg 1996). However, to date, no in-depth research on the trichome morphology of Mexican taxa has been carried out. Several regional taxonomic treatments of Mexican oak species have been conducted (Gonzalez 1986; Valencia 1989; Vazquez 1992, 2000; Spellenberg et al. 1998; Spellenberg 2001), but none of them has characterized trichome diversity. Only a few authors have discussed the potential utility of trichome types for identification of Mexican species (Mc Vaugh 1974; Rzedowski & Rzedowski 1979; Vazquez 1992). Accordingly, the objective of this study is to characterize the trichome morphol- ogy from twigs and leaves from thirty Mexican red oaks, mainly endemic to Mexico, us- ing scanning electron microscopy (SEM). Detailed examination of these species revealed the presence of ten trichome types: four of these trichomes have been previously described, three of them represent entirely new trichome types, and the remaining three are tri- chome types not previously documented for red oaks. Each species examined shows a particular trichome complement which, together with other vegetative and reproductive characters, provides useful information for species identification and characterization. MATERIALS AND METHODS Leaves and twigs from thirty species of red oak species chiefly endemic to Mexico were selected for examination of trichome morphology under SEM. Nixon (1993) has reported 41 red oak species endemic to Mexico, of which, twenty-one have persistent indument and were sampled in this study. Two additional species with persistent indument (Q. castanea and Q crassifolia) whose range extends to Central America were included in this study because they are widely distributed in Mexico. For comparison, six glabrous VAZQUEZ, TRICHOME MORPHOLOGY IN QUERCUS Tape 1. Red oak species (Quercus) examined with SEM for trichome morphological variation. Voucher speci- mens are deposited at Bailey Hortorium, Cornell University (BH = Taxa Voucher specimen Q. acherdophylla Trelease Q. affinis Scheidweiler Q. aristata Hooker & Arnott Q. candicans Nee Q. castanea Q. coahuilensis Nixon & Muller Q. coccolobifolia Trelease Q. conzattii Trelease Q. crassifolia Humboldt & ly Q. crassipes Humboldt & Bon plan Q. depressa Humboldt & :Sonpbnd Q. auido ia von Seem lysopt rylla Ber itt lalit QO ee Q. fulva Liebmann Q.gentryi Muller Q. hintonii Warbur Vdzquez et al. 3065 Muller 9716 Martinez 31, Breedlove 1640 Vdzquez et a Rzedowski 25361 Stewart 924 Vazquez & Phillips 3084 Muller 9420 Vazquez et al. 3049 Vazquez et al. 3037, 3070 Vdzquez & Tenorio 118 Muller 3579 Nixon s.n. (Oct. 1999) Nixon 4034, Vazquez et al. 3011 Vazquez et al. 3097 McVau eee 256 . Vazquez al.: cron ne 2336 Vazquez et al. 3069, 3109 Ven & Fetrqdgq 22322 Q. hintoniorum Nixon & Muller Q. hirtifolia Vazquez, Valencia & Nixon Q. hypoleucoides Camus Q. hypoxantha Trelease Cowan et al. 5402 Q. mevaughii Spellenberg Bacon et al. 5284 mexicana Humboldt & Bonpland Rzedowski 6374 Q. planipocula Trelease McVaugh12136 Q. radiata Trelease Rzedowski 23018 Q. salicifolia Né McVaugh 25456 Q. scytophylla Liebman Vazquez et al. 3035 Q. sideroxyla ree & Bonpland Vazquez et al. 3089, Bacon et al. 5304 Q. tarahumara Spellenberg, Bacon & Breedlove Bacon et al. 5307, 5375 Q. urbanii Trelease Vazquez et al. 3027 red oak species endemic to Mexico were included. Samples of the materialexamined were gathered from field collections and from herbarium specimens deposited at the Bailey Hortorium, Cornell University (BH) (Table D. Twenty to thirty specimens per species were initially examined under dissecting and light microscopy to survey the intra- and interspecific variation in trichome morphol- ogy. A more detailed examination was carried out under SEM in representative subsamples. About one square cm of the abaxial surface of a dry leaf or one cm of twig was mounted on an aluminum stub with no prior treatment. Previous studies have shown no difference in the preservation of nonglandular trichome structure using either fresh or dry tissue (Hardin 1976). Specimens mounted on stubs were sputter coated with 30 nm of gold palladium using a BAL-TEC sputter coater. Samples were observed under a Zeiss electron microscope, model LEO DSM 960, using working distances between 12 and 18 mm, voltages of 3.0 or 4.0 kV, and apertures of 300 um or 400 um depending on the sample and magnification. Most trichome micrographs were taken at magnifications of 200, although smaller trichomes required magnifications of 450-500. 1094 BRIT.ORG/SIDA 22 2) Tasve 2. Trichome types on twigs of Mexican red oak species (Quercus). Species Glandular Nonglandular Simple Branched Fasciculate Fasciculate Multiradiate Simple uniseriate sessile simple —_stipitate Stellate Q. acherdophylla x Xx Q. affinis X Q. aristata xX Q. candicans Xx Xx x Q. castanea Xx Q. coahuilensis x x Q. coccolobitolia xX x Q. conzattii x X Q. crassifolia xX Xx x Q. crassipes X X Q. depressa X Q. durifolia xX Xx Q. x dysophylla X Xx Q. eduardii x Xx Q. fulva x X ). gentryli Xx Q. hintonii X X Q. hintoniorum X X Q. hirtifolia Xx x Q. hypoleucoides Xx x Q. hypoxantha X xX Q. mevaughil Xx Xx Q. mexicana X Xx Q. planipocula Xx Q. radiata X Q. salicifolia x Q. scytophylla Xx x Q. sideroxyla Xx Xx Q. tarahumara X Q. urbanii X X RESULTS Trichome nomenclature follows Hardin (1976), where each cell composing a nonglandular trichome is called a “ray” and trichome types are named based on the degree of ray fu- sion, ray orientation, and overall morphology. However, additional terms have been in- troduced to describe trichome types not previously reported in the literature. Two general trichome categories are used to indicate secretory functions and nature of the cell wall: 1) glandular trichomes have a glistening appearance and thin cell walls that collapse upon drying, and 2) nonglandular trichomes lack a glistening appearance and possess thick cell walls. Examination of twigs and abaxial leaf surfaces of thirty red oak species resulted in two types of glandular trichomes and eight nonglandular trichome types (Tables 2 and 3). Glandular trichomes 1. Simple uniseriate (Fig. 1A, 1B).—This type consists of 2-7 united cells with slight constrictions along the trichome length. This trichome type was commonly found on —_— Taace 3. Trichome types in oak species (Quercus) with a persistent indumentum on the abaxial leaf surfaces. Species Glandular Nonglandular Simple Branched Fasciculate Fasciculate Fasciculate Fasciculate Multi- Simple Fused Multiple uniseriate sessile sessile sessile stipitate radiate stellate stellate stellate simple contorted crested Q. acherdophylla XV Q. affinis Xv Q. aristata X Xx Q. candicans Xx x Xx X Q.castan X Xx Q. coahuilensis xX x Q. coccolobifolia x Xv Q. conzattii X Q. crassifolia x x X Q. crassipes X x Q. depress Xv Q. durifolia X x . x dysophylla x x Q. eduardii xX Q. fulva x x Q.gentryi Xx X Xs Q. hintonii x x x Q. hintoniorum xX Xv x Q. hirtifolia x xX Q.hypoleucoides =X xX Q. hypoxantha x x Q. mevaughii X X Q. mexicana xX X Q. planipocula x x Q. radiata xX x Q. salicifolia XS XS Q.scytophylla x X Q. sideroxyla x x X tarahumara xX x X Q. urbanii xX X s=scattered v=trichomes on secondary veins SNIYINO NI ADOTOHdYOW IWOHDIIYL ‘ZINDZVA S60L 1096 BRIT.ORG/SIDA 22(2) \ Fic. 1.A. Thi | Wi | 1] +t tn lobifolia. x 40.B. Glandul . 1 leaf surface of Q. hintoniorum, x 500. C. Glandular t hed trict | surf f Q. salicifolia, x 500. D. A variation of the glandular I hed trict the abaxial | f Q. salicifolia, x 500. the abaxial leaf surfaces of twenty-one species, forming a sparse and deciduous ~matu- indumentum. In most species examined, trichome density decreased during lea ration and senescence; however, Quercus coccolobifolia showed persistent simple uniseriate trichomes on the abaxial leaf surface. This trichome type was found on the twigs of only three species: Q. acherdophylla, Q. coccolobifolia, and Q. crassifolia (Table 2). 2. Branched (Fig. 1C, lD).—This type consists of 2-3 cells united in either a V or a Y shape; however, under the dissecting microscope this trichome type looks like two uni- cellular simple trichomes lying next to each other. This trichome type was found exclu- sively on abaxial surfaces of Q. salicifolia leaves. Nonglandular trichomes Fasciculate sessile trichomes.—These e consist of several rays fused at the point of attach- ment to the epidermis. The rays are either short and completely erect (Fig. 2A), or long VAZQUEZ, TRICHOME MORPHOLOGY IN QUERCUS 1097 rc . nl eee oe Fic. 2. A t trichome on a. 4 | th |} oe | | chome on the abaxial leaf surface of Q. crassifolia, < 200. C. F mexicana, 200 iculat il torted trict the abaxial leaf surf. f Q. eduardii < 200. f Quercus aristata, X 200. B. Fasciculate sessile simple tri- a 1 1 th L a i | £ £. [ph Q. and tortuous diverging in different directions (Fig. 2B). Three types of fasciculate sessile trichomes are recognized in the species examined. 1. Fasciculate sessile simple (Fig. 2A, 2B).—The fasciculate sessile trichome described by Hardin (1976; 1979a) and Jones (1986) is termed here fasciculate sessile simple to dis- tinguish it from the two additional subtypes listed below. This trichome type was found on twigs and/or on the abaxial leaf surfaces of several species examined. It occurs on twigs of the following species: Q. candicans, Q. castanea, Q. coahuilensis, Q. coccolobifolia, QO. conzattii, Q. crassifolia, Q. crassipes, Q. durifolia, Q. x dysophylla, Q. eduardii, Q. fulva, Q. hintonii, Q. hintoniorum, Q. hirtifolia, Q. hypoleucoides, Q. hypoxantha, Q. mcvaughii, Q. mexicana, Q. planipocula, Q. radiata, Q. scytophylla, Q. sideroxyla, Q. tarahumara and Q. urbanii. Species bearing this trichome type on abaxial leaf surfaces are Q. aristata, Q. castanea, Q. coahuilensis, Q. crassifolia, Q. hintonii, Q. hirtifolia, Q. hypoxantha, Q. mcvaughii, Q. radiata, and Q. tarahumara (Table 2). 1098 BRIT.ORG/SIDA 22(2) 2. Fasciculate sessile contorted (Figs. 2C, 2D).—This study describes this trichome type for the first time. Contorted trichomes are similar to the fasciculate sessile simple trichome described above except that the rays curl down and then twist around the tri- chome axis. This ray arrangement gives the indumentum a punctate appearance when observed with the naked eye or at low magnification. This trichome type has been found on the abaxial leaf surface of only two species of Mexican red oaks: Q. mexicana and Q. eduardii. 3. Fasciculate sessile crested (Fig. 3A).—This is a new trichome type that consists of rays united along a longitudinal axis oriented parallel to the epidermis. The united rays result in a structure that resembles a crest. Trichomes of this type were found only on the abaxial surface of some Q. sideroxyla specimens. 4. Fasciculate stipitate (Figs. 3B, 3C).—Morphologically, this trichome type is similar to the fasciculate sessile simple except that the basal parts of the ae fuse, forming a stipe. The degree of ray fusion varies among species from 1/12 to 1/4 the total ray length. Ray length and degree of divergence are also different; some species display short and straight rays (Fig. 3B) while others show long curly rays that become intertwined with adjacent rays (Fig. 3C). Ray number is variable within and among species, although in- traspecific ranges are low. Fasciculate stipitate trichomes form a persistent indumentum on the abaxial leaf surface of the following species: Q. conzattii, Q. crassifolia, Q. crassipes, Q. x dysophylla, Q. hintonii, Q. hirtifolia, Q. planipocula, Q. tarahumara,and Q. urbanii. In Q. aristata, fasciculate stipitate trichomes are deciduous and scattered on the abaxial leaf surface. In many otherwise glabrous species, such as Q. acherdophylla, Q. affinis, Q. coccolobifolia, Q. depressa, and Q. hintoniorum, only a few fasciculate stipitate trichomes remain attached to the epidermis at the secondary vein axils. Fasciculate stipitate tri- chomes on twigs were less common and found only in Q. aristata, Q. coahuilensis, Q. crassifolia, Q. x dysophylla and Q. hirtifolia. 5. Multiradiate (Figs. 3D, 4A, 4B).—This trichome type is characterized by the diver- gence of rays in different directions from the central axis. In the species examined, the rays differ not only in length but also in thickness. Multiradiate trichomes were found on both twigs and abaxial leaf surfaces of Q. candicans, Q. durifolia, Q. fulva,Q. hintoniorum, Q. hypoleucoides, Q. scytophylla, and Q. a This trichome type was found only on the twigs of the following species: Q. affinis, Q. conzattii, Q. crassipes, Q. depressa, Q. eduardii,Q. gentryi, Q. hintonii,Q. hypoxantha, Q. mcvaughii, Q. mexicana, and Q. urbanii. Multiradiate trichomes occurring on abaxial leaf surfaces usually form a persistent indument, however, in Q. salicifolia these trichomes are found scattered sparsely across the lamina. 6. Simple stellate (Figs. 4C, 4D, 5A).—This trichome type consists of rays appressed to the lamina, usually fused at their base, forming a structure that resembles a starfish. Simple stellate trichomes are composed of seven to fifteen rays, which vary in length, thickness, and size. This trichome type was previously described by Hardin (1976) and Jones (1986) under the term stellate. Here the term simple stellate is used to set it apart from the multiple stellate and fused stellate also found in this study. Simple stellate tri- chomes were found on the twigs of Q. acherdophylla, and Q. candicans, as well as on the abaxial leaf surfaces of Q. candicans, Q. durifolia and Q. gentryi. 7. Multiple stellate (Fig. 5B).—This new trichome type consists of 2-3 stellate tri- chomes joined together forming a trichome cluster. It was found only on one species, scat- tered on the abaxial leaf surface of Q. gentryi. — VAZQUEZ, TRICHOME MORPHOLOGY IN QUERCUS 1099 Fic. 3. A. Fasciculat il tedt ial leaf surface of Q id la, * 450.B. Fasciculate stipitate trichome on nie abaxialleatutiace sche Ce aie C. Fasci ipitate trichome on ial leaf surface of Q. j 0.D. f Q. salicifolia, x 500. 8. Fused stellate (Fig. 5C).—This trichome type previously described by Hardin (1979a) and Jones (1986) consis a of rays fused about one sixth of the ray length. This trichome type, not previously reported for red oaks, was found exclusively on the abaxial leaf sur- face of O. candicans. DISCUSSION The study of trichomes with SEM revealed significant differences in trichomes that ap- peared similar when viewed with dissecting microscopy. These differences have allowed a more accurate characterization of trichome types and have also led to the discovery of three new nonglandular trichome types: fasciculate contorted, fasciculate crested, and multiple stellate. Fasciculate contorted trichomes, although recognized asa different type by several authors (Mc Vaugh 1974; Gonzalez 1986), had been included under the stellate 1100 BRIT.ORG/SIDA 22(2) Fic. 4.A. Multiradi oa ne eee peer Ree Jurifolia, x 300. B. Mul ej as eisai of Q. scytophylla, x 500. C. Simple stellate trich the adaxial leaf surf f Q. eduardii < 500. D. Simple stellate trichomes on the twigs of Q. acherdophylla 800. type. The second new trichome type, fasciculate simple crested, is found only on the abaxial leaf surface of some Q. sideroxyla specimens. The uncommon distribution of fas- ciculate crested trichomes suggests that they could represent a variation of the multiradiate type, which is an abundant trichome type in the indumentum of Q. sideroxyla. The third new trichome type, multiple stellate, is found ina scattered pattern exclusively on the abaxial leaf surface of Q.gentryi. Under the dissecting microscope this trichome resembles a multiradiate type but close examination with SEM reveals that it consists of two to three stellate trichomes fused together (Fig. 5B). Therefore, trichomes that appear multiradiate must be examined carefully to avoid inclusion of multiple stel- late types into this category. For example, Hardin (1979b) described a trichome type asa dense multiradiate, however, his corresponding illustration (Fig. 30) shows a morpho- logical resemblance to the multiple stellate type described here. In addition to the new trichome types found in this study, three types represent new VAZQUEZ, TRICHOME MORPHOLOGY IN QUERCUS 1101 Fic. 5. A. Simple stell ict the abaxial leaf surface of Q jurifolia X 250. B. Multiple stellate trichomes on lower leaf surface of Q. gentryii_450.C. Fused stell ict he abaxial leaf side of Q. candicans, < 200.D. Fasciculate sessile trichomes lleafetd f 0. coahuilensis. 150. reports for red oaks: stellate, fused stellate, and glandular branched. Previous research had indicated that stellate and fused stellate types occurred only in white oaks (Hardin 1979a); however, this study found these trichome types in Q. candicans. Jones (1986) stated that stellate trichomes are “usually restricted to large veins or sometimes only the peti- ole.” In this research, stellate trichomes were found forming a dense and persistent indumentum on the abaxial leaf surfaces of the aforementioned species. The glandular branched type was rare and only found on the abaxial leaf surface of Q. salicifolia. This result supports Hardin's statement (1976) that this trichome type is not very common in red and white oaks. SEM was also useful in the accurate identification of previously known trichome types. For example, at low magnifications, trichomes on the abaxial leaf surface of Q. coahuilensis appear fasciculate stipitate, but examination under SEM has revealed that they are fasciculate sessile: the stipe-like structure is formed by the twisting of rays near 1102 BRIT.ORG/SIDA 22(2) the trichome base (Fig. 5D). Similarly, detailed study of the trichome type denoted as “starred” in several descriptions of Mexican oak species (Bello & Labat 1987; Valencia 1989; Vazquez 1992, 2000; Romero 1993) showed that these categories correspond to four different trichome types: fasciculat ile, fasciculate stipitate, fasciculate contorted, or multiradiate trichomes. Acknowledging the trichome variation included under the term “starred” will facilitate future characterization of Mexican oak taxa. Study of twigs under SEM has also revealed remarkable characteristics regarding the nature of the indumentum. Several taxonomic treatments of Mexican oak taxa (Gonzalez 1986; Valencia 1989; Vazquez 1992) imply that the twig indumentum consists of only one nonglandular trichome type. However, the results of this study indicate that the indumentum of nineteen of the species examined is composed of two to three tri- chome types arranged in overlapping layers (Fig. 6A). Of the thirty species examined, only nine have a twig indumentum composed of one nonglandular trichome type (Figs. OB, 6C). In general, species bearing more than one nonglandular trichome type on twigs and/ or leaves show several combinations of glandular and nonglandular trichomes, termed trichome complements. Glandular trichome types are not included in the trichome complements because, if present, they usually are deciduous. Trichome complements on twigs The most common trichome complement on twigs is multiradiate/fasciculate sessile simple and occurs in the following species: Q. conzattii, Q. crassipes, Q. durifolia, Q eduardii, Q. fulva, Q. hintoniti, Q. hinloniorum, Q. hypoleucoides, Q. hypoxantha, Q. mevaughii, Q. mexicana, Q. scytophylla, Q. sideroxyla, and Q. urbanii. Twigs with a tri- chome complement consisting of fasciculate sessile simple/fasciculate stipitate trichomes are found in Q. codhuilensis, fa) crassifolia, Q. x Cane and Q. hirtifolia. A species- specific trichome complement was found on twigs of Q. candicans (fasciculate sessile simple/multiradiate/simple stellate) (Table 2). Trichome complements on the abaxial surface of leaves The distribution of glandular trichomes on leaves showed different patterns. On one hand, the simple branched glandular trichome type is found exclusively on the abaxial leaf surface of Q. sdlicifolia in a scattered fashion (Fig. 6D). On the other hand, the simple uniseriate type is found on the abaxial leaf surface of most of the species examined. Gen- erally, this trichome is deciduous and usually not found on mature leaves: however, in Q. radiata and Q. coccolobifolia, these glandular trichomes are persistent in the form of mucilaginous droplets, which according to Uphof (1962) are the result of decaying glan- dular ae enn The common presence of simple uniseriate glandular trichomes has also been documented for many US and European oak species (Hardin 1976, 1979a; Lla- mas et al. 1995), and has also been found in the red oak species examined here. Trichome complements on the abaxial leaf surface are composed of two to four tri- chome types (Table 3). The trichome complement fasciculate sessile simple/fasciculate stipitate is found in the species Q. aristata, Q. crassifolia, Q. hintonii, Q. hirtifolia,and Q. tara pelcake aes epee tr ae eee oy en on the abaxial leaf surface ), Q.durifolia (multiradiate/simple a ate), QO. hintoniorum (fasciculate stipitate/multiradiate), and Q. sideroxyla (fascicu- late sessile crested/multiradiate), (Table 3). Although the occurrence of more than one nonglandular trichome type on the abaxial surface has been documented for European —" VAZQUEZ, TRICHOME MORPHOLOGY IN QUERCUS 1103 6.6 A r . 1 yi Il L L 4 tn FY hd 200 R Cc = i: | = | l1nf4+\ L 1 tate (right) trichomes on pa: at en 200.C. Multiradi ict the twigs of Q. affinis < 200.D. Scattered branched glandular trict f Q. salicifolia, < 200. (Safou & Saint-Martin 1989) and North American (Thomson & Mohlenbrock 1979) oak species, it has only been described for a few Mexican taxa. For example, Nixon and Muller (1993) described the unique trichome complement found in Q. hintoniorum and related species. The abaxial leaf surface of six of the included species is essentially glabrous except for clusters of fasciculate stipitate trichomes in the vein axils, or scattered multiradiate or stellate trichomes distributed throughout the lamina. Species with fasciculate stipi- tate trichomes restricted to vein axils are: Q. acherdophylla, Q. affinis, Q. depressa, and Q. coccolobifolia, although the later species also has abundant simple uniseriate trichomes throughout the abaxial surface. Scattered multiple stellate and multiradiate trichomes have been found only in Q. gentryi while dispersed multiradiate and branched glandular trichomes are found in Q. salicifolia. 1104 BRIT.ORG/SIDA 22(2) Trichome morphology and species identification There are several instances where trichome morphology can be a valuable tool in the identification of red oak species, especially of those with similar overall features. For ex- ample, Q. crassipes and Q. mexicana (Figs. 7A, B) are frequently confused because they have similar leaf morphology. However, examination of trichomes on the abaxial leaf surfaces ie on ae fferences: while Q. crassipes has fasciculate stipitate trichomes, Q. mexic ile contorted trichomes. Reproductive characters (e.g. fruit morphology) agree with their distinctiveness based on trichome morphology. Another example of the utility of trichome morphology for species identification is found in the Crassifoliae complex (sensu Vazquez 2001, which consists of fourteen species charac- terized by the presence of a pale-yellow indumentum on the abaxial leaf surface, and by a morphological resemblance to leaves of Q. crassifolia. Two species often confused with Q. crassifolia are Q. mcvaughiiand Q. fulva. While the abaxial leaf surface of Q. crassifolia has distinctive lfasciculate stipitate trichomes, Q. mcvaughii displays fasciculate sessile trichomes and Q. fulva shows multiradiate trichomes on their abaxial leaf surfaces. In fact, trichome differences between Q. crassifolia and Q. mevaughii, together with other morphological and reproductive characters were used in the segregation of these mor- phologically similar species (Spellenberg 1992). A potential problem in trichome classification is that there are cases when there is not a clear-cut distinction among types. One case involves Hardin’s (1976) glandular rosulate trichome type, characterized by the rosette pattern of the rays and, most impor- tantly, the rays having a thin cell wall. In practice, it is difficult to determine the nature of the cell wall unless ultrastructural studies are carried out. In the absence of such stud- ies and taking into account ray arrangement, these trichomes should be classified as multiradiate. In this study, all trichomes with rays originating from a single point and diverging in different directions were considered within the multiradiate type A second problem in the identification of trichome types is the transition of one type into another. In the specimens examined, morphological transitions were observed between fasciculate sessile and fasciculate stipitate trichomes, and from fasciculate sessile to stellate. These examples agree with Jones’ (1986) statement that sometimes there are no discrete boundaries between trichome types and that “.. most forms intergrade with at least one other type...” Despite these potential limitations, SEM has been a key factor in the discovery of three new trichome types, as well as in the finding of three trichome types not previ- ously reported for red oaks. Furthermore, study of trichomes under SEM has revealed key morphological differences in apparently similar trichome types. Detailed examination of trichomes from Mexican oak taxa revealed that four different trichome types had been included under the “starred” category in the taxonomic literature. The trichome comple- ments of mature leaves are useful in the identification of morphologically similar oak species when taken together with other vegetative and reproductive features, as indicated in the dichotomous key below. This work PEpyinee a standard terminology that could be used in future taxonomic treatments to aid in sf : iption of Mexican oak taxa. = DICHOTOMOUS KEY FOR IDENTIFICATION OF MEXICAN RED OAK SPECIES . Abaxial leaf surface either completely devoid of trichomes or appearing glabrous except for scattered glandular or nonglandular trichomes. o suborbicular or pandurate; abaxial leaf surface with abundant simple 2. Leaf shape ovate VAZQUEZ, TRICHOME MORPHOLOGY IN QUERCUS eink: 2 Wier year ; KE hs ea BLEUE . o: + (a ict the abaxial leaf surface, X 74.C.—D. Q. mexicana L ae oe | _ m4 «74, Fic.7. A.—B. Quercus crassipes, left i i i F y TIGH, I + | H + €. H lo¢ H | = uniseriate glandular trichomes and a few fasciculate stipitate trichomes restricted to the Q. coccolobifolia secondary vein axils 2. Leaf shape elliptic, lanceolate, ovate-lanceolate, or oblanceolate; abaxial leaf surface devoid of trichomes or with scattered stellate or multiradiate trichomes. 3. Leaf margin entire and undulate 4. Leaves 4-6 times long as wide; abaxial leaf surface with sparse multiple stellate tri- chomes or multiradiate trichomes distributed throughout the lamina and vein axils; fruits with biennial maturation. 5. Secondary veins 10-15; two-thirds of the acorn included in the cup; epidermis on Q. gentryi the abaxial leaf surface bullate 5. Secondary veins 15-25; one-third of the acorn included in the cup; epidermis on Q. salicifolia the abaxial leaf surface smooth Leaves 2—2.5 times long as wide, abaxial leaf surface lacking trichomes on the lamina but with fasciculate stipitate trichomes on the vein axils; fruits with annual maturation Q.acherdophylla aN 1106 BRIT.ORG/SIDA 22(2) 3. Leaf margin entire and straight or with short teeth. 6. Leaf base attenuate, cuneate or decurrent; mature leaves 3.6-4.2 times long as wide Q. affinis 6. Leaf base round or obtuse; mature leaves 2.4—2.6 long as wid Q. depressa “Abn leaf surface with abundant and persistent nonglandular trichomes, sometimes trichomes 7. Abaxial leaf surfac e sparingly pubescent or glabrescent; if sparingly pubescent, at low mag- nifications the epidermis could be seen through the trichomes 8. Trichomes on axial leaf surface have a punctat ey trichomes fasciculate con- 9. Leaves elliptic-oblong, always entire, veins 6-16 9. Leaves elliptic-lanceolate, elliptic-oblanceolate, oothed, veins 5-8 Q. mexicana ometimes obovate or ovate, entire or Q. eduardii Trichomes on abaxial leaf surface have a floccose or arachnoid appearance; trichomes ion stipitate, fasciculate sessile simple, or multiradiat es orbicular, orbicular-pandurate, or broadly es twigs 4-8 mm thick. (ae leaves with 4-8 teeth; infructescence up to 6 mm long bearing 1-2 fruits Q. tarahumara 11. Mature ik with 13-19 teeth; infructescence from 4.7 to 15 cm long bearing 3-9 fru Q. radiata 10. Leaves sii ,lanceolate, ovate or obovate; twigs 0.75-4(-5) mm thick. 2. Fruit cupule with the margin involute; cupule diameter 20-28 mm; secondary veins 10-19 Q. planipocula 2. Fruit cupule with the margin straight; cupule diameter 7-15 mm;secondary veins = 12, 13. Fruits with annual maturation; bullate epidermis on the abaxial leaf surface; secondary veins 5-12 14. Leaves obovate to elliptic; trichomes on the abaxial leaf surface fascicu- late sessile or fasciculate shortly stipitate; leaf margin toothed; distributed y SUIp C in the Sierra Madre Oriental . hirtifolia 4. Leaves elliptic-lanceolate, elliptic-oblanceolate, sometimes obovate or ovate; trichomes on the abaxial leaf surface fasciculate distinctly stipitate; leaf margin entire or entire-aristate; distributed in the Sierra Madre Occidental Q. aristata 13. Fruits with biennial mat th epidermi the abaxial leaf surface; secondary veins 4-5 15. Aba xial leaf siintack appearing glabrous but with two types of glandular nontrichomes visible at 10:small multiradiate of short rays distributed on the abaxial lamina and fasciculate stipitate trichomes confined to the secondary vein axils Q. hintoniorum 15. Abaxial leaf surface pubescent but the trichomes detach with age; tri- chomes fasciculate sessile simple with long intertwined rays distributed enn e the lamina Q. hypoxantha 7. Abaxial leaf surface densely aie the epidermis completely covered with trichomes. 16. Leaves elliptic, lanceolate, obla | e. 17. Leaf margin always en a arista 18. Abaxial leaf surface Suess eee cent, the indumentum whitish, trichomes multiradiate or | t appearing fasciculate sessile at 10x: ;apex acute 9. Leaf margin flat; mature leaves 2.6-3 times long as wide; petiole 2-6 mm g lon Q. durifolia 19. Leaf margin revolute; mature leaves 3.75—5 times long as wide; petiole 5-15 mm long hypoleucoides 18. Q. Abaxial leaf surface loosely pubescent, the indumentum pale yellow, trichomes clearly fasciculate stipitate at 10x; apex rounded oro obtuse 20. Mature leaves about 2.5 times longer than wide; cupbie hemispheric or tur- VAZQUEZ, TRICHOME MORPHOLOGY IN QUERCUS 17. Leaf margin mainly toothed, rarely entire with aris 16. Leaves obovate, suborbicular, nes or very broadly elliptic. 25. The indumentu 25. Thei 1107 binate, the cupule margin not involute; sessile and fasciculate stipitate trichomes 20. Mature leaves 4—-4.5 times longer than wide; cupule hemispheric with invo- lute margin; current year twigs with fasciculate sessile and multiradiate tri- chomes year twigs with fasciculate dysophylla Q. crassipes 21. Twigs 3-5 mm thick; abaxial leaf surface with ake stipitate trichomes;cup very shallow up to 5 mm tall, enclosing 1/4 of the nut 21. Twigs 1-3 mm thick; avant leaf surface with Tescicilane ee simple, multiradiate, or fasciculat rested trict - enclosing 1/3 to 1/2 of the n 2. Abaxial leaf surface with ee sessile trichomes of 6 to 10 long inter- twined rays. 23. Petioles 3-5(—7) mm long; fasciculate sessile trichomes twisted at the e and detaching with age; leaf margin slightly revolute Q. coahuilensis 23. Petioles 5-25 mm long; fasciculate sessile trichomes not twisted at the base and pocisting with age; leaf margin flat 22. Abaxial leaf surface with multiradiate ee lat il of 20-22 short rays. Q. hintonii ’ a Q. castanea omes 24. Leaf base decurrent or cuneate; leaf tapering toward the apex; leaf mar- gin not revolute Q. scytophylla. 24. Leaf base rounded, subcordate or cordate; leaf apex acute or obtuse; leaf margin revolut Q. sideroxyla n the abaxial leaf surfaces detaches with age; twigs 0.75- thick; petioles eae fegaleh indumentum on the f surf petioles densely pubescent pubescent or - glabrescent, 26. Petioles 0.4-25 mm long; cup 7-9 mm wide, 4-5 mm tall; acorn 7-11 mm long o hypoxantha hed;twigs 2-13 mm thick; m wide Q. conzattii 26. Petioles 25-37 mm long; cup 8-24 mm wide, 5-15 mm tall; acorn 10-24 mm long, 7-21 mm wide. 27. Twigs fines season 8-13 mm thick; infructescences 3.5 to 10 cm long bear- ing 6-10 Q. urbanii 27. Twigs ee season 2.5-6.0 mm thick; infructescences less than 2 cm long bearing 1-2 fruits 28. indumentum on the apaalal leaf surface white or pale yer, leaves with 1 teeth; sp glandular trichomes on the abaxial leaf surface Q.candicans 28. Indumentum on the abaxial leaf surface yellow or yellow-brownish; leaves with 1-11 teeth; abundant glandular trichomes on the abaxial leaf surface. 29. Abaxial leaf surface with a felted indumentum composed of multiradiate trichomes with straight rays; rim of cupule involute Q. fulva 29. Abaxial leaf surface with a woolly indumentum composed of fas- ciculate trichomes with long tortuous rays; rim of cupule straight. 30. Mature leaves deciduous; trichomes on the abaxial leaf surface ee stipitat Q. crassifolia 30. Mature leaves Se Serre on the abaxial leaf surface fasciculate sessile simple Q.mcvaughii 1108 BRIT.ORG/SIDA 22 2) ACKNOWLEDGMENTS | wish to thank Kevin Nixon for sharing his knowledge on oak systematics, the Bailey Hortorium, Department of Plant Biology, at Cornell University, for financial assistance for the completion of this research, Richard Spellenberg for his great suggestions for the improvement of this manuscript, Brian Phillips and Amy McEuen for their comments on the manuscript and testing the dichotomous key, and Jeff Bacon, Hector Vazquez Villagran, Liliana Vazquez Villagran,and Rebeca Ocampo for their assistance in the field. Lalso want to thank Richard J. Jensen (NDG) and Paul S. Manos (DUKE) for their helpful suggestions as reviewers for Sida. REFERENCES Bacon, J.R.and R. Speccensers. 1996. Hybridization in two distantly related Mexican black oaks Quercus conzattii and Quercus eduardii (Fagaceae: Quercus: Section Lobatae). Sida 17:17-41 Brito, M.A. and J.N. 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Species of the genus Quercus in Italy: characterization by means of leaf surface observation. Giorn. Bot. Ital. 126:481-504 Gonzalez, L.M. 1986. Contribuci6n al conocimiento del género Quercus (Fagaceae) en el estado de Jalisco. Instituto de Botanica. Universidad Autonoma de Guadalajara. Mexico Haroin, JW. 1976. Terminology and classification of Quercus trichomes. J Elisha Mitchell Sci Soc 92: 151-161. Harbin, J.W. 1979a. Patterns of variation in foliar trichomes of eastern North American Quercus.Amer. J.Bot.66:576-585. Harbin, J.W. 1979b. Atlas of foliar surface features in woody plants, |. Vestiture and trichome types of eastern North American Quercus. Bull Torrey Bot Club 106:313-325 INAMDAR, J.A. AND N.V. Rao. 1983. Light and scanning electron microscopic studies on trichomes of some Brassicaceae. Feddes Repert. 94:183-190. JENSEN, R. 1997. Quercus section Lobatae.|n: Flora of North America Editorial Committee, eds, Fl.North Amer. 3:445-506 Oxford Univ. Press, New York and Oxford. Jones, H. 1986. Evolution of the Fagaceae: The implications of foliar features. Ann. Missouri Bot. Gard. 73:228-275. Kissuine, P.1977.Les poils des quatre especes de chenes du Jura (Quercus pubescens, Q. petraea, Q. robur et Q. cerris). Ber Schweiz Bot Ges 87:1-18. Kissuin, P. 1993. Heteroindumentum: Des poils de taxons differents sur la meme plante. Bot Helv 103:83-99, Liamas. F., C. Perez-Morates, A. Acebo, and A. Penas. 1995. Foliar trichomes of the evergreen and VAZQUEZ, TRICHOME MORPHOLOGY IN QUERCUS 1109 semideciduous species of the genus Quercus (Fagaceae) in the Iberian Peninsula. Bot.J.Linn.Soc. 117:47-57, Manos, PS. 1993a.Cladistic analyses of molecular variation of“higher” Hamamelididae and Fagaceae, and systematics of Quercus section Protobalanus. Ph.D. diss. Cornell University, Ithaca, NY. Manos, PS. 1993b. Foliar trichome variation in Quercus section Protobalanus (Fagaceae). Sida 15: 391-403 Manos, P.S., Nixon. K.C., Doyte, J.J. 1999. 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Otsson, U.1976.The structure of stellate trichomes and their taxonomic implication in some Quercus species (Fagaceae). Bot. Notiser. 128:412-424 Oostuuizen, L. 1983. The taxonomic value of trichomes in Pelargonium (Geraniaceae). J. S. Afr. Bot. 49:221-242. Romero, S. 1993. El género Quercus (Fagaceae) en el estado de México. Master's thesis. Universidad Nacional Auténoma de México. Mexico Rzepowski, J. 1965. Relaciones geograficas y posibles origenes de la flora de México. Bol. Soc. Bot. Méx. 29:121-177. Rzepowski, J. and G.C. Rzevowski. 1979. Flora fanerogamica del Valle de México. CECSA, México. Sarou, O. AND M. Saint-Maatin. 1989. Foliar trichomes of some perimediterranean oaks. Bull. Soc. Bot. France Lett. Bot. 136:291-304. SHANMUKHA, Rao S.R. 1987. Structure, distribution and classification of plant trichomes in relation to taxonomy: Sterculiaceae. Feddes Repert. 98 (1-2):127-135. SPELLENBERG, R. 1992. A new species of black oak (Quercus, subg. Erythrobalanus, Fagaceae) from the Sierra Madre Occidental, Mexico. Amer. J. Bot. 79:1200-1206. SPELLENBERG, R. 1998. Quercus lesueuri, un miembro meridional del complejo Q._ undulata (Fagaceae, subgenero Quercus). Acta Bot. Mexicana 42:25-33. SpeLLENBERG, R. 2001. Taxonomic overview of the oaks of La Frontera. In:G.L.Webster and C.J. Bahre, Changing Plant Life of La Frontera, Univ. New Mexico Press, Albuquerque. Pp. 195-211. SPELLENBERG, R., J.R. BACON, AND M.S. GonzALez. 1998. Los encinos (Quercus, Fagaceae) en un transecto sobre la Sierra Madre Occidental. Boletin Instituto Botanica, Universidad de Guadalajara, Mexico 9:357-38/7. THomson, P.M. and R.H. Monvensrock. 1979. Foliar trichomes of Quercus subgenus Quercus in the East- ern United States. J. Arnold Arbor. 60:350-367. Tucker, JM. 1952. Evolution of the California oak Quercus alvordiana. Evolution 6:162-180. Tucker, JM.and C.H.Mutter. 1957.A reevaluation of the derivation of Quercus margaretta from Quercus gambelii. Evolution 12 -17. Upnor, J.C. 1962. Plant hairs. Gebrtider Borntraeger. Berlin-Nikolassee. a 1110 BRIT.ORG/SIDA 22(2) tadn da CGiier DtdGO Qe WUE! VALENCIA, S. 1989.Contribucidn al conocimiento del género Quercus (Fagaceae) | rero, México. Undergraduate thesis. Universidad Nacional Aut6noma de México. VAzquez, M.L. 1992. El género Quercus en el estado de Puebla, Mexico. Undergraduate thesis. Univer- sidad Nacional Autonoma de Mexico. VAzauez, M.L. 2000. Flora del Valle de Tehuacan-Cuicatlan. Fagaceae. Instituto de Biologia, Univer- sidad Nacional Autonoma de México, Fasciculo 28. Mexico. VaAzauez,M.L. 2001 Molecular and morphological studies on Mexican red oaks (Quercus sect.Lobatae). Ph D. Dissertation. Cornell University, New York. THOMAS WALTER TYPIFICATION PROJECT, L. OBSERVATIONS ON THE JOHN FRASER FOLIO Daniel B.Ward Department or Borany I fap #C]/ UTIIVET OF ity GIT lorida Gainesville, Florida 32611, U.S.A. ABSTRACT The Natural History Museum, London, is the repository of a large volume containing plants collected by John Fraser in the southeastern United States in the 1780s and commonly known as the : ANalter Herbarium.” The importance of this collection is that mony a its ceed were seen and petheps ued Py Pom as Walter, au- thor of Flora Caroliniana, the first A | pt Linnae an | t Ade escription is provided here of this folio and the collection it contains. RESUMEN El Natural History Museum de Londre | | | John Fraser en el sudeste de los Estados ee en los 1780s y cenesido seeaalnente como el‘ “Walter He ee La importancia de esta coleccion radical en que muchos de sus especimenes fueros vistos y quizas usados por Thomas Walter, autor de la Flora micas aa: la ae flora americana en adoptar la oe Linneana y la nomenclatura binomial. Se hace un le este volumen y la coleccion que contiene The Thomas Walter Typification Project is the term now given of an effort, extended in- termittently over forty years, to bring understanding and nomenclatural precision to the many plant names published by Thomas Walter in his pioneer Flora Caroliniana (1788). Walter, an English rice-plantation owner and amateur but skilled botanist, resident of South Carolina, was the first to prepare a flora of American plants utilizing Linnaeus’s binomial nomenclature and sexual system of classification. A brief but close relation- developed between Walter and John Fraser, a Scottish plant explorer who in the late 1780s gathered numerous specimens from the southeastern United States into a large folio herbarium now held by the Natural History Museum, London. The specimens of this herbarium were seen and may in part have been used by Walter in preparation of the diagnoses of his new species. The present report is an account of the physical character- istics of this historic collection. The information of this report is derived from notes taken during a two-week ex- amination of the folio by the present author in July 1984, and from 8x10 glossy photos enlarged from negatives made of the folio by Dr. Bernice G. Schubert in November and December 1946 (Fernald & Schubert 1948). (Commercial microfiches distributed in 1985 were of lesser clarity. Digital images prepared in 2005 were not yet available.) Though much remains unknown, many aspects of the interrelated lives and works of Thomas Walter and John Fraser have been documented by Britten (1921), Coker (1910), Hogg (1852), Maxon (1936), Rembert (1980), Simpson et al. (1997), Small (1935), and Ward (1962 1977). The specimens are mounted on the pages of a large book, a folio, 54.5 cm tall, 38 cm wide, about 8 cm thick, bound in green cloth and half-leather with 5 transverse raised bands and several thin gold lines. The pages are 52 cm by 35.5 cm, of heavy linen-rag watermarked paper, bound permanently along a lateral margin. The book is of commer- SIDA 22(2): 111- 118.2006 2006 1112 BRIT.ORG/SIDA 22(2) cial origin, manufactured in advance of its present use; it was not made by binding to- gether separate sheets to which the plants had been attached. Two initial pages are un- numbered; each subsequent page has been numbered by hand near its upper margin. The folio consists of 119 numbered pages (not 117, as reported); two numbers (61 and 62) were used twice. The frontispiece, the unnumbered first page of the folio, bears an attached sheet (right- diagonals indicate new lines; left diagonals reflect superscripted letters): “The Herbarium of Tho\s Walter Esq. of South Carolina. / Author of the Flora Caroliniana, pub. 1788 / Presented May 23, 1849, to the / Linnean Society of London, by John Fraser, / son of John Fraser, the indefatigable / North American Botanical Collector, / from the years 1786 to I811. He died / in London, the latter year.” This slip (as believed by James Britten 1921) is in the hand of Fraser’s son, John Fraser fil. A second, unknown writer has restated these words by inscription onto the page itself: “The Herbarium / of Thomas Walter / author of the / “Flora Caroliniana.” / Presented to the Linnean Society, 23 May 1849, / and pur- chased at the sale of the Society’s / Surplus Collections in 1863 (for 15/.).’ The frontispiece also bears a detailed listing (probably by Britten) of the persons, into the 20th century, whose annotations are to be found within the folio. The title page of the folio, also unnumbered, bears the boldly written words, “Walter's Herbarium,” the dates “1786-1788,” and the initials, “JE” The hand is that of John Fraser. The dates encompass the frenetic eighteen months—from September 1786 until March 1788—during which Fraser traveled and collected in the American Southeast. The plants of the collection are mounted (counterintuitively) on the left-hand pages of the opened folio. The herbarium consists of 690 specimens (of which three are repre- sented only by their labels and by stains on the page to which they had been attached), The specimens are mounted with as few as 1 and as many as 12 ona single page (an aver- age of 5.8 specimens per page). For clarity of reference in present and future study, images (xeroxes from photos) of the specimens have been enumerated by two-part designators: each individual specimen is given the number of the page on which it is mounted, followed by a capital letter. (The two pages with duplicated numbering, p. 61 and p. ©2, are distinguished by suffixes “a” and “b.”) The specimens, insofar as their arrangement on the page permits, have been assigned a designator in alphabetical order, from upper left to upper right, second-row left to second-row right, down the page. A copy of the images showing the assigned des- ignators has been provided to the Natural History Museum, for placement with the folio. These designators are suggested for use in specimen citation, to replace the use of page numbers or other accompanying numbers seen on many labels, as resorted to by other authors. The designators are employed in the present discussion where appropriate. Nearly all specimens are accompanied by a small label. (A very few have no label; a few have either two labels, or a label clearly belonging with another specimen.) The la- bels vary in proportion and dimension, but are mostly of horizontal rectangles, 5 to 8 cm long, 2 to5 cm high. All have clearly been individually and somewhat roughly cut from other sheets. A few (e.g., 41-E, 51-A, 65-A, 71-A, 80-A, 83-A, 83-E) have been folded trans- versely, the lower half tucked behind the upper half which bears the writing. Two dis- tinct paper stocks were used for the labels, both of “laid paper,” formed froma linen slurry: the one thin, tan, with laid-lines spaced +9 per cm; the other heavier, whiter, with laid- lines +7 per cm. Most of the specimens are attached to the label by one slit or more often two parallel — — R IT COLLECTION 1113 slits cut in the label, with the plant stem passed under the resulting strap. At times the slits are in the form of shallow V’s (e.g., spms. 24-B, 31-C, 32-C, 34-B, 6la-F), indicating they were cut by folding the label, then making the two cuts with two snips of a scissors. Frequently part of the writing on the slip is beneath part of the specimen, obviously having been inscribed prior to attachment of the plant. A few of the specimens have been attached to the label by slender pins (e.g., 46-B, 60-E 61la-A, 92-J, 105-C). Others have been given firmer mounting, either to the label or to the underlying sheet, by conventional narrow tape strips. A significant achievement of the Project has been the determination of nearly all of the handwritings on the labels. Yet, for reasons explained elsewhere (Ward, in press), this information is of less value than might be assumed for determination of the collector of the accompanying specimen or for prediction of its use by Thomas Walter in forming the diagnoses of his Flora. Two handwritings predominate on the labels: Thomas Walter (ca. 1740-1789), and John Fraser (1750-1811). The hand of Fraser's son, John Fraser (1780-1861) [not “1799- 1860?”—Fernald & Schubert 1948] also appears to be present, though rarely. External evi- dence of these hands is provided by a 158-word letter written by Walter in 1783 and re- produced by Ewan (1979), and by a 48-word letter written by Fraser’s son in 1818 in the Autograph Collection of the Natural History Museum. Fraser's hand is established by the few words and dates of the title page and by occasional brief, initialed notes accompany- ing the specimens (e.g., 67-A, 78-E). The third hand probably of Fraser fil. differs mark- edly from that of Fraser and also somewhat from that of Walter. tis found on a few labels (ge GF 26) 676 lbp 105-G, 115-G). Numbering of the pages of the folio may also have been by Fraser fil; these numbers are not in the hand of Fraser sr. Once determination was made that two writers were responsible for the great major- ity of the labels, most of the handwriting could be assigned. Past authors have referred in general terms to similarities with that of Walter, Fraser, or Fraser's son, but few authors have claimed which hands were found in association with specific specimens, and none are known to have based nomenclatural arguments on such identifications. [Blake (1915:130) correctly identified Walter's hand on four labels, but wrongly concluded “most of the remaining labels are in another hand, which from its agreement with some writ- ing on the title page of the volume may be affirmed with considerable certainty to be that of John Fraser himself | Yet close—and tediously prolonged—examination of the hand- writing of the labels and of the available handwriting samples does usually permit cer- tain identification John Fraser’s hand in most cases is unambiguous. It is a smoothly flowing script, with the letters consistently formed and widely spaced. His capital “A” is of the classic “inverted V” and uniformly shows the cross-bar sweeping far left across the ascent. His capital “C” is always shaped with both its beginning and terminus well to the left of the figure. His capital “P” is initiated by a strong upstroke, sharply reversed to a downstroke, again reversed to form a clockwise loop. Many of his other capital letters also show dis- tinctive features, though his lower case letters are largely of standard form. He is unfail- ing in his practice of using capitals for the initial letters of both genera and epithets. His hand is of an extrovert—if this term can be applied toa writing style—fitting the man of confidence and courage his travels and ventures show him to have been. Thomas Walter’s hand is far more cautious, cramped, and poorly, unevenly formed. His capital “A” is at times an “inverted V,” but his cross-bar is formed separately and 1114 BRIT.ORG/SIDA 22(2) scarcely touches the ascent; for many of his plant names his capital “A” is merely lower case writ large. His capital “C” is of the same form as Fraser's, but only the beginning is from the left; the lower terminus invariably links with the next letter. His capital “P” is of a smooth clockwise swirl, with the initial stroke directed downward. With few excep- tions his epithets are not capitalized. His hand in general is that of a busy, quick thinking person, not overly concerned with the niceties of precise letter formation; in modern days his script might have been that of a doctor. A readily available sample of the two handwritings is given in photographs shown by Faden (1989:46-47). There, the single word “Commelina” is written on the label of a specimen of C. erecta by Walter, and on the label of a specimen of C. caroliniana by Fraser. The Walter label also bears “305,” a number written by Fraser. John Fraser fil’s hand is poorly understood because of its infrequent appearance. Though it is unlikely to be mistaken for that of John Fraser sr, brief sequences of letters (as in plant names or brief notes) may not show marked contrast with that of Walter. A character that may be useful is the “square r” here attributed to Fraser fil., while the lower- case “r” of both Fraser sr. and Walter is usually an abrupt simple upward pip More than half of the labels (371, or 54%) bear three-digit (rarely two- digit) numbers ina darker ink anda coarser pen, and seem to have been written prior to other markings. The hand forming these numbers is that of John Fraser; his distinctively malformed “8” appears also as part of the date given on the title page of the folio. (Representative speci- mens so marked: 1-H, 27-C, 34-D, 51-B,54-D, 59-E, 66-C, 101-C, 103-F 107-A.) Britten (1921) was unable to discover the use to which Fraser put these numbers, nor have others sug- gested any logical purpose. The numbers correspond neither to the present arrangement of the specimens nor to the sequence of Walter's Flora (nor presumably to its parent manu- script). Many of these numbers have been struck by a single diagonal slash; most of the rest are accompanied by a small checkmark. No numbers are repeated. The numbers may differ in size between two adjacent specimens (e.g.,62a-B and 62a-C, 94-D and 94-E), dem- onstrating that specimens now adjacent in the herbarium were numbered at different times. Frequently the numbers are partially cut away at edge of the label (e.g, 38-1, 50-E, 80-D, 84-A, 86-C, 92-A, 105-B, 108-F). These characteristics support the speculation that Fraser's bers served as field references, perhaps coupled toa now-lost separate record of source and date, the pages later cut to their present dimensions and used as permanent labels. The few two-digit numbers of this series (six exceptions, but not clearly in the same hand), together with knowledge that Fraser spent earlier years (1780-1784) in New- foundland where he also studied plants (Fraser 1789; Hogg 1852), suggests that these num- bers may have been a continuing enumeration of his collections begun before he first came to the Carolinas. Certainly, aside from their initial purpose, their presence on the label indicates that the specimen is of Fraser's collection, whether or not it was later seen by Thomas Walter Another series of numbers seems not to have been noted. Each number of this small class (20 numbers) is preceded by a capital “F” or “FC.” All are clear references to the num- bered genera in Flora Caroliniana. Examples: L00-C is named “Staphylea Trifolia” and numbered “F 132,” while genus 132 is Staphylea; 106-A is named “Verbena Urticifolia” and numbered “F 236,” while genus 236 is Verbena with the single species V. urticifolia; 116-Cand 116-D both bear the name “Syntherisma” and are numbered “F 35,” while genus 35is Syntherisma (= Digitaria). Further examples: 32-A, 32-B, 32-C, 32-D, 38-], 48-C, 115- G, 117-A, 117-D, 117-E.) Most of the numbers occur in clusters (e.g., 32-A thru 32-D, 101-A 1115 thru 101-G, 115-G thru 117-E), indicating they were assigned after the specimens were arranged in the herbarium. The hand inscribing this series of numbers is probably that of Fraser (e.g., 38-J, 48-C, 100-C), though at times the “F” seems unlike his. Possibly it is that of his son; it is not that of Walter. These numbers were certainly assigned after pub- lication of Flora Caroliniana, or at least after access to the completed manuscript. This second series of numbers appears to have been a partial, never completed attempt to match the specimens collected by Fraser to the appropriate souls as Su blal aice by Walter. Within the folio the specimens are arranged in l sequence, as determined by the names borne by the original labels. These names, in turn, are largely those used in Walter's Flora Caroliniana(1788).[The few exceptions are usually in the hand of John Fraser and beara name from Linnaeus that was not used by Walter (e.g., 37-G, “Cornus canadensis’; 52-E, “Globularia nudicaulis”, 58-C, “Helonias bullata”), or a comment by Fraser or Walter in place of a name le.g., 38-D, “Corypha arborea” (= Sabal palmetto), 43-B, “Erythronium Dens Canis” (= E.americanum),55-C, “Hedysarum Flore magnus’ (= Desmodium cuspidatum), 67-D, “Lupinus affinis” (= Crotalaria rotundifolia), 72-A, “Mespilus Large Cock Spur’ (= Crataegus crus-galli); 98-D, “Smilax Baccis albidis” (= Smilax laurifolia)], Of the specimens bearing an identification on the label (either complete, or only to genus), 345 (50.1%) bear Walter's hand, 230 (33.4%) bear Fraser's. Often the hands of Walter and of Fraser are found on the same label. On 49 labels Walter identified the specimen to genus and Fraser completed the naming by adding the species. On none of the labels did Fraser identify the specimen to genus, with Walter adding the species. Handwritings by other persons have been added over the years, usually in the form of identifications. They may be on separate slips, on the adjacent sheet, or on the original label itself. Those writers identified on the frontispiece of the folio (by James Britten) are: Oakes Ames, Sidney F Blake, Nathaniel Lord Britton, Alva A. Eaton, Asa Gray, Albert S. Hitchcock, Charles S. Sargent, James E. Smith, and Anna M. Vail. Others include James Britten, James E. Dandy, Merritt Lyndon Fernald, William T. Gillis, A.J. Kosterman, John Lewis, Mildred EF. Mathias, and Charles A. Weatherby. More recent annotations (since 1983) are by Joseph Ewan, David K. Northington, David H. Rembert, Daniel B. Ward, and others. Identification of the specimens is no easy task. As noted in 1839 by Asa Gray (J.L. Gray 1893), many of the specimens are “mere bits,” small vegetative fragments, single leaves or single flowers, often wholly unidentifiable from photographs and scarcely less so by direct examination without painstaking comparison to known materials. Signifi- cant efforts to identify more than single genera by direct examination are few. Hitchcock (1905) reviewed Walter’s grasses and matched many of their names with Fraser's speci- mens. Blake (1915) discussed 22 Walter names and their corresponding specimens and made new combinations where indicated. Fernald & Schubert (1948), working from pho- tographs, identified—not always correctly—19 specimens in the herbarium. Others have examined the species of individual genera and published their conclusions. Still others, notably Oakes Ames with the orchids, and Asa Gray and David H. Rembert with large parts of the entire herbarium, have given significant time to identifying the specimens, but have left no record other than their annotations and notes. At the present writing perhaps one third of the specimens has been given a credit- able identification, one third a tentative judgment, and one third no more thana cautious assignment of genus. Complete identifications must await opportunity for study by per- sons who have simultaneous access to the Fraser specimens and to a convenient refer- ence collection. 1116 BRIT.ORG/SIDA 22(2) Only two of the 690 specimens bear a date on the original label. Spm. 78-E, probably Platanthera flava, is atypically instructive: “Orchis / found near the / Table Mountains / llth June / 1787 /JE” The hand, confirmed by the initials, is that of John Fraser. Spm. 100- F, Gillenia trifoliata, is briefly informative though lacking the year of collection: “16 June / Indian Emetic,” written by Fraser, followed by “Spiraea trifoliata,” written by Walter. Equally few specimens bear a geographic source on the original label. Only two la- bels give locations; both are in Fraser's hand. As noted, spm. 78-E, probably Platanthera flava, was collected near the “Table Mountains,” perhaps Table Rock, Pickens Co., South Carolina (less likely: Table Rock, Burke Co., North Carolina). Spm. 67-A, Lupinus villosus, was labeled “Blue Flowering / Lupinus / Pilosus / Found on y\e / Border of y\e / Altamayhaw / J.E” This of course is the Altamaha River, which flows through southeast- ern Georgia. The first of these locations is about 360 km northwest of Fraser’s base in Charleston; the second is about 240 km to the south. The distance between these loca- tions, together with the certainty that these summer-flowering species must both have been collected in 1787, isa measure of John Fraser's energy and dedication. Most of the specimens (624, or 91%) are labeled with an identification—a genus and species, or a genus only. 328 (48%) of the specimens are fully identified (to genus and species), by either Walter or Fraser or by both (where Walter wrote the genus and Fraser added the epithet). Nearly half of the specimens (296, or 43%) are identified only to ge- nus, 183 of them (27%) by Walter, 113 (16%) by Fraser. 64 (9%) of the specimens bear la- bels, but were not identified. (ea ped ~ an the identification. Walter frequently appeared not to recognize the plant, but wrote a brief Latin diagnosis which reflects his familiarity with Linnaeus’ sexual system of classifica- tion. Some examples: 4-C, Ptilimnium capillaceum—“Ammi majus sp. nova?”, 55-C, Desmodium cuspidatum-“Hedysarum Flore magnus”, 60-E Silene stellata—“ignota”:; 61b- B, Iva imbricata-“lva ? nova”; 67-D, Crotalaria rotundifolia““Lupinus affinis”; 75-F. Nelumbo lutea-“The Great Nymphaea’; 86-B, Myriophyllum pinnatum-“Potamogeton monoicum”; 87-D, Physostegia virginiana-“Prasium nov”, 89-A, Cynanchum angusttifolium—“PentandIria] Digynlial bifollic”; 90-B, Quercus michauxii-“Quercus nova’: ae Brunnichia ovat d-“Rajania (monoica)” 97-C, Bumelia lanuginosa-“fol. angustis ”98-D, Smilax laurifolia-“Smilax Baccis albidis”, 106- E, Phyla nodiflo ra-“Verbena nodiflora Didynamia gymnosperm disperma’”, 108-D, Viola villosa-“Viola Fol. pubescent”; 114-E, Juncus effusus-“an Juncus”; 117-A, Decodon verticillatus-“Decand monogyn nov G. aquatic”; 117-B, Cynoctonum sessilifolium—“Genus nov Pentand digyn Fraser’s distinctive script usually appears to be of observations made of the plants in the field. When he ventures into Latin he is clearly in unfamiliar country. Britten (1921) gently noted Fraser's limited linguistic skill where “nova genera” was used “asa singular,” trusting his classic-trained readers to understand. Some of Fraser’s notations: 1-D, Vibur- num dentatum-‘not in y\e Flora”, 11-B, Asclepias sp.-“Dove Coloured”; 12-G, Aster concolor-“Aster Dark Blue Flower”, 22-C, Psoralea canescens—“Buchnera americana na- tive name Buck root”, 38-G, Croton punctatus-“Croton Maritimum Nova”, 52-1, (unidenti- fied)-“Granadilla not in y\e flora”, 62a-A, Aesculus parviflora—‘Juglans Alba nova”; 67- A, Lupinus villosus-“Blue Flowering Lupinus Pilosus Found on y\e Border of y\e Altamayhaw J.F”; 67-C, Lupinus perennis~“Lupinus Perrenis [sic]; 72-A, Crataegus crus- galli-“Mespilus Large Cock Spur”; 72-B, Crataegus uniflora-“Mespilus Small Cock Spur’; The labels of occasional specimens carry comments or information other t WARD 1117 92-K, Rhododendron minus-“Leaves like y\e Kalmia? Latifolia grows on y\e Mountain in decid. y\e Est\n & Wes\n Waters”, 93-C, Salix humilis-“Salix Minor? of Fraser”, 100-K, Arenaria uniflora-“No Name”; 100-F, Verbena tenuisecta-“Nova Genera”, 108-C, Viola primulifolia-“Viola White Flowers Downy Leaf”, 1L0-A, Vitis labruska-“Vitis? Labruska Fox Grape”; 116-B, Eriogonum tomentosum-“Sophoranthus nov genera”; 117-C, Lachnanthes caroliniana-“Nova Genera”; 117-E, Carphephorus sp.-“Genera Nova.” Comments ona few labels are not in Fraser’s hand nor in Walter’s, and show involve- ment of a third person: 8-C, Stylosanthes biflora-“Arachis ground nut”, 12-F, Aster sp.- “Aster grows 16 feet high”; 46-C, Conoclinium coelestinum—‘Bright violet blue: said to bea specific for the venereal complaint”; 105-C, Vaccinium arboreum-“A Vaccinium 5 miles from Cranberry measured 50 feet high”; 115-G, Planera aquatica-“Monoecia Triandria G. nova.” The handwriting on these last-cited labels, and a few others, poses a puzzling, unre- solved question. The content of most suggests they were written at the time of collection. Their subject matter is wholly different from the technical comments recorded by Walter. And the hand is quite incongruous with that of Fraser sr. The “square r” of some (8-C, 12- EF 27-D, 38-J, 46-C, 105-C) would seem to be that of Fraser fil. Yet John Fraser, the son, did not accompany his father to the Americas until 1800 (Hogg 1852), more than a decade after Fraser's 1786-1788 trip, and eleven years after Walter’s death. Could these notations have been made by Fraser fil. on his fatlier S ee after ey were brought to England? Or is it possible these specimens (and d,and annotated, by Fraser fil. on his 1800 trip to the prcienien: with his father? The labeling of one species confirms an involvement of a third person in identifica- tion and possibly a fourth in mounting of the specimens. A mint, Trichostema dichotomum, is represented by two collections, 61b-D and 103-F Both are labeled with its name. The label of 103-F is in Walter’s hand, and is spelled conventionally. But 61b-D is in another hand, neither Walter's nor Fraser’s. The style of the capital “T” is so different that the mounter of the specimens, intending to arrange them in alphabetical order (as was done with 103-F), but seemingly misreading the convoluted script of the initial letter, has placed 6lb-D between “Iva” and “Illicium.” Further, the writer of that label transcribed its epi- thet as “dichotoma,” suggesting one of sufficient education to perceive an apparent error of gender mismatch between epithet and genus, yet without adequate classic background to understand that “-stema” is a neuter root. Even more, the mounter himself must have been of limited experience not to have recognized the distinctive specimen as one al- ready found elsewhere in the folio. With certainty, Fraser sr. neither formed this label nor was involved in its mounting. One specimen is wholly aberrant. Spm. 102-A isa branched structure mounted alone ona full page. It is a gorgonian (Gorgoniidae-coral, s.L.). Enlarged bodies on its branches are barnacles (Archaeobalanidae; Conopea sp.). It was labeled “Sea Plant’ in Fraser’s hand. The physical processing of the specimens into the present folio remains poorly un- derstood. Walter, of course, was not involved. Fraser, perhaps soon after his return to En- gland in March 1788, must have acquired the empty folio and, as indicated by his writing on the title page, established its goal. It is often forgotten that the present herbarium of 690 plants is only a subset of the “upwards of thirty thousand dried specimens of plants” Fraser (1789) claimed to have gathered during his 1786-1788 trip to the Americas. An unknown number, perhaps the greater part, were sold by Fraser (1789) to Charles Louis LHeritier, a wealthy French botanist, and are now in the Lamarck herbarium, Paris. The 1118 BRIT.ORG/SIDA 22(2) basis is unknown by which Fraser, or his sons during his absence on later trips to the Americas, Cuba, and Russia (Hogg 1852; Simpson et al. 1997), selected from this larger collection the plants to be retained in the “Walter Herbarium.” The historic importance of this early sampling of the plants of Georgia and the Caro- linas is unquestioned. America is fortunate the fates have preserved this fragmentary glimpse of its vegetation as it appeared before the full impact of modern civilization. Even so, the larger value of the John Fraser folio and the specimens it contains lies in the degree it supports the writing of Thomas Walter and his Flora Caroliniana. The nomen- clatural basis for Walter's work and its relation to Fraser’s herbarium is to be discussed in subsequent reports of the Thomas Walter Typification Project. ACKNOWLEDGMENTS [remember with gratitude the courtesy shown my wife and me in 1984 by N.K.B. Robson, Mike Mullin, and other members of the staff of the Natural History Museum, London, and especially the salutation given me by William T. Stearn with his pleasure in meeting the “Anonymos botanist.” | have been immeasurably aided by the Bernice Schubert pho- tos whose negatives were loaned me in 1977 by Carroll E. Wood, Harvard University. I have been given assistance in matters of history, linguistics, and taxonomy by Charles E. Jarvis, Natural History Museum; Alexander Krings, North Carolina State University; James S. Pringle, Royal Botanic Gardens, Hamilton, Ontario; David H. Rembert, University of South Carolina; and Robert L. Wilbur, Duke University. REFERENCES Buake, S.F.1915. Some neglected names in Walter's Flora Caroliniana. Rhodora 17:129-137. Britten, J. 1921. Thomas Walter (1740?-88) and his grass. J. Bot. 59:69-74. Coker, W.C. 1910.A visit to the grave of Thomas Walter. J. Elisha Mitchell Sci. Soc. 26:31-42. Ewan, J. 1979. Addenda on Thomas Walter (c. 1740-1789). Taxon 28:345-346. Faben, R.B. 1989. Commelina caroliniana (Commelinaceae): A misunderstood species in the United States is an old introduction from Asia. Taxon 38:43-53. FerNatp, M.L.and B.G. ScHuserr. 1948. Studies of American types in British herbaria. Part IV:some spe- cies of Thomas Walter. Rhodora 50:190-208, 217-229. Fraser, J. 1789. A short history of the Agrostis Cornucopiae: or, the new American grass. Chelsea, England. Gray, J. L., ed. 1893. Letters of Asa Gray. Boston. HiicHcock, A.S. 1905. The identification of Walter's grasses. Missouri Bot. Gard. Ann. Rep. 16:31-56. Hoc, R. 1852. Life of John Fraser. Cottage Gardener 8:250-252. Maxon, W.R. 1936. Thomas Walter, Botanist. Smithsonian Misc. Collect. 95:1-6. RemBert, D.H. 1980. Thomas Walter, Carolina botanist. South Carolina Mus. Comm., Bull. No. 5:1-33. Simpson, M.B., S. Moran, and S.W. Simpson. 1997. Biographical notes on John Fraser (1750-1811): plant nurseryman, explorer, and royal botanical collector to the Czar of Russia. Arch. Nat. Hist.24:1-18. Smait, J.K. 1935. Thomas Walter's botanical garden. J. New York Bot. Gard. 36:166-167. Walter, T. 1788. Flora Caroliniana. London. Warb, D.B. 1962. The first record of the Fraser Fir. Castanea 27:78-79. Ward, D.B. 1977. Nelumbo lutea, the correct name for the American lotus. Taxon 26:227-234. — EVIDENCE FOR HYBRIDIZATION BETWEEN TWO SYMPATRIC VIOLET SPECIES, VIOLA GRAHAMII AND V. HOOKERIANA (VIOLACEAE), INCENTRAL MEXICO Aurea C. Cortés-Palomec'! Harvey E. Ballard, Jr. Department of Environmental and Plant Biology Department of Environmental and Plant Biology i iversity hio University Athens, Ohio 45701, U.S.A. Athens, Ohio 45701, U.S.A ac399295@ohio.edu ballardh@ohio.edu ABSTRACT Viola hookeriana and V. grahamii (Violaceae) are closely related species in section Viola, subsection Mexicanae occurring in POENTAO Ue habitats across much of central Mexico. In various locations, these species grow sym many individuals with intermediate morphological enasctes has ee jah eee and een between these species. Using a combination of nao POE ecological, and Oeulg evidence we evaluated two pmined populalions of V. ce ahamiiand V.} Lake Patzcuaro in the state of N I tential of detec hiv bud morphologies, the strength of phenologi- cal differences and ecological Re aen among the species and putative hybrid, and to Saini the level of gene flow between the two species. Our results indicate that hybridization is equ and that hybrids are morphologically distinct from the parental taxa. Pre-mating isolation is weak and has favored extensive hybrid- ization due to largely overlapping blooming times, weak ecological isolation and absence of spatial isolation. Hybrids are morphologically and genetically closer to Viola grahamii, but do show unique alleles as well. yy, Pp g y g y 8 q RESUMEN AA rr J | Viola hooke iy Vgral ii (Viol ) | pecies f la Subseccién la Seccion Viola del género Viola. E | | México y la presencia de individuos con eee nent ee ha sugerido la existencia de hibridizacion entre las especies. Usando una combinacion de datos morfolégicos, ecolégicos y moleculares evaluamos dos eee mixtas de V.grahamii y V. hookeriana en i. montanias cercanas al lago de Patzcuaro en el estado de Michoacan en México para determinar la presencia de morfologias caracteristicas a los hibridos, el grado de diferencias fenoldgicas y ecologicas entre las especies y el nivel de flujo génico entre ellas. Nuestros resultados indican la presencia de hibridos y que estos son mor sol Opicamente sae a bi boos Los mecani le aislamiento + 1] ]} entre las especies parenta tales noson muy f fuertes y tiempos de floracién similares, muy poc ient logico y la ausencia de separacion espacial entre las especies. Los hibridos son eee y genéticamente mas similares a Viola grahamii, pero muestran aleleos que son unicos a ellos. Hybridization isa common phenomenon in nature, as evidenced by an estimated 70,000 natural interspecific plant hybrids worldwide (Rieseberg & Ellstrand 1993; Judd et al. 1999), and the fact that between 16 and 37% of the plant families reported in different floras contain at least one hybrid taxon (Ellstrand et al. 1996). In some cases, hybridiza- tion can lead to the creation of hybrid species and/or intr ogressants between hybrid de- rivatives and the parental taxa (Arnold 1992; Rieseberg 1995). The genus Viola L. (Violaceae), which comprises about 525-600 species (Clausen 1964; Ballard et al. 1999), is well known for its taxonomic problems due to hybridization and introgression, as well as complex patterns of variation in individual traits (Brainerd 1924, Anderson 1954; Rus- 'Current Address: C Ecosistemas, Uni idad Naci | tc Je Méxic , Antigua Carretera a Patzcuaro No. 8701, Col. Ex-Hacienda de San José de la Huerta, Morelia, 58190 Michoacan, MEXICO, SIDA 22(2): 1119- 1133. 2006 1120 BRIT.ORG/SIDA 22(2) sell 1955; Ballard 1994; Krahulcova et al. 1996; Gil-ad 1998; Neuffer et al. 1999; Marcussen & Borgen 2000; Calderon de Rzedowski 2001; Harmaja 2003). Analysis of herbarium specimens from central Mexico has indicated that hybrid- ization is likely occurring among species belonging to Viola subsection Mexicanae (Bal- lard 1994). Viola subsection Mexicanae W. Becker (sensu Ballard et al. 1999) is a mono- phyletic group consisting of eight stoloniferous or rosette-forming species: Viola hemsleyana G. Calderon, V. grahamii Benth., V.oxyodontis H.E. Ballard, V. humilis H.B.& K., V. hookeriana H.B.& K,, V. beamanii G. Calderon, V.guatemalensis W. Becker, V.nannei Polak. (Ballard & Sytsma 2000; Ballard et al. 1999), and an additional undescribed spe- cies (Ballard, per. obs.). Instances of hybridization have been identified from several lo- calities in central Mexico occurring between Viola grahamii and four other members of the group, V.guatemalensis, V. hemsleyana, V. hookeriana and V. humilis. Occurrences of these putative hybrids appear to be restricted however to the immediate areas where both parents occur sympatrically. Viola grahamti (known in Mexico as “hoja de pasmo” or “pensamiento del cerro”) is a frequent to locally abundant perennial distributed across much of Mexico. Specifically, it occurs in mountainous regions from northern Mexico south to northern Guatemala, and is one of the most widely distributed members of the subsection Mexicanae (Fig. 1). It thrives in dry to mesic sandy loam under varied forest canopy and along stream banks at elevations of 1950-3600 meters (Ballard 1994; Calder6n de Rzedowski 2001), and often forms large mats connected via aboveground stolons. It produces chasmogamous flow- ers at the beginning of the rainy season June) for approximately four weeks and cleisto- gamous flowers for several months afterward June-December) until the onset of the dry season (Cortés-Palomec 2005). As previously mentioned, one of the four species with which Viola grahamii appears to hybridize is V. hookeriana (known in Mexico as “violeta”). Viola hookeriana is an infre- quent to locally common perennial generally growing in small isolated populations across northern and central Mexico (Fig. 1). It grows in mesic loam under mixed deciduous and coniferous forest canopy at elevations between a 2500 meters oe 1994; Calderon de Rzedowski 2001). Reproduction occurs via both ¢ d clei flow- ers and follows a similar pattern to that seen in V.grahamii (Cortés- Palomec, pers. obs.) Viola hookeriana and V. grahamii are divergent in several characters: growth habit, flower structure, and foliage, and can be separated without difficulty in both herbarium specimens and living populations (Table 1) (Ballard 1994). The putative hybrids have been identified in areas of overlapping distribution of the two species (Fig. 1), and are recog- nizable as exhibiting intermediate morphologies (Ballard 1994). The goal of our study was to assess the presence and extent of hybridization between V. grahamii and V. hookeriana in natural populations, and identify intrinsic pre-mating isolation mechanisms that may be functioning between the two species. We examined morphological and phenological differences between the two species, characterized the ecological factors governing where individual species and putative hybrids occur, and sur- veyed genetic patterns to identify gene flow between species in two study sites in central Mexico where V. grahamii and V. hookeriana intermingle extensively. MATERIALS AND METHODS Site Establishment and Sampling Two study sites were established in the mountains to the north of Lake Patzcuaro in the CORTES-PALOMEC AND BALLARD 1121 A Viola grahamii A Viola hookeriana © V. grahamii x hookeriana Study sites Su Northern Michoacan os 0 500 Km Fic. 1. Distribution of Viola grahamii, V. hookeriana, and V. grahamii < hookeri Mexi 1 I la. Distribution from Ballard (unpublished data). municipality of Quiroga, Michoacan, Mexico (Fig. 1). The sites were established in early June 2000 and observations were made weekly over a period of five weeks during the summer rainy season when chasmogamous flower production was most extensive. The two specific sites were chosen due to the presence of abundant populations of the two species and their putative hybrids as identified through a survey of the area conducted earlier in the spring. The first site (Site A) [19°41'57" N; 101°32'27" W] was situated on the lower southern slopes of Mt. Zirate, above the town of Santa Fe de la Laguna. The second site (Site B) [19°42' N; 101°35' W] was located 8 km to the west of Santa Fe de la Laguna (Km 48 on the road from Quiroga to Zacapu (MX-15)). While both sites were located in superficially similar environments, they differed in the level of disturbance and abun- dance of violet species. Viola grahamii was abundant in both sites, but areas of visually pure V. hookeriana separated from V. grahamii were found only in Site A. In Site A, four 20 m long transects were established. Two transects were arbi through the middle of a visually “pure” population of the parental species (A-T1VG ina V. grahamii population and A-T4VH ina V. hookeriana population). Two additional transects (A-T2HY and A-T3HY) were placed through areas where putative hybrid plants inter- mingled with the parental taxa. Ten quadrats (0.5 m2) were randomly established along each transect with position determined using a random number table. In Site B the same procedure was followed except that only three transects were positioned resulting in a to- tal of 30 quadrats (B-T1VG for V. grahamii, B-T3HY of mostly V. hookeriana and a mixture of V.grahamii and hybrids, and B-T2HY consisting mostly of putative hybrids). ily placed 1122 BRIT.ORG/SIDA 22(2) Taste 1.Comparison of morphological characters used to distinguish between Viola grahamii and V.hookeriana (on the basis of Ballard 1994). The characters of hybrids are not included since they are not consistent (i.e, hybrids show varying combinations of traits, making them difficult to characterize). Character Viola grahamii Viola hookeriana Stolon position above ground underground Stolon nodes present absent Stipule adnation free or adnate free Pubescence on petioles and present [abundant] absent abaxial surface of leaves Leaf shape ovate-oblong to elliptic broadly ovate to reniform Length of pedicels (cm) 3-12 5-21 Calyx pubesence ciliate glabrous Corolla color white white, sometimes purplish Elevation 1950-3600 m 1700-2500 m For transects representing “pure” parental taxa at each study site, three plants that were not connected via stolons were randomly selected in each of the 10 quadrats. One leaf was removed from each of the three plants and dried in silica gel for later DNA ex- traction. A second leaf, the largest of each individual, was collected and labeled to corre- spond to the DNA sample. This leaf was pressed and used for subsequent morphological analysis. The same methodology was used for plants in the putative hybrid/mixed-taxon transects, except that 5 individuals were sampled per quadrat. Phenology and Pollinator Visitation In each of the 70 quadrats (40 from Site A, 30 from Site B), the number of individuals present per quadrat and the number of individuals with open chasmogamous flowers were recorded weekly during the five weeks of field observations to interpret phenology. The percentage of flowering individuals, those bearing chasmogamous flowers at some point during the study, was calculated for each visit for the two species and for the puta- tive hybrids to allow for a comparison of phenology among the parental taxa and puta- tive hybrids at both sites. ae with more than one taxon were recorded separately for each taxon and analyzed as te observations. A Kolmogorov-Smirnov two-sample test was performed in NCSS (Hintze 1999) to compare phenological patterns among taxa at each of the two sites. Pollinator visitation was assessed during weekly visits to Site A, which was the only site to have clearly differentiated populations of V.grahamii and V. hookeriana. A total of 40 hours of direct observations of the flowering individuals were recorded with special emphasis made to observe instances of pollinator movement between or among differ- ing taxa. The hours of pollinator observation were equally divided between two daily periods relating to potential insect activity based on the times when previous observa- tions in the area suggested the potential pollinators would be more active, between 8:00 and 11:00 am, and between 12:00 and 3:00 pm. as Ecological Characterization To determine if the taxa grow in modally different microhabitats, infer whether ecologi- cal isolation might reduce gene flow between the species and the putative hybrids, and understand the local distribution of putative hybrids relative to the parents, an ecologi- cal characterization of microhabitat in the 70 quadrats was conducted. Soil N, P, K con- CORTES-PALOMEC AND BALLARD 1123 centrations and pH were evaluated using a LaMotte™ Combination Soil Test Kit (LaMotte, Chestertown, MD). Values were given in exact units for pH while values for N, P and K were determined via a colorimetric test procedure from which concentration could be estimated within a set range, 5-75 ppm for N, 5-100 ppm for P, and 50-200 ppm for K. Due to the inability of obtaining exact concentration values for N, P, and K, these values were range standardized and analyzed in rank form. Percent soil moisture was recorded using the gravimetric method described by Hadley and Levin (1967), with the modification that soil mass was determined after one week of air drying. Light availability as a direct means of inferring canopy closure was measured using a Model-C spherical densiometer (Vora 1988). Quadrats with more than one taxon were recorded separately for each taxon and analyzed as separate observations. Data were standardized and an analysis of variance was performed in NCSS to test for differences among V. hookeriana, V.grahamii and putative hybrids. Morphological Analysis From the pressed leaves, measurements were taken of the 0°, 30°, 60°, 90°, 120°, 150° and 180° radii, following the procedure outlined by Ballard and Wujek (1994). The center of the leaf was considered to be the point along the midrib opposite the widest part of the leaf blade. Presence or absence of leaf pubescence on the abaxial surface of the leaf, a characteristic considered diagnostic for separating V. hookeriana (glabrous) and V. grahamii (pubescent), was also recorded. Measurements were standardized and a dis- criminant analysis (Manly 1994) was performed, both including and excluding visually identified putative hybrids to determine overlap in leaf characters among the two taxa and putative hybrids. The analysis was performed in NCSS using an automatic variable selection procedure. Measurements at 30° and 120° were excluded from the analysis due to colinearity of the data. The canonical scores generated from these analyses were used in an analysis of variance (ANOVA) with a Fisher’s LSD pairwise comparison analysis (Zar 1996). Genetic Analysis To test the presence of genetic differences among the parental taxa and identify potential gene flow within the putative hybrids, biparentally inherited nuclear markers, inter- simple sequence repeats (ISSRs) were used. ISSR data have proven to be highly effective in detecting hybridization and/or gene flow among closely related species, and to test the hybrid speciation hypothesis (Wolfe et al. 1998a, b, Wolfe & Randle 2001; Archibald et al. 2004; James & Abbott 2005). Additionally these markers are useful in studies of differentiation among closely related species for they are able to utilize some of the vari- ability present at microsatellite loci without the need to develop species-specific microsatellite primers, an advantage when working with species which have not been well-studied genetically. Genomic DNA was extracted from the silica gel-dried leaves using a Wizard Genomic DNA purification kit (Promega, Madison, WD). Five different ISSR primers (McCauley & Ballard 2002) were initially screened to identify primers which would produce consis- tent and scorable polymorphic bands among the taxa. Two primers were selected: Wolfe #99B [(CA)6GG] and HB#15 [(GTG)3GC]. These were used for amplification of ISSR prod- ucts in replicated 25 ul reactions consisting of 1 wg wl! diluted genomic DNA, 19 pl auto- claved distilled water, 2.4 n110x PCR Buffer (Gibco BRL), 2 41 MgCl2 (50 mM, Gibco BRL), 2 ul of dNTP mix (10 mM, Fisher), 0.5 ul of BSA (Bovine Serum Albumin, 4 ug pl Fisher), 1124 BRIT.ORG/SIDA 22(2) 0.25 wl primer and 0.25 wl of Taq polymerase (5 U 11, Gibco BRL). The polymerase chain reaction was performed in a Stratagene RoboCycler 96 with hot-top (Stratagene Inc, La Jolla, CA) and programmed for 2 min. at 94°C; 40 (primer Wolfe #99b) 44 (primer HB #15) X 30 sec. at 94°C, 45 sec. at 44°C, 1 min. 30 sec. at 72°C; 20 min at 72°C. PCR products were electrophoresed in a 1.3% agarose gel in 0.5 x TBE buffer with flanking 250 bp ladders (Gibco BRL). Gels were stained with a solution of ethidium bro- mide in 0.5 x TBE buffer for 20 minutes and imaged under UV light. Gel images were analyzed with BioMax ID image analysis software (Version 2.0.3, Eastman Kodak Com- pany, Rochester, NY) to identify and size fragments. Fragments were scored as present (1) or absent (0), with fragments comigrating at identical rates (+ 10 base pairs, the general limit of resolution for agarose gels) considered equivalent. Statistics concerning fragment occurrence, including total fragment number, distribution across taxa, polymorphic and fixed fragments, and number of fragments shared among taxa per site were calculated using TFPGA 1.3 (Miller 1997). Analysis used three discrete methods to evaluate relationships and similarity among the taxa and putative hybrids. UPGMA cluster analysis and Principal Coordinates Analy- ses (PCoA) were performed both within and between the two sites using NTSYS ver. 2.02} (Applied Biostatistics Inc.). An Analysis of Molecular Variance (AMOVA) was addition- ally performed ina hierarchical fashion among species and sites using WINAMOVA 1.55 (Excoffier 1993). All analyses were performed according to the methods described in McCauley and Ballard (2002). RESULTS Phenology and Pollinator Visitation The parental spec d divergent but non-significant tendencies in blooming time; in fact, in Site A V. hookeriana bloomed first, whereas in Site B it bloomed later (Fig. 2). The Kolmogorov-Smirnov two-sample test showed no significant differences among the blooming times of the hybrids and the parental taxa in either site (p values = 1 in some pairwise comparisons) (Fig. 2). No pollinators were observed visiting violets at any time during this study. Ecology All three taxa inhabited modally different microhabitat conditions influenced by light and certain nutrients, in all cases where there were significant differences between the taxa, the hybrids grew in intermediate conditions to the parental species. The ANOVA of the ecological factors revealed significant differences in light (p= 0.02), P (p = 0.04) and K (p = 0.03) among V. hookeriana, V.grahamii and the putative hybrids; moisture (p= 0.32), pH (p=0.86) and N (p= 0.62) did not differ significantly. Viola hookeriana grew in higher concentrations of P, lower concentrations of K and more shaded environments than V. grahamii (Table 2). Leaf Morphology Two hundred and eighty eight individuals were classified into three groups correspond- ing to the two parental taxa and the putative hybrid individuals (Fig. 3). The discrimi- nant analysis indicated three marginally distinct groups (Fig. 4), with the GLM ANOVA and the Fisher’s LSD Multiple Comparison analysis indicating that they were well-sup- ported and significant (p < 0.05) groups. CORTES-PALOMEC AND BALLARD % of flowering individuals % of flowering individuals 70 Site A V. hookeriana AT4 KZA hybrids AT2 60 + hybrids AT3 SS V. grahami AT1 “| i. AT1 AT2. _AT3 me AT2 77 AT3 1 1 AT4 77 22 HE 40 + SO" 20 + i : ] 0 md 4 July 6 July 12 Visits 70 . Site B (—] V hookeriana BT1 V-ZiZA hybids BT2 60 4 RSs9_:V grahami BT3 BT1._ BT2 50 - BT2 .873 BT3 1.873 40 4 30 4 20:4] 1c 0 ; iA June 25 ! July 12 July 18 Fic. 2. Histogram of phenology for fl 1125 J t Vu Vb L als sampled are as f rs JTHITTOY ollows (num twU ber of reproductive individu VG(AT4)=146(77). Site B VG=35(19); HY=95(55); VH =142 (33). Ve. FIUURCHIGH als): Site A: VH(AT jana, VG=V.g 1)=24(12); HY(AT2)=151(76); HY(AT3)=232(134); 1126 BRIT.ORG/SIDA 22(2 Taste 2.Means and SE for < Se. lected eG ologic al parameters aMONng V. hookeriana (VI 1), \ ahamii (\ G) a and h ybr ids (HY). Superscript letters indicate a significant difference for a particular parameter among the taxa. Moisture light are expressed as percents. P N and K values are along a scale of | to 7 (l=very i 7= high). anc Taxon Moisture Light pH P N K VH 26(+£0.1) 16(£0.4)" 6.40(+0.14) 5.32(£0.49) 1.28(+0.10) 6.44(+0.16) VG 24(+1.1) 29(£0,3)’ 6.51(#0.12) 3.82(+0.37)° 1. 14(+£0.09) 7, O(+0.13)° HY 25(+0.1) 29(£0.3) 6.43(+0.10) 4.40(+0.33) 1.18(+0.08) 6.81(+0.12) Genetic Data A total of 242 Viola individuals were examined genetically with ISSRs: 33 Viola hookeriana in Site A and 19 in Site B, 65 V. grahamii individuals in Site A and 35 in Site B, and +8 putative hybrid individuals in Site A and 42 in Site B. In Site A, 70 ISSR fragments were resolved vs. 53 in Site B. The majority of these were identical between the two sites, resulting in a total of 75 unique and scorable fragments for the study sites together. The relative frequency of individual fragments ranged from less than 10% of individuals (rare to infrequent) to 100% (ubiquitous). Distribution of fragments across taxa (Table 3) showed that some were specific to one of the parental taxa (e.g. five fragments were species-spe- cific for V.grahamii,and three were specific for V. hookeriana) whereas others were shared among taxa. Most of the shared fragments occurred between V. grahamii and the puta- tive hybrids (e.g., 17 fragments in Site B). Nine fragments were specific to the putative hybrids. UPGMA cluster analysis (Fig. 5) showed strong separation of the two study sites. All individuals diverged substantially between Site A and Site B, regardless of taxon. The PCoA confirmed these discrete site differences (Fig. 6A). Additional PCoA analyses were carried out separately on the two sites [Site A (Fig. 6B) and Site B (Fig. 6C)]. In both sites, V. grahamii and the putative hybrids showed the most extensive genetic overlap. In Site A, V. hookeridana mostly segregated from the other taxa, although some individuals clus- tered with putative hybrid derivatives, and others with V. grahamii. In Site B (Fig. 6C) two distinct groups of V. hookeriana were evident, with some putative hybrids occurring within each of them. In all cases, putative hybrid individuals were widely placed and substantially overlapping in distribution with the parental taxa, together expressing a greater level of genetic diversity. The AMOVA (Table +) showed that most of the variation was within morphologi- cally defined taxa (69%), although significant differences were detected between sites and among taxa. Variation between sites accounted for 27% of the total variance. Differ- ences among parental taxa and hybrids accounted for only 3.72% of the variance. The same pattern was observed when the two sites were analyzed separately. In Site A, 94.5% of the variance was due to variation within the taxa; 95.3% in Site B. If the hybrids were eliminated, variation was slightly reduced within taxa (90.32% and 89.17%); however, lev- els of variation within the species themselves remained high. DISCUSSION Our results show conclusively that hybridization is occurring between V. grahamii and V. hookeriana. Phenetic analysis of leaf shape characters in the species, more exploratory than exhaustive, showed separation among V. hookeriana, V. grahamii and the hybrids. Hybrid individuals formed a distinct group from the parental taxa suggesting that leaf 1127 CORTES-PALOMEC AND BALLARD iu hookeriana, b Fic. 3. Rey Axis 1 PET] Oa ee Pe en Pa Pee \ i‘ a) s 4 -t “ ~ al —d So sc Pa oO pes L=3) = S = = xg cs YY a ba) =) ae = 1 by tk rs = trea g L bh ie re ep os, ee Pe! L Jk 1 J \ 1V.g LL ii x hookeriana r J (gray dots). 1130 BRIT.ORG/SIDA 22(2) Fase 4. Analysis of Molecular Variance (AMOVA) of 242 individuals collected from both sites (A and B), based on 2000 permutations. Mean squares (MDS), variance component and percentage of variance as well as P values are presented. A nested design was performed. Source of variation df MDS Variance component Percentage p Between sites | 286.41 2.30 26.79 < 0.0005 Among taxa 4 18.38 0.32 Se < 0.0005 Within taxa 236 5.98 5.98 69.49 < 0.0005 24) more likely that hybridization and subsequent gene flow have proceeded to the point that many morphologically “pure” parental individuals are actually products of extensive gene flow, bridging the gap between parents and hybrids and obscuring taxon boundaries. Additional genetic evidence suggesting that the hybridization between V grahamii and V. hookeriana has likely been occurring over a long period of time is the presence of ISSR fragments unique to the hybrids. It has been suggested by various authors that later- generation backcrosses might contain only a small number of the fragments from the parental species (O'Hanlon et al. 1999). If this is the case, the development of fragments that are “specific” to the hybrids are best interpreted as the result of later generation re- combination and long-term post-hybridization genetic reorganization, as suggested by O'Hanlon et al. (1999). Studies of introgression in German violets conducted by Neuffer et al. (1999) using RAPDs describe a similar phenomenon in the V. riviniana Rchb—V. reichenbachiana Jord. ex Bor. complex, but the fact that no unique hybrid bands were found led them to conclude that the hybrid complex hada recent origin. Directional back- crossing has also been suggested towards V. lutea subsp. sudetica (Willd.) Nyman in hy- brids of this species and V. tricolor L (Krahulcova et al. 1996). In this study, the presence of many “hybrid-specific” bands suggested a significant time component in the forma- tion of hybrid derivatives as well as differentiation in some. Our results suggest that in- trogression is occurring between V. grahamii and V. hookeriana, and that hybridization might have taken place thousands of years in the past or may occur only rarely, but high evels of interfertility and appropriate site conditions have allowed the hybrids sufficient time to backcross with the parental taxa. Hybridization has likely been favored by the absence of a temporal reproductive iso- lation between the two species. Despite a minor, non significant, shift in blooming times of the two species (between populations) they have greatly overlapping blooming times, which provide extended opportunities for inter-taxon crossing. This small shift in bloom- ing may be related to differences in microsite characteristics and levels of human pertur- bation such as wood cutting for local uses, which were more pronounced in Site B than in Site A. The differences in canopy openness between the two sites due to human distur- bance can also explain why V. hookeriana was not common in Site B, while V. grahamii was very abundant. Open environments with more light favor V. grahamii, which grows best in open-canopy forest or open sites while V. hookeriana is most abundant in more shaded environments, mainly under the forest canopy. Other differences were found in terms of soil requirements; V. hookeria nd SrOWs in higher concentrations of Pp and lower concentrations of K than does V. grahamii. Soil nutrients in the area however are highly heterogenous (Cortés-Palomec 2005), and while they may be affecting the distribution —" — CORTES-PALOMEC AND BALLARD 1131 of V. hookeriana and V. grahamii, the hybrids appear to survive in various soil condi- tions, accounting for their wider local distribution in each site, especially in areas with moderate to heavy levels of disturbance. It is commonly suggested that since hybrid in- dividuals have partial genotypes from both parental species they can potentially grow in both parental habitats as well as in their own “intermediate” niche and even lead to the extinction of the parental taxa if one of them is rare (O'Hanlon et al. 1999; Levin et al. 1996). In fact, hybrids between V.tricolorand V. lutea subsp. sudetica have been suggested to have a wider range of distribution than both parental species (Krahaulcova et al. 1996). In addition, disturbance can lead to “hybridization of the habitat” (Anderson 1954; Rieseberg & Ellstrand 1993), resulting in a mosaic of microhabitat conditions that favor hybrids and in extreme cases may cause the local extinction of one or both of the paren- tal taxa (Rieseberg & Ellstrand 1993; Rhyme & Simberloff 1996). Hybrids in this case can be compared to alien plants in the sense that they are new elements in the flora that can threaten the original flora of the region (Neufer et al. 1999). Perturbation was high in both study sites, mainly due to human activity (ie. livestock grazing and timber har- vest). If we consider our sampling of the study sites as representative for the region, then the hybrids may be said to be as abundant or even more abundant locally in the zone of sympatry than the parental taxa. The heavy genetic overlap of hybrids with V.grahamii may threaten that species with local genetic swamping, but since it is widely distributed and occurs as pure populations across large regions of Mexico and northern Guatemala that is probably not a problem. Given the large number of hybrid individuals between the two species at the sites, and the comparatively small number of V. hookeriana plants at any given site, however, the latter species may indeed become imperiled (Levin et al. 1996) Sharing of similar flowering times has likely favored pollen movement between the two species, however pollinator activity would play a key role in successfully moving the pollen. During the year this study took place, several potential pollinators were seen in the area of study, however none of them were observed visiting Viola, a finding which seems to be abnormal. Similar observations on blooming individuals of one of the spe- cies (V. grahamii) over the next two subsequent summers did however reveal the pres- ence of the golden banded skipper (Autochton cellus Boisduval and Le Conte, Hesperiidae) and an undescribed species of bee of the genus Dianthidium (Adanthidium) in the Megachilidae (Griswold, T. pers. com.). The skipper does not seem to be very specific and visits several species showing white flowers (Cortés-Palomec 2005), but the bee does ap- pear to only frequent Viola and it could be favoring gene flow between these two species. More specific studies of the pollinators would be needed to better understand their be- havior. In conclusion, we have shown that hybridization occurs between Viola grahamiiand V. hookeriana, and that the hybrids are morphologically distinct and intergrading from the parental taxa. Genetically, despite a larger similarity to V.grahamii, the hybrids have unique alleles. Ecologically, hybrids have a wider distribution and environmental toler- ance than any of the parental taxa, and at least in the area of study they are much more abundant than V. hookeriana. Similar flowering phenologies and effective pollen move- ment between the species has likely favored hybridization over a long period of time between the two species. 1132 BRIT.ORG/SIDA 22(2) ACKNOWLEDGMENTS ACP would like to thank the Consejo Nacional de Ciencia y Tecnologia (CONACyTF Mexico) for financial support awarded under grant #128098. The authors also thank the Secretaria de Recursos Naturales y Pesca (Semarnap) for granting the collecting permits for our work. Thanks are also extended to Ricardo Wong and Juan Carlos Gonzales for providing field assistance and to Ross McCauley for help with data analysis. We would additionally like to thank Susan E. Yost, Nir L. Gil-ad, Mary J. Haywood, and Rebecca A. Peters for their useful comments and suggestions on this manuscript. 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Wo re, A.D.and C.P.Ranoit.2001.Relationships within and among species of the holoparasitic genus Hyobanche (Orobanchaceae) inferred from ISSR banding patterns and nucleotide sequences. Syst. Bot. 26:1 20-130. ZAR, J.H. 1996. Biostatistical analysis. Prentice Hall, Englewood Cliffs. 1134 BRIT.ORG/SIDA 22(2) Book REVIEWS Davip Fross and Dirter WILKEN. 2006. Ceanothus. (ISBN 0-88192-762-7, 978-0-88192-762- 7, hbk.). Timber Press Inc, 133 $.W. Second Avenue, Suite 450, Portland, OR 97204- 3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax.). $39.95, 272 pp., color photos, b/w illustrations, Teo As noted on the jacket: “Ceanothus, with so many plants that tolerate sun and shade, that thrive in arid conditions ae and thus conserve water, that bear a profusion of beautiful, fragrant flowers, deserves a book that will bring more of them into gardens and encourage greater appreciation of them in the wild, and this is that book.” Plenty of color photos of these spectacular shrubs and small trees. Statements of the geographic ranges are given; the only thing Lcan think of in improvement of this book, “The Compleat Ceanothus,” would] ave been generalized range Oo, maps. The text is divided into two substantial and readable sections (chapters in parenthesis): Ceanothus in the Garden and Landscape (Natural Distribution; History of Culti g Ceanothus, Disease and Pests; Cultivar Development; ee oes Ceanothus Species and C itats for an Garden and Landscape [94 pages| and Ceanothus Selec suide eer ns in th wis (an evoOnety and taxonomic overview) ] = Distribution; E Orm anda Function in Ceanothus: ¢ les and descr intend and Other Ceanothus names) —Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, 1X 76102-4068, U.S.A. David D. Stuart. 2006. Buddlejas. (ISBN 0-88192-688-4, 978-0-88192-688-0, hbk.). Royal Horticultural Society Plant Collector Guide. Royal Horticultural Society and Tim- ber Press Inc, 133 5.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327- 5680, 503-227-3070 fax.). $34.95, 192 pp., color ne 7" x 9" A guid I Buddlej | | gard rith plenty of detailed information. “Buddlejas 5 are ihe most aocgmiadieoend eeneuing of garden pk ne ies acceptance of challenging soil conditions anc climate extremes and their resistance to pests (including deer, hares, and rabbits) are legendary.” The genus in- cludes about 100 species — the author lists and discusses 58 species of them (noting that he is concerned only with the woody species, vs. the herbaceous perennials). A middle s section provides beautiful color photos of most of the 58, along with many of the distinct cultivars of B. davidii. The super-hardy B. davidii (butterfly bush) is commonly used by breeders because of its abundant and nectar-rich flowers. “The major down side of the species is its ability to colonize inhospitable wasteland sites; the tiny winged seeds, blown far and wide, are capable of germinating wherever they settle, be it old tarmac, aioe or a crevice high on a wall. During World War II it besa known as the bombsite plant in Britain, as it was always the first plant to establish itself after an attack.” The North American natives B. marrubiifoli 1B. ulahensis are included in the book but not B. racemosa, B. scordioides, and B. sessiliflora. Chapter 1. Species Chapter 2. Hybrids and Cultivars Chapter 3. Care and Cultivation Chapter 4. Propagation Chapter 5. Wildlife Attracted to Buddlejas —Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, 1X 76102-4068, U.S.A. SIDA 22(2): 1134. 2006 HERBIVORY OF FERAL GOATS ON ESPIRITU SANTO ISLAND, GULF OF CALIFORNIA, MEXICO José Luis Leon-de la Luz and Reymundo Dominguez-Cadena Programa de Pl ion Ambiental y Conservacion G t At au ; R; logi dol NI. tf, (CIBNOR). Mar Bermejo 195, Col. Playa Palo de Santa Rita, La Paz, B.C.S. 23090, MEXICO jlleon04@cibnor.mx ABSTRACT Island fragil d iall luced ies, particularly large herbivores. In Mexico, ee are some dramatic cases. We ep alnated the influence of feral pone as well as the natives black jackrabbit and desert woodrat on the vegetation of Espiritu Santo Island, a land-bridge island in the southern part of the Baja California Peninsula. We found that goats consumed 36 vascular plant species, three times more species than natives jackrabbits and almost seven times more than woodrats. Also, goats consume all plant or- gans except roots, but reiterate browsing gradually affect vegetative and reproductive plant fitness. Our results support plans for the eradication of goats to spontaneously and progressively restore of the natural vegetation structure and composition. Further, a monitoring plan is recommended. Key Worps: goat herbivory, Capra hirca eradication, goat eradication, Baja California Islands RESUMEN Las islas son consideradas ecosistemas muy fragiles. Estas son sensibles a la introduccion de especies exoticas, icul herbi Fn Méxi ict | 44 EF te-trahoi laavebat, | if J la vegetacion en complejo insular Espiritu Santo, Baja California Sur, México, de las cabras silvestres contra el causado por las oe nativas, ra liebre negra y la rata de campo. Las cabras consumen 36 especies de plantas yeeculare es, tre mas esp que las HEDIES y casi 7 veces mas que las ratas. También, se encontr6 que las todo tipo d , pero el ramoneo reiterado afecta tanto la aptitud vegetativa y neprodilcnye de es pesca el -OS Eitedes apoyan el plan de erradicacion de las cabras con la finalidad de que espontanea y progresivamente se restaure la vegetacion en estructura y composici6n. Adicionalmente se requiere de un plan de monitoreo. INTRODUCTION Extinction and extirpation of native plants and animals from islands are significantly correlated with the introduction of exotic grazing mammals. Prevention of introduction and establishment of exotics on islands is vital, so that a program of eradication or con- trol of existing exotics is required on many islands around the world, and eradication of new introductions, as soon after Secccctne as possible, should be a high priority for con- servation natura by means of early aa and rapid response (McChesney & Tershy 1988; Burbidge 1999; enlan et al. 200 Feral domestic goats (Capra hircus L.) are implicated in habitat ie and al- teration of species composition in sensitive insular ecosystems around the world (Scowcroft & Hobdy 1987; Jennings 1987; Parkes 1990). In the absence of population con- trols, goats become the ecologically dominant species on many islands, with the result that numerous native and endemic plant species have been extirpated or are threatened with extinction (Coblentz 1978; Keegan et al. 1994). A dramatic case is Isla Guadalupe in the Pacific Ocean near the state of Baja California, Mexico. This island was devastated y SIDA 22(2): 1135- 1143. 2006 1136 BRIT.ORG/SIDA 22(2) grazing goats and erosion caused by soil trampling, to the extent that some native species and entire plant communities have vanished (Leon de la Luz et al. 2003; Oberbauer 2006). Espiritu Santo is a small archipelago of two major land bridge islands and several islets in the southwestern Gulf of California (Fig. 1). They are included in the Gulf of California island protection program covered by Mexican environmental laws, and ad- ministered by the Natural Protected Areas Commission (CONANP). The largest island is Isla Espiritu Santo is separated on the north by a 10 m wide channel from the second island, Isla Partida Sur. Additionally, there are the islets of La Ballena, El Gallo, La Gallina, and Los Islotes (Fig. 1). An undetermined number of domestic goats were introduced to Isla Espiritu Santo around the middle of the 20th Century to serve asa supply of fresh meat for local fisher- men. Currently, there are hundreds of feral goats on the island that are gradually destroy- ing the vegetation, eventually leading to extinction of native species. An informal ani- mal census by the authors in spring 2001 provided an estimate of 1,000 to 1,200 goats occupying Isla Espiritu Santo. Field observers could not be more precise because the gen- eral relief, escarpments, and steep slopes made conventional counting methods impos- sible. Because these islands are narrow and rocky, runoff is very rapid after the relatively rare rains. With little percolation into the soil, water is collects in occasional drainage pools. Thus, plants on the islands seem to have more profound water stress than on the main peninsula, a condition that is reflected by poor reproductive response, including less blooming, setting of fruit, and seed production than populations of the same species on the peninsula (author, pers. obs.). These conditions critically reduce the recovery ca- pacity of the plant community, and consequently, the dependent animal guilds. One of the greatest challenges for administrators of protected areas is maintaining biodiversity of native species and habitat loss. In Mexico, most of the islands have been protected to keep their biodiversity. This island, which is federal property and a protected area, is being considered for a program of goat eradication by the administrators (SEMARNAT, Baja California Sur). The administrators are familiar with the catastrophic effects of grazing activity on islands around the world and in Mexico. Based on field transects, this study compared and evaluated differential consumption of plants by feral goats and the endemic black jackrabbit (Le pus insularis Bryant) and desert woodrat (Neotoma lepida Thomas subsp. vicina Goldman) on Isla Espiritu Santo, com- paring their impacts on the vegetation. Following goat eradication, our data could pro- vide support for prediction of the recovery of the plant community. METHODS Site description The Espiritu Santo archipelago (24°24' to 24°36'N, 110°18' to 110°27' W) is part of the state of Baja California Sur. The nearest approach to the coast of the Baja California Peninsula is barely 8 km, northwest of the port at Pichilingue. The longest axis of the archipelago (NW-SE) is 19 km; the widest reach (E-W) is about 3.5 km. The archipelago covers 110 km2, 87 km2 on Isla Espiritu Santo and 20 km? on Isla Partida. The highest point, near the midpoint of the big island is 540 m (Fig. 1). Geologically, the archipelago is composed of sedimentary rocks of volcanic origins (sandstones and conglomerates) of the Comondu Formation. Variations in resistance to weathering lead to a rugged topography. Except in a few areas, most of the surface has FERAL 1137 110°20°W Los Islotes = x a’ GULF OF / CH) CALIFORNIA Y Partida aa La Ballena / poe, oe 7 El Gallo a, aes La Gallina 2 oe ° ey a Espiritu Santo Pacific Ocean 2Kkm Fic. 1.6 g phi I] i f th + ly 1 tl ri Many: bth Calf o£ FA10L, Sa zs, A y over 200 m. C (+) indi he hia + at then tcl 1 (540 \ very steep rocky slopes where some soil is present beneath rocks. The uplands are table- lands where goats concentrate at nights. The west side of the island is highly dissected by ravines descending from the uplands to sea level. On the shore, salt flats form behind mangrove stands and sandy pocket beaches at the end of coves. The east side of the is- land is largely wave-cut cliffs and pebble beaches. Weather data are not available for the island, but an approximation, based on records from the city of La Paz, suggests annual mean rainfall <170 mm and potential evapora- tion of 2500 mm. Rainfall is erratic, with fall-winter-spring of 8 to 10 months without any rainfall. Occasional hurricanes in the late summer could drop up to three times the annual mean. Vegetation is typical of the peninsular Sonoran Desert, the sarcocaulescent scrubland (Shreve @ Wiggins 1964) is characterized by perennials with semi-succulent stems, in- cluding cacti and deciduous woody shrubs. According to the last inventory, 254 taxa are found (Rebman et al. 2002). Families include Compositae (28 taxa), Leguminosae (25), Euphorbiaceae (24), Graminae (20), and ee (15), these five cabae 44% of all the taxa. Some of the tarborescent forms f ically dominating the slopes are: “palo blanco” Lysiloma candida Brandegee, “salate” Ficus petiolaris HBK subsp. palmeri S. Wats., “cacachila” Karwinskia humboldtiana (Roem. & Sch.) Zucc., and “cardén” Pachycereus pringlei(S. Wats.) Britt. &@ Rose. Constant and strong winds noticeably contribute to lignosity in branches and a more 1138 BRIT.ORG/SIDA 22(2) dwarflike appearance than examples of the same species growing on the Baja California Peninsula. With a north-south mountain ridge axis, the canyons run east and west, and the isolation generates different exposure-based plant associations on north-facing (shadow-side) and south-facing (sun-side) slopes. North-facing slopes having significantly more canopy and plant eon than the intensely exposed south-facing slopes. Sur Wor by the author, based on 400 m¢ * sampling plots, provide plant canopies oe 220m? on south-facing slopes and 305 m? on north-facing slopes, on opposite sides of the same ravine (author, unpublished data). ling In March 2001, ten straight-line transects of approximately 3 km each, were designed so that each would traverse several environments. Transects included beaches and man- grove (one site), arroyos and washes (three sites), slopes (four sites), and tablelands (two sites). Information recorded included: 1) plant and Olean consumed or injured, 2) evi- dence for the responsible herbi s species, based on characteristics of common sense, including type of bite on the plant and supporting evidence of droppings (pellets) beside the damaged plants. Droppings of goats are barrel form, approx. | cm long; those of jack- rabbits are elliptic-lenticular form, approx. 1 cm long, and those of desert woodrats are linear, approx. 1 cm long. Because the three herbivores typically defecate next to the browsed plant during feed- ing, we considered such occurrences related and definitive. Figure 2 shows a foraged plant and pellets of goats and black jackrabbits. Intensity of feeding activity was determined by the number of old scars on plants and the abundance and age of pellets around these plants. — RESULTS Table | assembles the information by sampled sectors, showing elevation, browsed spe- cies, consumed or injured plant organs, evidence of the herbivore species, and intensity of consumption based on scat abundance. From the transect surveys, we found that brows- ing by goats was concentrated on the more elevated mountains of the island near La Ballena and Los Candeleros Coves, and on uplands above +00 m Discussion In general terms, abundance and distribution of plants on the island is definitely depen- dent on terrain conditions, such as slope angle and exposure, as well as soil quality (soil deep and rockiness). These factors contribute to vegetation heterogeneity on the land- scape. We believe that many native plant species have been modified by goat browsing over the past several decades, knowing that goat herbivory directly affects fitness and reproductive success of mature plants and also contributes to consumption of juvenile plants. Donlan et al. (2002) documented positively changes in plant coverage on several Gulf of California islands after extirpation of exotic herbivores. Some actively browsed plant species are the perennial herbs Bebbia juncea (Benth.) Greene, Justicia californica (Benth.) Gibson, and Ditaxis lanceolata (Benth.) Pax & K. Hoffm. and the sub-shrubby Aeschynomene nivea Brandegee. We found that browsing of terminal branches promotes sprouting from basal nodes, leading to a branching archi- tecture of unusually short basal trunks and numerous branches. Several individual “palo blanco” Lysiloma candida, one of the few tree-like species, had trunks stripped of the high- tannin bark, as well as shrubby species of ‘lomboi” Jat ropha cinerea (Ortega) Muell-Arg. 1 ICANTCIA Sane, ©, foes Mt fst te oo hac) i la Pe 1140 BRIT.ORG/SIDA 22(2) Taste 1. Perennial plant species affected by herbivory of feral goats, black jack-rabbits (BJR), and wood rats on Isla Espiritu Santo, Baja California Sur, Mexico, based in ten 3-km transects during a field survey in spring 2001. Plant species (asl) Consumed organ Herbivore Intensity Beaches and mangrove (0-2 m) Rhizophora mangle L. Leaves, Bark Goat Low Avicenia germinans (L.) L. Leaves, Bark Goat Low Monantochloe littoralis a Leaves JR Low Jouvea pilosa (Presl.) Scrib Leaves Goat Low Arroyos and washes (5-15 m) Viscainoa geniculata (Kell.) Greene Leaves Goat Low Forchammeria watsonii Rose Leaves Goat Low Euphorbia magdalenae (Benth.) Millsp. Buds Goat Low aiadin candida Brandegee Flowers, Leaves, Bark Goat Low Melochia tomentosa L. Buds Goat Low sais ene (Engelm.) Gibson & Horak Stems BJR, Rat Low Is prin i (S.Wats.) Britt.& Rose Flower, Bark Goat, BJR Low rdsia chinensis (Link) Sch. Leaves oat Low Col ibaa viridis M. E. Jones Bark Goat Low Bebbia juncea (Benth.) G Leaves Goat ow ochemia poselgeri ener Britt.& Rose Stems BJR, Rat Hig Prosopis articulata S. Wats. Stems, Leaves Goat, BJR Medium Slopes (10-150 m) Justicia californica (Benth.) Gibson Stems, Leaves, Buds Goat High Ditaxis lanceolata (Benth.) Pax & K. Hoffm. Stems, Leaves Goat High Desmanthus fruticosus Rose Stems, Leaves Goat High ue nivea Brandegee Stems, Leaves Goat High Agave sobria Brandegee var. roseana (Trel.) Gentry — Flowers Goat High anes ia pacensis nas & carte Buds Goat High rtus (L.) Beauv.ex Roem. Leaves Goat Medium Cyiindropunts ia ciel a Weber Flowers, Fruit BJR, Rat Low Macrosiphonia hesperia |.M. Jhtn. Stems, Leaves Goat High Ficus petiolaris subsp. palmeri S. Wats. Leaves, Buds Goat High Bouteloua reflexa Swallen Leaves Goat Medium Solanum hindsianum Benth. Buds Goat Medium Tableland (150-450 m) cia Bases nee & Carte Stems Goat, BJR Medium bria Brandegee var. roseana (Trel.) Gentry Flowers, Leaves Goat Medium Buea eoinnata (Rose) Engler Buds Goat Medium Caesalpinia placida Brandegee Stems, Leaves Goat Medium Parkinsonia florida Benth. ex A. Gray Stems, Leaves, Buds Goat Medium subsp. peninsulae (Rose) Carter Fouquieria burragei Rose Buds Goat Medium latropha cinerea (Ortega) Muell.-Arg. Buds Goat Occasional hvaloua eandice Brandegee Bark Goat Medium Olneya tesota A. Gray Buds Goat Medium ea ia eis Engelm. Stems Goat, BJR Medium f hinensis (Link) Sch. Stem, Leaves Goat Medium Eushertia sneulenae (Benth.) Millsp. Buds Goat Occasional Supnorele DIOS (S.Wats.) Millsp. Buds Goat Occasional a (Benth.) G.L. Nesom Stems, Leaves Goat, BJR Occasional FERAL 1141 Table 1. continued Plant species (asl) Consumed organ Herbivore Intensity Tableland (150-450 m) (cont.) Stenocereus gummosus (Engelm.) Gibson & Horak Stems BJR High Maytenus phyllanthoides Benth. Stems BJR, Rat Low Pedilanthus macrocarpus Benth. Stems BJR Occasional Opuntia cholla Weber Stems Goat, BJR Medium Hyptis laniflora Benth. Buds Goat, BJR Medium Pachycereus pringlei (S.Wats.) Britt. & Rose Stems Goat, BJR Occasional Echinocereus brandegeei (Coult.) K.Schum. Stems Rat Medium (see Fig 3). Some plants of “cardon” Pachycereus pringlei, a large columnar-arborescent cacti, have old scars on their trunks (>20-40 cm above the ground), evidence of brows- ing by goats and probably also jack-rabbits during periods of severe drought. Our obser- vations suggest that only two species of “choya” cacti (Cylindropuntia cholla Weber and G alcahes Weber) could benefit from goat browsing pecause sexual reproduction in these wo very limited and their jointed st asily, adhere to goat hair, and are easily dispersed and eventually they could settling by vegetative processes. Black jackrabbits and desert wood rats have a significantly lower impact on the veg- etation because, as smaller browsers; individually they consume less plant material and their population biomass seems to be less than the hundreds of goats. From the data assembled in Tables 1 and 2, we found that the most intense location of browsing by goats is on stems and, at least during spring, is frequent and intense. This browsing habit almost entirely consumes the floral and leaf shoots, the juicy and less woody tissues of the plants. After the rainy season, the goats concentrate on browsing the newly emerging foliage. The evidence indicates that a set of plant species display consumption of or injury to the meristematic apex of the branches, that is, the fresh and fluid-filled tissues, rather than woody tissue and stem bark. Clades of Opuntia tapona Engelm. are specially con- sumed by all the animals because the high water content. With freshwater in very short supply and stored in “tinajas” after rainfall, goats manage to prosper by drinking salty water (Burke 1990). During our field work, we observed a few goats drinking seawater at the beach at sunset, which suggests daily or close to daily migration from the uplands to the shore. At the shore, goats also consume halophytic plants, including mangrove and species of salt grasses. Our findings demonstrated that goats are the most significant herbivores, capable of consuming almost all plant parts. While the native black jackrabbit and desert woodrat have the same feeding habits as goats and inhabited the island for millennia, their den- sity and biomass have far less impact on the vegetation. Finally, there was no direct evidence that goats consume an entire plant; however, high intensity herbivory strongly affects the fitness of each grazed plant, and can even- tually lead its death. We conclude that the natural balance of vegetation on Isla Espiritu Santo is deleteriously affected by goat herbivory, which combined with the impact of the native foragers, increases the deleterious effects of browsing on a large group of plant species. Our data supports plans for the total eradication of goats from the archipelago. BRIT.ORG/SIDA 22(2) Fic. 3. An exemonplar of Jat, } I | I t te hark 1 brok I } r f J ~ ad re i Taste 2.Concentrated data of plants consumed by herbivores on Isla Espiritu Santo, Gulf of California, Mexico. Herbivore Plant species Plant organs Sector of activity consumed consumed Goats 36 Flowers, buds, leaves, young stems, bark Slopes, high tablelands Jackrabbits 13 Stems Arroyos, high tablelands Wooadrats 5 Stems, fruit Arroyos, slopes On an ecological basis, black jackrabbit and woodrats are foragers that control the veg- etation growth. After eradication a monitoring plan is strongly recommended. We hope that the same plants indicated in Table | will increase their density and size. In fact, eradication of goats is currently underway by the administrators of the island. The project is expected to be completed by winter 2006/07, with only radio-collared and sterile sentinel goats remain- ing (Donlan et al. 2002). ACKNOWLEDGMENTS We appreciate the English editing of Ira Fogel at Centro Investigaciones Bioldgicas del Noroeste, as well Miguel Dominguez for field expertise during our surveys of the island. FERAL AND 1143 REFERENCES Bursince, A.A. 1999.Conservation values and management of Australian islands for non-volant mam- mal conservation. Australian Mammalogy 21:67-74. Burke, M.G. 1990. Seawater consumption and water economy of tropical feral goats. Biotropica 22: 416-419. Costentz, B.E. 1978. The effects of feral goats (Capra hircus) on island ecosystems. Biol. Conservation 13: 279-286. Dontan, C.J., B.R. TersHy, and D.A. Croit. 2002. Islands and introduced herbivores: conservation action as ecosystem experimentation: applied issues with predators and predation. J. Appl. Ecol. 39: JeNNincs, G.Y. 1987. Guadalupe: An island refuge under siege. Oceans 20(5):40-45. Keecan, D.R., B.E. Costentz, and C.S. WincHEeLL. 1994. Ecology of feral goats eradicated on San Clemente Island, California. In: Halvorson, W.L., GJ. Maender, eds. The Fourth California Islands Symposium: update on the status of resources. Santa Barbara Museum of Natural History, Santa Barbara, CA. Pp. 323-330. Leon ve ta Luz, J.L., J.P. Resman, and T. Osersauer. 2003. On the urgency of conservation on Guadalupe Island, Mexico: Is it a lost paradise? Biodivers. Conservation 12:1073-1082. McCuesney, G.J.and B.R. TersHy. 1988. History and status of introduced mammals and impacts to breed- ing seabirds on the California Channel and northwestern Baja California Islands.Colon.Waterbirds 21:335-347. Osersauer, T.A.La vegetacion de isla Guadalupe. Entonces y Ahora. 2006. In: K. Santos del Prado and E. Peters. Isla Guadalupe: Restauracion y Conservacion. Instituto Nacional de Ecologia (INE- SEMARNAT), México. Pp. 39-53. Parkes, J.P. 1990. Eradication of feral goats on islands and habitat islands. J. Royal Soc. New Zealand 20:297-304, Resman, J.P, JL. LEON bE LA Luz, and R.V. Moran. 2002. Vascular plants of the Gulf Islands (Appendix 4.1). In:T.J. Case, M.L. Cody and E. Ezcurra, eds. A new island biogeography of the Sea of Cortés. Oxford University Press. New York. Pp.465-512. Scowcrort, P.G.and R. Hospy. 1987.Recovery of goat-damaged vegetation in an insular tropical mon- tane forest. Biotropica 19:208-215. SHreve, F.and ILL. Wicains. 1964. Vegetation and flora of the Sonoran Desert. Stanford University Press, Stanford, CA. 1144 BRIT.ORG/SIDA 22(2) Book REVIEWS Doucias D. Stokke and Leste H. Groom (ed.). 2006. Characterization of the Cellulosic Cell Wall. (ISBN 0-8138-0439-6, 978-0-8138-0439-2, hbk.). Black well Publishing. 2121 State Ave., Ames, [A 50014-8300, U.S.A. and 9600 Garsington Road, Oxford, OX4 2DQ, UK (Orders: orders@ames.black wellpublishing.com, 515-292-0140, 515-292-3348 fax I- 800-862-0657, www.blackwellprofessional.com). $149.99, 274 pp., color photos, nu- merous b/w illustrations, 7" X 10' ing ... but so a more. This collection of (oy “Crack” as the twig breaks, the slightly elastic wood suddenly yielc studies, fundamental and applied science, represents the proceedings of a work : sored by the USDA Forest Service, the Society ‘ Wo a Science and Technology, aus lowa State U niversity — “the eeu i collabora- tion between three institutions witha ae: history of res g lignoc cel- lulosic substances, primarily wood.” The book is “intended for scientists, university faculty, sec trte students, and applied researchers in the fields wood science and technology, cellulose science, and biomaterials.” The book is arranged in three sections: Cell Wall Assembly and Function: New Frontiers (5 chapters, e.g. tracheid and sclereid differentiation in callus cultures of Pinus radiata; mechanics of the wood cell wall; prediction of wood structural patterns by using eco- logical models of plants water relations) Probing Cell Wall Structure: Advances in Analysis (6 chapters, e.g: determining xylem cell wall properties by us- ing model plant species; near infrared spectroscopic monitenng of the diffusion process of deuterium-labeled molecules in wood; wood stiffness by X-ray diffractometry) Mesostructure dnd Applications: Science in Practice (8 chapters, e.g: variation in kink and curl of longleaf pine fibers; wood structure and adhesive bond strength). —Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A. Marti J. GRoom, Gary K. Merve, and C. RoNALD CARROLL. 2005. Principles of Conservation Biology, Third Edition. (ISBN 0-87893-518-5, hbk.). Sinauer Associates, Inc., 23 Plumtree Road, Sunderland, MA 01375-0407; U.S.A. (Orders: orders@sinauer.com, wwwsinauer.com, 413-549-4300, 413-549-1118 fax). $92.95, 699 pp., 369 illustrations, S172 Sr This is a major update and revision of the 1997 Second Edition, with much new information and guided by a survey of “over 60 instructors on their needs for teaching conservation biology.” The text is divided into three major units: Conceptual Foundations for Conservation Biology (5 chapters); Focuson a yl hreatsto Biodiversity ] t (6 chapters), Approaches to Solving Conservation Problems (7 chapters). A “companion sa variety of study materials and supplemental resources to accompany the textbook.” On the first page of Chi ices I: “the cumulative impacts of 6.4 billion people (Figure 1.1) have stressed the many ecological support systems of the planet. ... All prope should neCOB Ize the degree to irae human impacts affect the natural world, and in turn, | } al diminish our biologists conservation should have this book at hand (or one like it, if there’s anther: so seed) —Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A. SIDA 22(2): 1144. 2006 A DEMOGRAPHIC AND ECOLOGICAL ANALYSIS OF GEOCARPON MINIMUM (CARYOPHYLLACEAE): A FEDERALLY THREATENED SPECIES IN SOUTHWEST MISSOURI Tim Smith! Joseph S. Ely Missouri ee Department of Biology PO, Central Missouri State University Jefferson City, iaissonan see 0180, U.S.A. Warrensburg, Missouri 64093, U.S.A. ABSTRACT A Geocarpon minimum population growing on a sandstone glade in Dade pei! Missouri was monitored for ten years (1994-2003). Plants were counted annually in eight permanent plots at two scales (0.01 and 1.0 m2) to determine population fluctuations through time. Soil depths as well as cover of eee ce lichens, vascular plants, and bare soot were recorded in subplots between 1994 and 2000 to determine any ae association with G. minimum mbers. The number of G. minimum plants in the eight plots varied from 1 to 4230, a mean of 1332. Mean soil — aa of subplots containing G. minimum was 19 mm. There was a weal association (1, = “0 16, P<0O. ool) seubibidae soi depth and Geocarpon density. Regressic e soil depth to about 20 mm and ee ee In addition, there were significant differences among vegetation - al that G types (P < 0.05) with the exception of lichens (P > ac There was a ag pious difference between G. minimum W population numbers and year (P < 0.05). There was 1 scale and the Spearman rank correlations, a however, when ecological Broups were cgunpined into oie eatteoas (ie. lichens and bryophytes were lumpe into 5), patterns E bar are tock areas indicates a dynamic process ] Be | a ]; of soil oT eae into Banc out of plots, | lar pla nte s Hee | gay I ] y ee i ] a a | f= crease In cove ies of moss, Vasc u- disturbance and with minimal competition from pb onaytes or perenaal vasa plants Although seed kau was L I {.]] ] 4 LOLLOVY ing yeAalo W ith VII tually ] node Teeter hi : of RESUMEN Una Poe de soles ble i m is mopireraca cuante ae anos caeaiipicns enun nenoueente bas aL enisca y numero de pate en ar Reach durante el periodo de estudio, las plantas se contaron anualmente en ocho parcelas permanente las (0.01 y 1.0 m2). Durante los anos 1994 hasta el 2000 fueron evaluados datos sobre ee de stele coe de briotitas, liquenes, plantas vasculares y area de tierra o rocas sin q P y plantas en subparcelas con el { ito ded la posible asociacion de estas ae con el numero de plantas de G. minimum. El numero de ae de G. minimum en las ocho parcelas vario de 1 a 4230 con un promedio de 1332. La oes poe del suelo eeu las ven con G. minimum fue “ 19 mm. La asociacion entre la y densidad de pl ‘pon fue minima (r, = 0.16, P < 0.001). El andalisis de regresion demostr6 que el numero de G. minimum parece aumentar a Rene re que la profundidad de suelo alcanza los 20 mm y disminuye a mayor profundidad. Ademas, la diferencia entre tipos de vegetacion (P< 0.05), con excepcion de los liquenes (P >= 0.05), fue significativa. La diferencia fue significativa también entre el numero ue oe de G. minimumy el ano del muestreo (P < 0.05). Se observ6 una discontinuidad entre escala y le rango de Spearman, sin embargo, al combinar los grup oldgi n categorias mayores (por ee los aie y las pbloritas se unieron en e eatseCHe de las criptgaas) los patrones fueron mas claros a través de las escalas. La as indica la existencia de un proceso dinamico de movimiento del suelo dentro y fuera de las parcelas, pero is ne sobre cobertura no indi- can un crecimiento constante eae Selo es eos pleats eo o liquenes durante los diez anos del } -] estudio. I - tienen Acie ell i & A Fig L oO 'Tim Smith, corresponding author: Telephone: (573) 522-4115; Fax: (573) 526-5582; E-mail: tim.smith@mdc.mo.g SIDA 22(2): 1145- 1157. 2006 1146 BRIT.ORG/SIDA 22(2) en este estudio, su importancia fue evidente. En base a los datos anuales, esta poblacion de G. minimum tiene la capacidad ae reproducirse considerablemente en anos siguientes aunque no exista produccion de semilla en anos anterio INTRODUCTION Geocarpon minimum Mack. (Caryophyllaceae) isa tiny, succulent winter annual that grows on sandstone glades in Missouri, on saline soil prairies in Arkansas and Louisiana (U.S. Fish and Wildlife Service 1993; McInnis et al. 1993; Orzell & Bridges 1987), and on saline barrens in Texas (Keith et al. 2004) (Fig. 1). The diminutive height of the plant (up to 4 cm), its inconspicuous flowers, and its limited distribution are probably responsible for it not having been described until 1914 (Mackenzie 1914). Taller plants are usually multi-stemmed and many-flowered. The smallest plants are only a few mm in height and produce just one flower. Both morphologies are often present in a single population, requiring surveyors to acquire a broader search image to locate the full range of plant forms. Plants turn froma whitish-green to wine-colored to light brown as they mature and senesce. Geocarpon minimum was designated by the U.S. Fish and Wildlife Service as a feder- ally threatened species in 1987 because of low population numbers (U.S. Fish and Wildlife Service 1987). It is known today from fewer than 35 sites (Nienaber 2005). Twenty-four ex- tant Missouri populations are known from Pennsylvanian-age sandstone glades in seven southwestern counties, giving it a state rank of S2 (Missouri Dept. of Conservation 2006). The sandstone glades occur in 1 the Springfield Plain subsection of the Ozark Highlands ecological section and ar idered imperiled (a rank of S2) because of their rarity within the state (Nelson 2005). G. minimum is considered extirpated from its type locality, Jasper County, Missouri, where suitable habitat may have been lost due to land-use changes. Prior to this study, little was known about the management requirements for main- taining populations of G. minimum or the range of year-to-year population fluctuations. Several sites for the species occur on public lands where lack of information has led toa lack of management action. There was concern that some periodic disturbance might be necessary to maintain suitable microhabitat for G. minimum. On many of the glades where the species occurs, the thin soil near the rock/soil interface is dominated by dense mats of moss, except where trampling, off-road vehicle use, or water flow have maintained areas of loose sand. The purpose of this study was to gather long-term monitoring data that could provide needed insights for effective management of G. minimum populations. Specific objectives of this study were to: 1) provide a long-term census of G. minimumat the Flint Hills Glade site, 2) determine the correlation of bryophytes, lichens, vascular plants, and soil depth on the G. minimum population, 3) provide management recommendations for the conservation of this species. MATERIALS AND METHODS Study Site The study site, Flint Hill Glades, is located in Dade County, Missouri, near Stockton Lake, on land owned and managed by the U.S. Army Corps of Engineers (Fig. 2). Geocarpon minimum was discovered at Flint Hill Glades during a 1989 survey for the species (Thurman & Hickey 1989). The site is atypical in one respect; the impoundment of Stock- ton Lake inundated the creek valley below the glade and floods the lower portions of the glade during extremely high lake levels, which occurred during the summer of 1993. SMITH AND ELY, ECOLOGICAL ANALYSIS OF GEOCARPON MINIMUM IN MISSOURI 1147 Fic. 1. Geocarpon minimum plant. Photo by Rick Thom It is estimated that less than five-percent of the six-acre glade area is suitable habitat for G. minimum, which grows in greatest abundance in shallow sand- and pebble-filled depressions in the sandstone bedrock that are often devoid of other plants (Fig. 3). Based on research at another Dade County site, Morgan (1986) reported that optimum habitat was found in zones of shallow soils (1-5 cm) over sandstone bedrock. These microhabi- tats are maintained by flow of water over the bedrock that deposits sand and pebbles into the depressions. Water flow can also carry away previous deposition and prevent lichen or vascular plant succession. An intermittent drainage flows through the study site carrying runoff from adjacent grazed pasture on private property into Stockton Lake at the lower edge of the glade. Much of the surface bedrock is covered by a thick mat of mosses (principally Polytrichum ohiense Ren. & Card.) intermixed with lichens, liverworts, and a few vascu- lar plant species. Areas of deeper soil within the glade support grasses, shrubs (e.g. Vaccinium arboreum Marshall) and dwarfed trees (e.g. Quercus marilandica Muenchh. and Amelanchier arborea (FE Michx.) Fernald). Geocarpon minimum plants were observed in the grazed portion of the glade on adjacent private land during the course of this study. They are found only near a bedrock exposure where pasture grasses are not established. The dense beds of moss found on the study site are not present in the pasture. The tram- pling from cattle may help to maintain the loose sand particles that accumulate to form G. minimum microhabitats. The pasture area may be a source of G. minimum seed for the monitoring area. No monitoring plots were tablished in the pasture, primarily because cattle grazing is not viewed asa realistic management tool for most publicly-owned sites. Data Collection Ten plots were established at the site in April 1994. The 1.0 m x 1.0 m quadrats were sub- 1148 BRIT.ORG/SIDA 22(2) , Stockton Lake, Dade County, Missouri jectively located in areas containing G. minimum in close proximity to areas dominated by mosses, lichens, or perennial vascular plants. Two opposite corners of each plot were marked by masonry nails driven into holes that were drilled into the bedrock. Two of the original ten plots eventually were dropped from the study because they — — SMITH AND ELY, ECOLOGICAL ANALYSIS OF GEOCARPON MINIMUM IN MISSOURI 1149 Fic. 3. Habitat at Flint Hill Glades. Photo by Tim Smith. did not support any G. minimum after 1994. Inundation of part of the glade in 1993 cre- ated temporary habitat in these plots due to silt and sand deposition with little vegeta- tive competition. At one plot, sand and silt was deposited into a patch of bryophytes on bedrock, temporarily providing habitat that supported G. minimum. The bryophytes had been killed by the prolonged inundation and were soon gone, leaving only bare rock at subsequent surveys. The other plot was located near the lake shore and included essen- tially bare sand after the inundation, although it had deep enough soils to support pe- rennial vascular plants. Although G. minimum was found in the plot when it was estab- lished in 1994, this plot thereafter became covered with debris from subsequent rises in the lake level and no longer contained exposed sand or G. minimum. Alm X 1m (1.0 m?) wooden frame was used to monitor the permanent plots. Fly- fishing line was strung on the inside of the quadrat to further subdivide the 1.0 m2 plots into 100 subplots of 10 cm x 10 cm (0.01 m2). The number of G. minimum plants was recorded in each subplot. Soil depth was measured at the center of each subplot by using a nail to probe to bedrock and then measuring the length of the nail that had entered the soil. Bare rock was recorded as zero soil depth. Vegetative cover was estimated using seven cover classes (a modified Daubenmire cover class) and dominant cover was character- ized as mosses, lichens, vascular plants or bare rock/soil (Daubenmire 1959) The seven cover classes were: 1 (O-1%), 2 1-5%), 3 (5-25%), 4 (25-50%), 5 (50-75%), 6 (75-95%) and 7 (95-100%). In the spring of 1995, three additional plots were established within the study area, in moss-dominated areas that did not contain G. minimum. These plots were created by removing a 1m X 1m section of dense moss to expose underlying mineral soil or bed- 1150 BRIT.ORG/SIDA 22(2) 2500 2000 1500 Numbers of plants 1000 1994 1995 1996 1997 1998 4999 ae 2000 Monitoring year 2001 5n92 2003 Fic. 4. Numbers of Geocarpon minimum plants i lots from 1994 to 2003. rock. These three plots were subsequently monitored for G. minimum, but soil and cover measurements were not taken. Data Analysis The subplot (0.01 m2) values of G. minimum numbers, soil depth, and cover classes for each vegetation type (see above) were summed within each large plot (iec., 1.0 m*) and were subsequently used for all analyses (Spearman rank correlation, analysis of vari- ance, and regression analyses). The independent variables were year, soil depth (mm), and cover values of bryophytes, lichens, and vascular peu Lichens and bryophytes were bined into a single category (cryptogams) and vascular plants and cryptograms were also combined into a single category of total vegetation. Spearman rank correlation coefficients, rs, (Zar 1999; Sokal &@ Rohlf 1995) were cal- culated on both subplots and large plots in order to determine strength of association among G. minimum, year, soil depth, bryophytes, lichens, cryptogams, vascular plants, and total vegetation at two spatial scales (0.01 m2 and 1.0 m2). In addition, partial correla- tions were also determined on ranked data controlling for the effect of year to determine potential associations among the vegetation types and soil. Model Il regression analyses were performed with the number of subplots that con- tain G. minimumas the dependent variable and soil depth as the independent variable. The regression analyses were performed on subplots only. Note that both soil depth and the number of subplots that contain G. minimum follow a normal distribution, as indi- SMITH AND ELY, ECOLOGICAL ANALYSIS OF GEOCARPON MINIMUM IN MISSOURI 1151 cated by a Kolmogorov-Smirnov normality test (P > 0.05). Specific regression analyses were performed independently on soil depth range of 1 to 20 mm and 21 to 110 mm. The justification for the latter was to show that the subplots containing G. minimum increase with increasing soil depth to 20 mm and declines thereafter. A Friedman repeated measures (FRM) analysis of variance (ANOVA) (a non-para- metric equivalent to the parametric repeated measures ANOVA) was used to determine differences among population numbers annually. Natural log transformed G. minimum numbers was the dependent variable, year was the repeated measures factor, and plot was the subject factor (Zar 1999; Sokal & Rohlf 1995). Because some of the plots had zeros and ones, it was necessary to add a count of 2 (a constant) to each observation for natural log (in) transformation (Zar 1999; Sokal & Rohlf 1995). The transformation was done in order to improve statistical models (Quinn & Keough 2002) and to maximally normal- ize the data. Independent variables, with the exception of year and plot, were treated as random in all models because they were not specifically controlled (Zar 1999; Sokal & Rohlf 1995). The year 1996 was excluded from all analyses because only one G. minimum plant was observed for that year and thus it was considered as an outlier. Additional FRM ANOVAs were performed on cover classes of bryophytes, lichens, vascular plants, cryptogams, total vegetation, and soil depth. There were three cover classes for bryophytes, lichens, vascular plants, cryptogams, and total vegetation while soil depth was placed into six. FRM ANOVA was performed on the large plots only. Independent variables were natural log transformed as needed. Additional repeated measure ANOVAs (Kruskal-Wallis) were performed without 1999 data in order to ascertain the relation- ship between soil depth and vegetation groups without the possibility of an interacting effect or confounding influence for the year 1999. The year 1999 was considered an out- lier similarly to 1996 but with large numbers of Geocarpon observed for that year. The alpha level for all analyses was 0.05. RESULTS AND DISCUSSION Population Fluctuations Winter annual glade plants commonly exhibit annual fluctuations in population size. Available moisture for fall germination and spring growth can vary greatly, with low moisture making shallow-soiled areas hostile for plant growth. Long-term monitoring is important in order to document the range of fluctuations in population size. Knowledge of this range improves our ability to interpret population data from sites where long- term data are not available. The ability of a species’ seed bank to repopulate a site after a poor seed production year has obvious implications for long-term conservation. Results of annual G. minimum counts in eight plots are depicted in Figure 4. The cumulative annual totals from all eight plots ranged from one plant (1996) to 4230 plants (2003). Although all of the plots are the same size, the amount and quality of the G. mini- mum habitat within each plot varies, with some plots containing more bare rock or denser vegetative cover. Plots 6 and 7, which often contained the most Geocarpon, provide a greater percentage of thin soil with little competition from bryophytes, lichens, or vas- cular plants. The small number of G. minimum plants at the site in 1996 was probably due toa prolonged dry period during early spring, prior to monitoring conducted in April. Only one additional G. minimum plant was observed outside of the plots during that year’s monitoring. More substantial numbers of G. minimum were found at other sites in the region that spring, indicating the critical effect of very local drought on the popula- 1152 BRIT.ORG/SIDA 22(2) tion at the study site. No insect herbivory of G. minimum was observed at the study site or at any other Missouri sites. There was a moderate positive correlation (1; = 0.44, P< 0.001) of G. minimum with year at the 1.0 m? scale (Table 1). Plot to plot variation was significant between 1997 and 1999 (RM: X? = 1L7, P = 0.045; Tukey: q = 4.16, P < 0.05). Although the population fluctuated from year to year depending on local conditions, there was an overall increase in the population over the years. Soil Depth Considerations The stream and sheet flow of water over the glade due to precipitation creates a dynamic environment for G. minimum, with the depth and location of the loose, sandy soil with small pebbles changing dramatically through time. It was not uncommon to find G. mini- mum plants that had been uprooted and probably displaced by water and soil movement. It seems likely that the succulent stems are at least somewhat tolerant of displacement and can become established at new locations, especially when partially buried in sand. The mean soil depth for subplots containing G. minimum was 19 mm (Fig. 5a, b). In comparison, Somers et al. (1986) found the shallowest soil depths on xeric communities of the limestone glades of Middle Tennessee to be (35 + 17 mm), with such areas domi- nated by annuals and mosses, including several endemic species. In this study, soil depth was measured at the center of each subplot rather than at the actual location of any G. minimum plant. Therefore the high number of subplots with zero soil depth does not imply that plants were growing on bare rock but rather that they grew within about 7 cm (the distance from the center to a corner of a subplot) of the bare rock/soil interface. Nevertheless, there was a significant association between soil depth and G. minimum numbers (rs = 0.16, P< 0.001 at 0.10 m? scale; r;= 0.41, P < 0.001 at 1.0 m2 scale)(Fig. 5b and Table 1). Note that the Spearman rank correlation between soil depth and the fre- quency of subplots containing G. minimum was -0.46 (Fig. 5a). It was observed that in areas of deeper soil, perennial vascular plants were more often present. Where vascular plants become numerous, fallen leaves from trees can lodge between stems and accumu- late, whereas G. minimum-rich areas of the glade are windblown and/or washed by mov- ing water and remain largely free of leaves. Shallow soil, per se, is not a requirement for G. minimum growth. Rather, it is the mechanism by which plant competition is minimized on the sandstone glades. This was borne out by the growth of G. minimum on deeper soils following removal of other veg- etation and litter by prolonged inundation and wave action on the lower portions of the study site in 1993. This is also apparent with the observation that G. minimum grows on deeper soils in the saline prairies of Arkansas, Louisiana and Texas, where high sodium levels prevent the growth of competing vegetation. In addition, there were significant differences with soil depth and the distribution of bryophytes, lichens, and vascular plants at the 0.01 m? scale as indicated by the Kruskal-Wallis analysis of variance and/or the Spearman rank correlation coefficients (bryophytes: H4 = 19.96, P < 0.001, rs = -0.12, P< 0.001; lichens: H4 = 19.10, P = 0.002, r; = -0.110, P > 0.05; and vascular plants: H4 = 17.18, P= 0.002, rs = 0.445, P < 0.001)(Fig. 5c, d, and Table 1). Both soil depth and the number of subplots that contain G. minimum passed the normality test (kolmogorov-Smirnov P > 0.05). The results of the model Il regression analysis show that G. minimum increases with increasing soil depth until about 20 mm (F2, 18 = 4.45; P < 0.05) albeit with a low adjusted r? (15.4%). The regression model ac- counted for fifteen percent of the variation and is consistent with the results of the Spearman rank correlations (Fig. 5a, b and Table 1). An additional regression analysis SMITH AND ELY, ECOLOGICAL ANALYSIS OF GEQCARPON MINIMUM IN MISSOURI 1153 Taste 1.Spearman rank correlation coefficient for Yea ini bryophytes, lichens, vascular plants, cryptogams, total vegetation, and soil at the ane 0. 01m: 2, n = 862, (above the diagonal) and large plot (1.0 m’),n = 50 (below diagonal) scales. Correlations between large plot and subplot data (along the diagonal) were not calculated because the data are not paired. * = significance at the 0.05 level. Year G.minimum Bryophytes _Lichens Vascular Cryptogam _sTotal Soil Plants Vegetation Year _ 0.440* “0.23” -0.317* 0.022 0331" SO: 53i4" 0.038 G. minimum 0.190 = -0.230* -0.168* 0.186* -0.305* -0.236* 0.162* Bryophytes -0.161 -0.098 - -0.160* -0.577* 0.729% 0.362* -0.118* i -0.184 -0.045 0.367* - -0.442* 0.550* 0.298* -0.027 Vascular Plants -0.024 0.204 -0.154 -0.273 ~ -0.781* 0.070* 0.165* Cryptogam -0.283* -0.099 0:727* 0.829% -0.346* - 0.542* -0.111* Total Vegetation -0.312* -0.041 0.663* 0.755* 0.120 0.871* - 0.043 Soil 0.042 0.408* -0.199 -0.256 0.471* = -0.222 0.448* - shows that G. minimum numbers decrease with increasing soil depth beyond 20 mm (F2, gg = 68.45; P < 0.0001) with an adjusted r? of 43.1% (Fig. 5a, b). These results suggest that G. minimum numbers increase with increasing soil depth where CoMpe Non isata mini- mum but decreases thereafter. The latter observation is probably due to th petition from perennial vascular plants, which are more prevalent on deeper soils. Vegetation Correlations Large areas of the glade’s sandstone bedrock are covered with dense bryophytes and it was expected that cover measurements would demonstrate that bryophytes were spread- ing into G. minimum populations. Although bryophytes were negatively correlated with G. minimum numbersat the subplot scale (1; = -0.23, P < 0.001), they were not at the large plot scale. Similar results were observed with lichens and G. minimum numbers in that there was a negative relationship at the subplot scale but not at the large plot scale (Table 1). Although there is some discontinuity among scale with the latter two groups, there is a strong negative association (r; = -0.78, P < 0.001) between cryptogams (combined cat- egory of bryophytes and lichens) and G. minimum numbers (Fig. 5f and Table 1). The latter may suggest that cryptogam competition is a factor in depressing G. minimum populations. A positive association occurred with coverage by vascular plants and G. minimum (r5= 0.19, P< 0.001) at the small plot scale but not at the next scale (Fig. 5e and Table 1). It may be that through time G. minimum has persisted only on those areas of the glade that are subjected to the same periodic disturbances that make it difficult for mosses and vas- cular plants to become established. Lichen cover declined steadily from 35.1% in 1994 to 10.9% in 1998. Perhaps this aspect of the monitoring would show better statistical trends if conducted over a longer period and with more detailed data collection with respect to the species of vascular plants and cryptogams across population sites. Certainly the movement of soil over the bedrock would explain the erratic variation in numbers of subplots dominated by bare soil or bedrock. There are similarities between G. minimum population dynamics and winter desert annuals in that there are large population num- ber changes from year to year. Guo et al. (2002) discussed dramatic fluctuations in abun- dance, productivity, and diversity among desert winter annuals. Furthermore, the popu- lation dynamics of Vulpia ciliata Link. has been well studied (Watkinson et al. 2000; Watkinson 1990; Watkinson & Davy 1985; Watkinson & Harper 1978). This annual grass, 1154 BRIT.ORG/SIDA 22(2) ¢ ] 60 | . < 1 2 & ie a | € | - 1, = 0.16, p< 0.00 b- 2 & | & 50 | | Qa iS) 3 il e° 28 40 E 40 a © 5 5 oD 30 ° | 2 . | = 0 | c 8 g 20 & 20 Ec 3 360 10 | Q 10 zo | 8 0 0 l Soil Depth (mm) Soil Depth (mm) 80 : 80 _ ; | | r, = 0.46, p< 0.05 Cl | r= -0.12, p< 0.05 d | | | ° ej. wo 60 | oom ce 6 oe @ c a | a o& 40, - 40 | ee @ Gpareee @ eo ec aupe e | Pal > 7 CUED? ER ee eee e | Ss no 2 3 =O a5 | R 20 8 Of SE oeoe | wo (eo) | > 0 | F 0 | Soneeeene = aoe e e be om oe = : 0 40 60 80 100 0 40 60 80 100 Soil Depth (mm) Soil Depth (mm) | = t 60 ETS —Vv. < . > . = 60 r, = 0.19, p< 0.001 e € r, = -0.31, p =0.014 f 50 E 50 ! E & E 40 & 40 *. £ E |, ! . E 30 sia i 5 2 27H, : Q 20} |e , : o o oS H ° ‘ ra) 1 S 105 | | ° ye Qo 10) Bs 8 : o 0 | | j sos O 95! . 0 0 40 60 80 10 0 20 40 60 80 Total Vascular Plant Cover Total Crytogam Cover 1,=-0.24,p<0.001 | § q@ a r. = -0.24, p< 0. # 60° | g 500 = FlintHills = h + se fe Collins E 50 | = i E | * § 400 \ © 40; *. ' iS nN \ 3) E ua \ Ea ; ‘ 8 300 / \ . \ 9° 8 oO / \ J: \ 8 / \ 7 » & 20 8 : : = 200 Lg ir : - i] . ry s / vA \ V/ \ Pace 10 : : 1° 100 ; \ 8 |i —i—_+__ Yi 0 . + 4 d aap ey spss : 0 eee near See ner anne es eee —_ 0 20 40 60 80 00 1998 1999 2000 2001 2002 2003 2005 Total Vegetation Cover Year Fic. 5. Scatter plots of G. minimum, soil depth (millimeters), other vegetation types, with ponding Sy rank correlation Site, Dade and St. Clair counties, Missouri). t indicates significance only at the large (1.0 m’) plot scale. SMITH AND ELY, ECOLOGICAL ANALYSIS OF GEOCARPON MINIMUM IN MISSOURI 1155 occurring on sandy heaths and warrensalong coastal England, has similarities to G. mini- mum populations in that population dynamics probably occur at very small scales. It has been shown that other vegetation types depress V. ciliata populations significantly. The coupled effect of all vegetation upon G. minimum populations further exempli- fies that the persistence of this taxon may be dependent upon periodic disturbances. There was a significant negative correlation with G. minimum numbers at the subplot level (rs =-0.24, P< 0.001) with the combined vegetation cover (cryptogams and vascular plants) (Fig. 5g and Table 1). Colonization Plots The three plots that were created in 1995 by removal of dense moss were not colonized by G. minimum during the study. Instead they were eventually recolonized by the moss, al- though there were several years following establishment when the habitat appeared suit- able for G. minimum. The fact that these plots were nearly surrounded by undisturbed areas of dense moss may have precluded the water-transport of G. minimumseeds to the plots. Krieger et al. (2003) studied the temporal dynamics of ephemeral plant communi- ties in seasonal rock pools on Ivorian inselbergs, monolithic outcrops of granites and gneisses along the Ivory Coast of Africa. These rock pools or depressions are typically several hundred years old and are seasonally wet with local edaphic that may be typically xeric with a thin soil layer. They observed that plant communities in sea- sonal pools tend to follow the ‘turnover hypothesis’ which states that species that occur in discrete subpopulations tend to be at dynamic equilibrium with extinction and colo- nization. The turnover hypothesis is a significant part of island biogeography theory (MacArthur & Wilson 1967). The authors also noted that the turnover rate of an annual, Cyanotis lanata Benth.(Commelinaceae), was higher than other species. Although turn- over was not studied here, the fluctuation in G. minimum population numbers from year to year may suggest that turnover does occur. The extent to which it may occur needs further study. Furthermore, Krieger et al. (2003) also examined physical characteristics of depressional pools including area, mean depth, volume, water depth, and precipita- tion for these rock pools. It seems that distance between pools limits the ‘rescue effect’, or the migration of a BeEWeen poole G. minimum gene flow among and between populations and be ascertained to determine if these populations are indeed isolated or have metapopulation characteristics To effectively establish G. minimum in additional plots, it is possible to move seeds into plots following soil disturbance. This has been done successfully at a highway miti- gation site in St. Clair County, Missouri (Smith & Gardner 1999) by moving soil in the summer from areas where G. minimum had grown during the spring. It would seem pru- dent, however, to select establishment sites that are not subject to rapid recolonization by competitive plant species or offset recolonization by competitors with sufficient distur- bance. The current study casts doubt on the efficacy of relying on an existing G. mini- mum seed bank in the created plots or on the rapid, natural dispersal of seeds into the newly-available habitat. Climatic Effects An aspect of this study that needs further work is the relationship between local and regional weather conditions and G. minimum populations. A graphical comparison of G. minimum populations between Flint Hill Glades and a relocated population, St. Clair County, Missouri, (Smith & Gardner 1999) shows a high degree of symmetry among popu 1156 BRIT.ORG/SIDA 22(2) lation numbers among years, indicating that precipitation and or temperature may play a key role (Fig. 5h). The differences between populations in particular years (2001 to 2002) may indicate that local precipitation and or temperature events may also influence geo- graphically distinct populations. Weather data should be collected on each site for most accurate correlations because very local drought conditions at presumed critical life stages (fall germination of seed and early spring growth) would not be captured by available regional weather station measurements. Management Implications The long-term monitoring of G. minimum at Flint Hill Glades ecosystem provided in- sight on effective management of this threatened species. There is a negative association of cryptogams and the population of G. minimum as evidenced by the Spearman rank correlations (Fig. 5f and Table Ll). Furthermore, G. minimum seems to utilize habitats where there is periodic soil disturbance and minimal competition from mat-forming mosses or perennial vascular plants. The level of soil disturbance necessary to keep water-carried sand moving into bedrock depressions will be difficult to quantify. Geocarpon minimum population bers appear at least partially dependent upon the amount of soil present (Fig. 5a, b; Table 1) as indicated by the results of the analysis of variance and type II regression, but there are other unknown factors that affect G. mini- mum numbers. Some level of grazing by cattle may benefit the plant by maintaining ar- eas of loose sand with minimal plant competition, but excessive trampling or nutrient deposition would be a concern. Although it seems that many of the moss-dominated areas of the glade are currently unsuitable for G. minimum, we have little information on how abundant or widespread G. minimum was historically on Missouri’s sandstone glades. Fire suppression following settlement of the region has nearly eliminated a critical pro- cess that may have previously altered the ratio of bare soil to heavily-colonized soil. Management recommendations include providing periodic soil disturbances to the glade to assure that loose, sandy soil is available for movement by surface water. The peri- odic movement and accumulation of sand into shallow bedrock depressions should facili- tate the development of suitable G. minimum habitat. Further studies are needed to address the frequency and type of disturbance required to achieve this result. More work is needed on the autecology and synecology of this taxon and associated species at other G. mini- mum suitable habitats. In addition, the physical envi ent needs to be characterized, as well as local and regional climate. It is encouraging that G. minimum was more abundant in the plots at this study site in 2003 than during any of the previous nine years and that the population is capable of rebounding following a year with little seed production. ACKNOWLEDGMENTS We thank the following persons for their assistance with monitoring: Mike Skinner, Ann Wakeman, John Hays, Dorothy Butler, Craig Tenbrink, Indi Braden, Deborah Renek, Matt Parker, Teresa Cochran, and Chris Wieberg. We also would like to acknowledge the U.S. Army Corps of Engineers, Stockton Project Office for permission to study Geocarpon at Stockton Lake. George Yatskievych, Paul McKenzie, Harold W. Keller, and Lori A. Dameron provided invaluable comments on a draft of the manuscript and Brad Jacobs and Nadia Navarrete-Tindall provided the Spanish translation of the abstract. SMITH AND ELY, ECOLOGICAL ANALYSIS OF GEOCARPON MINIMUM IN MISSOURI 1157 REFERENCES Dausenmire, R. 1959. A canopy coverage method of vegetation analysis. NorthW. Sci. 33:43-64. Guo, Q., J.H. Brown and TJ. Vatone. 2002. Long-term dynamics of winter and summer annual commu- nities in the Chihuahuan Desert. J. Veg. Sci. 13:565-574. Kerry, E.L., J.R. Sincuurst, and S. Cook. 2004. Geocarpon minimum (Caryophyllaceae), new to Texas. Sida 21:1165-1169. Kriecer, A., S. Poremeski, and W. BarTHLoTT. 2003. Temporal dynamics of an ephemeral plant community: species turnover in seasonal rock pools on Ivorian inselbergs. Pl. Ecol. 167:283-292. MacArtHur, RH. and E.O. Witson. 1967. The theory of island biogeography. Princeton University Press, Princeton. Mackenzie, K.K. 1914. A new genus from Missouri. Torreya 14:67. McInnis, N.C., L.M. Smith, and A.B. Pittman. 1993. Geocarpon minimum (Caryophyllaceae), new to Louisi- ana. Phytologia 75:159-162. Missourl Dept. oF ConservaTION. 2006. Missouri Natural Heritage Database. Resource Science Division. Jefferson City, Missouri. Moraan, S. 1986. A study of a population of G. minimum in Missouri. Unpublished manuscript. Mis- souri Dept.of Conservation, Jefferson City. Newson, PW. 2005. The terrestrial natural communities of Missouri. Revised Edition. Missouri Natural Areas Committee. Jefferson City. Nienaser, M.A. 2005. Geocarpon Mackenzie. In: Flora of North America Editorial Committee, eds. Flora of North America north of Mexico, Vol.5 Magnoliophyta: Caryophyllidae, part 2.Oxford Univ. Press. New York. Pp. 148-149. Orzett, S.L.and E.L. Brioces. 1987. Further additions and noteworthy collections in the flora of Arkan- sas, with historical, ecological and phytogeographical notes. Phytologia 64:81-164. Quinn, G.P. and MJ. Keouct. 2002. Experimental design and data analysis for biologists. Cambridge University Press, United Kingdom. Smit, T.E. and J.E. Garpner. 1999. Geocarpon successfully relocated at a western Missouri site. Ecol. Restoration 17:91-92. Sokal, R.R.and F.J.Ronir. 1995. Biometry. 3 Ed. W.H. Freeman & Co. New York. Somers, P, L.R. SmitH, PB. Hamet and E.L. Brioces. 1986. Preliminary analyses of plant communities and seasonal changes in cedar glades of Middle Tennessee. ASB Bull. 33(4):178-192. THurMAN, C.and E.E. Hickey. 1989. Status report on G. minimum Mackenzie. Provided under contract to the U.S. Fish and Wildlife Service, Southeast Region, Atlanta, Georgia. U.S. Fish NO Wicowire Service. 1987. Endangered and threatened wildlife and plants; determination of threatened status for G. minimum (Geocarpon). Fed. Register 52(115):22930-22933. U.S. FisH AND Witoure Service. 1993. Recovery plan for G. minimum Mackenzie. Atlanta, Georgia. WATKINSON, A.R., R.P. FreckteTon, and L. Forrester. 2000. Population dynamics of Vulpia ciliata: regional, patch, and local dynamics. J. Ecol. 88:1012-1029. WATKINSON, A.R. 1990. The population dynamics of Vulpia fasciculata: a nine year study. J. Ecol. 78: 196-209. Warkinson, A.R.and AJ. Davy.1985.Population dynamics of salt marsh and sand dune annuals.Vegetatio 62:487-497. Warkinson, A.R.and J.L. Harper. 1978. The demography of a sand dune annual: Vulpia fasciculata. 1.The natural regulation of populations. J. Ecol. 66:1 5-33. Zar, J.H. 1999. Biostatistical analysis. 4"" Ed. Prentice Hall, NJ. 1158 BRIT.ORG/SIDA 22(2) Book REVIEW Ria Loonuizen. 2006.On Chestnuts: The Trees and Their Seeds. (ISBN 1-903019-832-3, pbk.). Prospect Books, Allaleigh House, Blackawton, Totnes, Devon, TQ97DL, U.K. (Orders: www.oxbowbooks.com, 1-800-791-9354). $17.95, 136 pp., 51/2" x 71/4". “Chestnuts roasting by an open fire...” We are all familiar with the Bing Crosby song White Christmas, but few of us are familiar with the history and lore of the chestnut. Growing up in North Texas, I did not see a chestnut tree until | was an exchange student in Northern Ireland. The huge tree with its gigantic nuts made quite an impres- sion when I came upon it in the dark Irish woods. Ria Loohuizen, a translator living in Amsterdam and the au- thor of The Elder, has written a lyrical account of the horse ose and the Sweet chestnut, complete aus Wee oy 1 i } rifal f -y L delight. T] ite book is divided i ti he eecistae Aes¢ ulus hippocastanu and cee i on the European sweet chestnut, Castanea sativa. It is full of interesting facts. For example, the horse chestnut may have gotten its common name from the horseshoe-like shape of its leaf scars a 7. genus name Castanea is probably derived from the name of a city in Turkey. The sweet chestnut is native to southern Europe, the Caucasus, and North Africa. It is believed to have been brought to Europe by the Celts. A significant section of the book is devoted to the culinary applications of pes sweet bee ees is fous only nut — nalen levels of vitamin C, according to the author. TI ts, pumpkin chestnut soup, Roman-style csinnd with i ntils chestnut sie chestnut cookies, and te chest- nut poultry stuffings. T] recipe for a hair conditioner made from horse chestnut leaves. Other interesting tips abound: Stich as using crushed horse chestnuts to keep insects out of potted plants. Recently, the horse chestnut has gained attention as the source of a treatment for chronic venous insufficiency (Suter et al. 2006). The American chestnut tree, Cast dentata, has been a beloved fix- ture in the forests of the east coast of the United States, but it has been decimated by chestnut blight, a fungal disease imported from Asia. Let us hope that this book is not a eulogy, but rather an ode to the remarkable chest- nut.—Marissa Oppel, MS, Collections and Research Assistant, Botanical Research Institute of Texas, Fort Worth, TX, 76102-4060, U.S.A REFERENCES Suter, A., S. Bommer., and J. RecuNner. 2006. Treatment of patients with venous insufficiency with fresh plant horse chestnut seed extract: a review of 5 clinical trials. Adv Ther. 23:179-190. SIDA 22(2): 1158. 2006 MOUANTITATIVE STUDY OF STYLES AND AGEIENES OF TERMINAL AND BASAL FLOWERS OF SCHOENOPLECTUS HALLII (CYPERACEAE), A RARE PLANT SPECIES OF TRANSIENT WETLAND HABITATS Marian Smith, Sara Ammann, Nancy Parker — Paige Mettler-Cherry Department of Biology ee of Biology Southern Illinois University indenwood Universi se Edwardsville Illinois 62026, U.S.A. St. Bis Missouri 63301, ABSTRACT Schoenoplectus hallii (A. Gray) S.G. Smith (Cyperaceae), a rare plant restricted to wetland habitats, is of conser- vation concern throughout its range. ee of the species recently has been clarified; however, quantitative i Because of descriptions of achene and flower morpholo | d life history its scarcity and the transient nature of ee any ee -scale study of the species will require the recovery of achenes from bulk soil samples and the identification and separation of the dimorphic achenes. The objec- tives of this study were to eves, gency and photograph Ge two mechs pes to crane we size pane mor- phological differences that wi I g f vena in styl phology and ach ithin and amcne, 12 populations in four states. ee each achene type ifi ly in size among populations land basal achenes are statistically ane and the range of sizes within each achene type is ae than has been previously reported. Terminal paid are significa smaller in length, width, beak length, mass and surface area than basal achenes, and not in surface ridging. Differences in style morphology are distinct: terminal flower styles are ae bifid and consistent in shape, while basal flower styles, which are six times longer Een me spies are trifid witha wide variety of branching pea Terminal and basal achenes can be sey ly from bulk soil ples using ase oc soil sieves. The vis | provided in this study will facilitate the collection ot ntification of eae and ey achenes of S. hallii from plants, soil and wildlife. RESUMEN Sch lectus hallii(A. Gray) S.G. Smith (Cyperaceae) ] ingida a habi le] humed es paras ser consetvadaie en ieee su area. La taxonomia de la oad se ha devneads recientemente; sin embargo, y la morfologia floral son incompletas y falta informacion de su ciclo vital. penis a su escasez y ala earnyeleaa transitoria de sus poblaciones, as estudio a ee a de especie requerira la ee de ee a de grandes muestras de suelo y u identificacion y separacion de | Por lo tanto, los objetivos a este estudio Pein separar, identificar y topralae os dos tipos de aquenio: cuantilica el tamano us eHocnGa mo PICeIee que seran utilesen | basales; y 1en la morfologia del mn y mans del cogeae en y entre 12 poblaciones en cuatro estados. Aun cada tipo de aquenio varia significativamente en tamano entre poblaciones, ms diferencias de tamanio entre aquenios terminales y basales son significativas estadisticamente, y el rango de tamano para cada tipo d e lo que se habia indicado previamente. Los aquenios terminales son sa teaeganicne mas ee en longitud, anchura, longitud el pico, masa y area de su superficie que los aquenios basales, y hay diferencias notables en las costillas de B supetticie Las enerenclas en ae morfologia del estilo son oe oe So de las flores Se son f] bifidos ey onstante | basales. que son seis veces ] +f: 1 Fee | ] 7] f] 7 ] rs | iB a ee Le | vValLICQGad ae } precision y de un modo ee a partir de muestras de af suelo Slee usando una serie de Subse para suelos. Las visuales y cuantitativas aportadas en este estudio facilitaran la recoleccion e identificacion de aquenios recarnales y basales de S. halliia partir de plantas, suelo y de la naturaleza. SIDA 22(2): 1159- 1173. 2006 1160 BRIT.ORG/SIDA 22(2) INTRODUCTION Schoenoplectus hallii (A. Gray) S.G. Smith is a member of the family Cyperaceae. Asa Gray (1863) described the species as Scirpus hallii A. Gray, based on specimens collected in Illinois. Recent work by Smith (1995) and Smith and Yatskievych (1996), however, has emphasized the breakup of the polymorphic supergenus Scirpus sensu lato and resulted in the acceptance of Schoenoplectus hallii as the appropriate binomial for the species (Smith 1995, 2002). The known distribution of S. hallii prior to 1973 extended from Massachusetts to Wisconsin and lowa on its northern boundary, westward to Kansas and Nebraska, and south to Georgia. The number of states in which populations existed declined from nine in 1973 to six in 1997 (McKenzie 1998). It has apparently been extirpated from Massachu- setts and Georgia, and is now restricted to the Midwest. In 2000, S. hallii was reported from 14 sites in Oklahoma (Magrath 2002); however, the conservation status of the spe- cies at these sites is unclear due to the recent discovery of putative hybrids (S. hallii ~ S. saximontanus) at some of the sites (Smith et al. 2004). In 2002, surveys for S. hallii re- vealed new populations in several states, including Indiana (M. Homoya, pers. comm.) and Missouri (pers. obs.) and Texas (O’Kennon and McLemore 2004). In years not suit- able for germination, the species may persist only in the seed bank (McKenzie 1998); there- fore, it is difficult to obtain accurate estimates of the mE! of wae populations in any given year. Although population number an gra tion are not clearly defined, S. hallii is considered to be of conservation concern in every state in which it occurs (Beatty et al. 2004). Schoenoplectus hallii is restricted to wetland habitats (Swink & Wilhelm 1994; McClain et al. 1997) in areas characterized by fluctuating water levels (Ostlie 1990; Ostlie & Gottlieb 1992; Robertson et al. 1994). The species is thought to have a persistent seed bank (Ostlie 1990; Ostlie & Gottlieb 1992; Robertson et al. 1994) that may contribute to its potential for population regeneration. Achenes germinate sporadically, depending on the availability of wet, exposed habitat (McKenzie 1998), resulting in wide fluctuations in population number and size from year to year (Chester 1988; Robertson et al. 1994, McClain et al. 1997). It has been suggested that because of loss of suitable wetland habi- tat, management of S. hallii sites is necessary to ensure the species’ continued existence (Bowles et al. Cee Robertson et al. 1994). Scl tus halliiexhibit hicarpy (Bruhl, 1994), as do other members of the section Sumpini (ie, S. erectus and S. saximontanus in North America), and achenes pro- duced in multi-flowered spikelets near the tip of stems (hereafter designated as “termi- nal” achenes or flowers) have been illustrated and/or described by various authors (Gleason & Cronquist 1963; Mohlenbrock 1963; Steyermark 1963; Radford et al. 1964; Mohlenbrock 1976; Kolstad 1986; Yatskievych 1999; Smith 2002). Although achene size is estimated in the taxonomic literature, and varies among authors, a quantitative measure of variation in achene size from a representative sample of populations across the range of the species is lacking. Basal achenes, which are produced by solitary pistillate flowers enclosed within the leaf sheaths at culm bases, are less adequately illustrated and de- scribed than terminal achenes. With the exception of a photograph of one basal achene (Schuyler 1969) no photographs or illustrations have been published. In addition, no quan- titative description of variation in basal achene size within and among populations ex- ists. Quantitative data establishing the size and morphology of the basal achenes are im- portant for the correct separation and identification of seed bank components and are SMITH ET AL., SCHOENOPLECTUS HALLI] ACHENES 1161 essential for the development of life history studies that estimate the relative reproduc- tive contribution of basal achenes to population size. In early summer before achenes have been produced, style lobe number is the mor- phological character most useful in separating S. hallii from S. saximontanus, because of their similar vegetative appearance. As far as we are aware, there has been no large-scale study of style size and morphology in S. hallii. Recently, S. hallii and S. saximontanus were found growing in mixed populations in Oklahoma (Magrath 2002); therefore, it would seem prudent to determine if style lobe number is constant throughout a number of widely distributed populations of S. hallii in areas where S. saximontanus is not sym- patric to confound the issue. The objectives of this study were 1) to develop a simple procedure for separating, collecting and identifying achenes from bulk soil samples; 2) to photograph the two achene types; 3) to quantify the size and morphological differences that will be useful in separating terminal and basal achenes; and 4) to determine the range of variability in style morphology and achene size within and among 12 populations in four states (IL, IN, KY, and MO) MATERIALS AND METHODS When possible, living plants were collected in situ and transferred to the greenhouse at Southern Illinois University, Edwardsville, IL. Plants were potted in a 75%-sand:25%-pot- ting soil mixture in 10 x 10 x 11cm pots, covered with a plastic bag to decrease transpira- tional water loss, and placed in flats in 5cm of standing water. The organic content of the mixture (%OM = 2.5%) was approximately that of soil at the study sites (mean %OM = 2.6+0.5%, analyses conducted by Alvey Labs, Belleville, IL). Plants were given identifying numbers and mature terminal acl (and basal achenes, when present) were collected and saved for measurement. Otherwise, basal achenes were collected as they matured. Site names, acronyms, approximate locations (exact locations are not listed to pro- tect population sites) and dates of collection are listed in Table 1. At three sites that were without plants (SP FO and KY), but were known to have had populations of S. hallii within the past 5 years, soil cores were collected to provide a seed source. In all cases, plants or cores were collected at regular intervals along transects from across the known extent of the population. To ensure accurate identification of achene types, initial collections were of terminal and basal achenes that were attached to the parent plant. After achene iden- tification was established, others were collected from soil samples. Whenever possible, pairs of terminal and basal achenes were selected, using a random number table, from 15 plants and photographed, measured and weighed in the laboratory. For sites without plants, soil sieves (U.S. Standard by Fisher Scientific Co) #16 (1.18 mm mesh), #18 (1.00 mm) and #20 (0.841 mm) were used to separate basal and terminal achenes. Identifica- tion was verified using a dissecting microscope. As some achenes that appeared to be healthy and mature fragmented when pressed between the thumb and forefinger, all achenes were given this preliminary test before being measured (Baskin et al. 2003). Achene photographs were made using a Nikon CoolPix 995 digital camera mounted on a Nikon dissecting microscope (Model SMZ800) fitted with a fiber optic ring-light. Length, width, aud beak measurements were taken at a magnification of 40X using a Leitz comy pe (Model Laborlux S) anda Bausch and Lomb ocular microme- ter that was calibrated with a Leitz stage micrometer. Achene beak length was measured from the point where tangential lines drawn at the top of the achene and alongside the ~=— 1162 BRIT.ORG/SIDA 22(2) Tasie 1.Names, acronyms and county and state locations of 12 Schoenoplectus hallii sites and types of samples collected for this study. Site State County Date and Sample Collected Ebken (EB) IL Mason 2000 dist and soil Fornoff (FO) IL Mason 2000 Soil only Sand Pond (SP) IL Mason 2000 a | Wemker (Wk) IL Mason 2000 Plants and soil Indiana (IN) IN Daviess 2002 Plants and soil Kentucky (KN) KN Christian 2000 Soil only Baptist Camp (BC) MO Scott 2000 Plants and soil Howell County (HC) MO Howell 2002 Plants and soil Petite Isle (Pl) MO Scott 2002 Plants and soil Sherer (SH) MO Scott 1999-2001 Plants and soil Waterman (WM) MO Scott 2002 Plants and soil West Vaco (WV) MO Scott 2002 Plants and soil beak intersected (Fig. LD. As beak length varied considerably, but contributed little to achene area, achene surface area was calculated as width x (achene length-beak length). Mass was determined using a Mettler analytical balance (Model AT 261 Delta Range). To illustrate the contrast in size, one pair of terminal and basal achenes were photographed with attached styles (Fig. 2A). To illustrate variability in basal and terminal achene mor- phology, three of each achene type were photographed in the same field-of-view (Fig. 2B). Also, three achenes of each type were photographed in cross-section (Fig. 2C) and one pair of achenes was photographed showing the achenes in frontal (Fig. 2D) and pro- file (Fig. 2E) views. Styles were excised from flowers of each type, floated on water and measured. For were made on terminal styles from 15 plants from eight populations, but for basal styles, which are scarce and difficult to collect intact, measure- ments were limited to 15 plants from one population. Others from eight additional popu- lations were observed for style morphology, but not measured. To illustrate some of the aera in style morphology observed in basal flowers, drawings were made to scale (Fig. 3B- rae data sets for terminal and basal achenes were combined for analysis, vari- ances were significantly different (P<0.001) between achene types for all dimensions; therefore, analysis by two-way ANOVA was rejected and data for each achene type were analyzed separately by one-way ANOVA. Equal variance was verified using Levene’s Test and data were log transformed, when necessary, to ensure normal distribution. Compari- sons of mean values for achene characteristics among sites were determined by Bonferroni's t-test. Terminal style measurements, for which data could not be success- fully transformed, were analyzed by Kruskal- Wallace one-way ANOVA on ranks, followed by Tukey’s t-test for comparison of means. Pooled differences in dimensions between achene types were determined by t-test, assuming unequal variances. When matched pairs of terminal and basal achenes were available within a population, statistical com- parisons were made using a paired-samples t-test. All statistical procedures were calcu- lated using SPSS 11.4 (SPSS, Inc. 2002) and are in accordance with Sokal and Rohlf (1981). Principle ANOVA statistics, means, and P-values) were reported when differences were statistically significant. statistical analysis SMITH ET AL., SCHOENOPLECTUS HALLII ACHENES 1163 1mm Fic. 1. Method for determining beak length in basal (left) and terminal (right) achenes of Schoenoplectus hallii. Drawn to scale as indicated in the figure. RESULTS Comparisons among populations Terminal achenes.—Terminal achenes varied significantly in every dimension (length, width, beak length, mass and surface area) among populations (Table 2, Fig. 4+A-E). Achenes from HC were significantly larger than those from all other populations, with the exception of beak length (Table 3). There was a wide range in terminal achene size and mass (Table 4), even with the removal of the extremely large mean values for HC achenes, which, when included, raised the upper limit for every character. Terminal styles.—All terminal styles examined were bifid, with the exception of the one 4-parted style from HC. There was a much wider range in style length compared to the range for any achene character (Table 4). There were significant differences in style length among populations (Table 2): the styles of the WK population were longer than those from any other site and the styles from the SH site significantly longer than all except those from WK (P<0.05). There were no significant differences in style lengths among the other six sites. Basal achenes.—Basal achenes were more varied in shape than terminal achenes (Fig. 2B); however, there were significant differences among populations for every basal achene character measured (Table 5). Unlike the case with terminal achenes, in which those from HC were significantly larger than those from all other sites, no single population had consistently larger basal achenes (Fig. 5A-E, Table 6). Comparisons between terminal and basal achenes and styles.—Pooled means for every 1164 BRIT.ORG/SIDA 22(2) ean didi Fi. 2A-E. A. Basal (too) and Saeaer ar ; ere iiaiceylee Headlavial view or nasalad terminal achenes ofS. hallii; C. cross sectiona d basal ac f S. hallii; D. close-up of terminal (left) and basal achenes of 5. hallii; and E. profile view of basal (left) and terminal achenes of S. Aallii. Bars on all photographs are 1 mm in length SMITH ET AL., SCHOENOPLECTUS HALLII ACHENES 1165 mm Fic. 3 AD. Representative styles of A. terminal and B-D. basal flowers of Schoenoplectus hallii. dimension measured were significantly larger for basal compared to terminal achenes (Table 4, P<0.001, df = 289 for each comparison). Within-population comparisons for the nine sites for which achene pairs were available indicated that in seven populations (EB, WK, IN, BC, SH, WM, WV), basal achenes were significantly larger for all dimensions. At HC, area was not significantly different between terminal and basal achenes (P<0.536, df 1166 BRIT.ORG/SIDA 22(2) Tasie 2. Summary ANOVA statistics for terminal achenes and styles of Schoenoplectus hallii from 12 population sites Character Source of Variation $s df MS F P Length Between pops 2457 11 0.223 28.656 <0.001 Within pops 1.310 168 0.008 Width Between pops 3.24] 11 0.295 45,384 <0.001 Within pops 1.091 168 0.006 Beak length Between pops 0.056 1] 0.005 10.136 <0.001 Within pops 0.085 168 0.001 Area Between pops 23.54 1] 2132 61.447 <0.001 Within pops 5.829 168 0.035 Mass Between pops 4.092 1] 0.372 26.263 <0.001 Within pops 2.380 168 0.014 Style Between pops 24.163 1] 3.452 15.980 <0.001 Within pops 27.193 168 0.216 = 11), and at PI, width for terminal and basal achenes did not differ significantly (Ps0.348, = 14) All terminal achenes examined had prominent transverse ridging extending the width of the achene surface as recently reported for S. hallii achenes from Oklahoma populations (Magrath 2002). In contrast, transverse ridging was less conspicuous and did not extend the entire width of basal achenes, which had obvious vertical ridges. The cross-sectional shape of achenes was generally as described by Yatskievych (1999) (“un- equally biconvex in cross-section, sometimes slightly concave on 1 side” and “unequally 3-angled,” for terminal and basal achenes, respectively); however, both achene types oc- casionally varied from these descriptions. Some terminal achenes were plano-convex as illustrated in Figure 2 C. Styles of basal flowers were approximately 6 times longer than those of terminal flowers (Fig. 2A, Table +), and exhibited a wide variety of structural morphologies. Styles of all terminal flowers examined were bifid (Fig. 3), with the exception of one style from HC that had four style lobes (not shown). All styles from basal flowers were trifid with a variety of branching patterns, some of which are illustrated in Fig. 3B-D. DISCUSSION AC. Martin,a U.S. Fish and Wildlife Service biologist stated that when identifying achenes encountered in the stomachs of wildlife, his staff found the description of Scirpus achenes in the taxonomic literature was unhelpful in separating species (Martin 1943). Although he stated that achenes of some species are distinctive enough to present no problem in identification, “the difficulty and danger involved in attempting to distinguish some of the other species important to wildlife frequently necessitated noncommittal conserva- tism.” These troublesome species included what is now known as Schoenoplectus hallii, and, although descriptions of its achenes are far more comprehensive now aes in 1943, no taxonomic reference adequately describes the variation of achene size and shape ex- isting within the species. Our data clarify and illustrate the distinctions between achene types and expand the range of achene dimensions cited in the current published litera- ture. While several publications (Beetle 1942; Magrath 2002; Schuyler 1969; Smith & Yatskievych 1996; Smith 2002) stated clearly that S. hallii exhibits amphicarpy, many SMITH ET AL., SCHOENOPLECTUS HALLII ACHENES 1167 A. : é aia D 3.0 2.5 Se 1.5- — E 2.0 “Ee Ps = = 1.0 4 1.5 Pi x 15-4 pret = 08 G&D £ £ — — = w S 10- Oe <8 = = 0.4 0.5 0.2 0.0 0.0 0.16 4 5 0.14 4 7 0.12 4 4 = 0.104 = <= = a £ 5 0.08 + 2 <~ 0.06 — 20 s : co 0.04 4 0.02 0.00 tT syle Tose ay T Lyd aaa EB FO SP WK IN KN BC HC PI SHWMWV EB FO SP WK IN KN BC HC PI SC WMWV Site Site £ Ch. J Wes €, 12 populations, Fic. 4 A-F. A. Length, B. width, C. beak length, D. area, and E. mass of | dF. style length of inal f] f S. hallii from 8 populati E + SE of 15 measurements. did not (Mohlenbrock 1963; Radford et al. 1964; Steyermark 1963; Kolstad 1986; Rolfsmeier 1995). Of those that recognize variation in floral types, only three provided estimates of basal achene size (Schuyler 1969; Yatskievych 1999; Smith 2002), and none presented data to quantify differences between terminal and basal achenes, nor any measure of the vari- ance in size within achene type across a range of populations. Although there is eclee apreenent between our achene-size data and published reports, there are notable differences. With the exception of the three cases where size of basal achenes was specifically aesed (ie., Schuyler 1969; Yatskievych 1999; Smith 2002), we assumed that all published estimates referred to terminal achenes. Six authors listed the following estimates of length for terminal achenes: 1.3-1.7 mm (Yatskievych 1999), 1.5-2.0 mm (Mohlenbrock 1976); “up to” 1.5mm (Kolstad 1986); 1.3-1.5 mm (Gleason & Cronquist 1963); “less than” 0.4 mm (Robertson et al. 1994); and 1.3-1.7 mm (Smith 2002). There are three estimates of achene width in the literature: 1 mm (Beetle 1942), 1168 BRIT.ORG/SIDA 22(2) Taste 3. Significant differences in mean length, width, beak length, beak length and mass of terminal achenes among 12 populations of Schoenoplectus hallii. All P values calculated by Bonferroni's t-test. Length P Beak P Area P Width P Mass P HC vs All <0.001 INvs FO <0.001 HC vs All <0.001 HC vs All <0.001 HC vs All <0.001 FOvsSP <0.001 vs SP <0.001 FOvs SP <0.001 SHvs SP S0.014 SPvsAll 0.040 vs SH <0.002. = vs SH <0.001 vs WK <=0.019 SHvsWV S0.018 INvsBC 0.007 0.002. - - INvs SP <0.001 HC vs EB S0.002 vs BC vs PI =0.009 vs BC <0.006 vsFO <0.001 vs Pl SO001I6 - - vs SH <0.001 vs SH <0.001 vs SP <0.001 vs SH <0.002 - vs WM <0.001 vs Pl <0.018 vs WK <0.003 vs WM =0.002 ~ - = vs WV <0.011 vs SH 0.001 vs WV 0.001 - - - - Taste 4. Pooled means + SE and range of values for length, width, beak length, area and mass for terminal (T) and basal (B) achenes, and styles of terminal achenes of Schoenoplectus hallii from 12 population sites. *Omit- ting extreme values from Howell County **Mean values of basal styles from SH only. T Mean T Range T Range* B Mean Brange Length 1.52+0.01 1.14-2.7 1.14-1.76 2.30+0.03 1.20-3.11 Width 1,32+0.02 0.88-1.88 0.88-1.50 1.43+0.01 1305-1.93 Beak length 0.11+0.00 0.04-0.22 0.04-0.18 0.34+0.03 0.11—-0.90 Area 1.88+0.03 1.12-3.32 1.12-2.42 2.81+0.04 1.64-4.32 Mass 0.60+0.01 0.23-1.13 0.23-0.95 1.68+0.04 0.85-2.83 Style 3.254+0.06 1.30-6.00 1.30-6.00 17.07+0.89** 12.00-22.00** 1.2-1.5 mm (Radford et al. 1964) and 1.2-1.3mm (Smith 2002). With the exception of the measurement reported by Robertson et al. 1994), which undoubtedly refers to beak length, as previously stated in Schuyler (1969), rather than achene length, all measures of terminal achenes are within a reasonable range; however, the incomplete dimensions given in any single source create uncertainty in identification or confirmation of achene identity. As we have demonstrated, size can vary significantly between sites, and in ex- ceptional cases (e.g., HC) the variation is so extreme that the achenes of S. hallii could be misidentified if size, as described in the existing taxonomic literature, was included in the criteria for identification. As it is not customary for authors of taxonomic literature to indicate the sample sizes upon which their estimates are made, it is possible that some are based ona single achene, although most cite a range in size, which implies more than one measurement. In any case, it is time that a clearer delineation of the range of dimen- sions in achenes becomes available in the published literature. Relying solely on information in the taxonomic literature, it is possible that some- one examining the seed bank of S. hallii could fail to recognize basal achenes as belong- ing to the species, and thus underestimate the potential population size or the contribu- tion of basal achenes to it. Only three authors described basal achene size: Yatskievych (1999) states that they are “slightly larger” than terminal achenes, Smith (2002) stated a range from 1.7-2.5 mm long and 1.0-1.3 mm wide, and Schuyler (1969) published a photo- graph of one terminal and basal achene pair. While the photograph is the most useful representation of the appearance of basal achenes currently available in the published literature, neither the verbal nor the visual description is adequate to understand the SMITH ET AL., SCHOENOPLECTUS HALLIT ACHENES 1169 Length (mm) in Width (mm) Beak length (mm) Area (mm?) Mass (mg) z 9 lati Each bar Fic. 5 A-E.A. Length, B. width, C. beak length, D.area, and E f basal act f Sct I t represents the mean + SE of 15 measurements. 1170 BRIT.ORG/SIDA 22(2) Taste 5.Summary ANOVA statistics for basal achenes of Schoenoplectus hallii from nine population sites. Character Source of Variation SS df MS F P Length Between groups 2.5755.929 8 0.322 5.537 <0.001 Within populations 102 0.058 Width Between groups 0.4141.969 8 0.052 2.680 0.010 Within populations 102 0.019 Area Between groups 0.3411.401 8 0.043 3.106 0.004 Within populations 102 0.014 Area Between groups 7.246 8 0.906 5.416 <0.001 Within populations 17.066 102 0.167 Mass Between groups 8.584 8 1.068 11.204 <0.001 Within populations 9.727 102 0.096 Taste 6. Significant differences in mean length, width, mass and beak length of basal achenes among nine populations of Schoenoplectus hallii. All P values calculated by Bonferonni's t-test. Length P Mass P BC vs IN <0.003 BC vs EB <0.001 Vs HC <0.001 vs IN 0.014 vs Pl <0.001 vs HC <0.001 Width Pe vs Pl <0.001 Pl vs WV =0.009 vs WV <0.001 Beak P Pl vs SH <0.001 EB vs HC <0.006 vs WV <0.001 magnitude and constancy of the differences between achene types, nor the potential of using size to separate achenes. Mean size and mass data, including values for the unusu- ally wide terminal achenes at the HC population, indicate that basal achenes average 15x longer, Ll« wider, 1.5~ greater in area and 2.8x heavier than terminal achenes (Table 4). Although data in the present study indicated that there was significant variation in morphology among basal achenes, they easily can be distinguished from terminal achenes, and at any site lacking S.saximontanusor S. erectus they can be identified easily to species. Except for the mixed populations in OK and KS, there are no known sites where S. hallii co-occurs with the species with which its basal achenes might easily be con- fused. Although we processed from 20-100 soil cores from each of the 12 sites to deter- mine the potential seed bank for S. hallii, we did not find any mature achenes of any other species that resembled S. hallii basal achenes. Various authors have noted that transverse or horizontal ridges are present on the surface of achenes of S. hallii (Beetle 1942; Gleason & Cronquist 1963; Magrath 2002; Radford et al. 1964; Schuyler 1969; Mohlenbrock 1976; Kolstad 1986; Robertson et al. 1994; Rolfsmeier 1995; Yatskievych 1999). Our observations confirm the presence of prominent horizontal ridges on all terminal achenes examined; however, they are absent or incom- plete on basal achenes, which have conspicuous vertical ridges. Although prominent horizontal ridges are limited to terminal achenes, this is universally attributed to achenes in S. hallii. Cross-sectional shape is often used to separate terminal achenes in S. hallii,described as “unequally biconvex in cross section” by Yatskievych (1999), from terminal achenes in SMITH ET AL., SCHOENOPLECTUS HALLII ACHENES 1171 S. saximontanus, which have a distinctly 3-angled shape (Yatskievych 1999). Our study corroborates the general regularity of this feature in S. hallii throughout the 12 popula- tions studied; however, as is the case with basal achenes, there are exceptions (Fig. 2C). Because all published studies of S. hallii are confined to the species’ taxonomy or occurrence aan a and rarity or abundance in various regions), which required no large ollection, no authors have suggested any methods for separating achenes from bulk soil ae The results of our study indicate that it is possible to separate terminal and basal achenes from soil samples using a series of soil sieves. Surface litter and a large soil fraction can be separated from achenes by using sieves of various sizes, depending upon Be Dip of soil and the nature of the litter. For example, if a sample ntain ic matter, one can use a #5 sieve to remove litter, a #14 sieve to remove coarse maarieles if the soil is sandy, and then sift the soil through sieves #16, #18 and #20 to separate achenes by type. The majority of the basal achenes will not pass through the #16 sieve, and the few that do are always retained by the #18 sieve. A small proportion of the terminal achenes remain in the #18 sieve (as was the case with many of HC achenes) with the majority of achenes passing through into the #20 sieve. In our study, none of the achenes passed through the #20 sieve into the next sieve. If the soil is predominantly silt or fine loam, all soil passes through the #20, which facili- tates the final collection and counting of achenes. If soil is composed of coarse sand, larger grains can be filtered out of the sample using the #14 sieve and finer particles of sand will pass through the #18 sieve; however, it is impossible to remove all coarse sand from achenes using sieves. Final separation of achenes from sand must be done manually. Al- though this final process requires some manual separation of achenes and soil using a dissecting microscope, the processing of bulk samples of soil for seeds is greatly facili- tated using the graduated sieve method described. For future seed bank and germination studies requiring large numbers of achenes, this method will be useful. Based on assump- tions of a long-lived seed bank, populations of S. hallii are often listed as “extant” if plants existed at the site during the previous 25 years (McKenzie 1998); therefore, seed bank stud- ies may be essential to the correct assessment of the species’ status. Unlike terminal flowers, which have uniformly bifid styles, with the exception of the single 4-parted style from HC, basal flowers have trifid styles as shown in Fig. 2A and 3B-D. Most references to the number of divisions present in S. hallii styles indicate the presence of bifid styles (Steyermark 1963; Kolstad 1986; Rolfsmeier 1995; Smith 2002), with only Yatskievych (1999) noting “stigmas 2 (rarely 3 in basal spikelets).” Although all basal styles had trifid branching, this only became apparent in some cases when the styles were floated on a film of water, which allowed shorter style lobes to separate from the main branch of the style (Fig. 3-D). Styles of basal flowers, as previously indicated by Yatskievych (1999), are longer than those of terminal flowers (Table 4, Figs. 2A and 3A- D). Our data indicate that this difference is significant (statistics not shown, see Table 4 for mean values) with no overlap in size between the two style types. Insummary, significant variation in achene size and shape occurs within and among populations; identification and separation of terminal and basal achenes can be accom- plished rapidly using a series of sieves; transverse ridging is not a regular or prominent feature of basal achenes; and in the vast majority of cases, a bifid style is a consistent character in the terminal flowers (we noted one exception). All basal styles examined were trifid. Data from this study provide visual and quantitative information that will facilitate the collection and identification of terminal and basal achenes of S. hallii from 1172 BRIT.ORG/SIDA 22(2) plants, soil and wildlife and contribute to an accurate assessment of the species’ conser- vation status. ACKNOWLEDGMENTS We thank the Missouri Department of Conservation, U.S. Fish and Wildlife Service and the Graduate School at Southern Illinois University, Edwardsville, IL, for financial sup- port for this study. We are also grateful toa number of colleagues and students who helped in making this study possible: Chairo Shaffer, Dan Free, Bethany Meinardii, Josh Knolhoff and Rasheda Johnson who helped with achene collection and identification; Xiaowen Wang who helped measure achenes; Michael Homoya who provided achenes and plants from Indiana, C.C. Baskin, J.M. Baskin and E. W. Chester for collecting soil samples in Ken- tucky; reviewers Paul McKenzie, Larry Magrath and Galen Smith for their helpful com- ments; and Paul McKenzie for comic relief and help in field collection of plants and achenes. REFERENCES Baskin, C.C,,J.M. Baskin, E.W. CHester, and M. Smit. 2003.Ethylene as a possible cue for seed germination of Schoenoplectus hallii (Cyperaceae), a rare summer annual of occasionally flooded sites. Amer. J. Bot. 4:620-627. Beatty, B.L., W.F. JENNINGS, and R.C. Rawunson. 2004. Schoenoplectus hallii (Gray) $.G.Sm. (Hall's bulrush):a technical ee aon assessment. USDA Forest Service, Rocky Mountain Region. http:// 222.fs.fed.us/r2/projects/sep/a nents/schoenoplectushallii.pdf. BeetLe, A.A. 1942. Studies in the genus or L.V. Notes on the section Actaeogeton Reich. Amer. J. Bot. 29:653-656. Bow es, M.L., M.M. DeMauro, N. Pavotic, and R.D. Hiesert. 1990. Effects of anthropogenic disturbances on endangered and threatened plants at the Indiana Dunes National Seashore. Nat. Areas J. 10 187-200 BauHt, J.J. 1994. Amphicarpy in the Cyperaceae, with novel variation in the wetland sedge Fleocharis caespitosissima Baker. Aust. J. Bot.42:44-448. Cuester, E.W. 1988. An unusual habitat type for three rare Kentucky plants. Trans. Kentucky Acad. Sci. 49:1-102. Gieason, H.A.and A. Cronauist. 1963. Manual of the vascular plants of the northeastern United States and Canada. D. Van Nostrand Co., Princeton, New Jersey. Gray, A. 1863. Manual of botany of the northern United States. 3" edition. New York. Koistap, O.A. 1986. Cyperaceae Juss, the sedge family. In: T.M. Barkley and R.L. McGregor, eds. Flora of the Great Plains. University Press of Kansas, Lawrence. Pp. 1059-1113. Macratn, L. 2002. Schoenoplectus hallii and S. saximontanus: 2000 Wichita Mountain Wildlife Refuge Survey. Oklahoma Native Plant Record 2:54-64. Martin, A.C. 1943. Achene characters in the genus Scirpus. Amer. Midl. Naturalist 31:250. McCiain, WE, R.D.McCiain, and J.E. Esincer. 1997. Flora of temporary sand ponds in Cass and Mason Counties, Illinois. Castanea 62:65-73. McKenzie, RM. 1998. Hall’s bulrush (Schoenoplectus hallii) status assessment. U.S. Fish and Wildlife Ser- vice. Columbia, MO. Mon ensrock, R.H. 1963. The Cyperaceae of Illinois: Scirpus. Amer. Midl. Naturalist 70:1-46. Montensrock, R. H. 1976. The illustrated flora of Illinois: Sedges—Cyperus to Scleria. Southern Illinois University Press, Carbondale. SMITH ET AL., SCHOENOPLECTUS HALLII ACHENES 1173 O’Kennon, RJ. and C. McLemore. 2004. Schoenoplectus hallii (Cyperaceae), a globally threatened spe- cies new for Texas. Sida 21:1201-1204. Ostuie, W.R. 1990. The Nature Conservancy's element stewardship abstract for Scirpus hallii (Hall's bulrush). The Nature Conservancy, Arlington, Virginia. Ostue, W.R. and S. Gortues. 1992. The Nature Conservancy's element global ranking form for Scirpus hallii. The Nature Conservancy, Arlington, Virginia. Raprorb, A.E.,H.E. AHLes, and C.R. Beit. 1964. Manual of the vascular flora of the Carolinas. University of North Carolina Press, Chapel Hill. Rosertson, K.R., L.R. PHituipre, and S.M. GEHLHAUSEN. 1994. The current status of Scirpus hallii_A. Gray, Hall’s bulrush, in Illinois. Illinois Natural History Survey Technical Report. Champaign. Rorsmeier, S.G. 1995. Keys and distributional maps for Nebraska Cyperaceae, Part 1:Bulbostylis, Cyperus Dulichium, Eleocharis, Eriophorum, Fimbristylis, Fuirena, Lipocarpha,and Scirpus. Trans. Nebraska Acad. Sci. 22:27-42. Scuuyter, A.E. 1969. Three new species of Scirpus (Cyperaceae) in the southern United States. Notul. Nat. Acad. Nat. Sci. Philadelphia 423:1-12. Smith, M., P. McKenzie, P. Mettter-CHerry, and G. SmitH. 2004. A putative hybrid of Schoenoplectus saximontanus and S. hallii (Cyperaceae) from Oklahoma. Sida 21:475-479. Smitu, S.G, 1995. New combinations in North American Schoenoplectus, Bolboshoenus, Isolepis and Trichophorum (Cyperaceae). Novon 5:97-102. Smit, S.G. 2002 enoenanicgas In: Flora of North American Editorial Committee, eds. Flora of North America, Vol.23.Mag | linidae (in part): Cyperaceae. Oxford University Press, New York. Pp. 44-60. Smith, S.G. and G. YatskievycH. 1996. Notes on the genus Scirpus sensu lato in Missouri. Rhodora 98: 168-179. Sokat, R.R.and FJ. RoHir. 1981. Biometry. 2nd Edition.W.H. Freeman and Co., San Francisco. SPSS, Inc. 2002. SPSS for Windows. SPSS, Inc. STEYERMARK, J.A. 1963. Flora of Missouri. The lowa State University Press, Ames. Swink, F.and G. WitHetm. 1994. Plants of the Chicago region. 4th Edition. Indiana Academy of Science, Indianapolis. Yarskievych, G. 1999. Steyermark’s flora of Missouri. Revised edition, Vol. 1. Jefferson City. 1174 BRIT.ORG/SIDA 22(2) Book REVIEWS Davip A. SLePER and JOHN M. POEHLMAN. 2006. Breeding Field Crops, Fifth Edition. (ISBN 0- 8138-2428-1, hbk.). Blackwell Publishing. 2121 State Ave., Ames, [A 50014-8300, US.A. and 9600 Garsington Road, Oxford, OX4 a UK. Otc: ed — QIAO, 515-292- 3348 fax 1-800-862- “6657, www.blacl I] Lcom, blackwell publishing.com,). $84.99, 392 pp., 448 b/w illustrations and photographs, 7" x 10". This book is intended for beginning students of plant breeding as well as more advanced students and profes- sional Plant prEedels The ns oe was published in oak “and since that time it has been one of the most One of the most significant wees ay in plant popular plant | the world.’ Daeeding since edition 4 (1995) has been cea ation - ens ne Be nomics”—using “tools developed by molecular biologists such as DNA marker I g p ple son ae genomes . —. of our crop plants.” Section L: What is Plant Breeding? Section II: The Genetic Basis of Ent ‘Breecig Section IIL: Tools of the Plant B gcl ber, ion, fertility-regulating mechanisms, biotechnology) Section IV: Methods in Plant Breeding Section V: Germplasm Resources for Breeding Crop Plants Sections VI through IX: Applications: Breeding field crops that are self-pollinated or vegetatively propagated; Utilizing hybrid and other miscellaneous breeding procedures Section X: Maintenance and Seed Production of Improved Cultivars —Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A. Rosin Wuite. 2006. Daphnes: A Practical Guide for Gardeners. (ISBN 0-88192-752-X, 978- 0-88192-752-8, hbk.). Timber Press Inc, 133 S.W. Second AVES Suite +50, Portland, OR 97204-3527, US.A.(Orders: www.timberpress.com berpress.com, 503- 227-2878, 1-800-327-5680, 503-227-3070 ao $34.95, 232 pp., color photos, 7" x 9". The book is written from the author's 30 years of experience in propagating and growing daphnes, both as a commercial horticulturalist and as a home gardener (in his “daphnetum”) in southern England. It is arranged in seven chapters: History, Taxonomy, and Morphology; Daphne Species; ao Hybrids; Propagation; Cultiva- tion; Pests oo“ Diseases; and aoe Use. Nei any he au a White's book is given to discussion of 36 species, each with i ti d with subheadings or ed ‘and “botani- cal variants’ tandyor’ ‘named clones.” Various cultivars of the two North A aureola L. (na- tive to North Africa and Europe) and D. mezereum L. (native to Central Asia and aaa are included. Excellent color ees of many of the species and variants < eee any the Daphne includes 70 apecies ne ees J. Halda in 2001) mainly found in Europe and temperate and subtropical Asia. The shrubs, from prostrate and mat-forming to erect and up to 5 meters tall. Plomers are 4-petaled and vary it creamy white to pink, purple, lilac, yellow, and green. The au- thor notes that “if they are given the right environment and treatment, daphnes are long-lived plants worthy of a place in every garden. What is true is that daphnes are unforgiving of mismanagement ... [but] ... To most of us, growing daphnes is a challenge which makes success that much sweeter.’—Guy Nesom, Botanical Research Insti- tute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A SIDA 22(2): 1174. 2006 ANNOTATED CHECKLIST OF THE VASCULAR FLORA OF THE LANCE ROSIER UNIT OF THE BIG THICKET NATIONAL PRESERVE, HARDIN COUNTY, TEXAS Larry E. Brown Houston Community College Barbara R.MacRoberts and Michael H. MacRoberts Bog Research Holman 740 Columbia Houston, Texas 77004, U.S.A. Shreveport, Louisiana 71104, U.S.A. and an Herbarium, see Branch Science Center Herbarium, Museum of Life Sciences Westview Drive Louisiana State University in Shreveport 88 Houston, Texas 77055, U.S.A. Shreveport, Louisiana, 71115, U.S.A. Paul A.Harcombe, Warren W. Pruess, |. penile Elsik rcs | a Department of Ecolo gy Stanley D. Jones Dp | Research Center Rice University PO. Box 6717 Houston, Texas 77005, U.S.A. Bryan, Texas 77805, U.S.A. Biology Herbarium ABSTRACT An annotated, vouchered checklist is provided of the vascular plant taxa of the 10,100 hectare Lance Rosier Unit, Big Thicket National Preserve, Hardin County, in southeastern Texas. Documented for the unit are 694 taxa of which 71 (10%) are exotics. RESUMEN 1: 1.] ] ee | 100 hectareas de la reserva Lance . Rosier Unit, Big Packet) Nacional ve, del condado de Hardin, en el sureste de Texas. Se documentan para la unidad 694 taxones de los que 71 (10 INTRODUCTION This is the fourth part of an inventory, based upon available voucher specimens, of the vascular flora of the Big Thicket National Preserve, in southeastern Texas. In three previ- ous papers, we provided an annotated checklist of the Hickory Creek, Turkey Creek, and Big Sandy Creek units (MacRoberts et al. 2002; Brown et al. 2005; Brown et al. 2006). In this paper, we provide an annotated checklist of the Lance Rosier Unit. SITE AND METHODS The Big Thicket is located mostly within the longleaf pine region of the West Gulf Coastal Plain in southeastern Texas (Parks & Cory 1936; McLeod 1971; Harcombe & Marks 1979; Marks & Harcombe 1981; Harcombe et al. 1993). The Big Thicket National Preserve con- sists of 12 units scattered over seven Texas counties. The units range from 222 to 10,100 hectares and total about 34,000 hectares (Peacock 1994). The Lance Rosier Unit consists of 10,100 hectares and is located in southern Hardin County (Fig. 1). It is relatively flat with poor drainage over most of the area. Elevation ranges from about 14 to 30 meters. The Little Pine Island Bayou runs through its western and southern sections. The Lance SIDA 22(2): 1175— 1189. 2006 BRIT.ORG/SIDA 22(2) 1176 VN I \ | secon o> al ALABAMA - COUSHATTA WOODVILLE 5am INDIAN RESERVATION _— —f UPPER NECHES RIVER CORRIDOR UNIT HILLISTER (BS & WARREN (DB gt Y i TURKEY CREEK UNIT MENARD CREEK CORRIDOR UNIT aan NECHES BOTTOM & JACK GORE BAYGALL UNIT KOUNTZE LA f SILSBEE 2} py yh EVADALE 287 LOWER NECHES RIVER 69 . LANCE \ | ORRIDOR UNIT ROSIER UNIT \ LITTLE PINE ISLAND A a aes ag BAYOU CORRIDOR UNIT SY celia a i cae Co)— .. }SOUR LAKE Men 2 BEAUMONT Che a ——_— ey lee Oe ce Fug, IK Soe 5 Fic. 1. Location of the Lance Rosier Unit, Big Thicket National Preserve, Texas. Rosier Unit was chosen for inclusion in the Big Thicket National Preserve because of its wide range of plant communities (Ajilvsgi 1979; Harcombe & Marks 1979; Watson 1979; Harcombe & Marks 1981; MacRoberts & MacRoberts 1998). It is dominated by lower- slope hardwood pine forest and wetland pine savanna, but also has flatland hardwood forest, floodplain hardwood forest, and baygall thicket (Harcombe & Marks 1979; Marks S& Harcombe 1981). Mima mounds are common in the wetland pine savanna sections. Fire suppression over the past 30 years has resulted in extensive midstory development and shading out of herbaceous vegetation. A map of community types can be found in Harcombe and Marks (1979), and details regarding soils, topography, and climate can be found in Deshotels (1978), Watson (1979, 1982), Harcombe and Marks (1979), and Marks and Harcombe (1981). BROWN ET AL., FLORA OF PRESERVE 1177 Like all units of the Big Thicket National Preserve, the Lance Rosier Unit had not previously been the subject of a detailed floristic inventory. Larry Brown collected about 750 specimens during 15 field days beginning May 23, 2004 and ending May 14, 2006. The MacRoberts’ collected 73 specimens during a study of wetland pine savanna in 1997 and 1998. Stanley Jones collected 317 specimens during 2002 and 2003. Geraldine Watson collected 87 specimens over several years in the late 1970s and early 1980s. Other collectors (Sandi Elsik, Warren Pruess, William Carr, Paul Harcombe, Donna Streng, C. Liu, and Eric Keith) gathered 198 specimens. A search of the on line TAMU herbarium consortium database found some collections at TEX which were annotated and added to the list. Because our purpose is to produces a net of taxa known to occur on the Lance Rosier Unit, a vouchered specimen was con ed to be the only evidence acceptable for inclu- sion in the list. In all, more than 1331 herbarium specimens form the data for this report. All specimens at Rice University have been annotated by Larry Brown. A complete set of these mounted specimens are to be donated to TAES, the Tracy Herbarium, at Texas AGM University. Duplicates collections will be distributed to other herbaria. In most cases, nomenclature follows Jones et al. (1997) and Kartesz and Meacham (1999). For Dichanthelium we followed a pre-publication key in Diggs et al. (2006). Some recent literature sources were followed, especially some nomenclature in the published Flora of North America volumes. In cases of multiple collections, no more than four are listed for each taxon. RESULTS The following is an annotated list of the Lance Rosier Unit taxa. Abbreviations used are given below. CL =C. Liu. His specimens are at Rice University. DS = Donna Streng. Her collections are at Rice University. DW = David Wolf. His one specimen is at TEX. EK = Eric Keith. His collections are at Rice University. GW = Geraldine Watson. Her specimens are at Rice University. LB = Larry Brown. His specimens are at Rice University except for those few at SBSC. MM = Barbara and Michael MacRoberts. Their specimens are at TEX, VDB, and BRIT. Also, as noted, one or two are at SBSC. PH = Paul Harcombe. His collections are at Rice University RJ = Ray Jordan. His collections are at TEX. SEWP = Sandra Elsik and Warren Pruess. Their specimens are at Rice University. SJ = Stanley Jones. His specimens are at Rice University. WRC = William Carr. His specimens are at TEX. * = exotic + =see note at end of list. RGA NTHACERS ACERACEAE vata (Walter) Lindau pees (Chapm.) — Acer rubrum L., SISEWP, 15042, 15041; LB 30287 RWL ong, LB 30165; SEEKWP 3 Ruellia caroliniensis (J.-.Gmel.) Sate : 30574: SEEKWP AGAVACEAE Manfreda virginica (L.) Salisb.ex Rose, LB 31083 3098 Ruellia humilis Nutt., SJ 14095, 14069; LB 30030 1178 ALISMATA Sagtoapop Buchenau,LB 29375, 29363, 30041; SJ 1375 Suna ate a (Engelm.) J.G.Sm., LB 3017 ANACARDIACEAE Rhus copallinum L., SJ 13762;LB 29863 Toxicodendron radicans (L.) Kuntze, LB 29837, SEWP ANNONACEAE Asimina parviflora APIACEAE Centella erecta (Lf) Fernald, SJ 17780 erophyllum tainturieri Hook. var. tainturieri, LB 30407; SJSEWP 1 Cicuta maculata L., LB 30666; SJ 14091 *Cyclospermum leptophyllum (Pers.) Sprague ex Brit- ton &Wilson, LB 30596 Eryngium integrifolium Walter, LB 30231, 30101,30596 30697 — Michx.) Dunal, SJ 13806;LB 31181 fron ae petal Nutt. ex 7. . ae SJ 13906 | ly d) / rticillata Thunb, Lb Onpol ator Le a . ae 30016;MM ' Pea capi ae (Michx.) Raf,, LB 29344; SEEKWP 3094 Ptilimnium tor n (Elliott) Raf, LB 30254 is L., LB 29915; SJ 14073; GW 3102 Can; / | APOCYNACEAE Trachelospermum difforme (Walter) A. Gray, LB 30246 AQUIFOLIACEAE llex ambigua (Michx.) Torr. SJ 14048 llex coriacea (Pursh) Chapm., LB 30678, 3011 1a; SJ 13866,13773 llex decidua Walter, LB 29329; : regi ss longipes Chapman, LB 310 a Soland., LB 29838; ar ies ene Aiton, LB 29983; SJ 13759 ARACEAE Arisaema triphyllum (L.) Schott, SJ ] 3843 ARALIACEAE Aralia spinosa L., LB 29854a ARECACEAE Sabal minor Uacq.) Pers., SJ 14022;LB 30037 ARISTOLOCHIACEAE Aristolochia serpentaria L., GW 156-2 ASCLEPIADACEAE heer es ae ti subsp. longifolia, SJ 13992; 29379; MM 387 asclepic perennis aes ad SEEKWP 3104 Ascle 1S VUOFIE GUL L LB Pp g ' Matelea gonocarpos omen ‘ace LB 30592, 29862; BRIT.ORG/SIDA 22(2) ASPLENIACEAE lee eal cele (L : ae Sterns & Poggenb., LB 29910; SJ 13848, 1 pene! SUE OTIG (Lamarck ) RJ. Jansen var. repens “alien RJ. ren LB 29919: SJ 14245 Ageratina altissima (L.) King & H.E. Rob., GW 2006 Ambrosia artemisiifolia L., LB 30044, 30872 Ambrosia psi ee ze WP 068; LB 30872, 30891 Ambrosia trifida L., L Amphiachyris Nie ides (DC.) Nutt, GW 3142 Arnoglossum ovatum (Walter) H.E. Rob., SEEKWP 3115 13 Baccharis halimifolia L., GW 3150 Bidens aristosa oe Britton, LB 30141, 29397; SJ 14 Bidens bipinnat ay 3136, 3149 Bidens Ronieee L.,, LB 30224, 30213:GW Boltonia diffusa Elliott, LB 29998, a ee 3652; SJ 14027 Chaptalia tomentosa Vent., MM 3733;LB 31058 Chloracantha spinosa (Benth.) Nesom, GW 3409 en horri idulum Michx., LB 30391, 30617;SEWP 3052 iNiu tin (L.) DC., SJ 14088 con oe (L.) Cronquist var. canadensis, SJSEWP 15090 Conyza canadensis (L.) Cronquist var. glabrata (A. Gray) Cronquist, LB 3006 +Coreopsis gladiata Walter, LB 30229, 30901;:MM 3662; 901127 +Echinacea pallida (Nutt.) Nutt. var. sanguinea (Nutt.) Gan D.Thomas, SJ 14098 Ec epost the LB 30158 Elephan Raeusch., LB 30053 (opus Elephantopus nudatus A. Gray, LB 30239, 30772; GW 2412 Elephantopus tomentosus L., LB 30060, 30050; 30049;WP a Erechtites hieraciifolia (L.) Raf.ex DC., an SJ 13890 Erigeron philadelphicus L., SJSEWP. 1 Erigeron procumbens (Houst. ex ai — WP 088 Erigeron strigosus Muhl. ex Willd. SJ 13819 Erigeron tenuis Torr.& A. Gray, SEWP 3073 Eupatorium capillifolium (Small) Lam.,LB 30193;WP 070 oan aaa folium L.,LB 30735, 30257;RJ s.n. is (DC) Torr.& A. Gray, MM 3641 Fupatorum chil Guene LB ee PH 901146 torium perfoliatum L., a 30635, — mpaonan rotundifolium L.,LB 2936 +Eupatorium semiserratum cae - 66, 29979:RJs.n. Eupatorium serotinum Michx., LB 30142; RJ s.n. Eurybia hemispherica (Alexander) Nesom, LB 30120 Euthamia leptocephala (Torr. & A.Gray) Greene ex Por- ter & Britton, LB 29847, 30143; GW 3139 PRESERVE 1179 BROWN ET AL., FLORA OF Gaillardia pulchella Foug. var. pulchella, LB 30625, 29854; Gamochaeta coarctata (Willd.) Kerguelen, LB 30623, 30397 Gamochaeta pensylvanica (Willd.) Cabrera, LB 30601, Gamochaeta purpurea (L.) Cabrera, LB 30398, 30590; SEWP 3058; SJ 14264 Helenium amarum (Raf.) H.Rock var.amarum, LB 29887. 29878; SJSEWP 15048 Helenium drummondii H.Rock, SJ 13787;MM 3784, 3786 Helenium flexuosum Raf.,,LB 29957, 30059; SJ 14170;GW 101 Helianthus angustifolium L., MM 3735; CL 901125 Heterotheca subaxillaris (Lam.) Britton & Rusby, LB 30194, SJ 14240 Hieracium gronovii L., LB 30720 *Hypochaeris Hee (Sch. Bip.) Cabrera var. albiflora (Ku Cabrera, SJ 13932 lva angustifolia nue ex DC.,LB 30071 va annua L.,WP 062; LB 30249, 30249a ue oe (A. Gray) K.L. Chambers ex KJ. Kim, LB Liatris acidota Er coe nN. Gray, LB 29996; WRC 17233; M 4005, SJ 1 Liatris a ae MM 3648b; LB 30238, 3023 Marshallia graminifolia (Walter) Small LE. Watson, LB 30008; MM 3378; WRC 10838 Mikania scandens (L.) Willd., LB 30149, 29976 Pityopsis graminifolia (Michx.) Nutt., LB 30118; WP 059 +Pluchea baccharis (Mill.) Pruski, LB 29871, 30692; WRC 17707 Pluchea camphorata (L.) DC., LB 30183, 30058 Pluchea foetida (L.) DC., LB 29848; SJ 14511 Pseudognaphalium obtusifolium (L.) Hillard & Burtt, LB 30076 eas ca (Walter) DC., SEEXWP 3074; LB 29864 Pyrhepappus eae (D. Don) DC., GW 2004; LB - ia hirta L.,LB 30652 Rudbeckia texana (Perdue) P. Cox & Urbatsch, LB 30582; SEEKWP 3106 Silphium gracile A. Gray, CL 901124 Smallanthus uvedalia (L.) Mack. ex Small, LB 31054 Solidago canadensis L. var scabra Torr. & A. Gray, LB 30139 idago odora Aiton var. odora, LB 30255, 30221, WP wn Qs 061 Solidago rugosa Mill.subsp.asper (Aiton) Cronquist, LB 29988 Solidago rugosa Mill. subsp. rugosa, LB 30140, 30884 Solidago sempervirens L. var. mexicana (L.) Fernald,LB 62 Solidago tortifolia Elliott, RJ s.n. *Soliva sessilis Ruiz & Pavon., SEEKWP 3077; LB 31168 *4Soliva anthemifolia Jussieu) Sweet, LB 31233 *Sonchus asper (L.) Hill, SJSEWP 15092;LB 31160 Symphyotrichum divaricatum (Nutt.) Nesom,GW 1574; Symphyotrichum dumosum (L.) Nesom, LB 30235, 30240; SJ 14257; MM 3648 Synenyonientin lateriflorum (L A. we S Love, WP 058 h LB3 30881, 30880 Vernonia missurica Raf., LB 29833, 30889 12 Xanthium strumarium L., LB 299 *Youngia japonica (L.) DC., SJ 14021;LB 31049 AZOLLACEAE Azolla caroliniana Willd., LB 30902 BERBERIDACEA Podophyllum pram L., GW 3048 BETULACEA Carpinus ee Walter, SJ 1 Ostrya virginiana (Mill) K. te 7 on 14, 14173; LB 30093 BIGNONIACEAE Bignonia capreolata L., LB 29328, 30091; WP 091 Campsis radicans (L.) Seem. ex Bureau, LB 29832; SJ 14250 Catalpa bignonioides Walter, LB 29340 BLECHNACEAE Woodwardia areolata (L.) T. Moore, LB 30777 Woodwardia virginica (L.) Sm., LB 29396 BORAGINACEAE a procunpel Mill., LB a ae Et 1 ell Vey SEW g BRASSICACEAE Cardamine bulbosa (Schreb. Ex Mull) Britton, Sterns, oggenb., GW 3049 *Cardamine debilis D.Don, LB 30292a *Cardamine hirsuta L., LB 30292 Lepidium virginicum L., LB 30603 BROMELIACEAE Tillandsia usneoides (L.) L., LB 29904; SJ 13845 BUDDLEJACEAE Palynremi tm pr rmbens [ea LB 29896 pb BURMANNIACEAE Burmannia capitata UU.F.Gmel.) Mart., LB 30107 CACTACEAE Opuntia stricta (Haw.)Haw., GW 1639 CALLITRICHACEA Callitriche heterophylla Pursh, SJSEWP 15076 Callitriche fineie Nutt., SJSEWP 3107;LB 31228 CAMPANULACEA Lobelia cardinalis L., = 30167 1180 Lobelia flaccidifolia Small, LB 30636, 29362; MM 3920 Lobelia puberula Michx., SJ 14261;LB 30241 Lobelia reverchonii B.L. Turner, LB 30230; PH 901144 Triodanis biflora (Ruiz & Pavon) Greene, LB 30604 Triodanis perfoliata (L.) Nieuwl., GW 3061 CAPRIFOLIACEAE “Lonicera japonica Thunb., SEWP 3065 Lonicera sempervirens L., SJ 13929 Sambucus nigra L. subsp. canadensis (L.) R. Bolli, LB 303 3105 Viburnum dentatum L., SEEKWP 3078;LB 2986 1a, 29853 Viburnum nudum L. var. nudum, SJ 13998; LB 29401, CARYOPHYLLACEAE *Cerastium glomeratum Thuill., SJSEWP 15066 Sagina decumbens (Elliott) Torr. & A. Gray, LB 30400; SEEKWP 3108 *Stellaria media (L.) Vill., SISEWP. 15063 CISTACEAE +Lechea minorL., LB 30003 COMMELINACEAE Com: / elir IG virgit 1iCa is LB 301 53 CONVOLVULACEAE Dichondra carolinensis le a 30075; SJ 13849 Nah vulus sericeus = LB 3 LG ~ a fain 10ea cordatotriloba Dennst. var. cordatotriloba, LB 4235 CORNACEAE Cornus florida L., SJISEWP 15043;LB 31047 Cornus foemina Mill. 8 29938; SJ 13758 Nyssa aquatica L., SJ 1 Nyssa biflora Walter, : ae 17365 Nyssa sylvatica Marshall., LB 29336, 29978 CRASSULACEA Penthorum oe L., LB 30671 CUCURBITACEAE Cayaponia quinqueloba (Raf) Shinners, SJ 14515 CUPRESSACEAE Taxodium distichum (L.) L. C. Rich., LB 30207, 29917 SEEKWP 3099 CYPERACEAE Carex abscondita Mack., SJ 13851 Carex albicans Willd. ex Spreng. subsp JH. Rettig, SEEKWP 3097;LB 30291 Carex annectens E.P. Bicknell, LB 30382 Carex basiantha Steud., SJ 13696;LB 30610 Carex blanda Dewey, SJ 1391 Carex bulbostylis Mack., LB 31163 Carex caroliniana Schwein, LB 30612, 30575 Carex cherokeensis Schwein., SJSEWP. 15049 = lis (Bailey BRIT.ORG/SIDA 22(2) Carex complanata Torr. & Hook., LB 30381, 29330; SJ 2, 13864 Carex crebriflora Wieg., SJ 14553, 13597 arex crus-corvi Shuttl. ex Kunze, SEEKWP. 3102 Carex debilis Michx. var. debilis, SJ 13860;LB 31075 Carex digitalis Willd. var. asymetrica Fernald, SJ 13852 Carex flaccosperma Dewey, SJ 13803;LB 30388a; SEWP 3054 Carex frankii Kunth, LB 30598; SEEKWP 3090 Carex Ss Rudge, SJ 14177, 14020 Carex Elliott, CL 901119;SJ 1 Carex intumescens Rudge, SEEKWP 31 a ae LB 31077 Carex joorii Bailey, LB 30781;SJ 14016 Carex leavenworthii Dewey, SJSEWP. 15079 Carex longii Mack., SJ 13863; LB 30581, 2934] Carex louisianica Bailey, LB 29845 Carex oxylepis Torr. & Hook. var. oxylepis,, SJ 13917 Carex retroflexa Muhl. ex Willd., LB 30406, 30602; SJSEWP ] 5080 ba AACS triatula MUCH I Car J] ne iene ia eee SJ 13862 Carex texensis Bailey, LB 30395; SJSEWP 15056 Carex triangularis Boeck., SJ 13908;LB 30578 Carex tribuloides Wahlenb. var. sangamonensis Clokey, B 30607; SEEKWP 3088 Carex verrucosa Muh, SJ 14024 arex vulpil oidea Mich + LB 3 30383 Cladium mariscus J. Pohl subsp jamaicense (Crantz) ae LB 30898, 19238b (SBSC), 19421 (SBSC): orr. & Hook. ex Torr, LB 29891a pene articulatus : LB 30154, SJ 14058 *Cyperus entrerianus Boeckl., LB cae Cyperus erythrorhizos Muhl.,LB Cyperus has LB 30262, a 7 14041 Cyperus ochraceus < Vahl, LB 30885 tie odoratus LLB 30712 [ N Steud, SJ 14060, 14260 opens, Rottb., SJ 14059; GW 3165 Cyp Steud., SJ 1 a 14076, 14010 eae retrorsus ae fea 1427 Cyperus strigosus L., SJ 1 Cyperus surinamensis sek - 1426 Cyperus thyriflorus Jungh.,LB 29905; i 14174; SEEKWP al s virens _ - J 13896; LB 2933 orr.) Chapman, LB 30387, 30163 Eleocharis ae ae Roemer & J.S. Schultes, LB 30212, 29946 Eleocharis microcarpa Torr., LB 30654, 30643; SJ 14550; IC I | Eleocharis montana Roem. & Schult., LB 30890 Eleocharis montevidensis Kunth, LB 29333; SJ 14070 Eleocharis tuberculosa R.Br., MM 3858;LB 30216, 29961; WRC 17230 BROWN ET AL., FLUAA UE PRESERVE 1181 Fimbristylis aqutumnalis Roem. & Schult. LB 30157, 30036; GW 3166b *Fimbristylis miliacea (L.) Vahl, LB 30157a, 29989 Fimbristylis puberula Vahl var. puberula Vahl, SJ 14101; LB 31182 Fuirena breviseta ae a LB 30727,29948;MM 3480; WRC 1 Fuirena bushii Kral, iB ne 30010, 19239; SJ 14296 Isolepis carinata Hook. & Arn.ex Torr., LB 30409 Kyllinga brevifolia Rottb., LB a Kyllinga odorata Vahl, SJ a Kyllinga pumila Michx,, L Cada caduca ee hi 3479b , 3540 LB Rhynchosp ata (L.) H. Pfeiffer, LB 29880, 29417 SJ 13770 Rhynchospora corniculata (Lam.) A.Gray, LB 29369 Rhynchospora divergens Chapm.,LB 294 Rhynchospora elliottii A.Dietr., LB 19280;MM 3927, 3948; Rhynchospora fascicularis (Michx.) Vahl, LB 30217 Rhynchospora filifolia A. Gray, LB 30658, 30020; MM 85,3851 Rhynchospora globularis eu ) Small, var. globularis LB 15998; MM 3864, Rhynchospora globularis oo Small var.pinetorum (Britton & Small ex Small) Gale, SJ 14074 Rhynchospora ae : ) Vahl, LB 30260; MM 3646; WRC 17703a; SJ 14 jure ee : Gray, MM 3844c; LB 30645, 1;5J 14613 mcg sen (Michx,) Vahl, LB 30079, 30073; 1; SJ rhychospo pie (Baldw. ex Elliott) Thomas, SJ 4077, MM 394 Rhyne chespera mixta Britton, LB 29895; SJ 14106 Rhynchospora perplexa Chapm.ex M.A.Curt.LB 30633, 354 Rhynchospora plumosa Elliott, LB 30236; WRC 17355; MM 3488b; SJ 13936 Rhynchospora pusilla ee ex M.A. Curtis, LB 30665, 30024a, 20417;MM 3 Rhynchospora rariflora lee LB 30630, 29383, 29353; MM 3867 iychoso ee (Gale) Kral, LB 29416, 30647, 29390; SJ 1408 Scirpus cyperinus an LB 30696 Scleria georgiana Core, MM 3386 (SBSC),3488; SJ 13768; LB 15999 (SBSC) Scleria oligantha Michx., LB 30608, 30576; SEEKWP 3110 Sclera pauciflora Muhl. ex ae 30015; SJ 13797 Scleria reticularis Michx., MM 6 Scleria Hee Michx., ie ae 29355, 30061; SEE 111 CYRILLACEAE Cyrilla racemiflora L.,LB 30006; SJ 14089 DENNSTAEDTIACEAE Pteridium aquilinum (L.) Kuhn, WP 090;LB 30045 DROSERACEAE Drosera brevifolia Pursh, SJ 13769; MM 3781 Drosera capillaris Poir., LB 29418; SJ 14043; MM 3860; GW 3064 DRYOPTERIDACEAE Polystichum acrostichoides (Michx.) Schott, LB 30614, 30056, 29984 Woodsia obtusa (Spreng.) Torr.subsp obtusa, SJ 14018 EBENACEAE Diospyros virginiana L., LB 29937a;SJ 14051 ERICACEAE eee canescens (Michx.) Sweet, SJSEWP 15044; LB 31080 ese viscosum (L.) Torr. SJ 141 Vaccinium arboreum Marshall, SJ 13785; LB 29953, , 14248 30280 Vaccinium corymbosum L., LB 30412, 30223; SJ 13799, 14167 ERIOCAULACEAE Friocaulon decangulare L., LB 29398; SJ 14156; WRC 7697; 3100 anceps Morong, LB 29415; SJ 13774 EUPHORBIACEAE Acalypha gracilens A. Gray, GW 3138;LB 30210, 30715 lachnocaitlon 7 Acalypha virginica L., SJ 14165 *Caperonia palustris (L.) St.-Hil. GW 3138 Chamaesyce cordifolia (Elliott) Small, SJ 14198 Chamaesyce humistrata (Engelm.) Small GW 3162 Chamaesyce maculata (L.) Small, LB 30201, 29935, 30713 Chamaesyce nutans (Lag.) Small, GW 3106, 3168; LB J pit atric NAj hx. WP 069; LB 2992 Croton glanduoss L. var. glandulosus cael -Arg. LB 307 oa monanogynus Michx., LB 30886 a L.,LB 30732a, 30252, 30099 Phy ni caroliniensis Walter, LB 30896, 30198, GW ee urinaria L., LB 29868; GW 3105, 3140 *Triadica sebifera (L.) Small, SEEKWP 3096 *Vernicia fordii (Hemsl.) Airy-Shaw, SEEKWP 3109; LB 30090 FABACEAE *Albizia julibrissin Durazz., SEEKWP 3083; LB 30248, 051 Amorpha fruticosa L.,.LB 30618, 29926 Baptisia bracteata Muhl.ex Elliott var./aevicaulis (A. Gray ex Canby) Isley, LB 31085 Centrosema virginianum (L.) Benth., LB 29851; GW 3145 1182 Cercis canadensis L. var. canadensis, SISEWP 15054 Chamaecrista fasciculata (Michx.) Greene, LB 29849 Chamaecrista nictitans (L.) Moench, GW 3137 peo nies lig Mac. Mill., LB 30667 m (Michx.) DC., LB 30721 Desm sel aes (Nutt.) DC. GW 3144 Desmodium paniculatum DC., LB 30185, 30043; SJ nee GW 3152 Erythrina herbacea L.,LB 3 Galactia volubilis (L.) ce 29860; GW 3148 aes tricanthos L., GW 2717 ttidium vesicarium Jacq.) Harper, LB 30716, 29943, 9925 Lespedeza virginica (L.) Britton, GW N20 SUES syecamienys Hook., GW 3074;LB 31052 puli LB aie laal 308] “Medicago Sarees L., LB 3 & Rose) B.L. Turner, MM 3660 *Melilotus indicus (L (L ) au ue eae me 31048 LB 29387, 29373; MM 3660; SJ 13779 Mimosa strigillosa Torr.& A. Gray, te oak een Puneet Benth., LB 312 usifolia (L.) Irw peers LB 29918 Sesban crummondi faydb) Cory, SEEKWP 3103; LB ae iene (Mill.) McVaugh,LB 30710; WP 065 oe umbellata (Muhl. ex Willd.) Britton, LB 57;SJ 14086, 14263 lps pean Nutt., SJ 1 te LB 29999 reb., SISEWP 15050 “Trifolium dubia sibihrce. LB 30403 Trifolium repens L., SEWP 3062; SJSEWP 15096 “Vicia hirsuta (L.) S.F.Gray, LB 30402, 31053; SEWP 3067 Vicia ludovici Nutt. subsp. ludoviciana, GW 3073;LB 3105] Vicia minutiflora D. Dietr., SJSEWP. 15091 ata sativa L.subsp. nigra (L.) Ehrh., SEWP 3066 0 PEF LB 30 Wisteria eee. (L.) Poir., LB 30619 FAGACEAE Castanea pumila (L.) Mill, SEEKWP. 3116; SJ 13784 Fagus grandifolia ate WP 178;LB 31078 Quercus alba L., LB Quercus falcata ae LB 30584, 29374, 29977; SJ 13801 Quercus hemisphaerica W. Bartram ex Willd., LB 30282; J 141 3 Quercus incana W. Bartram, SJ 14082 Quercus laurifolia Michx., LB 30591, 30174, 29922 16466 Quercus lyrata Walter, LB 29928; SJ 13783; GW 514 Quercus ee a (Ashe) Ashe, rs 31173 Quercus m ndica Muench., SJ 1379 Qu ich i Nutt., LB 29840; SJ 13850 WN BRIT.ORG/SIDA 22(2) Quercus nigra L., LB 29385, 29331; SJSEWP. 150843 Quercus pagoda Raf., SJ 13844, 13838, 14116 Quercus phellos L.,LB 29391, 30025, 29975 Quercus stellata Wangenh., SJ 13793 Quercus velutina Lam., SJ 14056 Quercus virginica Mill., WP 180 GENTIANACEAE *Centaurium floribundum (Benth.) B.L. Rob., LB 30583 *Centaurium pulchellum (Sw.) Druce, LB 30597, 30094; SEEKWP 30 Sabatia arenicola Greenm., SJ 13870 Sabatia calycina A. Heller, LB i Sabatia campestris Nutt., SJ 1 Sabatia gentianoides Elliott, ae = GERANIACEAE ani lini L., SEWP 3075;LB 30285 GROSSULARIACEAE ltea virginica L., SJ 14118, 14023 HALORAGACEAE Myriophyllum pinnatum (Walter) Britton, Sterns, Poggenb.,, SJ 13921 Proserpinaca palustris L,SJ 13913;LB 30644 Proserpinaca pectinata Lam., LB 30644; GW 3108; MM 3475; DS 090 HAMAMELIDACEAE Liquidambar styraciflua L.,SJSEWP 15071 EE ROPNYLLACEAE t.ex Choisy, LB 29958; GW 3109 HYPERICACEAE Hypericum crux-andreae (L.) Crantz, LB 29412, 29994; SJ 13788; MM 3389 Hypericum galioides Lam., LB 30728, 29944; S) 14514 MM 4001 Hypericum mutilum L., LB 30002; SJ 14160 Triadenum walteri (J.G.Gmel.) Gleason, LB 30180, 29892, 30780; GW 3093 IRIDACEAE Sisyrinchium angustifolium Mill. GW 3054; LB 29358; LLB s.n. (SBSC) Sisyrinchium teas E.P. Bicknell, SJ 13760, 14255; MM 387 ie LB ala SisvriNCAhIM Be) ae O ae LB 31 Sisyrinchium rosulatum E. P. he ae 3061; SEWP 3061 Sisyrinchium sagittiferum E. P. Bicknell, GW 3082 LB JUGLANDACEAE Carya aquatica (Michx.) ee — 30085 Carya illinoensis K. Koch, LB 3 Carya texana Buckley, LB ve i 13970 Carya tomentosa (Lam.ex Poir.) Nutt., LB 30611,30579 Juglans nigra L., LB 29913 BROWN ET AL., FLORA OF THE LANCE ROSIER UNIT, BIG THICKET NATIONAL PRESERVE 1183 JUNCACEAE Juncus acuminatus Michx., SJ 13898, 14012 Juncus brachycarpus Weigel, LB 29408; 2: 3899, 14099 Juncus coriaceus Mack.,LB 29932, 29945;SJSEWP 15068 Juncus dict Elliott, LB 31167 Juncus diffusissimus Buckley, SJ 13900, 14008 Juncus effusus L.,LB 30155; SEWP 3071 sein elli MONEE: SJ 14061; LB 29342, 31180C Rostk,, SJ 14002, 14065; LB 29347 Juncus nodatus Coville, SJ 13901 Juncus repens Michx., SJ 14013 LB 30641 Juncus tenuis Willd., LB 30086; SJ 14074a, 13897 Juncus validus Coville, LB 29334 LAMIACEAE edeoma hispidum Pursh, LB 31166 Hyptis alata (Raf) Shinners, LB 29885; MM 3484 lLycopus rubellus Moench, LB ae PH 901159 Monarda punctata L., LB 2985 *Perilla frutescens (L.) Britton, i“ 29890; GW 3052 Prunella vulgaris L.,SJ 13919 ycnanthemum albescens Torr. & A. Gray, LB 30196, 29954 = Salvia coccinea Buchoz ex Etlinger, SJ 14259 Salvia lyrata L.,LB 30054; SJSEWP 15047 Scutellaria integrifolia L., SJ 13789, 14549, 14052; MM 3876 Stachys crenata Raf.., LB 30396; SEWP 3068 LAURACEAE Persea palustris (Raf.) Sarg., LB 30211, 29866, 29381, 3 Fy oO foto eee (Nutt.) Nees, LB 30222 LEMNACEAE ea punctata (G. Meyer) Les & DJ. Crawford, LB 1069 ae brasiliensis Wedd., LB 30878, 070 Wolffia aH. Karst., LB 310 Wolfflla eee (Hegelm.) in. is 31068 LENTIBULARIACEAE Pinguicula pumila Michx., SJ 13702; MM 3761 LILIACEAE MELD aurea welsh LB 30028, 29991;SJ 14094;MM 3639 L. var. canadense, SJ 13765, 13874 Hymenocallis liriosme (Raf.) Shinners, SEWP 3072 Hypoxis rigida Chapm.LB 30233, 311 oe Hypoxis wrightii (Baker) Brackett, LB 311 *Narcissus tazetta L., WP 098 Nothoscordum bivalve (L.) Britton, ete 5046 *Ophiopogon jaburan Lodd., LB 3 cic: croceum (Michx.) men SJ 14548; MM 3783, LB 31057 *Zephyranthes candida Herb., LB 29911;SE 4581 LINACEAE Linum medium (Planch, Britton var. texanum (Planch.) Fernald, LB 303728a, 30657; SJ 14055; MM 3546 Linum striatum Walter, LB 30653, 30639; SJ 14169: GW 3075 LOGANIACEAE Gelsemium a (L.) Aiton f,, LB 29386, 30279; SJSEWP 15038 sare pel (JF. Gmel.) Torr. & A. Gray, LB 29881; Mi eal sessi sto te F.Gmel.) G. Don, LB 30012; MM 3483; PH 9 LYCOPODIACEAE een appressa a ) Cranfill, GW 3435; WRC 7228; MM 3 LYGODIACEAE *Lygodium japonicum (Thunb.ex Murray) Sw., LB 29888, 29856; SJSEWP 15045 LYTHRACEA ee cathagenes (Jacq.) J.F. Macbr., GW 3076 a L.,LB 30600 LU faerie ee Push var. lanceolatum (Elliott) Torr. & A. Gray, LB 30042; SJ 14270 Rotala ramosior (L.) ees LB 3016 MAGNOLI Magnoli sen L., LB 29857;SJ 14239 Magnolia virginiana Ee SJ 1 4038 MALVACEAE “Abutilon theophrasti Medik., GW 3134 Hibiscus moscheutos L., LB 30173; GW 3094 Malvastrum coromandelianum (L.) Garcke, GW 3088 Modiola caroliniana (L.) G. Don, SEWP 3064 Sida rhombifolia L., LB 30694 Sida spinosa L., GW 3163 MELASTOMATAC lutea hates - 30663; MM 3388; WRC 17227;SJ 1, GW 3090 4054 rei mariana L., MM 3489; SJ 14155, 14015;GW 3126 MORACEAE Morus rubra L., LB 29939 MYRICACEAE Morella caroliniensis (Mill.) Small, SJ 14117, 14062 Morella cerifera (L.) Small, LB 30047; WRC 17356 OLEACEAE SH oUanNys virginicus L., WP 087; LB 29335, 31071 aroliniana M Mill, S 4175 Fraxinus eee es hese LB 29924 *Ligustrum sinense Lour, LB 29865; SJ 14068 ONAGRACEAE Gaura lindheimeri Engelm. & A. Gray, LB 30007; WRC 17701; SJ 14510; DS 124 Cua ugg Span LB 30146 a L., LB 29879; = 14509 Walter, LB 3015 Froayint of lL PpAiasicay, { os: 1184 Ludwigia glandulosa Walter, LB 29987; SJ 14234; GW 3104 Ludwigia grandiflora (Michx.) Greuder & Burdet, GW 3180 Ludwigia hirtella Raf.;LB 2995 1,29367;SJ 14171;DS 123 Ludwigia linearis Walter, MM 3538; LB 29959, 29992 Ludwigia microcarpa Michx., LB 30684 Ludwigia octovalvis Jacq.) PH. Raven, LB 30725 Ludwigia palustris (L.) Elliott, LB 30669, 30159 Ludwigia peploides (Kunth) PH. Raven, LB 3014 Ludwigia pilosa Walter, LB 30774, 30122a, a CL 01157 sabia sulle L. Oe iniata Hill, SEWP 3069, SJSEWP on Denotieas: speciosa Nutt., LB 30624; SEEKWP 310 OPHIOGLOSSACEAE Botrychium biternatum (Sav.) Underwood, GW 3051 Opioalessiinn crotalophoroides Walter, LB 31050 Ophioglossum petiolatum Hook., LB 30394 ORCHIDACEAE Calopogon tuberosus (L.) Britton, le Poggenb., LB 29406, 29392; SJ 14079; MM 3 Listera australis Lindl. SJ 13698 Spiranthes cernua (L.) L.C. Rich., WP 060 Spiranthes floridana Metre re LB 2041 +Spiranthes longilabris M Spiranthes odorata a a LB 30894 Spiranthes praecox (Walter) S. Watson, MM 3863; LB 178 ,LB 30247 8(SBSC) Spiranthes tuberosa Raf., SJ 14093 Spiranthes vernalis Engelm. & A. Gray, MM 3919 OSMUNDACEAE smunda regalis | 29360; WP 098 OXALIDACEAE Oxalis corniculata L. var. wrightii (A. Gray) B.L. Turner, LB var spectabilis (Willd.) A. Gray, LB 30 3077 ee debilis Kunth var. oo (DC.) Lourteig, LB 0250; SEWP 305 ae i Pursh, (1) ee 15055 Beate RCEDE L., LB 29907; SJSEWP. 16461 eaaiicie Gee L. in 30613, 30067 ee L., SJ 14272 PINf a echinata Mill.,LB 30 Pinus palustris Mill, LB 2 Pinus taeda L.,LB 30144, oo SEEKWP 3092 PLANTAGINACEAE *Plantago major L., LB 30289, 30284 Plantago virginica L., LB 30404, 29882,; SEWP 3060 PLATANACEAE Platanus occidentalis L., LB 29852 BRIT.ORG/SIDA 22(2) POACEAE Agrostis hyemalis (Walter) Britton, Sterns, Poggenb., SE 3059; SJ 13868 Andropogon glomeratus (Walter) Britton, Sterns, Poggenb., LB 30188; WP 063 Andropogon virginicus L., LB 30882, 30172; CL 901126 Anthaenantia texana Kral, LB 30234; WRC 10840; MM 1 3 6 Aristida longispica Poir. var. geniculata (Raf.) Fernald, LB Aristida longispica Poir. var. longispica, LB 30236a Aristi dee sae oo ) Vasey, LB 30730, 30729;WRC 2;MM 354 Aristida purpurascens Poir. var. virgata (Trin.) Allred, LB 30237, 30228, 30227, 29956 Arundinaria gigantea (Walter) Muh. eeaen 5072 Axonopus fissifolius (Raddi) Kuhlm., SJ 14113; MM 36 LB 30680, 30674 Axonopus furcatus (Fluggé) A. S. Hitche., LB 29349 PH 61;5) 14115 *Briza minor L., SJISEWP 15053 “Bromus Seis mo LB 30599; SEEKWP 3095 Cha m (Michx.) Yates, SJ 14252 Chasmanthium laxum (L.) Yates var. laxum, LB 29839 Chasmanthium laxum (L.) Yates var. se Wipff & S. Jones, LB 30717 — rugosa (Nutt.) Nash, LB 30736; MM 3721, ied HETEPOPP EET (P oir. i oe n nda (L.) Pers., LB 30621; SEEKWP 3086 Dichanthelium aci aie (Desv. ex Poir.) Gould & Clark var. angustifolium (Elliott) S.L. Hatch, LB 30588, 31184, 30035; SJ 14459 Dichanthelium acuminatum (Sw.) Gould & CA. Clark var.acuminatum SJ 14556;LB 29357, 30695 Dichanthelium acuminatum (Sw,) Gould & C.A.Clark var. oe (E.L.Rand & Redfield) Gould & CA.Clark, ae thea. acumin oe ey ean & CA.Clark var. C.A.Clark, LB 30626, GUIUCUTTT MING! 1) IOUT 346. MM 3482. CL 901 136 Dich oe boscii (Poir.) Gould & Clark, LB 30609, 29908, 30876 nee commnutatuin (Schultes) Gould subsp. omm LB 29883, 29982: SJSEWP 15058 pon commutatum (Schultes) Gould subsp. equilaterale (Scribner) Freckman & Lelong, LB 30063 (SBSC) Dichanthelium consanguineum (Kunth) Gould & CA. Clark LB 30650; MM 3862 Dichanthelium dichotomum (L.) Gould var. dichotomum, LB 30586, 30585,3064 2a, 29897 Dichanthelium dichotomum (L.) Gould & C.A. Clark subsp. microcarpon (Muhl. ex Elliott.) Freckman & eee LB 30642, 29897, 29986 Dichanthelium dichotomum (L.) Gould subsp. nitidus (Lam.) Freckman & Lelong,, LB 31162 BROWN ET AL., F LYUNA VE PRESERVE 1185 Gould a Dichanthelium ensifolium (Baldw. ex Elliott subsp. ensifolium, LB 30089, 29897 ae ee laxiflorum (Lam.) Gould, LB 29842; SJSEWP 15057 fe ae ovale (Elliott) Gould & C.A. Clark WRC 17362 Dichanthelium ovale (Elliott) Gould & CA. Clark subsp. villosissimum (Nash) Freckman & Lelong, LB 30660, 30651;SJ 14554; WRC 17366 Dichanthelium polyanthes (Schult.) Mohlenbr., LB 30589, 31319 Dichanthelium ravenelii (Scribner & Merr.) Gould & CA. Clark, LB 30699 Dichanthelium scabriusculum (Elliott) Gould & CA, Clark, LB 30034, 29980, 29348 Dichanthelium scoparium (Lam.) Gould, LB 30659, 30261, 29370; SJ 14042 (Elliott) Gould, LB 29345, 30004; SJ 14460 Dichanthelium tenue (Muhl.) Freckman & Lelong, LB 29377 Dichanthelium wrightianum (Scribn.) Freckmann, MM 3385 Digitaria ciliaris (Retz.) Koel., LB 30226, 30190, 29903, 30072 *Digitaria ischaemum (Schreb.) Schreb. LB 30189 Echinochloa walteri (Pursh) Heller, SJ 14290 *Fleusine indica (L poe eS 30078 Elymus virginicus les SJ Eragrostis elliottii ed ie 30187;PH 901148 Fragrostis hypnoides (Lam.) Britton, Sterns, Poggenb,, LB 30177 Fraor tis refracta (Muhl.) Scribn., GW 2400; MM 3654 Eragrostis secundiflora (J.Presl) subsp. Oxylepis (Torr) S. Dsoen LB mais: SJ 14075; PH 901143 le ides (Munro) Hack., LB 30065 Hordeum pie tien Nutt.,LB 30405; SEEKWP 3085 Leersia hexandra Sw., LB 30038 Leersia lenticularis Michx., LB 30166a; SJ 14172 Leersia virginica Willd., LB 30197; SJ 13856 *Lolium arundinaceum (Schreb.) S.J. Darbyshire, SJ 15095 Melica mutica Walter, SJ 13697 ees capillaris (Lam,) Trin., Cl 901128;MM 3656; LB 3090 Pears expansa (Poir,) Trin., GW 2408a; MM 3724 (SBS Oplismenus hirtellus (L.) Beauv., LB 30169, 29916 Panicum anceps Michx., LB 29899; PH 90114 Panicum brachyanthum Steud.,LB 30187; CL 901156 Panicum dichotomiflorum Michx., ee 1, Panicum Oe Elliott, LB 3 Pan he Sera eae mitomon J *Panicum repens L., LB 2 343 Panicum rigidulum ee ex Nees, LB 30175;PH 901163, 901167, MM 3732 Panicum see re ex Trin, LB 19343, 30019; WRC 17698; MM 3 Panicum verrucosum nM ies 3666; LB 30184, 30259 Panicum virgatum L., LB *Parapholis nea) G : cee 3865 *Paspalum m Poir., LB 29884 Paspalum floridanum Michx., LB 30192, 30123, 30770 Paspalum laeve Michx., LB 30775, 30171, 29963; MM 3663 Paspalum langei (Fourn.) Nash, LB 29902a; SJSEWP 16460 Paspalum lividum Trin., LB 30162 Paspalum minus Fourn., LB 30879, 30640, 30096, 19238 (SBSC) *Paspalum notatum Fluggé, LB 29964, 29405, 19238 (SBSC) Paspalum plicatulum Michx., SJ 14067 aoe ae a LB 30026; CL 901158; SJ 05; MM 3 Psp ee ens LB 29876, 29870a pal teud., LB 29942; GW 3023; SEEKWP "3093. *Poa annua L., SJSEWP 15061 Poa autumnalis Muhl. ex Elliott, LB 31055 *Polypogon monspeliensis Muhl. ex Elliott, LB 30606 Saccharum baldwinii Sprengel. LB 30168, 29931, 30773 Saccharum giganteum (Walter) Pers., LB 30147, 30892 Schizachyrium scoparium (Michx.) Nash var. di eu (Hack.) Gould, PH 901145; CL 901122; DW s. Sorghum halepense (L.) Pers., LB 30594, 29941; eu 3084 Sphenopholis obtusata (Michx.) Scribn., SJ 13915; GW 2246b; LB 31164 Sporobolus indicus (L.) R.Br, LB 30622, 29909 Sporobolus st cael LB14907 (SBSC), 14702 (SBSC); PH 9011 Steinchisma ea ae Nash, SJ 13892 Tridens ambiguus (Elliott) J.A.Schultes, WRC 17234; CL 01120; GW 2406; MM 3544 Tridens flavus (L.) Hitche. var. flavus, LB 29923;WP 066 Tridens strictus (Nutt.) Nash,LB 30145,30258;CL 901121 m dactyloides (L.) L., SJ 14025 urocta wianvenylla a (Munro ex Wright) R.Webster, GW POLYGALACEAE Polygala cruciata var. cruciata L., LB 29995; SJ 14111, 14425, 14293 | hiaala incarnata ea Polygala leptocaulis Torr. . i - LB 30691, 30687; SJ 14161, 13993 Polygala mariana Mill, LB 29400, 30027; MM 3726 Polygala ramosa Elliott, LB 30017; SJ 14158; MM 3877 POLYGONACEAE Brunnichia ovata (Walter) Shinners, LB 29846 1186 ‘Polygonum cespitosum Blume var. longisetum (De oe yor aan LB al i jonu Vieisn., LB 30179; GW 3166a Beets udroniaawides Michx, SJ 13891 Polygonum punctatum Elliott, WP 064; LB 30150 30084a; GW NO2 Polygonum virginianum L.,LB 30088, 30209 *Rumex crispus L.,SJ 13859, Ree Rumex chrysocarpus Moris, LB 3 *Rumex pulcher L., LB 30605; eee 3089 POLYPODIACEAE Pleopeltis polypodioides Andrews & Windham subsp.michauxiana (Weatherby) Andrews & Windham, LB 30084 eee agalli is L., SISEWP 15051 Ana fea minima (L.) E.H.L. Krause, LB 31 Samolus valerandi L. subsp. parviflorus a File LB 29889; SJ 13831 RANUNCULACEAE Clematis crispa L.,LB 30723, 30580; WP 092:SJ 14262 Ranunculus hispidus Michx.var.nitidus (Chapm.) T. Dun- can, SJSEWP 15064 ie —! Ranunculus pusillus Poir., SJSEWP 15074 RHAMNACEAE Berchemia scandens (Hill) K.Koch,LB 29874; SEWP 3049 Rhamnus caroliniana Walter, WP 099 ROSACEAE Crataegus berberifolia Torr. & A. Gray, SJ 14249 Crataegus brachyacantha Sarg. & Engelm., LB 30205, 29930; GW 3041 Crataegus marshallii Eqglest., SJ 14244 Crataegus opaca Hook. & Arn., SJ 13761; LB 30206, 9836, 19244 Crataegus rhe ae hx., LB 30679 Crataegus viridis L., SJ 14253 “‘Duchesnea indi os .) Focke, SISEWP 15062 Photinia pyrifolia (Lam.) K.R.Robertson & J.B.Phipps,LB 29402; SJSEWP 15039; SJ 13996 Prunus caroliniana (Mill.) eh SJ 13909 Prunus serotina pie SJ 14 08 “Pyracantha } rita Rehder, SEWP 3055 “Rosa bracteata J.C.Wendl., SJ 14063 Rubus argutus Link, LB 30873, ee SEEKWP 3082 i trivialis Michx., SJISEWP 15052 ea cantoniensis Lour., LB 30398a RUBIACEA one sree 5 - SJ 14258;LB 30384 Diodia teres Walte Diodia virginiana L,, ce 2300 sas 3475 Galium aparine L., SJSEWP 1 Galium obtusum Bigelow, - Galium pilosum Aiton, LB co. 14241; SJSEWP 16462; GW 3066 => BRIT.ORG/SIDA 22(2) Galium tinctorium (L.) Scop., SJSEWP 15073 Gardenia augusta Merr., SJ 14143 Houstonia pusilla Schoepf, LB 30393: aa 15060 Mitchella repens L., LB 29841, 30074; SJ 13 Olden landi a boscii Oe Eu LB en on *Sherardia arvensis L., ae oe SJSEWP 15082 Stenaria nigricans (Lam.) Terrell, SJ) 14039; PH 901162; 31226 RUTACEAE *Poncirus trifoliata (L.) Raf.,SJ 14254 SALICACEAE Salix nigra Marshall., LB 29949 SARRACENIACEAE +Sarracenia alata Wood, LB 30726; SJ 13777; WRC SAURURACEAE Saururus cernuus L., LB 29835; SJ 14168 CROPHULARIACEA ae fasciculata ie Raf.,SJ 13861, Agalinis heterophylla (Nutt.) Small ex ae i: ee W 3135, 3401 Agalinis purpurea (L.) Pennell, LB 30875, 30214,30144a; => 9 Agalinis viridis (Small) Pennell, LB 30218 Aureolaria flava (L.) Farw. var. flava LB 29955, 30001; SJ 14045, 14236 Bacopa caroliniana (Walter) Robins.LB 30778 Bacopa monnieri (L.) Pennell, LB 30203, 29886 Gratiola brevifolia Raf., LB 29364; SJ 14003; GW 3117 3916 Grati Pigpiona Michx,, LB ee ha 3127;PH 901155 Gratiola virg! nian al. SJISEWP J *Lindernia crustacea, (L.) F. He | 29900, 30080 Mecardonia acuminata (Walter) Small,LB 30200, 30100; 901142 Mecardonia procumbens (Mill.) Small, SJSEWP. 15089: Micranthemum umbrosum (J.G.Gmel.) Blake, LB 30178, 30164 Scoparia dulcis L.,LB 30719, 30204, 29901;SJ 14072 “Verbascum thapsus L., GW 3047 *Veronica arvensis L.,LB 30401 Veronica peregrina L., LB 30401a, 30386 “Veronica polita Fries, LB 30290 SELAGINELLACEAE Selaginella apoda (L.) Spring, SJSEWP 15077; LB 30281 SMILACACEAE Smilax bona-nox L., SEWP 3050, SJ 14066 Smilax glauca Walter, LB 30182, 30022;5J 14007 Smilax laurifolia L., SJ 13778; LB 29389, 30113, 30874 Smilax pumila Walter, LB 30225, 29894; SJ 13904 Smilax rotundifolia .,LB 29365 BROWN ET AL., FLORA OF THE LANCE ROSIER UNIT, BIG THICKET NATIONAL PRESERVE 1187 milax smallii Morong, LB 30040 Smilax walteri Pursh, GW 2405 SOE NASERE Physi is seat L. are 30399; GW 3 L., SEEKWP ae. : 29875 Solanum elaeagnifel um Cav., SJ 14246; GW 3146 STYRACACEAE Styrax americanus Lam.LB 30384a, 30243; SJ 13910 Styrax grandifolius Aiton, SJ 14005 SYMPLOCACEAE Symplocos tinctoria (L.) Her, LB 29834, 29378, 30095; WRC 17360 TEE IERID A SENE hii (Desv.) Morton, LB 29898; SJ 13933 TYPHACEAE Typha latifolia L., LB 31073 ULMACEAE Celtis laevigata Willd., LB 29861 Planera aquatica J.F. Gmel., LB 30156, 29932; SJSEWP Lantana urticioides Heyek, GW 3141 Phyla nadiflora (L.) Greene, SEEKWP. 3079; LB 29869; SJ 14097 *Verbena brasiliensis Vell., LB 30722; SEEKWP 3080 Verbena halei Small, a se SEEKWP 3114 Verbena scabra Vahl, LB 29902 VIOLACEAE Viola lanceolata subsp. vittata (Greene) Russell., WRC 17357; SJ 13699; MM 3762; LB 30296 iola palmata L., SJSEWP 15059; SJ 13700; GW 30469; LB 30294 Viola sororia Willd. var missouriensis (Greene) McKin- ney, GW 3050; LB 31067 VISACEAE Phoradendron serotinum (Raf) M.C. Johnston, $J 14505; VITACEAE Ampelopsis arb p a (L.) Koehne, WP 067;LB 29934 Pa ened) iInquetolia (L.) Planch., LB 29850 Vitis aestivalis Miche LB 30724, 30062; SJ 14105 Vitis cinerea (Engelm) Millard. var. cinerea, LB 29326; SJ 141 16a 15067 Ulmus alata Michx., LB 29920; SJSEWP 15067 Vitis rotundifolia Michx., SEEKWP 3087; LB 30219 Ulmus americana L., LB 29858, 30288 00 XYRIDACEAE Vien hi. R | yris ambigua Bey.ex Kunth,LB 29419, 30116;SJ 14083: GW 3446 Xyris baldwiniana Schult.,LB 29413; SJ 14045a URTICACEAE Boehmeria cylindrica (L.) Sw., LB 30616, 29891, 30052 VALERIANACEAE Valerianella radiata (L.) Dufr., SJSEWP 15088 Xyris difformis Chapm. var. difformis, ol a geal Xyris supicai L.C. Rich., LB 30218a; SJ 1 L.,WP 179 Xyris laxifolia Mart.var.iridifolia cae mn LB 30083; ream id io (Sweet) Troncoso, SJ 14247; GW 3107a EWP 1 Xyris stricta Chapm. var. obscura Kral, LB 30109, 29950, 29981; MM 3535 Notes Coreopsis gladata.—We are merging C. linifolia with this species following Strother Dichanthelium ovale.— WRC 17362 has the papilose-based hairs of subsp. villosissimum but the longer (up to 3 mm) spikelets of subsp ovale (refer to Diggs et al. 2006). Thus we are unable to assign a subspecies to this collection. Echinacea pallida var. sanguinea.—This globally rare (G3) species was once on the Texas rare plant list (Pool et al. 2002; Wildlife Diversity Program 2004). It occurs in many east Texas counties (Turner et al. 2003). Eupatorium semiserratum.—A|though not mapped in Texas by Turner et al. (2003) nor by Siripum and Schilling (2006), plants matching the key characters and description in Siripum and Schilling (2006)are found in east Texas. Lechea minor.—This is the first report of this species for Texas.; details are to be provided in a future publication. Pluchea baccharis.—An older name for P. rosea. Sarracenia alata.—This species was introduced to the Lance Rosier Unit in the 1960s or 1188 BRIT.ORG/SIDA 22 2) 1970s where it has thrived. It occurs naturally and is common in other parts of the Big Thicket region. Soliva anthemifolia.—T his name replaces S. mutisii in Flora of North America volume 19. Spiranthes longilabris.—T his globally rare (G3) species was once on the Texas rare plant list (Pool et al. 2002; Wildlife Diversity Program 2004). In Texas, it occurs in wetland pine savannas only in Hardin and Newton Counties (Liggio W Liggio 1999). Vicia hirsuta.—Jerry Flook (1975) reported this introduced legume from Shelby County and Turner et al. (2003) mapped it only in this county. It is a frequent weed near the site of the former Teel residence along the road to Little Pine Island Bayou. DISCUSSION There are 114 families (113 native) and 352 genera GLI native) for the 694 taxa (612 na- tive) on the Lance Rosier Unit list. Ten percent of the taxa are non-native. Eight families: Poaceae (96 taxa), Cyperaceae (87 taxa), Asteraceae (84 taxa), Fabaceae (38 taxa), Scrophulariaceae (19 taxa), Onagraceae (16 taxa), Rosaceae (15 taxa), Euphorbiaceae (15 taxa), and Rubiaceae (14 taxa), account for 56 % of the taxa collected. The largest genera are Carex (31 taxa), Dichanthelium (21 taxa), Rhynchospora (19 taxa), Cyperus (15 taxa), Quercus (15 taxa), and Juncus (12 taxa). We compared the present list of species for Lance Rosier Unit with a sample of the species reported by Turner et al. (2003) in their Atlas of the Vascular Plants of Texas and found that only 74% of the species on the Lance Rosier Unit list are reported as occurring in Hardin County in that source. This is not surprising considering that, until recently, little focused collecting has been undertaken in the Big Thicket region. Basic floristic collecting and documenting in the Big Thicket region are still needed. While it is the case that our list is incomplete, as are all floras, and a few taxa re- ported here may no longer exist on the unit. Most taxa that have grown in the Lance Rosier Unit during the past quarter century have been collected (probably 85+ %). We estimate that the Lance Rosier Unit has about 620 to 705 native species/taxa. ACKNOWLEDGMENTS This study was supported in part by a National Park Service Cooperative Agreement No. CA 14001004 to Paul Harcombe who also provided the publication funds. Thanks to Tom Wendt and Lindsay Woodruff at TEX for the prompt loan of some critical specimens and to Eric Keith of Raven Environmental who reviewed the paper. REFERENCES Avivsal,G. 1979.Wild flowers of the Big Thicket, east Texas, and western Louisiana. Texas A.& M.Press, College Station. Brown, L.E., B.R. MacRoserts, M.H. MacRoserts, PA. Harcomee, WW. Pruess, LS. Evsik, and D. JOHNSON. 2005. Annotated checklist of the vascular flora of the Turkey Creek Unit of the Big Thicket National Preserve, Tyler and Hardin counties, Texas. Sida 21:1807-1827. Brown, L.E., B.R. MacRoserts, M.H. MacRoserts, PA. Harcomee, W.W. Pruess, IS. Evsix, and S.D. Jones. 2006. Annotated checklist of the vascular flora of the Big Sandy Unit of the Big Thicket National Pre- serve, Polk County, Texas. Sida 22:705-723. Desuotets, J.D. 1978. Soil survey for the Big Thicket National Preserve, Texas. U.S.D.A. Soil Conservation Service, College Station, Texas. BROWN ET AL., FLORA OF PRESERVE 1189 Dices, Jr., G.M., B.L. Liescome, M.D. Reeo, and RJ. O’Kennon. 2006. Illustrated flora of East Texas. vol 1. Sida Bot. Misc. 26:1-1594. FLook, JM. 1975. Additions and corrections to the flora of Texas. Sida 6:114. Harcomee, PA. and P.L. Marks. 1979. Forest vegetation of the Big Thicket National Preserve. Unpub- lished report: U.S. Park Service, Santa Fe, New Mexico. Harcomee, PA., J.S. Guitzenstein, R.G. Knox, $.L. Orzett, and E.L. Brioces. 1993. Vegetation of the longleaf pine region of the West Gulf Coastal Plain. Proc. Ann. Tall Timbers Fire Ecol. Conf. 18:83-103. Jones, S.D., J.K. Wiprr, and PM. Montcomery. 1997. Vascular plants of Texas. Univ. Texas Press, Austin. Kaatesz, J.T. and C.A. MeacHam. 1999. Synthesis of North American flora. Version 1.0. North Carolina Botanical Garden. Chapel Hill. Liccio, J.and A.O. Liccio. 1999. Wild orchids of Texas. Univ. Texas Press, Austin. MacRoserts, B.R. and M.H. MacRoseats. 1998. Floristics of wetland pine savannas in the Big Thicket National Preserve, southeastern Texas. Phytologia 85:40-50. MacRosrats, B.R., M.H. MacRoseets, and L.E. Brown. 2002.Annotated checklist of the vascular flora of the Hickory Creek Unit of the Big Thicket National Preserve, Tyler County, Texas. Sida 20:781-795. Marks, PL.and PA. Harcomer. 1981.Forest vegetation of the Big Thicket, southeast Texas. Ecol. Monogr 567-305: McLeop, C.A.1971. The Big Thicket forest of east Texas. Texas J. Sci. 23:221-233. Parks, H.B.and V.L. Cory. 1936. Biological survey of the east Texas Big Thicket area. Texas Agric. Exp. Sta., College Station. Peacock, H.H. 1994. Nature lover's guide to the Big Thicket. Texas A. & M. Press, College Station. Poot, J.M., J.R. SincHurst, D.M. Price, and W.R. Carr. 2002. List of the rare plants of Texas. Wildlife Diversity program of the Texas Parks and Wildlife Department and the Nature Conservancy, Austin, Texas. Unp;ublished January 2000 edition. Siripum, K.C. and E.E. ScHitinc. 2006. Eupatorium. In: Flora of North America, vol. 21. Oxford University Press. New York, NY. StROTHER, J.L. 2006. Coreopsis section Eublepharis. In: Flora of North America, vol.21.Oxford University Press. New York, NY. Turner, B.L., H.NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Sida Bot. Misc. 24: 1-888. Warson, G.E.1979.Big Thicket plant ecology:an introduction. Big Thicket Mus. Publ. Ser.,,No.5, Saratoga, Texas. Watson, G.E.1982.Vegetational survey of Big Thicket National Preserve. Unpublished report.Big Thicket National Preserve, Beaumont, Texas. Wivowire Diversity Procram. 2004. A list of the rare plants of Texas. Wildlife Diversity Program of Texas Parks and Wildlife Department and The Nature Conservancy of Texas. Unpublished report. 1190 BRIT.ORG/SIDA 22(2) Book REVIEW Kenbatt R. Lamkey and Micnari Ler. (Eds.). 2006. Plant Breeding: The Arnel R. Hallauer International Symposium. (ISBN 0-8138-2824-4, 978-0-8138-2824-4, hbk.). Black- well Publishing. 2121 State Ave., Ames, [A 50014-8300, U.S.A. and 9600 Garsington Road, Oxford, OX4 2DQ UK. (Ondiees: at 292- vane 55 — gia fax 1-800-862- 6057, wwwhlackwell lcom I hing.com,). $149.99, 379 pp., numerous illustrations, rel A total of 84 authors contribute a collection of 27 chapters, the first “Plant Breeding: Past, Present, and Furure.’ From the Preface: * The world of plant breeding has ee ienced dramatic changes during the [last 45 years]. At the institutional level, pternauena) centers of cro] nt have emerged and declined, legal and ethical issues have become the private Secor (national encarta: federal governments, universi- ies) | li ified and placed greater emphasis on basic res to varietal development. Changes in ines inucture (e.g., ons season nurseries, service eleborater ies) and tec ee (e.g., computers, machinery, ana- of plant breeders to sialuare more germplasm in more we ve in more environments ane to identily ge ane pes that exhibit optimal adaptation to the needs of s of nature, and the desires of the market. Nascent developments in basic biological and infor- > coronal sciences, as -exenmplil fied by the gradual annotation of entire genomes and their gene products, have provided additional tools and options for various aspects of plant breeding.”—Guy Nesom, Botanical Research In- stitute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A. Book NOTICE M. Nevin Smit. 2006. Native Treasures: Gardening with the Plants of California. (ISBN O- 520-24425-7, pbk.). University of California Press, California/Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08018, U.S.A. (Orders: www.ucpress.edu/, 609-883-1759, 609-883-7413 fax). $24.95, 278 pp., color photos, 7" x 10". From the author— This is not a ‘how-to’ gardening book, as practical subjects are addressed here in detail, and it is certainly not the comprehensive encyclopedia ... Rather, it combines my own personal thoughts, some- times maverick opinions, and experience with a woody measure of plainer facts on selected g cup of native plants and their culture. The plant genera described here include most of the major ones in cultivation, but also some more obscure but garden-worthy groups, thrown in as whim and personal interests dictate.” Smith divides his favorites into “trees” (oaks, madrono), “shrubs” (e.g, wild lilac, manzanitas, flannel bushes, giant poppies), “two broader genera” (sage and buckwheat), “subshrubs and herbaceous perennials” (e.g., penstemons, iris, lewisias, blue eyes, golden eyes), and “bulbs and corms” (lilies, brodiaeas and friends). He ends with “Small Matters—The trouble with annuals”: “We forget that Nature's best displays are occasional and unpredictable. We tell ourselves | that anything seen over such vast areas s [Heer of annual wildflowers] must be easy to grow. And so they are, ina well tended t ses pot. Creating ar splay in the open garden is more difficult—as | have learned through harsh experier Bae book is Saal at least, aimed toward California gardeners, but surely many of the species could be in North America. “California gardeners, like their counterparts the world over, can be woefully oO ignorant of the flora close at hand. They are influenced by a long and hallowed tradition of importing horticul- tural treasures from Europe, Asia, and other far-flung regions.” But .... “A fair number of native plants are now used eerenan ea ly with exotics of similar features even by g pater whose notion of geographic a te inthe local nursery.” Even for non-gardeners, this is a great book with lots of interesting biology, fun to read, even if youre not from California—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort a TX 76102-4068, U.S.A. SIDA 22(2): 1190. 2006 AN UPDATED, ANNOTATED VASCULAR FLORA OF CADDO PARISH, LOUISIANA, WITH NOTES ON REGIONAL PHYTOGEOGRAPHY AND ECOLOGY Barbara R. MacRoberts and Michael H. MacRoberts Bog Research, 740 Columbia Shreveport, Louisiana 71104, U.S.A. and Herbarium, Museum of Life Sciences Louisiana State University in Shreveport Shreveport, Louisiana /1 al Red River Watershed Management Institute Louisiana State University in Shreveport Shreveport, Louisiana 7 USA. Ga ABSTRACT It has been a quarter century since a checklist of the vascular flora of Caddo Parish, Louisiana, was published. Therefore, we revise and update the Caddo Parish checklist, accepting only vouchered species. We found 1405 species (1168 native, 237 non-native). Additionally, we examine the Caddo Parish flora in terms of its phytogeo- graphical affinities. It is closely associated with the eastern United States and particularly with the southeastern United States. Caddo Parish appears to be particularly species rich compared to other parishes and counties of approwimately the same size across the southeast. This may be: not only because Caddo I Pa oe is abieise put | well-collected Caddo Par are particularly ie represented and account for a bigh percentage of West Gulf Coastal a endemic and rare oy species Key Worns: Floristic checklist, Caddo Parish, Louisiana flora RESUMEN Hace ya un cuarto de siglo que se publico el catalogo de la flora vascular de Caddo Parish, Louisiana. Por ello, revisamos y ponemos al dia el catalogo de Caddo Parish, aceptando tnicamente especies con pliegos testigo. Eaconanie re is (1168 nativas, 237 no nativas). Ademas, examinamos la flora de Caddo Parish en Esta muy asociada con el este y particularmente con el sureste de los Eotades Unidos. Caddo een merece ser Se auencnenes rico en especies conn Datade: con otros iaseaael de aproximadamente el mismo tamano en el sureste | solo porque C porque ha sido ae bien muestreado ee las unidad aparecen en LCaddo Parish, estan muy bien representac do y aparece un alto porcentae de S} Sy endémicas de la Teas costera cel oeste ael golfo. INTRODUCTION Caddo Parish, Louisiana, is located in the Red River watershed in the center of the West Gulf Coastal Plain (Fig. 1). It is bordered on the north by Arkansas, the west by Texas, the east by the Red River, and the south by DeSoto Parish. The only boundary that could be considered ecologically significant is the Red River; the remainder are political. Approxi- mately equidistant from Caddo Parish are blackland prairies to the west, the Ouachita Mountains to the north, the Mississippi River and its extensive floodplain to the east, and the Gulf of Mexico to the south. Caddo Parish is located in the Oak-Pine-Hickory phyto- geographical region of eastern North America (Braun 1950; Skeen et al. 1993; MacRoberts SIDA 22(2): 1191— 1219. 2006 1192 BRIT.ORG/SIDA 22(2) ai SS AL che nae By penis cael pzaans Fic. 1. Location of Caddo Parish, Louisiana S& MacRoberts 2003a). McLaughlin (in press) places it in the Austroriparan floristic subprovince, with some characteristics of Comanchian and Illinoian subprovinces. The topography of Caddo Parish is relatively simple. The Red River floodplain cov- ers about 28 percent of the 2,283 km sq. parish; the remainder is uplands (Fig. 2). The {floodplain forms a continuous north-to-south border along the eastern edge of the par- ish. The floodplain can be narrow as when the river approachesa bluff, but it is generally several kilometers wide. At the edge of the river are natural levees and swales. Floodplain elevations range from approximately 60 meters at the northern edge of the parish to about +3 meters at the southern edge. The overall north-south slope is very slight. Sediments are almost entirely of Red River alluvial origin and are Pleistocene/Holocene (Edwards et al. 1980). Uplands flank the floodplain at its western edge. This often involves an abrupt el- evation transition generally in the range of 10 to 20 meters. Elevations rise to no more than 137 meters in rolling low hills and relatively flat terraces. The upland is geologically older than the alluvium and generally consists of Tertiary deposits. The geology, soils, and climate of Caddo Parish are described by Teague and Wendt (1994) and are a con- tinuum of east Texas (Diggs et al. 2006). MACROBERTS AND MACROBERTS, VASCULAR FLORA OF CADDO PARISH LOUISIANA 1193 . ARKANSAS C f gs (eA | dD Loe L . 1 J if] j ila rears. 4 1 7A\ Fig. 2 I I 1194 BRIT.ORG/SIDA 22(2) Land surveys conducted by the Government Land Office in the 1830s show that in presettlement times the uplands were dominated by oak, pine, and hickory and the flood- plains by cottonwood, sycamore, willow, pecan, box elder, ash, cypress, and hackberry, with essentially no overlap in species between upland and floodplain (MacRoberts & MacRoberts 2005). In some parts of the uplands, pine was nearly absent and oak and hickory were dominant; in others, pine reached about 50 percent of the canopy. There appeared to have been no monospecific pine forest in Caddo Parish. The distribution of woody species appears to be the same today as it was 170 years ago (MacRoberts & MacRoberts 2005). Historically, prairies were scattered across the region in both the uplands and flood- plain (MacRoberts et al. 2003; MacRoberts & MacRoberts 2004a). The large “Caddo Prai- ries” on the floodplain near modern day Gilliam and Hosston no longer exist, and we have little or no information about them aside from the brief descriptions Custis and Freeman made in 1806 and those in the Government Land Office survey maps and line notes (Flores 1984; MacRoberts & MacRoberts 2005). Canebrakes and cedar forests ap- parently were common in the early nineteenth century butare entirely gone today (Flores 1984; MacRoberts et al. 1997). The Red River Raft, an immense logjam long since cleared, caused extensive flooding and the creation of “raft lakes,” the water level of which rose and fell with the annual fluctuations of the river (Triska 1984). Wallace, Cross, and Caddo lakes are impounded remnants of the raft lakes and continue to be dominated Dy cy- press. Oxbow lakes are common on the floodplain. While the effects of Native Americans on the Caddo Parish landscape are unknown, the effects of Euroamericans have been extensive. The river, its floodplain, and adjacent upland ecosystems have undergone major modification over the past two centuries. Es- sentially the entire landscape has been converted to farms and plantations and, latterly, urban sprawl. The once continuous savanna/forest with its numerous plant communi- ties covering hundreds of thousands of hectares is now essentially gone with only frag- ments of the natural vegetation remaining and few areas, if any, in virgin condition. Bald- cypress swamps, black willow riverbank shrublands, and cottonwood forests hang on while some floodplain communities such as cedar forests, canebrakes, and prairies have vanished entirely. The upland has also been modified, mainly by urban sprawl, logging, fire suppression, and clearing for farms. Vast areas of forest were clear-cut in the latter part of the nineteenth and early part of the twentieth century and replanted as pine plan- tations. Caddo Parish is one of the best collected and reported Louisiana parishes and is cer- tainly the best collected and reported parish or county in the “Ark-La-Tex” region—north- eastern Texas, southwestern Arkansas, and northwestern Louisiana (Smith 1988: Tho- mas & Allen 1993-1998; Turner et al. 2003). Caddo Parish was first visited and collected by Peter Custis in 1806 (Flores 1984: MacRoberts et al. 1997; MacRoberts & MacRoberts 2004b). Josiah Hale collected in the parish in the mid-nineteenth century. N.F Peterson from Louisiana State University and Reginald WS. Cocks from Tulane University collected in Caddo Parish in the early twen- tieth century. FW. Pennell from the Academy of Natural Sciences in Philadelphia worked briefly in the area in 1913 when he collected the only known example of Agalinis caddoensis (MacRoberts 1978b; Canne-Hilliker & Debrule 1993). EJ. Palmer from Mis- souri Botanical Garden and D.S. and HB. Correll (1941), working out of Duke University, collected in the area in the late 1930s. Clair A. Brown from Louisiana State University — LAUUY PARISH, LOUISIANA 1195 sporadically collected in Caddo Parish through the mid-century. John W. Thieret, then at the University of Southwestern Louisiana in Lafayette (now University of Louisiana in Lafayette), collected in Caddo Parish in the 1960s. During the early 1970s, Roselie Overby (1974) collected the flora of Caddo Parish for her thesis at the University of Louisiana at Monroe (formerly Northeast Louisiana University) under the direction of R. Dale Tho- mas, who also collected regularly in the parish into the twenty-first century. Upon begin- ning the herbarium at Louisiana State University in Shreveport in the early 1970s, D.T. MacRoberts collected the parish for the next three decades. His collections and those of various students and faculty resulted in numerous contributions to the flora of Louisi- ana (MacRoberts 1977a, 1977b, 1978a, 1978b, 1979a, 1979b, 1980a, 1980b, 1980c, 1984a, 1984b, 1987, 1989; MacRoberts et al. 1997). Allen et al. (2004) collected mainly grasses in Caddo Parish through the late twentieth and early twenty-first centuries. Special studies have been undertaken such as those of xeric sandylands in northern Caddo Parish (MacRoberts & MacRoberts 1995) and the Bickham-Dickson Park/Red River Watershed Research Park on the Red River floodplain in Shreveport (MacRoberts & MacRoberts in prep.). Other notable collections are those of Larry Raymond from the Walter Jacobs Memorial Park, near Blanchard (Thomas et al. in press). The Louisiana Natural Heritage Program has been active in documenting the rare flora of Caddo Parish (Louisiana Natu- ral Heritage Program 2006; Reid & Faulkner in press). The purpose of this paper is to provide an updated, vouchered checklist of the vas- cular flora of Caddo Parish, put Caddo Parish in its phytogeographic context, summa- rize data on rare plants found in Caddo Parish, and summarize data on regional endemics. The construction of the list has been both aided and hampered by precedents. While previous lists often provided information on where specimens might be found, they did not do so invariably. Although we located vouchers for most reported taxa, many reported taxa could not be located, presumably because the sf were misfiled (lost), out on loan, initially misidentified and subsequently annotated to another taxon, or were lo- cated in an unspecified herbarium. Additionally, some authors reported taxa for which no voucher was collected (ie., “sight records”). METHODS Using published accounts (MacRoberts 1979a, 1984a, 1989; Gandhi & Thomas 1989; Tho- mas & Allen 1993-1998: MacRoberts & MacRoberts 1995; Allen et al. 2004; Louisiana Natural Heritage Program 2004, 2006; Thomas et al. in press; MacRoberts & MacRoberts in prep.), we compiled a checklist Gwith vouchers where possible) of species reported for Caddo Parish. We then checked the list against Kartesz and Meacham (1999) and Flora of North America (Flora of North America Editorial Committee 1993-2006) to eliminate synonyms and to correct nomenclature. We searched LSUS and NLU for vouchers in ad- dition to obtaining loans from BRIT, DUKE, LAF LSU, LTU, MO, NLU, NO, and US. Some authors included various waifs and garden and horticultural “escapes” (e.g., corn[Zea mays L.], and okra [A belmoschus esculentus (L.) Moench]) and we had to make a judgement as to their status in the flora. We decided to eliminate almost all of these un- less we had good reason to believe that they are propagating themselves in the wild or at least persisting. Our taxonomic viewpoint favors “lumping” and we do not concern ourselves here with taxa below the species level, the reason being that not only are authors inconsistent on the number of infraspecific taxa recognized (e.g., compare Smith 1994; Thomas & Allen 1196 BRIT.ORG/SIDA 22(2) 1993-1998: Turner et al. 2003) but also on how to divide (and identify) the often subtle morphological differences between certain species. Consequently, when comparing flo- ristic richness between various geographic areas, our list should be recognized as a “spe- cies’ list, not a “taxa” list. We do not include species on the vouchered list that we have not inspected except for some Cyperus determined by D. Ferguson (LSU, pers. comm.) and two specimens determined by C. Reid (Louisiana Natural Heritage Program, pers. comm.). Nomenclature and authorities follow Kartesz and Meacham (1999) in most cases. For acaulescent violets, we follow McKinney (1992) and McKinney and Russell (2002). For Gamochaeta, we follow Nesom (200, 2004b). In order to put Caddo Parish into a phytogeographic context, we randomly selected 417 native species from our list and, using Kartesz and Meacham (1999) and Thomas and Allen (1993-1998), determined their distribution by state or region across North America north of Mexico (see MacRoberts & MacRoberts 2003b for method). On the basis of this and other phytogeographical information, we developed a thematic map of the floristic affinities between Caddo Parish and the remainder of North America. Rare plants often indicate rare habitat and vice versa. Using the Louisiana rare plant list (Louisiana Natural Heritage Program 2006), we listed all rare plants found in Caddo Parish and their primary habitat. We then summarized these data by habitat. Regionally endemic plants also indicate unique habitat, as do singularities. Because Caddo Parish is in the center of the West Gulf Coastal Plain, we used a list of endemics in that region to determine how many occur in Caddo Parish and with which community they are asso- ciated (M.H. MacRoberts et al. 2002). Additionally, Louisiana plant species that occur only in Caddo Parish were determined using the Louisiana Natural Heritage Program (2006) rare plant list and Thomas and Allen (1993-1998). We listed these and their associated habitats. RESULTS AND DISCUSSION We documented 1405 species in Caddo Parish (1168 native and 237 non-native). The vouchered checklist of the vascular plants of Caddo Parish is given after the discussion. Figure 3 gives the results of the phytogeographic analysis of Caddo Parish species. Table I gives Caddo Parish rare plants/plant community associations. Table 2 gives West Gulf Coastal Plain endemic species/community associations that occur in Caddo Parish. Table 3 lists the Louisiana native species found only in Caddo Parish with their associated com- munities. The figure of 1405 species for Caddo Parish is higher than that for most parishes and counties of similar size throughout the southeastern United States (e.g., Nesom & Brown 1998; Leidolf et al. 2002; Neyland 2002; Marsico 2005). However, Caddo Parish has been better collected than most parishes and counties and additional collecting would bring up their species counts. North America north of Mexico has approximately 19,000 vascular plant species (Thorne 1993; Kartesz @ Meacham 1999). Caddo Parish has 7.4 percent of this total but constitutes only 0.01 percent of the land area. Likewise, while Caddo Parish constitutes only about | percent of the land area of the West Gulf Coastal Plain, it contains about 36 percent of the West Gulf Coastal Plain flora. The Caddo Parish flora is eastern and specifically southeastern. Caddo Parish shares 99 percent of its flora with Texas, 98 percent with the remainder of Louisiana, 98 percent with Arkansas, 91 percent with Mississippi and Alabama, 88 percent with Oklahoma, 50 MACROBERTS AND MACROBERTS, VASCULAR FLORA OF CADDO PARISH, LOUISIANA 1197 at el | ¥ Se. 30-40 Kf —Nee“apsnt Wir KT ene MN Gaxrem: ae percent with the New England states, but only 21 percent with New Mexico. While Caddo Parish has 99 percent of its flora in common with Texas, there is a sharp transition zone across Texas in which eastern species drop out and are replaced by western species (MacRoberts & MacRoberts 2003b). The transition is much less dramatic to the north and east. This pattern is also shown in McLaughlin (in press). Thirty-six percent of the Louisiana rare species listed by the Louisiana Natural Heri- tage Program (2006) for Caddo Parish are essentially specific to one community: xeric sandylands (MacRoberts & MacRoberts 1995; B.R. MacRoberts et al. 2002). The next ranked community for rare species for Caddo Parish is calcareous prairie with 16 percent (MacRoberts et al. 2003) and saline prairie with 14 percent, followed by pine-hardwood forest with 1] percent (Larke & Smith 1994). The remaining 23 percent of rare species are spread among six plant communities Caddo Parish has 45 of the approximately 100 West Gulf Coastal Plain endemic taxa (MacRoberts et al. 2002), of which 64 percent are associated with xeric sandylands. The next highest communities for endemics are baygalls and calcareous prairies with 9 and 7 percent, respectively. The remainder of endemics are spread among five communities Of the 15 native Louisiana species occurring only in Caddo Parish, 8 occur in xeric sandylands, once again pinpointing this habitat/community as unique. Since two ities account for the majority of rare and endemic species in Caddo Parish, perhaps some comments on these communities is desirable. Xeric sandylands are found in eastern Texas, southeastern Oklahoma, southwestern Arkansas, and western Louisiana (McBryde 1933; MacRoberts & MacRoberts 1994, 1995, BRIT.ORG/SIDA 22(2) 1198 Taste 1.Rare plants/community associations for Caddo Parish. Community Number of Rare Plants Percentage Xeric sandyland 27 36 Calcareous Prairie 12 16 Saline Prairie 10 14 Pine Harwood Forest 8 1] Calcareous Forest 5 7 Baygall 3 4 Swamp 3 4 Small Stream Forest 2 3 Batture Forest ] 1 Unknown e) 4 Total 74 100 Taste 2.Endemic West Gulf Coastal Plain plant species with community associations for Caddo Parish. Community Number of Endemic Plants Percentage Xeric sandyland 29 64 Baygall 4 9 Calcareous Prairie 3 7 Pine-Hardwood Forest 2 4 Floodplain/Bottomland 2 4 Mayhaw Pond 2 4 Small Stream Forest | 2 Barrens | 2 Unknown 1 2 Total 45 100 1996, 1997; B.R. MacRoberts et al. 2002; Diggs et al. 2006). They are open to sparsely wooded areas that typically occur on terraces or ridges composed of deep sands, generally of marine Tertiary origin. Water and air move through these sands creating a dry, desert- like environment. Soils, where undisturbed, are of ten cryptogamic. Lichens Cladonia spp.) may be common. Quercus incana is common, but stunted Q. stellata and Q. margarettiae are also present. Other characteristic species include Shreneie ry id, Cnidoscolus texanus, Coreopsis intermedia, oe. ton aneyr am bITa. Er iogonum longifolium, E. multifolium, Froelichia floridana, H Matelea cynanchoides, Mirabilis albida, Opuntia humifusa, Paronychia drummondii, Pediomelum digitatum, P. hypogaeum, Penstemon murrayanus, Phlox drummondii, Polygonella americana, Selaginella arenicola, Streptanthus hyacinthoides, Talinum rugospermum, Tetragonotheca ludoviciana, Thelesperma filifolium, Tradescantia reverchonii, Yucca louisianensis, and Zornia bracteata. Floristic richness is high in this community. Al- though widespread, xeric sandylands are not a dominant community across the West Gulf Coastal Plain and are generally found in relatively small patches. At the time of European settlement, it has been estimated that xeric sandylands accounted for less than one percent of western Louisiana (Lester et al. 2005) and, although there are no pub- lished figures, probably much less than this in either Oklahoma or Arkansas. In Texas, MACROBERTS AND MACROBERTS, VASCULAR FLORA Of LOUISIANA 1199 Taste 3. Native Louisiana species found only in Caddo Parish with community associations. Agalinis caddoensis (community unknown) Callirhoe digitata (prairie? Canes dno leiaee (oraie Carex nd forest, roadside, saline prairie?) Coreopsis intermed poe sandyland) Dalea ‘phle eoides (xeric sandyland) Dasistoma macrophylla (mixed pine-hardwood forest) Helianthemum rosmarinifolium (mima mounds in saline prairie) Loeflingia squarrosa (xeric sandyland) vai ed RAG CNg aS oe sandyland) eric sandyland) Prunus an (xeric Sa ie spermum (xeric sandyland) Thelespe filifolium (xeric sandyland) Trillium pusillum (baygall) a eer | -| especially along the Carrizo formation, xeric common, but for the whole of this region this y is unlikely to have amounted to more than 5 percent of the area. Today, most xeric sandylands have been destroyed. Again figures are not available, but probably less than 5 to 10 percent of the original remain in any natural condition. Most of this community has been converted to pine plantations, grazing land, and wa- termelon patches. In Caddo Parish, it is decidedly rare and is found only in the northern part of the parish near Mira, Ida, Rodessa, and Vivian. Over the last twenty years, we have watched considerable acreage of this community vanish under cultivation, urban sprawl, and habitat deterioration caused by fire suppression (MacRoberts & MacRoberts 1995, 1998). Generally speaking, therefore, it represents a minor community by area but is an important community because of endemism and locally rare species. Calcareous prairies were once widespread across the West Gulf Coastal Plain (MacRoberts et al. 2003; MacRoberts & MacRoberts 2004a; Lester et al. 2005). Virtually none of this community is left today and none in Caddo Parish although prairies were once fairly common not only in the uplands, but on the floodplain as well (MacRoberts et al. 1997: MacRoberts & MacRoberts 2004b). While prairies were common in central Texas (Diggs et al. 1999), far less than one percent of Louisiana (or Caddo Parish) was ever prairie land (MacRoberts et al. 2003; MacRoberts & MacRoberts 2004a). Today, Louisi- ana prairies can be counted only in the tens to hundreds of hectares. Another community that deserves mention because of its rareness is saline prairie. This community occurs in several places in the West Gulf Coastal Plain (McInnis et al. 1993; Keith et al. 2004: Lester et al. 2005). It has received some attention because Geocarpon minimum, which is listed as federally threatened, occurs in it. In Caddo Parish, there are two known saline prairies. One, about 0.5 ha, was discovere d by Overby (1974); the other, a prairie of 11 ha, was found in 2005 by C. Reid of the Louisiana Natural Heritage Pro- gram (Barron Road Saline Prairie). Overby found Talinum parviflorum in the prairie she located. We examined the Overby prairie several times, and relocated Talinum, but the last time we visited the site in March 2006, it had nearly been destroyed by all-terrain vehicles (ATVs). Barron Road Prairie is also damaged by ATVs. Saline prairie is mostly treeless (mima mounds may be present), with many open soil areas known as “slicks” with cryptogamic, high sodium soils, and sparse vegetation. 1200 BRIT.ORG/SIDA 22(2) — Characteristic plants include Anagallis minima, Claytonia virginica, Crassula aquatica, Eragrostis lugens, Evolvulus sericeus, Houstonia micrantha, Houstonia pusilla, Houstonia rosed, Krigia occidentalis, Lepuropetalon spathulatum, Plantago pusilla, and Sporobolus vaginiflorus. Ten Louisiana and one globally rare species have been located at the Barron Road Saline Prairie. These are Cooperia drummondii, Geocarpon minimum, Gratiola flava, Helianthemum rosmarinifolium, Lotus unifoliolatus, Minuartia drummondii, Minuartia muscorum, Phacelia glabra, Schoenolirion wrightii,and Talinum parviflorum. The two surviving plant collections from the 1806 Freeman-Custis’ Red River Expe- dition are Eustoma russellianum (Hook.) G. Don and Veronicastrum virginicum (L.) Farw. (Flores 1984, MacRoberts &@ MacRoberts 2004b). Eustoma russellianum does not occur anywhere in the region today, and Veronicastrum virginicum is very rare in the region and has not been found in Caddo or Bossier parishes since Custis’ original collection. We have noted this same pattern with subsequent early collectors in the area. Pennell found the only species endemic to Caddo Parish—Agalinis caddoensis—near Shreveport in 1913; it has not been found since. Hale, Cocks, Peterson, Correll, Palmer, and other early collec- tors found species in the area that are either very rare today or entirely missing (e.g., Buchloe dactyloides, Dasistoma macrophylla, Gratiola flava, Helianthus occidentalis, Helianthus petiolaris, Onosmodium bejariense, Panicum flexile, Prosopis glandulosis, Sophora affinis, and Taenidia integerrima). This suggests that in the last two centuries, a great deal of the original habitat has been lost and that the flora was once richer in native species than it is today. ~— VOUCHERED CHECKLIST OF VASCULAR PLANTS The following is a vouchered list of the vascular plants for Caddo Parish. We list one voucher for each species. Key to vouchers: DTM = D.T. MacRoberts. Unless otherwise stated, his specimens are at LSUS. RDT = R. Dale Thomas. Unless otherwise stated, his speci- mens are at NLU.MM = BR. and M.H. MacRoberts. Unless otherwise stated, their speci- mens are at LSUS. Raymond = Larry R. Raymond. The herbarium where his specimens are located is always given. Hardy = L.M. Hardy. His specimens are at LSUS. Haynes = Robert Haynes. His specimens are at LSUS. Barbour = Philip Barbour. His specimens are at LSUS unless otherwise stated. Other collectors are given full citations. An * (asterisk) indicates a non-native species. PTERIDOPHYTES Polystichum acrostichoides (Michx.) Schott; DTM 2197 Woodsia obtusa (Spreng.) Torr; DTM 835 faeces platyneuron (L.) B.S.P; DTM 829 ee AZOLLACEAE m hyemale L.; Barbour 932 < ; ae ie Clute; DTM 1700 Azolla caroliniana Willd.; MM 5935 BLECHNACEAE Doria eae : Isoetes melanopoda Gay & Durieu ex Durieu; RDT 89029 Woodwardi lata (L.) T. Moore; DTM 1546 LYGODIACEAE Sm.; RDT 33699 lA m fs 7 si te | VVYUUCUVV UT CICt VIFGTEIICCH (L.) *L ygodium japonicum (Thunb. ex Murr) Sw.; DTM 1090 eo incdtes eae (L.) Kuhn; DTM 898 DRYOPTERIDACEAE Athyrium filix-femina (L.) Roth; DTM 1221 *Cyrtomium falcatum (L.f.) K.Presl; DTM 113 (question- ably naturalized) Onoclea sensibilis L.; DTM 950 OPHIOGLOSSACEAE mala ll a atum (Sav.) Underwood; DTM 2012 preng.; Raymond 1600 NLU see Sw. RDT 148118 Botrychium virginianum (L.) Sw.; DTM 2222 Ophioglossum crotalophoroides Walt.; MM 6062 iecocfiy Bohn hy / ai. LOUISIANA 1201 ied ee Sng e snail Prantl; RDT 34526 LSU udicaule Lf; RDT 5 1996 Onhiogiesum petiolatum Hook.; RDT 33845 LSU Ophioglossum vulgatul re le MM 1 OSMUNDACEAE Osmunda cinnamomea L., DTM 2156 Osmunda oe L.; DTM 2157 POLYPO Pleopeltis eee oides (L.) Andrews &Windham;DIM 949 SELAGINELLACEAE Selaginella arenicola Underwood; MM 2840 THELY PTERIDACEAE Thelypteris kunthii (Desv.) Morton; DTM 1734 Thelypteris palustris Schott; DTM 1212 GYMNOSPERMS CUPRESSACEAE Juniperus virginiana L.; DTM 1609 odium distichum (L.) L.C. Rich.; MM 6514 PINACEAE Pinus echinata P. Mill; MM 6199 Pinus taeda L. MM 5982 ANGIOSPERMS MONOCOTYLEDONS AGAVACEAE Manfreda virginica (L.) Salisb. ex Rose; DTM 1861 Yucca louisianensis Trel., MM 241 ALISMATACEAE Echinodorus cordifolius (L.) Griseb.; DIM 77 Sagittaria graminea Michx, DTM 1315 Sagittaria latifolia Willd.; DTM 672 *Sagittaria montevidensis Cham.& Schlecht; DTM 1590 Sagittaria papillosa Buch.; DTM 1300 Sagittaria platyphylla (Engelm.) J.G.Sm.;DTM 1216 ARACEAE Arisaema dracontium (L.) Schott; DTM 377 Arisaema triphyllum (L.) Schott; DIM 2153 *Colocasia esculenta (L.) Schott; DIM 1306 Peltandra virginica (L.) Schott; DTM 2224 ARECACEAE Sabal minor (Jacq.) Pers.; MM 190 BROMELIACEAE Tillandsia ides (L.) L.; DTM 3002 BURMANNIACEAE Burmannia biflora L.; Reid 4639 LSU (Reid, pers.comm.) CANNACEAE *Canna indica L., DTM 1855 (questionably naturalized) COMMELINACEAE Gt lina communis L.; DTM 2329 Commelina diffusa Burm.f. DTM 1408 Commelina erecta L.; DTM 141 ommelina virginica L DIM 486 Tradescantia hirsutiflora Bush; DTM Tradescantia occidentalis (Britt.) fae DIM 473 Tradescantia ohiensis Raf; DTM 1 ae Tradescantia reverchonii Bush; DTM 1 CYPERACEAE Bulbostylis capillaris (L.) Kunth ex C.B. Clarke; RDT 155761 Bulbostylis ciliatifolia (Ell.) Fern.; MM 2833 pe ae icans wal ick ex Spreng.; Raymond 1722 NLU chwein.; Hyatt 8580 LSU ne arkansana (Bailey) ae ee 25936 LAF Carex atlantica ea RDT 95343 LS Carex austricina Mackenzie; nov 98507 Carex planed Dewey; RDT 17006 Carex Schwein; MM ae Carex is Schwein.; DTM 1736 Carex complanata Torr.& Hook.; Barbour 990 Carex crebriflora Wieg.; Raymond 1234 NLU Carex crinita Lam.; DIM 2231 Carex crus-corvi Shuttlew. ex Kunze; DIM 1353 Carex debilis Michx.; RDT 95345 Carex decomposita Muhl.; MM 7429 Carex festucacea Schkuhr ex Willd.; Thieret 22620 LAF Carex flaccosperma Dewey; Raymond 1213 LSUS Carex frankii Kunth; DTM 1281 Carex glaucescens Ell; Raymond 1359 LSU Carex glaucoidea be ex Olney; RDT 99544 Carex hyalina Boott; MM 3 ae Rae Steud.; ie 1430 NLU arex intumescens eee ea 979 ia Bailey; RDT Carex leavenworthii nes i Carex leptalea Wahl.; Reid a a (Reid, pers.comm.) Carex louisianica Bailey; D Carex ee ina Muhl. ex Re an D352 Ca arex lurida W Nahlenb.; DTM 1 236 Carex meadii Dewey; MM 2525 Carex muehlenbergii schkuhr ex Willd.; RDT 88719 Haynes 5697 Se anne Naczi; Raymond 1677 NLU s (Bailey) Small; ae 1214NLU Carex retrolexa DE ex ee rad LI UICA style HCAU DULKI Carex texensis (Torr.) Bailey; i 88223 Carex triangularis Boeckl.; RDT 99524 Carex tribuloides Wahlenb.; : 70062 Carex vulpinoidea Michx.; D 349 Cyperus acuminatus Torr. & Hess ex Torr.; DTM 2332 Cyperus compressus L.; RDT 32983 Cyperus croceus Vahl; MM 6834 1202 *Cyperus difformis L.; MM 6993 Cyperus echinatus (L.) Wood.; DTM 1931 Cyperus erythrorhizos role oo 5430 Cyperus esculentus L.; MM 6857 Cyperus filiculmis Vahl; 170174 (Ferguson, pers. comm.) ae os L.; CE. DePoe 6652-1 m.) eo ee ee Shireman 57 LSU ici iria L.; DTM Cyperus odoratus L.; nee Cyperus oxylepis Nees ex on RDT 37207 Cyperus plukenetii Fern.;Lynch 2639 LSUS Cyperus polystachyos Rottb.; Lynch 2674 LSUS Cyperus. pseudovegetus Steud.; RDT 166992 (Ferguson, pers.comm.) NLU (Fergu- Cyperus retroflexus Buckl.; MM 2725 Cyperus retrorsus Chapm.; MM 6630 *Cyperus rotundus L.; DTM 1361 Cyperus strigosus L; RDT 45424 (Ferguson, pers.comm.) Cyperus virens Michx.; Overby 85 NLU (Ferguson, pers. comm.) Eleocharis acicularis (L.) Roemer & J.A. Schultes; DTM 1458 Fleocharis obtusa (Willd.) J.A. Schultes; DTM 1238 Eleocharis palustris (L.) Roemer & J.A.Schultes; MM 6429 Eleocharis parvula (Roemer & J.A. Schultes) Link; Bar- bour 498 Eleocharis tortilis (Link) J.A.Schultes; RDT 33624 Eleocharis tuberculosa (Michx.) Roemer & J.A.Schultes; T 36268 Fimbristylis autumnalis (L..) Roemer & J.A.Schultes; DTM 1567 Fimbristylis miliacea (L.) Vahl: DTM 1542 Fimbristylis puberula (Michx.) Vahl; ete 2147LSUS Fimbristylis tomentosa Vahl; DTM Fimbristylis vahlii (Lam.) Link; a 2 Fuirena simplex Vahl: MM 6673 me valle Michx.; RDT 14068 Hook. & Arn. ex Torr,; ov 2829 Isolepis molesta (M.C. Johnston) S.G. Sm.; Shinners 28087 BRI Kyllinga cae Rottb., DIM 1928 yllinga odorata Vahl; DTM ae Kyllinga pumila Michx.; RDT 3 Lipo ae micrantha (Vahl) [ ae Shireman 56 ~_ ep caduca Ell; Raymond 2028 LSUS hospora corniculata (Lam.) A.Gray; DTM 1113 dee sic glomerata (L.) Vahl; Barbour 494 Rhynchospora harveyi W. Boott; MM 7427 Rhynchospora inexpansa (Michx.) Vahl; Haynes 5352 Scirpus atrovirens Willd.; Allen 7975 NO Scirpus cyperinus (L oe 1448 Scleri eG ata MCN: RDT Scleria tt ichx.; ne — LAF BRIT.ORG/SIDA 22(2) Sc leria paucinora Muhl. ex Willd.; Allen 9879 LAF Creel ChIQGUIOT TICE Michx.; RDT 92403 Websteria confervoides (Poir.) S. Hooper; Barbour 495 DIOSCOREACEAE Dioscorea villosa L; DTM 1288 HYDROCHARITACEAE *Egel ia densa Plat ich.; Barbour 483 Limnobium spongia (Bosc) L.C. Rich. ex Steud.; Haynes 5342 IRIDACEA Alophia ae mondii (Graham) R.C. fe oo 1762 tia lahue (Molina) Goldblatt; M ris brevicaulis Raf; Raymond 2094 ne : fulva Ker-Gawl.; French s.n. NLU Iris giganticaerulea Small; Overby 192 NLU “ris pallida Lam.; Laborde 44 *Iris pseudacorus L.; MM 172 Iris virginica L.. DTM 2291 Nemastylis geminiflora Nutt; DTM 369 eats aaa ifolium P.Mill; DTM 415 ickn.; DIM 340 Sisyrin ich jum langloissii aie MM 2558 Sisyrinchium minus Engelm.& A. ee a 25939 LAF Sisyrinchium rosulatum Bickn.; DTM JUNCACEAE Juncus acuminatus Michx.; RDT 81369 Juncus brachycarpus rae ;DIM 1112 Juncus bufonius L.;M oo capitatus ne oan 128670 LSU INCU M nzie; DePoe 6650 NLU Juncus dichotomus Ells D DTM 1272 Juncus diffusissimus Buckl.; DTM 1234 J Juncus marginatus Rostk.; DTM 1235 Juncus nodatus Coville; Raymond 1241 NLU Juncus polycephalus Michx.; RDT 629 Juncus repens Michx.; RDT 136859 Juncus scirpoides Lam.; RDT 33633 Juncus tenuis Willd.; DTM 1163 Juncus torreyi Coville RDT 128675 uncus validus Coville; DTM 126 Luzula bulbosa (Wood) Smyth & Smyth; Haynes 4568 LEMNACEAE Lemna aequinoctialis ew ; Overby 86 NLU Lemna minor L.; Hayne Lemna valdiviana Phil. ae Spirodela polyrrhiza (L.) Schleid.; Haynes 4155 Spirodela punctata (G.F.W.Mey.) C.H. Thompson; Haynes 422 Wolffia brasiliensis Weddell; Haynes 41 f Wolffia columbiana Karst.; Overby 227 Wolffiella gladiata (Hegelm.) Hegelm.; me 4153 LOUISIANA 1203 LILIACEAE Alliitm canandense| « TM 3058 ce nena L.; MM s.n (questionably natural- d). ze Erie scilloides (Raf) Cory; DTM 3015 Cooperia drummondii Herbert; a oi Erythronium ies Nutt; DTM 2 Habranthus tubispathus (L'Her.) ae MM 262 *Hemerocallis fulva (L.) L.; DTM 1723 (localized and persisting) Hymenocallis caroliniana (L.) Herbert; DTM 1843 Hymenocallis liriosome (Raf.) Shinners; DTM 853 Hypoxis hirsuta (L.) Coville; DTM 343 (includes H. rigida Chapman) *Narcissus pseudonarcissus L.; Barbour 559 (localized and persisting) Nothoscordum bivalve (L.) Britt; DTM 250 *Nothoscordum inodorum (Aiton) Nichols; DTM 1675 *Ornithogalum umbellatum L; DTM 3014 Polygonatum biflorum (Walt.) Ell; DTM 674 Schoenolirion wrightii Sherman; MM 7291 Stenanthium gramineum (Ker-Gawl.) Morong; DTM 811 Trillium recurvatum Beck; MM 144 Trillium pusillum Michx,; DTM 2185 Uvularia sessilifolia L.; DTM 2617 *Zephryanthes candida (Lindl.) Herbert; Barbour 467 nee nuttallii (Gray) S.Wats.; DIM 861 NAJADACEAE Najas guadalupensis (Spreng.) Magnus; DTM 1522 ORCHIDACEAE Corallorhiza wisteriana Conrad; DIM 11 pagcnata Tepes Nutt; ar 1947 i rid © ied og Ge cillata Raf. -DTM ea australis Lindl; DTM a Malaxis unifolia Michx.; RDT 43537 Platanthera ciliaris (L.) Lindl; RDT 42439 Platanthera clavellata (Micha Luer; MM 256 flava (L.) Lindl; Raymond 1483 NLU Platanthera lacera ae G.Don; DTM 1759 Spiranthes cernuus (L.) L.C.Rich.; MM 278 Spiranthes lacera (Raf.) Raf; Overby 262 NLU Spiranthes laciniata (Small) Ames; DTM 2013 Spiranthes odorata oe ee ie 5426 Spiranthes ovalis Lindl; Spiranthes praecox bevy - vo DTM 2313 LSU Spiranthes tuberosa Raf.; DT Spiranthes vernalis Engelm. & iy DIM 903 ae aria goer) NO aNens oe LSU R ; 1529NLU P latanthera => POACEAE Agrostis elliottiana J.A. Schultes; DTM 1632 *Agrostis gigantea Roth; DTM 12 Agrostis hyemalis (Walt.) B.S.P.; DTM 2029 Agrostis perennans (Walt.) Tuckerman; DTM 1416 Agrostis stolonifera L DTM 1309 *Aira caryophyllea L., DTM 3059 *Aira elegans Willd.ex Kunth; DTM 1659 Alopecurus carolinianus Walt.; DIM 2603 Andropogon gerardiiVitman; DTM 1458 Andropogon glomeratus (Walt.) B.S. i DTM 1498 A OPER ON ternarius Michx.,, DTM Andropogo ia virginicus L.; ‘DTM 1 ys el KCN, qustatae Boiss.; DIM a Trin. & Rupr; MM 28 Aristida dichotoma Michx.; DTM 2004 Aristida lanosa Muhl. ex Ell; MM 2928 Aristida ene eee fies DTM 1563 Aristida MM 2893 Aristida purpurascens ie DTM 1468 Arundinaria gigantea (Walt.) Muhl.; DTM 2193 *Arundo don 1569 *Avena fatua L.; DTM 1732 *Avena sativa. DIM 2615 Axonopus fissifolius (Raddi) Kuhlm.; DTM 1264 *Bothriochloa ischaemum (L.) Keng; RDT 68561 Bothriochloa languroides (DC.) Herter; DTM 1382 Bothriochloa longipaniculata (Gould) Allred & Gould; DTM 2079 se springfieldii (Gould) Parodi; Thieret 24481 res iza minor. DTM *Bromus catharticus ae DTM 2298 *Bromus diandrus Roth; RDT 65106 LSU Buchloe dactyloides (Nutt.) coer Snyder s.n.US Cenchrus spinifex Cav.; MM 2 Chasmanthium latifolium ies DTM 1735 Chasmanthium laxum (L.) Yates; DTM 1384 Chasmanthium ee florum (Poir.) Yates; DTM 2557 Chloris virgata Sw.;D 320 *Cynodon dact ctylon (L NY Pers.; one 4237 ad < aa iS) n cy = cS a tas) in aed = g mi ey Wn is *Dactylis glomerata L.,; DTM 1 *Dactyloctenum aegyptium He ine Overby 121 NLU Danthonia spicata (L.) Beauv.ex Roemer &J.A.Schultes; M 1655 Dichanthelium aciculare (Desv. ex Poir.) Gould & CA. Clark; DTM 1671 Dichanthelium acuminatum (Sw.) Gould & C.A. Clark; DTM 1210. Dichanthelium boscii (Poir.) Gould and C.A. Clark; DTM 1408 Dichanthelium commutatum (J.A. Schultes) Gould; DTM 1241 1204 Dichanthelium depauperatum (Muhl.) Gould; Lewis NLU Dichoanthelittm dic-hantamirim (I L.) Gould; DTM 1 182 m.) Gould; DTM 1239 sec sca linearifolium ee ex Nash) Gould; Allen Pe Sosa oligosanthes (J.A.Schultes) Gould; MM 2617 Dichanthelium ovale (Ell.) Gould & C.A.Clark; Lewis 3585 N Nichanthel lavifl nthes Schult.; MM 6491 a a ieveneln (Scribn. & Merr.) Gould; DTM ichanthelitim polya Dhan boasted item) Gould; DTM 1403 Dich Ell) Gould; DTM 1274 cane on (Retz) Koel: D M1414 Digitaria cognata (JA. aaa Pilger; DTM 1527 Digitaria filiformis (L.) Koel, DTM 1544 *Digitaria ischaemum (Schreb.) Schreb. ex Muhl; DTM *Digitaria violascens Link; DTM 137 a *Echinochloa colona (L.) Link; DTM 1 *Echinochloa crus-galli (L.) Beauv.; oe 277 SU Echino chloa muricata (Beauv.) Fern.; MM 6854 Echinochloa walteri i (P urs| 1) Heller; DTM 2553 *Eleusine indica (L (L.) Gaertn.; DTM 1596 Elymus nicus L.; DTM 2999 pri ees (All.) Vign. ex Janchen.; DTM 1420 *Eragrostis curvula (Schrad.) Nees; DTM 2312 Eragrostis hirsuta (Michx.) Nees; MM 2927 Eragrostis hypnoides (Lam.) B.S.P; DTM 1479 *Eragrostis japonica ea Trin; DTM 2706 Eragrostis lugens Nees; DTM 1643 *Eragrostis minor Host; ae 1377 *Eragrostis pilosa (L.) Beauv.; DTM 1510 Eragrostis refracta (Muhl.) Scribn.; DTM 1997 Eragrostis reptans (Michx.) Nees; DTM 1986 Eragrostis secundiflora J. Presl; DTM 1803 Eragrostis spectabilis (Pursh) Steud.; MM 2917 man trichodes (Nutt.) Wood; MM 2918 hloa ophiuroides (Monro) Hack,; DTM 1363 ie acuminata (J. Pres!) Kunth; DTM 1971 Eriochloa contracta A.S. Hitchc.; DTM 1481 Festuca paradoxa Desv., DTM 1277 te cisanana Fern. OM 2327 ichx.) B.S.P; DTM 1489 *Holcus lanatus L.; DIM 1346 Hordeum pusillum Nutt. DTM 1621 Leersia lenticularis Michx.; Haynes 5422 Leersia oryzoides (L.) Sw., DTM 1488 Leersia virginica Willd; DTM 1359 Leptochloa fusca (L.) Kunth; DTM 1325 Leptochloa panicea (Retz.) Ohwi; MM 6767 Leptochloa panicoides (J. Presl) A.S. Hitche. DIM 1 *Lolium arundinaceum (Schreb.) S.J. Derbyshire; 7 139] BRIT.ORG/SIDA 22(2) *Lolium perenne L.; DIM 1187 *Lolium temulentum L; DTM 1673 Luziola fli itans (Michx.) Terrell & Robins., DTM 1549 Melica mutica Walt.; DTM 1614 Muhlenbergia schreberi J.F.Gmel.; Nassella leucotricha (Trin. & Rupr.) an oa 3050 Oplismenus hirtellus (L.) Beauv.; ca 1305 Panicum anceps Michx,; DTM 1 Panicum brachyanthum net au 1998 Panicum capillare L.; DTM Panicum dichotomiflorum rie DIM 1386 Panicum flexile (Gattinger) Scribn.; Correll & Correll 10097 LSU Panicum eet Ell.; Haynes 5425 Panicum A.Schultes; RDT 140681 He sc.ex Nees; DIM 139 Panicum verrucosum Muhl.; ae 1529 Panicum uoene L; DTM atatum Poir.; a t m (L) L, rae fee Paspal listich L.; DTM 1970 Paspalum floridanum Michx.; DTM 1389 uncial ave lleNs DIM 1374 i (Fourn.) Nash; DTM ny Fluegg ge; DIM 1 Paspalum pli atriltim NAj hx.; DTM oo ss Pan j m rigid es ror palum TEPIOCOCUT YI Paspalum pubi florum Rupr. ex Fourn.; DTM 1586 Paspalum repen Bera. R 47 ean ara Bn . igs villei Steud.; DTM 1 phalaris gare iniana ae DIM ee *Phleum piaineh L.; RDT 65054 *Phyllosta a Carr ex A. Riv.& C. Riv, MM 7244 Pipt (L.) Parodi; DTM 2297 *Poa annua L.; DTM 1 Poa autumnalis the ex Ell; DTM 2186 Poa pratensis L.; Thieret 22666 LAF *Polypogon monspeliensis (L.) Desf; RDT 66455 LSU mean pe ranse ald (L.) Nutt. at 2959 haetitim aqvengq winii Spreng.;RDT 3727] ee. by evi arbe (Michx.) Pers. DTM 1469 h teum (Walt.) Pers.; DIM 1536 Sacci ek sia (L.) Nash; DTM 1431 Schedonnardus paniculatus (Nutt.) Trel.; DTM 2627 ee aoe scoparium (Michx.) Nash; DTM 1533 *Secale cereale L., DIM 2232 *Setaria faberi Herrm.; RDT 119026 Setaria parviflora (Poir.) Kerguelen; MM 2055 *Setaria pumila (Poir.) Roemer &J.A.Schultes; DTM 1573 *Setaria viridis (L.) Beauv.; RDT 45434 Sorghastrum elliottii (Mohr) Nash; MM 2931 Sorghastrum nutans (L.) Nash; DTM 2005 *Sorghum bicolor (L.) Moench; DTM 1411 *Sorghum halepense (L.) Pers.; Haynes 4228 Sphenopholis intermedia (Rydb.) Rydb.; RDT 88229 Sphenopholis nitida (Biehler) Scribn.; DTM 1626 — MACROBERTS AND MACROBERTS, VASCULAR FLORA OF CADDO PARISH, LOUISIANA 1205 Sphenopholis obtusata (Michx.) Scribn.; DTM 2632 Sporobolus clandestinus (Biehler) A.S. Hitche.; RDT 37044 Sporobolus compositus (Poir) Merr; DTM 1579 pee eee tus peana (Torr.) A.Gray; DIM 1578 I ;DT ! coerce us junceus Gesu) Kunth; DTM 1515 Sporobolus pyramidatus (Lam.) Hitche.; MM 7277 Sporobolus vaginiflorus (Torr.ex Gray) Wood; RDT 37396 LSU Steinchisma hians (Ell.) Nash; DTM 1 Stenotaphrum secundatum (Walt.) nc rs 308 Tridens ee (Ell.) A. Schultes; DTM Tridens flavus (L) A.S.Hitche.,MM 2915 oa Gee vai Hy chapmanii (Small) Chase 136 LSUS ve 2932 Trisetum interruptum Buckl.; Allen 3774 NLU *Triticum aestivum L.; DTM 2645 Urochloa platyphylla (Monro ex Wright) R.Webster; DTM 1378 *Urochloa ramosa (L.) Nguyen; DTM 13 Urochloa texana (Buckl.) R Webster; iene Burkett Co. s.n. LSU *Vulpia bromoides (L.) S.F.Gray; DTM 1228 ve a Leyines (L.) K.C.Gmel.; DTM 1650 (Walt.) Rydb,; DTM 2226 Vulpia a sciurea (Nutt). Henr.; MM 2537 Zizaniopsis miliacea (Michx.) Doell. & Aschers.; DTM 1158 PONTEDERIACEAE *£ichhornia crassipes (Mart.) Solms; DTM 240 Pontederi data .; DIM 88 POTAMOGETONACEA poor Nee ce Barbour 519 Potamoge 5L;DIM 1521 SMILACACEAE Smilax bona-nox L.; ale eae Smilax glauca Walt.; D Smilax laurifolia L: a 25 LSUS / smi ax rotund ifolia L.; ab 959 Smila eri Mii M T 155720 Smilax tamnoi des 2 ee 32815 SPARAGANIACEAE Sparganium americanum Nutt.; DTM 1830 gee is Pers; DTM 1758 Wee auiolia L.; DTM 1299 XYRIDACEAE Xyris caroliniana Walt.; Barbour 1099 Xyris jupicai L.C. Rich.; MM 263 DICOTYLEDONS ACANTHACEAE Dicliptera brachiata (Pursh) Spreng.; DTM 227 Justicia ovata (Walt.) Lindau; DT De *Ruellia brittoniana Leonard; RD Ruellia caroliniensis (J.F. ee ) ee a 64 Ruellia humilis Nutt; DTM 8 Ruellia pedunculata Torr. ex on DTM 447 ACERACEAE Acer barbatum Michx.; RDT 166956 Acer negundo L.; Haynes 4146 Acer rubrum L.; Haynes 4583 Acer saceniaenua ee Sais 2 LSU Ac AIZOACEAE esuvium it (Walt.) B.S.P; Thieret 26136 LAF Trianthema portulacastrum L.; DTM 174 AMARANTH “Aenanthea ice ies oo . Griseb.; DIM 84 Amaranthu i RDT 6 Amaranthus arenicola |.M. lens RDT 59959 Amaranthus aie idus LD Amaranthus palmeriS. ie oa 1964 pe spinosus L.; 924 s viridis La MM 6978 pa foeeee es ae DTM 1793 Froelichia gracilis (Hook.) Moq.; RDT 76696 ANACARDIACEAE Rhus aromatica Aiton; DTM 2181 Rhus copallinum L; DTM 1924 Rhus glabra L; RDT 33565 Toxicodendron radicans (L) Kuntze; Haynes 4140 ANNONACEAE Asimina parviflora (Michx.) Dunal; RDT 80926 Asimina triloba (L.) Dunal; Hardy 8685 APIACEAE Ammoselinum butleri (Enelm.ex S.Wats.) Coult.& Rose; RDT 88289 aaa incana Ruiz & Pavon; ca 7180 otundifolium L.; RDT res) ee tainturieri Hook, DTM Se cea (Pers.) ee ex Britt.& Wilson; Haynes 577 Cicuta maculata L,; eee ae ee *Conium maculatum L.; DTM SECU: eauaGense ve 2 “ 25312 cyn DTM 1190 *Daucus carota L. DIM an Daucus pusillus Michx.; MM 2653 Eryngium hookeri Walp.; DTM 143 Eryngium prostratum Nutt.ex DC; Haynes 4109 1206 Eryngium yuccifolium Michx.; DTM 142 *Falcaria vulgaris Bernh.; RDT 88299 aecayc ranunculoides Lf; sai 1097 yan Rous umbellata L.; “DIM Hydrocotyle verticillata Thu nb.; a 75 Limnoscadium pinnatum (DC.) Mathias & Constance; DIM 478 Oxypolis rigidior (L.) Raf.; ROT 7 Polytaenia nuttallii DC.; DTM 549 Prilimmnium capillaceum (Michx.) Raf.; DTM 1022 (Ell.) Raf; RDT 128660 im nuttallii (DC.) Britt.; Raymond 770 LSUS Sanicula canadensis L.; DTM 1040 — Dt ilire Prilingniy I Sanicula odorata (Raf.) K.M. Pryer & L.R. Philliope; RDT 33075 Sanicula smallii Bickn.; RDT 170095 Spermolepis divaricata (Walt.) Raf.ex Ser; DTM 2285 et a echinata (Nutt. ex DC) Heller; Allen 7999 Siar een (Nutt.ex DC.) Mathias & Constance; Taenidia integerrima (L.) Drude; Cocks s.n. NO *Torilis a is (Huds.) Link; DTM 29 *Torilis noueya ey DTM 1033 *Tre) rpu je Nutt.ex DC, DTM 464 Zizia aurea (L.) W.D i Koch; Raymond 732 LSUS APOCYNACEAE eee nae he he 838 LMM recs difforme i: ) " a DIM 454 *VINC ne minor Ls RDT ns AQUIFOLIACEAE llex ambigua (Michx.) Torr; RDT 140651 LSU llex decidua Walt.; Haynes 4093 Ilex opaca Aiton; Hardy 8616 Ilex vomitoria Aiton; DTM 1151 ARALIACEAE Aralia spinosa L.; DTM 1154 ARISTOLOCHIACEAE Aristolochia reticulata Jacq.; RDT cue Aristolochia serpentaria L; RDT 119103 Aristolochia tomentosa Sims; RDT 36969 ASCLEPIADACEAE Asclepias amplexicaulis Sm.; DTM 2294 Asclepias perennis Walt., DTM 523 fog ne tuberosal.; DTM a riegata Le -DIM8 Asclepias Vu Asclepias verticillata L.; DTM ve siete viridifiora Raf; DTM 556 idis Walt.; DIM 62 Asclepias Vil Cynanchum laeve (Michx.) Pers.; Hardy 6382 Matelea cynanchoides (Engelm.) Woods.; MM 2613 BRIT.ORG/SIDA 22(2 Matelea decipiens (Alexander) Woods.; RDT 51975 Matelea gonocarpos (Walt.) Shinners; RDT 36994 ASTERACEA ACNE mill flu i. seal 329 m.) R.K. Jansen; DTM 442 Ageratina ‘alti issima (L ) ce & E.Robins.; DIM 693 Ambrosia artemisiifolia L., DTM 9 Ambrosia psilostachya re si in Ambrosia trifida L DTM nears i Hayes nthemis cotula L.; D nen. cali a )Trel.; DTM 1771 be eal po neum Raf., DTM 407 lia L.; Hardy 6871 ” Berlandiera betonicifolia (Hook.) Small; DTM 1074 Berlandiera pumila (Michx.) Nutt. DTM 997 Bidens aristosa (Michx.) Britt., DTM 1993 Bidens bipinnata L.; Haynes 5419 Bidens discoidea (Torr. & ie Britt; DTM 1 Bidens frondosa L.; DIM Bidens laevis (L.) B.S.P; ae os Boltonia di Ell; DIM 718 *Calyptocarpus vialis Less.; DTM 1860 *Carduus nutans L.; DTM 460 Chrysopsis pilosa — oie 2720 #Cj horinm intyl ni 223 Cirsium alti es DIM 5 Cirsium eee (Walt.) Fern. & oe DTM 840 Cirsium engelimanill ae cen LSU ,DIM5 Cirsium Conoclinium coelesti (L.) DCD 639 *Conyza bonariensis (L.) Cronq.; a 321 Conyza canadensis (L.) Cronq.; DTM 589 Coreopsis grandiflora Hogg ex Sie DIM 434 Coreopsis intermedia Sherff; MM 2605 Coreopsis lanceolata L.; DTM 2 a Coreopsis tinctoria Nutt.; Raymond 768 LSUS *Crepis pulchra L.; DIM 842 Croptilon divaricatum (Nutt.) Raf.; DTM 592 Doellingeria sericocarpoides Small; DTM 1947 copis amplexicaulis (Vahl) Cass.; DTM 60 59 LAF ) Dra Echinacea pallida (Nutt.) Nutt; DTM 491 (includes £. angustifolia DC. and E. sanguinea Nutt.) Echinacea purpurea (L.) Moench; RDT 136864 Eclipta prostrata (L.) L., DTM 247 Elephantopus carolinianus Raeusch.; DTM 8 Elephantopus nudatus Gray; RDT 155792 diate oe es tosus L.; DTM 241 an Raf. ex DC; DTM 1524 DTM 1988 oe ae ie DIM 45 Erigeron pulchellus Michx.; Raymond 750 LSUS Erigeron strigosus Muhl.ex Willd, DTM 52 oe tenuis ae & A.Gray; DTM 360 (Lam.) Small; DTM 2007 Eriger Nannie (l .) Pe LOUISIANA 1207 Eupatorium compositifolium ea ;DTM 2052 Seana ad L;DTM indifolium L a ynoatoris 1m Eupatorium semiserratum DC.; ay ah (includes E. glaucecsens Ell. Eupatorium serotinum Michx,; DTM 172 Eurybia hemispherica (Alexander) Nesom; DTM 705 =e leptocephala (Torr. & A. Gray) Greene; DTM 7 i een & A. Gray) A. Gray; al a 1 LSU *Facelis retusa (Lam.) Schultz-Bip.; DIM 8 Fleischmannia incarnata (Walt.) King & ie aie RDT 33060 Gaillardia aestivalis (Walt.) H. ines DTM 1800 Gaillardia pulchella Foug.,, DTM Gamochaeta te (Urb.) eee DTM 400 Gamochaeta a a Nesom; Strout 112 LSUS Gamochaeta pers (Willd.) Cabrera; DTM 1751 Gamochaeta purpurea (L.) Cabrera; DTM 373 Helenium amarum (Raf.) H. Rock; DTM 24 Helenium flexuosum Raf.; DTM 27 BISNanENtS alc L;DIM 178 arbo a LE Helianthus eee L; DTM 1894 pea ae ee DTM 1756 a elianthus mollis Lam.; -RDT 362 lianthus occidentalis Riddell; oe. s.n.NO Helianthus petiolaris Nutt.; Se s.n.NO Helianthus strumosus Heterotheca subauians oe ) a & Rusby; MM 2777 Hieracium gronoviiL.; DTM 390 Hymenopappas a He DC.; Barbour 1193 Hymenopappas scabiosaeus L'Her,; DTM 17 *Hypochaeris glabra L.; DIM 2830 lva angustifolia Nutt. ex DC; DTM 2077 lva annua L.; DTM 2025 Krigia caespitosa (Raf.) Chambers; DTM 38 Krigia dandelion (L.) Nutt; Raymond 731 LSUS Krigia occidentalis Nutt.; DT Krigia virginica (L.) Willd; ae - 3 Lactuca eqnadens L;DTM 1 Lactuca floridana (L.) Gaertn.; ar 651 Lactuca Seine oe hia RDT 36155 */ actuca serriola L.; DIM 4 *Leucanthemum ae DTM 1 naturalized) Liatris aspera Michx.; DTM Liatris elegans (Walt.) Ante ane 204 Liatris pycnostachya Michx.; DTM 517 Liatris squarrosa (L.) Michx,, DTM 542 Hale sala is shinners; RDT 37304 ex DC.; DTM 351 Mba a scandens ( . 2 ae DTM 209 li rr. & A. Gray) Small; DTM 1 Olig igoneuron rigidum (| ic Small; RDT 93593 ye e 19 (questionably Packera gla bella (Poir. ) G, Jeffrey; DIM 41 Packera ahora (Muhl.ex Willd.) W.A.Weber & A. Love; 2 *Parthenium hysterophorus L.; DTM 1999 Parthenium intearifolium Ee *-DIM 13 Pityopsis ee (Michx.) Nutt.; MM 906 eacampho cE ;DIM 698 Pluchea fecal soe MM 284 Pluchea odorata (L.) Cass.; MM 277 Pseudognaphalium helleri (Britt.) A.Anderb.; RDT 41805 Pseudognaphalium obtusifolium (L.) Hilliard & Burtt; 20738 = a Pyrrhopappus carolinianus (Walt.) DC.; DTM 552 Pyrrhopappus pauciflorus (D. Don) DC; DTM 499 atibida pinnata (Vent.) Barnh.; DTM 94 Rudbeckia grandiflora (D.Don) J.F. Gmel. ex DC.; DTM Rudbeckia hirta L; DTM 1 Rudbeckia maxima Nutt,; ow 95 *eneck vulgaris L.; MM 5 aad um He ifolium Niche DTM 2326 inia 20 Inhitwym tum L.; *Sily bum marianum (L ) ce RDT 345 small anthus uvedalia (L.) Mackenzie ex ad DTM 1088 Solidago altissima L.; DIM 627 Solidago auriculata Shuttlew. ex Blake; DTM 1916 aie udoviciana \Gley) ee MM 2889 odora Aiton Soop eto ane Aiton: oe 96 1 Solid nes 4081 20 bse cages B Mil iime 686 hl ex Willd.; DTM 670 Soli iva eos Ruiz & Pavon; DTM 1014 *Sonchus asper (L.) Hill; DIM 424 *Sonchus oleraceus L.; DTM 896 si aes trichum divaricatum (Nutt.) Nesom; Haynes 4261 > a — richum drummondii (Lindl.) Nesom; DTM ee a trichum dumosum (L.) Nesom; MM 6957 Symphyotrichum laeve (L.)A.& D. Love; DTM 744 Symphyotrichum lanceolatum (Willd.) Nesom; DTM 2586 Symphyotrichum lateriflorum (L) A.& D.Love; DTM 2057 Sp euenum cee (Wieg. ) Neon RDT 69310 Riddell DIM ' b NANA BO00 570 Symphyotrichum patens (Aiton) N Symphyotrichum pilosum (Willd.) econ DTM 2569 Symphyotrichum praealtum (Poir.) Nesom; DTM 700 Symphyotrichum pratense (Raf.) Nesom; DTM 2080 Symphyotrichum racemosum (Ell. Symphyotrichum subulatum (Michx.) — Hardy 5297 NI -MM QOQ0Q a *Taraxacum officinale G.H.Weber ex Wiggers; DTM 283 1208 pa ero Cone (Torr. & A. Gray) A.Gray ex Hall; M Plea 4 a lium (Hook.) A. Gray; ony 88997 LSU Verbesina helianthoides Michx.; DTM 4 Verbesi Hi virginica L; ‘ Barbo sts Vernonia baldwinii Torr; MM 6 Vernonia missurica Raf; amet 773 LSUS Vernonia texana (Gray) Small; Ane 126 anthium strumarium L., DTM 1095 ie ica (L.) DC; DTM 3012 BALSAM ACEAE ae alee Meerb.; DTM 1094 BERBERIDACEAE *Nandina domestica Thunb; MM 6465 (questionably naturalized) Podophyllum peltatum L., DTM 47 BETULACEAE Alnus serrulata (Aiton) Willd.; DTM 2228 Betula nigra L.; Hard Carpinus carolin iana Walt; Haynes 5316 Ostrya virginiana (P. Mill) K. Koch; Haynes 4569 BIGNONIACEAE Bignonia capreolata L.; DIM 803 Campsis radicans (L.) Seem. ex Bureau; DIM 137 Catalpa bignonioides Walt.; Barbour 82 Catalpa speciosa (Warder) Warder ex Engelm.; Frank 134 LSUS BORAGINACEAE *Bugl Tal nsis s (L.) .M.Johnston; DTM 1071 baa 132 LAF 7 Holint He fotopium indicum i DIMI He jotropium pro rmbens ae a 624 Lith nerm! \A/a| citmospermum Cal nse (Walt. ex JF. Gmel.) M acM.:; MM 2534 Myosotis macrosperma Engelm.; MM 6155 (includes osotis verna Nutt.) Onosmodium bejariense DC. ex A.DC.; Cocks s.n. NO BRASSICACEAE *Arabidopsis thaliana (L.) Heynh.; RDT 70506 *Camelina microcarpa DC.; RDT 65101 LSU *Capsella bursa-pastoris (L.) Medik.; DTM 256 Cardamine bulbosa (Schreb. ex Muhl.) B.S.P; MM 6205 x.) Sw; MM 143 Cardamine COMCALEN GLA (Michx *Cardamine hirsuta L; DTM 261 Cardamine parviflora L.; Barbour 1133 Cardamine pensylvanica Muhl.ex Willd; DTM 1 *Coronopus didymus (L.) Sm. DIM 852 *Descurainia sophia (L.) Webb. ex Prantl; RDT 65110 ie brachycarpa Nutt. ex Torr.& A. Gray; DTM 2587 ifolia Nutt. ex Torr. & A. Gray; DTM 943 Lepidium den n Schrad.; RDT 65049 LSU Lepidium VIFGINIC um Ls ;Raymond oe LSUS *Rapistru mM rliqos mit L.) All; DIM ¥R ; cr ry PP ACU ie Hayek; DTM 2204 qatnatrictym AM-aqguauiculn BRIT.ORG/SIDA 22(2) Rorippa palustris (L.) Bess.; DTM 975 Rorippa sessiliflora (Nutt.) AS. Hitch.; Barbour 970 36 Sibara virginica (L eens as arvensis iL, DTM 27 L,; ieee *S) ffici (L.) Scop; RDT 83089 oe ae ae Hook.; MM 2656 *Thlaspi arvense L.; Barbour 1003 BUDDLEJACEAE Polynremitm nr / = mbens L.;DTM 154 CABOMBACEAE Cabomba caroliniana Gray; DTM 1721 CACTACEAE Opuntia humitu a (Ra f.) R Raf; MM 239 CALLITRICHACEAE Callitriche Metron yi Pursh; si 4577 Callitriche COhPOOU 1S Raf. 4 CAMPANULACEAE Lobelia appendiculata A. ae : TM 732 Lobelia cardinalis L.; D Lobelia puberula Michx.; ae a 5 Triodanis biflora (Ruiz & Pavon) rae — 387 [riodanis perfoliata (L.) Nieuwl,; DTM 9 CAPPARACEAE leome hassleriana Chod,; Haynes 5938 Polen ia dodecandra (L.) DC; RDT 64990 APRIFOLIACEAE *Lonicera japonica Thunb.; DTM 120 Lonicera sempervirens L.; DTM 402 Sambucus nigra L.; DTM 1284 Symphoriocarpus orbiculatus Moench; Raymond 865 [riosteum angustifolium L.; DTM 2287 Viburnum nudum L.; DTM 2286 aay enPHE NCH jum; RDT 25303 Raf.; MM 2823 CRORE IEENGE RE naria benthamii Fenzl ex Torr.& A. Gray; RDT 65108 LSU Arenaria lanuginosa (Michx.) oe DTM 1760 *Arenaria sey llifolia L.; MM 261 Cerastium brachypodum (Engelm.ex A. Gray) B.L. Rob- ins, DTM 2136 *Cerastium fontanum Baumg.; Strout 122 LSUS *Cerastium salad Thuill.; DTM 937 ackenzie sah 7284 ee squarrosa suite Reid 4 Minuartia drummondii aco McNeill; MM 7308 Minuartia muscorum (Fassett) Rabeler; MM 7309 Paronychia drummondii Tort. & A.Gray; DIM 2448 Paronychia fastigiata (Raf.) Fern,; Correll & Correll 10096 LSU Sagina decumbens (Ell.) Torr.& A. Gray; DTM 2723 LAUUU PARISH, LOUISIANA 1209 *Saponaria officinalis L, MM 237 *Scleranthus annuus L.; a 82790 LSU Silene antirrhina L. ae *Silene gallica L; DTM 3 *Silene Hociior Re a an Silene stellata (L.) Aiton f; DTM 2302 Stellaria media (L.) Vill; Barbour 33 *Vaccaria hispanica (P. Mill.) Rauschert; RDT 65131 LSU CELASTRACEAE vonymus americana L.; Hardy 8615 CERATOPHYLLACEAE C lum d um L.; DTM 1554 CHENOPODIACEAE iets ee (L.) AJ. Scott; RDT 65018 Chenopodium berlandieri Moq.; RDT 66452 *Che a Chenopodiu db.; RDT 65055 Se um si sinnles (Torr.) Raf; DTM 935 Chenopodium standleyanum Aellen; Overby 94 NLU Cycloloma atriplicifolium (Spreng.) Coult.,, Lewis 3663 NLU Monolepis nuttalliana (JA. alana Greene; DTM 2196 *Salsola tragus L RDT 65028 CISTACEAE Helianthemum carolinianum (Walt.) Michx.; DTM 161 Helianthemum georgianum Chapman; MM 2745 Helianthemum rosmarinifolium Pursh; DTM 1744 Lechea mucronata Raf.; MM 2715 Lechea tenuifolia Michx.; DTM 1785 CLUSIACEAE p reae (L.) Crantz; DTM 586 Hypericum densiflorum Pursh; RDT 65084 LSU 1ondii (Grev. & Hook.) Torr. & A. Gray; Hypericum drumn DIM 149 1 frondosum Michx.; DTM 1257 Byer icum pvaenuane ies: .) B.S.P; Allen 8539 LAF Hypericum gymnanthum Engelm. & A. Gray; RDT 165816 Hy peri um hypericoides (L.) Crantz; DTM 1 Ay peictlen mutilum L.; ae 5317 Hypericum nudiflorum Michx.ex Willd.; Raymond 1291 LSUS a prolificum L. RDT 42445 LSU Hypericum pseudomaculatum Bush; DTM 453 me cum mpunctotm Lam.; DTM 1066 Triadenum walteri iJ. G.Gmel. ) Gleason; RDT 36265 oct iadeladeiacaien *Col ulus arvensis L.; DTM 163 Di cHonaia caroliniensis eee DTM 308 Evolvulus sericeus Sw.; MM 3355 LSU !pomoea cordatotriloba Dennst.; DTM 696 Ipomoea hederacea Jacq.; DTM 1920 pomoea lacunosa L.; DTM 667 pomoea panderata (L.) G.F.W.Mey.; DTM 1805 pomoea purpurea (L.) Roth; MM 125 pomees auamgel it L. RDT 42043 */pomoea wrightii Gray; DTM 2693 Jacquemontia tamnifolia (L.) Griseb.,; DTM 640 Stylisma humistrata (Walt.) Chapman; RDT 92409 Stylisma pickeringii (Torr. ex M.A. Curtis) A. Gray; MM 2677 ee ornus drummondii C.A. Mey.; Hardy 8607 zee florida L; Graham 571 LSUS Cornus foemina P. Mill. Le ee Nyssa aquatica L; RDT 3 Nyssa eee ca Marsh.; ae 638 (includes WN. biflora T. Walte rigbenaminsane tica (L.) Schoenl.; RDT 34511 Penthorurn sedoides _ ae 683 LSUS RDT 83128 NLU CUCURBITACEAE Cayaponia quinqueloba (Raf) Shinners;RDT 25353 LSU Melothria pendula L., DTM 233 ee acene Cu Juss.ex Choisy; DTM 1934 Cuscuta gronovi ii Willd. ex. J.A. pi MM 696 Cuscuta indecora Choisy; MM Cuscuta pentagona Engelm.; ae 1788 DROSERACEAE Drosera brevifolia Pursh; DTM 832 EBENACEAE Diospyros virginiana ke DTM ] 176 ELAEAGNACEAE *Elaeagnus pungens Thunb.; DTM 2054 (questionably naturalized aa ERICACEAE Lyonia mariana (L.) D.Don; RDT 33683 Rhododendron canescens (Michx.) Sweet; RDT 33665 Rhododendron oblongifolium (Small) Mallais; RDT 88707 d| osum (L.) Torr; DTM 2278 Vaccinium ae Marsh.; DTM 1637 Vaccinium corymbosum L.; Haynes 4628 (includes V. elliottii Chapman, V.fus n,and V.virgatum Aiton Vaccinium stamineum L.,; DTM 1155 EUPHORBIACEAE fi at gracilens Gray; DTM calypha monococca jee aA Gray) L. Mill & Gan- dhi; ROT Acalypha es Riddell; DTM 1392 Acalypha rhomboidea Raf.; DTM 704 Acalypha setosa A. Rich.; DTM 1092 atum Ait 1210 Acalypha virginica L., DTM 1371 Chamaesyce cordifolia (Ell.) Small; DTM 1940 Chamaesyce humistrata (Engelm.) Small; RDT 36205 Chamaesyce maculata (L.) Small; DTM 1821 LSU Chamaesyce nutans (Lag.) Small; Haynes 4223 Chamaesyce prostrata (Aiton) Small; DTM 1830 Chamaesyce serpens (Kunth) Small; RDT 32893 Cnidoscolus texanus (Muell.-Arg.) Small; Haynes 5414 Croton argyranthemus Michx.; RDT 81395 LSU Croton capitatus Michx.; DTM 11 Croton glandulosus L.; DTM 188 Croton michauxii G.L.Webster; MM 2772 Croton monanthogynus Michx,; DTM 170. Croton willdenowii G.L Webster; DTM 532 aati corollata L.; DIM 71 us dentata Michx.; MM 31 72 Euphorbia tetraspora Engelm.; Moore s.n. LSU A abt us carolini iensis Walt., MM 698 1 aria L.; RDT 166975 *Rick inus communis L.; Thieret 21037 LAF (questionably naturalized) sia lingia sylvatica Garden ex L.; DTM 1 ragia eetonisioile Nutt; RDT 45419 ichx hee TGUGLU INI 4" tears urticifol lia Michx.;D 45 *Triadica sebiferum (L.) a a 1757 FABACEAE Acacia angustissima (P. Mill.) Kuntze; ta 74 *Albizia julibrissin Durazz., DTM 1 pibllene fruticosa L.; MM 6793 Amorpha paniculata Torr.& A. Gray; MM 255 Amphicarpaea bracteata fe “ ae DTM 650 Apios americana Medik. Astragalus canadensis ‘ ay an Astragalus distortus Torr.& A. Gray; DTM 260 => is soxmaniorum Lundell; DTM 1806 Baptisia alba (L.) Vent.; DTM 867 Baptisia bracteata Muhl. ex Ell; MM 176 Baptisia nuttalliana Small; DTM ed Baptisia sphaerocarpa Nutt.; MM 1 Centrosema virginianum (L.) pent a 169 Cercis canadensis L.; Barbour silica fasciculata ae Greene; DTM 86 maecrista nictitans (L.) on DTM 215 arene mariana. DTM 1 Crotalaria sagittalis L.; see a talaria fee ed Roth; ee z ex Willd; DTM ae phleoides a &A, ues eae MM 2657 Dalea purpurea Vent; DTM 534 Dalea villosa (Nutt.) Fie DIM 1797 LAF AAD KAT BRIT.ORG/SIDA 22(2) Desmanthus illinoensis (Michx.) MacM.ex B.L. Robins. & Fern.; 1969 (L.) DC; DTM 1973 (Muhl. ex Willd.) DC.; DTM 211 Desmodium glabellum (Michx.) DC.; RDT 37293 pons ium hes old ee DC.; RDT 136869 odium nudiflorum ( C.,;, Overby 292 NLU se Schub.; Raymond 2069 [) di Desmoditim ni itt hij LSUS oditnm pan iculatum (L.) DC; Raymond 782 LSUS Desmodium sessloliam (Torr) Torr.&A. se DIM 1084 Desmodium viridiflorum (L.) DC.; RDT 3 Dioclea multiflora (Torr. & A. Gray) C. a 3729] Sna herbacea L., DTM 439 Galactia volubilis (L.) Britt; DTM 591 Gleditsia ia. aquati ica Marsh,; Reid 4636 NLU leditsia triacanthos L.; Hardy 8612 Glottidium vesicarium Jacq.) Harper; Haynes 5328 eu striata (Thunb.) Schindl.; DTM 1505 L.;DITM 37 bans pusillus Ell; DTM 795 Lathyrus venosus Muhl. ex Willd.; DTM 2199 *Lespedeza cuneata (Dum.-Cours.) G. Don; RDT 36216 Lespedeza hirta (L.) Hornem.; DTM 678 Lespedeza procumbens Michx.; DTM 733 Lespedeza repens (L.) W. Bart.; re Lespedeza stuevei Nutt.; MM 2 eae a violacea (L.) Pers.; aa i le inica (L.) Britt., DTM ae uni oliolatis (Hook.) Benth.; ae 66462 LSU Be Inus texensis ae Newsom s.n. LSU (L.) Huds.; ae 362 ona) cago ating L; DTM 1 (L.) L. RDT oe “Media polymorpha L.; oo 530 Medicago sativa L.; DTM 1 *Melilotus albus Medik.; ane *Melilotus indicus (L.) All, DTM “ *Melilotus officinalis (L.) Lam. DTM 410 Mimosa nuttallii (DC.) B.L. Turner: “it 164 ee Mimosa Strigillosa Torr.& A. Gray; DIM 19 Neptunia lutea ese atek ie ;DTM 50 Neptunia pubescen h.; Lasseigne 1002 LAF Orbexilum pene (P. Mill.) Rydb,; DTM 385 Orbexilum simplex (Nutt. ex Torr.& A. Gray) Rydb,; DTM 1119 Parkinsonia aculeata L.; DTM 756 Pediomelum digitatum (Nutt. ex Torr. & A. Gray) Isely; MM 235 Pediomelum hypogaeum (Nutt.ex Torr.& A.Gray) Rydb,; M M 2549 Prosopis glandulosa Torr; Cocks s.n. NO *Pueraria montana (Lour.) Merr.; Hardy 8896 Rhynchosia latifolia Nutt. ex Torr.& A. Gray; DTM 520 Rhynchosia minima (L.) DC; MM 6991 Rhynchosia reniformis DC.; DTM 129 LAUUY PARISH, LOUISIANA 1211 Rhynchosia tomen t (L.) Hook. & Arn.; RDT 36931 ene isola i MM ia Robinia pseudoacacia L.; Haynes 5699 Senna iene (L.) Link; DTM 196 Senna obtusifolia (L.) Irwin & Barneby; DTM 1575 Sesbania drummondii (Rydb.) Cory; DTM 202 Sesbania herbacea (P. Mill.) McVaugh; DTM 1476 *Sesbania punicea (Cav.) Benth.; DTM 1215 Sophora affinis Torr.& A. Gray; Cocks s.n. NO Strophostyles helvula (L.) Ell; DIM 564 Strophostyles leiosperma (Torr.& A. Gray) Piper; DTM 544 Strophostyles umbellata (Muhl.ex Willd.) Britt; Raymond SUS Stylosanthes biflora (L.) B.S.P; MM 2713 ee brychoides Nutt.; DIM 1904 Te phrosia Migtolaaa (lis) Pers.; ‘DTM 46 rifolium a Seek 80865 *Trifolium campestre Schreb.; DTM 151 Trifolium carolinianum Michx.; DTM 2639 *Trifolium dubium eee a 289 ¥Trifolitsm Incarnatiim L.;: DIM Trifolium reflexum L.; in 2638 *Trifolium repens L.; DTM 136 *Trifoliun upinatum L., DTM 313 * Trifolium ee Savi; au 125 Vicia caroliniana Walt.; DTM 2137 Vicia ludoviciana bees a 1160 *Vicia lutea L.; RDT 1 Vicia anne FG. ies MM 152 *Vicia sativa L.; 5 *Vicia ee (li ) Schreb.; RDT 88360 LSU *VICIA villosa Roth; 394 Wisteria frutescens (L.) Poir; Haynes 5356 *Wisteria sinensis (Sims) DC.; DTM 1131 Zornia bracteata J.F.Gmel.; MM 2844 — FAGACEAE Castanea pumila (L.) P. Mill; DTM 1202 Fagus grandifolia Ehrh.; ne 208 LTU Quercus alba L.; Barbour 56 Sie qusagsaie ae MM 2872 Michx.; Haynes 4094 eee hemispherica Bart. ex Willd.; RDT 32931 aU incana Bat ne: MM 581 ee laurifolia Michx.; RDT 37236 Quercus lyrata Walt; RDT 42015 uercus i iaiaicelle ices: se nae LAF Qu ex Small; RDT 167037 Quercus manleaaee Mu enh. Foymone 1321 LSUS Quercus michauxii Nutt.; DTM Quercus Se eae ot 36212 Quercus nigra L.;M Tie Quercus phellos L.; eee 1292 LSUS Quercus rubra L.; Middlebrooks 206 NLU Quercus shumardii Buckl.; Hardy 8609 Quercus stellata Wangenh.; MM 5016 (includes Quercus imilis Ashe) Quercus velutina Lam. Raymond 145 NLU Quercus virginiana P.Mill.; Rowe 105 LSUS FUMARIACEAE Corydalis flavula (Raf.) DC.; RDT 88228 Corydalis micrantha (Engelm. ex A. Gray) A. Gray; MM L 00 ‘ | \A/\AS GENTIANACEAE Centauri a W.Wright ex Piper; MM 6435. Sabatia angularis (L.) Pursh; DTM 145 Sabati pestris Nutt; DTM 72 GERANIACEAE anium carolinianum L.; DTM 301 mL;DTM 990 GROSSULARIACEAE tea virginica L.; DTM 1196 Ribes curvatum Small; DTM 2129 HALORAGACEAE *Myriophyllum aquaticum (Vell.) Verdc.; Barbour 454 Myriophyllum heterophyllum Michx.; a 908 *Myriophyllum spicatum L.; Barbou Proserpinaca palustris L.; MM 205 HAMAMELIDACEAE Hamamelis virginiana Ly ; Manning 1 176 LTU Liquidambar he L;DTM 1071 HIPPOCASTANACEA Aesculus pavia L.; Baas 446 LSUS HYDROPHYLLACEAE Hydrolea ovata Nutt. ex ae DIM 12 Hydrolea uniflora Raf.; D Phacelia glabra Nutt.; a - Phacelia strictiflora (Engelm.& A.Gray) A.Gray; MM 2544 JUGLANDACEAE ee lba(L -) Nutt. ex Ell; Raymond ee mi quatica (Michx. f) Nutt; Barbour cane cor formis (Wangenh.) K. Koch; ee 1302 * raninnm diccect a. ia (P. Mill.) Sweet; RDT 170092 Carya illinoiensis (Wangenh,) K. Koch; MM 6470 ee f) ail 503 LSUS ane ovata (P. Mill Koch; MM Carya texana Biel: “ rdy 8611 Juglans nigra L.; Raymond 1679 NLU LAMIACEAE *Ajuga reptans L.; MM 202 *Glechoma hederacea L; RDT 43545 one Al eaee cain DIM 873 L;DIM 53 *Lamium purpureurn Ls DIM 2095 Muhl. ex W. Bart; DTM 1 7 1212 h; DTM 682 Lyc Opus ViIrginicus Lee -RDT soi *Mentha spicata L.; DTM 1 Monarda citriodora Cerv. ex e RDT 45372 Monarda clinopodioides Gray; DTM 1753 Monarda fistulosa L.; DTM 93 Monarda punctataL.; DTM 15 *Perilla frutescens (L.) Britt.;, MM 270 Physostegia angustifolia Fern.; 2 - Physostegia digitalis Small; DTM Ph ial intermedia (Nutt.) in & A.Gray; DTM a is vulgaris L.; DTM 3 Pycnanthemum albescens aa & A. Gray; DTM 194 Enea RIC n (Michx.) Pers.; iat 6/061 h M5 Pycnanthemur PLCRIUTIUilUreil Salvia azurea Which ex Lam.; DTM 214 Salvia lyrata L.; DTM 35 Seutellaria cardiophylla ceae : A.Gray; MM 2723 mained rea tiel ial; DIM4 Hill; DTM . san laria ie es ar a 323 Bet floridana se ie ex «Benth MM 214 sae sila we DTM L.;DTM i L.; DTM 653 Trick t Ait LAURACEAE Lindera kl eee Hardy 8617 Sassafras albidum (Nutt.) Nees; DTM 1068 LENTIBULARIACEAE Uticularia gibba .; Barbo eae Uticularia inflata Walt; DTM 2 Uticularia macrorhiza sees len 464 LINACEAE Linum medium (Planch.) Britt.; DTM 522 ‘inum striatum Walt.; RDT 170116 LOGANIACEAE Gelsernium sempervirens (L.) Aiton f; DTM 270 j—~< Mitreola petiolata (J.F. mel.) Torr. & A. Gray; DTM 201 Spigelia marilandica (L.) L.;MM 210 LY THRACEAE mmannia auriculata Willd.; ra . 7 Ammannia coccinea Rottb.; M orca te us (L.) Ell; sis on LSUS Lythrum alatum Pursh ; MM 6783 Rotalo ra ior (L.) | Koeht 1e; f laynes 5336 MAGNOLIACEAE Magnolia grandiflora L.; Burns 238 LSUS Magnolia virginiana L.; Barbour 1191 MALVACEAE *Abutilon theophrasti Medik.; RDT 119031 BRIT.ORG/SIDA 22(2) Callirhoe alcaeaides (Michx.) A. ay DTM 1049 a rhoe digitata Nutt; RDT 8837 allirhoe involucrata (Torr. & an A. Gray; Newsom Callirhoe papaver (Cav.) A. a MM 257 ;DIM9 Hibiscus laevis Al Hibiscus moscheutos L.; oe 168 *Hibiscus syriacus L.; Gamblin s.n. LTU (localized and ersisting) Malvaviscus arboreus Dill.ex Cav.; MM 123 Modiola caroliniana (L.) G. Don; DTM 379 Sida rhombifolia L.; DTM 186 Sida spinosa L.; DTM 521 MELASTOMATACEAE Rhexia mariana L.; DTM 243 MELIACEAE *Melia azedarach L.; MM 6183 sates a balsas (L.) DC; Barbour 473 MOT ene ene Mollugo verticillata L.; DTM 1373 MONOTROPACEA Monotropa ica L.; Thieret 24524 LAF Monotropa uniflora L.; MM 281 MORACEAE *Broussonetia papyrifera (L.) UHer.ex Vent. DTM 1145 Maclura ae (Raf.) aoe Barbour 570 *Morus Morus a L.; Haynes iD MYRICACEAE Morella caroliniensis (P. Mill) Small; RDT 33652 Morella cerifera (L.) Small: Haynes 4507 NELUMBON Nelumbo lutea ie Barbour 467 NYCTAGINACEAE Boerhaavia rite L. DTM 2573 ete ia abe L.; RDT 65066 LSU da (Walt.) Heimerl; MM 2610 NYMPHAEACEAE Nuphar advena (Aiton) W.T. Aiton; MM 208 Nymphaea odorata Aiton; DTM 9 OLEACEAE Chionanthus virginicus L.; Raymond 1307 LSUS Forestiera acuminata (Michx.) Poir; Barbour 1004 Forestiera clan ee Poir.; Raymond s.n.LSUS axinus analjH 613 Fraxinus a P. vey M252 Fraxinus pennsylvanica Marsh.; MM 6682 *Ligustrum japonicum Thunb.; DTM 1173 *Ligustrum lucidum Aiton f, RDT 32907 *Ligustrum sinense Lour; DIM 1171 a CADDO PARISH, LOUISIANA 1213 ONAGRACEAE Gaura longiflora Spach; DTM 221 Gaura mollis James; DTM 104 Gaura sinuata Nutt. ex Ser.; DTM 1790 Ludwigia alternifolia L, DTM 115 Ludwigia decurrens Walt., DTM 617 non a aa Walt.; DTM 1400 Ludwigia hi .; DTM 1301 Ludwigia leptocarpa ses DTM 226 Ludwigia linearis Walt; Ludwigia palustris (L.) i oe 4090 Ludwigia peploides (Ku oy hee Bes 4087 Ludwigia repens J.R. For Oenothera biennis L.; ie Oenothera heterophylla een aia 31220 LAF Oenothera laciniata Hill; DTM 7 Oenothera linifolia Nutt, DTM : ] Oenothera spachiana Torr. & A.Gray; DIM 989 Oenothera speciosa Nutt; DTM 122 OXALIDACEAE Oxalis corniculata L., MM 5947 *Oxalis rubra St.-Hil. DTM 492 Oxalis violacea L., DIM 417 PAPAVERACEAE PASSIFLORACEAE Passiflora incarnata L; DTM 133 Passiflora lutea L.; DTM 96 PEDALIACEAE Proboscidea louisianica (P. Mill.) Thellung; Hall s.n.LSUS PHRYMACEAE Phryma oa DTM 1290 PHYTOLACCA Phytolacca ie L.; DTM 61 Rivinia humilis L.; DTM 1490 PLANTAGINACEA ey DTM 1025 Plantago hookeriana Fisch. & C. = Mey.; MM 2580 *Plantago ignecolaea DTM 8 Plantago | fT see Plantago pusilla Nutt; 7 730 Plantago anes oe Dene; a 65015 ica L.; DTM 1026 na ADeneE DTM 913 i] - | D FICOge virgin re a MUGY VE ightia PLATANACEAE Platanus occidentalis L.; Haynes 4634 POLEMONIACEAE Phlox divaricata L.; Lawrence 17 LTU Phlox drummondii Hook.; MM 197 Phlox pilosa L.; Raymond 1200 LSUS POLYGALACEAE Palygala mariana P. Mill; RDT 170119 ke a polygama Walt; MM 2629 olygala sanguinea L., DTM 461 Polygala verticillata L., RDT 36193 POLYGONACEAE Brunnichia ovata (Walt.) Shinners; DTM 171 Eriogonum longifolium Nutt; MM 2323 Eriogonum multiflorum Benth.; MM 2870 Polygonella americana (Fisch. & C.A. Mey.) Small; MM aly lygonum avi DTM 1336 *Polgorum ey Blume; RDT 170115 , RDT 66461 LSU Polygonum densiflorum Meisn.; DTM 608 L; RDT 32876 ee hydropiperoides Michx.; DTM 113 *Polygonum lapathifolium L., DTM 601 Polygonum pensylvanicum L., DTM 213 *Polygonum persicaria L.; Haynes 4222 Polygonum punctatum Ell; Haynes 4224 Polygonum ramosissimum Michx.; DTM 643 Polygonum scandens L.; MM 292 ee setaceum Baldw.; DTM 239 nN virginianum fs a 775 SUS pare altss imus Wood; *Rumex crispus L.; DTM 81 - Rumex hastatulus Baldw.; DTM 798 *Rumex pulcher L.; DTM 865 Rumex verticillatus L.; DTM 907 PORTULACACEAE | Uyltr nia virginica L.; ‘DIM 11 Portulaca oleracea L.; DTM a Talinum parviflorum Nutt, MM 3354 Talinum rugospermum Holz.; MM 2757 PRIMULACEAE *Anagallis arvensis L.; Lynch 3861 LSUS ana minima (L.) Krause; a ies dec-atheon meadia | Lysimachia lanceolata Walt.; ee 565 LSUS Samolus valerandi L., DTM 374 RANUNCU Anemone reste Pritz., DIM 2590 Anemone caroliniana ae DTM 264 Clematis crispa L.; MM 2 Clematis reticulata Walt.; — 2680 *Clematis terniflora DC,; ey - Clematis virginia Chih lee MM 2 Delphinium carolinian um ae MM 2774 Myosurus minimus L.; Barbour 556 Ranunculus re L. DTM 953 Ranunculus fascicularis Muhl.ex Bigelow; DTM 773 Ranunculus laxicaulis (Torr.& A. Gray) Darby; DTM 292 1214 *Ranunculus marginatus d'Urv,; DTM 980 *Ranunculus muricatus L.; DTM 409 *Ranunculus parviflorus L.; DIM 878 Ranunculus pusillus Poir; OTM 775 urvatus Poir; DTM 1047 *Ranunculus sardous Crantz; DTM 1048 Ranunculus sceleratus L.; DTM 2019 Ee ce trilobus Desf; DTM 902 da m Fisch. & Ave-Lall.; DIM 901 RHAMNACEAE Berchemia scandens (Hill) K. Koch; DTM 1781 Ceanothus americanus L.; MM 2370 Ceanothus herbaceus Raf; RDT 93612 Frangula caroliniana (Walt.) A.Gray; Hardy 8646 *Ziziphus zizyphus (L.) Karst. RDT 33078 ROSACEAE Agrimonia microcarpa Wallr.; Haynes 5337 Agrimonia rostellata Wallr.; Haynes 5315 Amelanchier arborea ( Michx.f.) Fernald; Manning 485 Ranunculus re LTU *Aphanes microcarpa (Boiss. & Reut.) Rothm.; RDT 82 Aronia arbutifolia (L.) Ell; RDT 33632 Crataegus berberifolia Torr.& A.Gray; Graham 460 LSUS Crataegus brachyacantha Sarg. & Engelm.; Rowe 85 LSUS Crataegus crus-galli L.; RDT 165825 Crataegus engelmannii Sarg.; DIM 1966 LSU ig ee marshallii Egglest.; Haynes 4623 Crataegus opaca Hook. & Arn.; Haynes 4627 Caton apatiiiata Michx.; Haynes 4095 hh. MM 2578 sai viridis L.; Barbour 1164 *Duchesnea indica uae: foe DIM4 in Ansty, ~ Malus canes (Aiton) vee 88278 Porteranthus stipulatus (Muhl.ex le Britt.; DTM 1030 *Potentilla recta L.; MM 2620 Potentiilla simplex Michx., DTM 48 Prunus americana Marsh.; RDT 96045 Prunus angustifolia Marsh.; MM 2538 Prunus caroliniana (P. Mill.) Aiton; DTM 1121 Prunus gracilis Engelm.& A. Gray; MM 2539 Prunus mexicana S.Wats.; Raymond 69 LSUS Prunus serotina Ehrh.; Barbour 47 Prunus umbellata Ell; DTM 1603 M is 1.p A rd . | ' *Rosa bracteata J.C.W naturalized) Rosa carolina L.; MM 201 *Rosa laevigata Michx.; Barkley 103 LSU (questionably naturalize *Rosa multiflora Thunb. ex Murr; Lynch 2273 LSUS (questionably naturalized) BRIT.ORG/SIDA 22(2) Rosa setigera Michx.; MM s.n. Rubus argutus Link.; Raymond 2006 LSUS Rubus trivialis Michx.; Graham 444 LSUS RUBIACEAE Cephalanthus Heater is DTM 1296 Diodia teres Walt. di Ua virginiana las one _ Gah ium pee Le OT) 288 ful ans Michx.; DTM 922 Galiu ium abuse Bigelow; fa 994 *Galium parisiense L.; 323 Galium pilosum Aiton; DTM 915 Galium triflorum Michx.; RDT geen Galium uniflorum Michx.; RDT Hedyotis nigricans (Lam.) ase ae 227 oustonia micrantha (Skinner) Terrell; DTM 1979 Houstonia pusilla Schoepf; Raymond 741 LSUS Houstonia rosea (Raf.) Terrell; DTM 255 Mitchella repens L.. DTM 378 Oldenlandia boscii (DC.) Chapman; DTM 2996 Oldenlandia uniflora L.; ie ae nna scabra L T ans nee 9 Spermacoce glabia Michie! DIM 1 RUTACEAE *Poncirus trifoliata (L.) R a oie ata L.; RD ava ae if Raymond 13 a _ 19LSUS SALICACEA *Populus Ae L.; DIM 2230 Populus deltoides Bartr.ex Marsh.; Haynes 4633 Salix exigua Nutt; Haynes 4157 Salix humilis Marsh. RDT 88359 LSU Salix nigra Marsh.; Haynes 5700 SAPINDACEAE Cardiospermum halicacabum L.; Haynes 5292 Sapindus saponaria L.; RDT 88240 riniaanates 1Michx.; MM 2828 . RDT 33011 Sideroxvlon Ivcioides L SAURURACEAE Saururus cernuus L.,; DTM 30 SAXIFRAGACEAE Heuchera americana L.; Hay ea eke ies ai . 34504 SCROPHULARIACEAE Agalinis ae Pennell; Pennell 5658 PH Agalinis fasciculata (Ell.) Raf; DIM 755 Agalinis gattingeri (Small) Small; DTM 541 Agalini is heterophylla (Nutt.) ex Britt; DIM 736 Agalinis homalantha Pennell; DTM 748 Agalinis purpurea (L.) Pennell; a 2570 BERT. AAVUY TART, LOUISIANA 1215 Agalinis tenuifolia (Vahl) Raf; ate a Ag alinis Vil idis (Small) Penn ell: D Aureolaria grandiflora (Benth.) et DTM 605 Aureolaria pectinata (Nutt.) Pennell; Cocks s.n.NO Bacon g Kk poungiols en) Wettst., DTM 1592 aie indivisa ee Barbour *Chaenorhinum minus (L.) Lange; ae 65115 LSU Collinsia violacea Nutt.; Maynard 151 LTU Dasistoma macrophylla (Nutt.) Raf; Palmer 10611 MO Gratiola flava Leavenworth; MM 7285 Gratiola neglecta Torr; DTM 398 Gratiola pil osd Michx.; RDT 33636 Gratiola virginiana L.; DTM 1127 ee multifida (Michx.) Nutt.; DTM 1103 Lindernia crustacea (L.) F. Muell.; RDT 41803 Lindernia dubia (L.) Pennell; DTM 165 *Mazus pumilus (Burm.f) Steenis; DTM 964 Mecardonia acuminata (Walt.) Small; DTM 216 Mimulus alatus Aiton; DTM 603 Nuttallanthus canadensis (L.) D.A. Sutton; DTM 263 Nuttallanthus texanus (Scheele) D.A. Sutton; DTM 381 *Pqrentucellia viscosa (L) Caruel; MM 2627 Beale ial is ae s Ly DTM 37 is Nutt. ex Sims; ao 554 LSUS Penstemon iain Pennell; DTM Penstemon murrayanus ee MM o 7 Penstemon tenuis Small M557 Penstemon tubaeflorus ce DTM 81 Scrophularia marilandica L; RDT 88232 *Verbascum blatteria L. Hardy 6 *Verbascum thapsus L.; DTM 200 *Veronica arvensis L.. DTM 262 *Veronica hederifolia L; Burns 13 LSUS Veronica peregrina L.; DTM 303 *Veronica persica Poir.; MM 5966 *Ver UT Ca polita FI ies; DIM 273 SIMAROUBACEAE *Ajlanthus altissima (P. Mill.) Swingle; Lewis 2185 NLU SOLANACE fee stramonium L pie 930 Watd |: Physalis cineraesens (Dunal) ; S.Hitche,; RDT 32855 Physalis cordata P. Mill; RDT 32854 ede hederi i ia Gray; RDT 89017 salis f [| Nees; MM 2619 Physalis longifol ia Nutt. MM 6667 Physalis mollis Nutt.; MM 2608 Physalis pubescens L; DIM 1779 oe pu) Nutt; RDT 140326 eL.; DIM 99 Gc lanum i fj R af.; Petersen s.n.LSU Solanum elaeagnifolium Cav.; DTM 108 *Solanum pseudocapsicum L.; Raymond 2149 LSUS hanthum Dunal; DTM 118 Solanum nf\ | ia Salanum rostratum Dunal: RDT 81358 SPHENOCLEACE *Sphenoclea aoa ee Gaertn.; RDT 36943 STYRACACEAE Styrax americanus Lam.; MM 189 Styrax grandifolius Aiton; RDT 37239 SYMPLOCACEAE Symplocos tinctoria (L.) Her; Barbour 1166 TAMARICACEAE *Tamarix gallica L, DTM 1318 TILIACEAE Tilia americana L; DTM 1144 — = ULMACEAE Celtis laevigata Willd.; Haynes 4092 Celtis tenuifolia Nutt.; RDT 37283 Planera aquatica J.F. Gmel.; Haynes 8648 ie alata Michx.; Haynes 4508 us americana L. Haynes 4574 ae crassifolia Nutt.; Hardy 8610 Ulmus rubra Muhl.; Raymond 308 LSUS URTICACEAE Boehmeria cylindrica (L.) Sw, DTM 224 Parietaria pensylvanica Ae ex a DIM 1796 Pilea pumila (L.) A. Gray; M Urtica chamaedryoides ee oa 3 VALERIANACEAE Valerianella radiata (L.) Dufr; DTM 1028 VERBENACEAE Callicarpa americana L; DTM 1 Glandularia bipinnatifida (Nutt. ; ie RDT 99514 Glandularia canadensis (L.) Nutt; DTM 9 *Glandularia pulchella (Sweet) Troncoso; DTM 1355 *Lantana camara L; RDT 36934 Lantana urticoides Hayek; DTM 1570 Phyla lanceolata (Michx.) Greene; DTM 22 Phyla nodiflora (L .) Greene; DTM 9 *Verb 5 Verbena nate I; D I a 893 Verbena na [ss DTM 440 Verbena xutha Leh RRs *Verbel 1A *Vitex agnus-castus . MM 226 VIOLACEAE Viola bicolor Pursh; DTM 257 Viola lanceolata L., DTM 2106 Viola palmata L; DTM 772 Viola pedata L., DTM 43 Viola primulifolia L.; DTM 21 a He gee pus RDT 1 iola DTM aoe la sagittata Aito 1216 BRIT.ORG/SIDA 22(2) Viola sororia Willd.; DTM 788 Parthe issus quinquefolia (L.) Planch.; MM 6208 Viola villosa Walt. MM 141 Vitis aestivalis Michx., DTM 912 Viola walteri House; DTM 816 Vitis cinerea (Engelm.) Millard; Correll & Correll 10091 VISCACEAE Prati ane Phoradendron tomentosum (DC) Engelm. ex A. Gray; vitis palmata es RDT 25315 DTM 2102 Vitis riparia Michx., DTM 1159 Vitis rotundifolia Michx.; MM 6201 VITACEAE Vitis vulpina L.; RDT 42042 Ampelopsis arborea (L.) Koehne; DTM 152 Ampelopsis cordata Michx., DTM 905 ZYGOPHYLLACEAE Cissus incisa Des Moul.; DTM 1784 *Tribulus terrestris L., DTM 600 ACKNOWLEDGMENTS Stephanie Aamodt, former Chairman of the Biology Department, Louisiana State Uni- versity in Shreveport, and Al Vekovius, former Dean of Science, Louisiana State Univer- sity in Shreveport, provided the space for an herbarium where the work could be done. Guy Nesom, BRIT; Larry Brown, SBSC; Billie Turner, TEX, and Diane Ferguson, LSU, helped in determining several problematic specimens. Diane Ferguson, LSU; Kerry Heafner, NLU; Garrie Landry, LAF; Amanda Neill, BRIT; Eric Sundell, UAM; Milan Vavrick, LTU: Anne Bradburn, NO; Sherri Herndon, DUKE: George Russell, US,and James Solomon, MO, loaned specimens. Thanks also to Chris Reid, Louisiana Natural Heritage Program; Charles Allen, Botanist, Fort Polk; the late John Thieret, KNK; Larry Raymond, Caddo Parish Parks De- partment, and Gary Hanson, Patricia Peyton, and Scott Foord of the Red River Watershed Management Institute, Louisiana State University-Shreveport. Paul Sisson, Dean of Sci- ence, Louisiana State University in Shreveport, obtained funds for publication costs. Charles Allen, R.D.Thomas, and an anonymous reviewer provided helpful comments on the manuscript. REFERENCES Aiten,C.M.,D.A. Newman, and H. Winters. 2004. Grasses of Louisiana. 3rd Edition. Allen’s Native Ventures, Pitkin, Louisiana. Braun, E.L. 1950. Deciduous forests of eastern North America. Hafner Press, New York. Canne-Hivuker, JM. and M. Dusrute. 1993. A new species of Agalinis (Scrophulariaceae) from Grimes County, Texas. Sida 15:425-440, Corrett, D.S. and H.B. Corrtit. 1941. A collection of plants from Louisiana. Amer. MidI. Naturalist 26: 30-64. Diccs, G.M., B.L. Lipscome, and R.J.O’Kennon, 1999. Illustrated flora of north central Texas. Sida Bot. Misc. 16:1-1626, Diccs, G.M., B.L. Lirscome, M.D. Reep, and RJ. O’Kennon. 2006. Illustrated flora of east Texas. Sida Bot Misc. 26:1-1594., Epwarbs, J.P, P.G. Martin, J.W. Macoun, W.W. Kicpatrick, and C. Henry. 1980. Soil survey of Caddo Parish, Louisiana. U.S.D.A. Soil Conservation Service, Washington, D.C. Flora or Nort America Eptroriat Comirtee (eds.). 1993-2006. Flora of North America North of Mexico. Vols. 1, 2, 3,4, 22, 23, 25, 26. Oxford Univ. Press, New York and Oxford. Fiores, D.L. 1984. Jefferson and southwestern exploration: The Freeman and Custis accounts of the Red River expedition of 1806. Univ. Oklahoma Press, Norman. GANDHI, K.N. and R.D. THomas. 1989. Asteraceae of Louisiana. Sida Bot. Misc. 4:1—202. Be Ah ace RISH, LOUISIANA 1217 Karresz, J.A. and C.A. Meacham. 1999. Synthesis of North American flora. Version 1.0. North Carolina Botanical Garden. Chapel Hill. Ker, E.L,, JR. SincHurst, and S. Cook. 2004, Geocarpon minimum (Caryopyllaceae), new to Texas. Sida 21:1165-1169. Larke, J.O.and L.M. Switn. 1994. Rare plants of pine-hardwood forests in Louisiana. Louisiana Depart- ment of Wildlife and Fisheries, Baton Rouge. Leipotr, A., S.McDaniet, and T. Nuttie. 2002. The flora of Oktibbeha County, Mississippi. Sida 20:691—765. Lester, G.D,, S.G. Sorensen, PL. Fauckner, C.S. Reio, and .E. Maxi. 2005. Louisiana Comprehensive Wildlife Conservation Survey. Unpublished report. Louisiana Natural Heritage Program, Louisiana Depart- ment of Wildlife and Fisheries, Baton Rouge. Louisiana Naturat Heritace Procram. 2004. Rare plant species of Louisiana. Unpublished report. Louisi- ana Natural Heritage Program, Louisiana Department of Wildlife and Fisheries, Baton Rouge. Louisiana Naturat Heritace Procram. 2006. Rare plant species of Louisiana. Unpublished report. Louisi- ana Natural Heritage Program, Louisiana Department of Wildlife and Fisheries, Baton Rouge. MacRoseats, B.R. and M.H. MacRoserts. 1994. Floristics of a xeric sandyland in western Louisiana. Phytologia 77:414-424,. MacRoseats, B.R.and M.H.MacRoserts. 1995. Vascular flora of xeric sandhills in northwestern Louisiana. Phytologia 79:123-131. MacRoserts, B.R. and M.H. MacRoserts. 1996. Floristics of xeric sandhills in east Texas. Phytologia 80: desk. MacRoserts, B.R. and M.H. MacRoserts. 1997. Talinum rugospermum Holz. new to Louisiana with notes on terete-leaved Talinum in Louisiana. Phytologia 82:86-93. MacRoserts, B.R., MH. MacRoserts, and J.C. CatHey. 2002. Floristics of xeric sandylands in the post oak savanna region of east Texas. Sida 20:373-386. MacRoeerts, D.T. 1977a. Additions to the Louisiana flora. Sida 7:220-222. MacRoseats, D.T. 1977b. New combinations in Tradescantia. Phytologia 37:451-452. MacRoserts, D.T. 1978a. Notes on Tradescantia: T. diffusa Bush and T. pedicillata Celarier. Phytologia 38:227-228. MacRoserts, D.T. 1978b.The status of Agalinis caddoensis Pennell. Phytologia 40:1-6. MacRoserts, D.T. 1979a. Checklist ifthe plants of Caddo Parish, Louisiana. Bull. Mus. Life Sci,, Louisiana State University-Shreveport 1:1-54. MacRoserts, D.T. 1979b. Notes on Tradescantia Ill. Tradescantia ohiensis Raf. var. paludosa (Anderson and Woodson) MacRoberts, comb. nov. Phytologia 43:380-382 MacRoserts, D.T. 1980a. Notes on Tradescantia IV: The distinction between Tradescantia virginiana (Commelinaceae) and Tradescantia hirsutiflora. Phytologia 46:409-416. MacRoserts, D.T.1980b.Notes on Tradescantia (Commelinaceae) V. Tradescantia of Louisiana. Bull. Mus Life Sci., Louisiana State University-Shreveport 4:1—-15. MacRoserts, D.T. 1980c. Notes on Tradescantia VI. Observations on the mechanism of pigment depo- sition. Phytologia 46:41 7-418. MacRoserts, D.T. 1984a.The vascular plants of Louisiana. An annotated checklist and bibliography of the vascular plants reported to grow without cultivation in Louisiana. Bull. Mus. Life Sci., Louisiana State University-Shreveport 6:1-165. MacRoserts, 0.1.1984b.Helianthemum rosmarinifolium and Oenothera triloba in Louisiana. Phytologia 55:343. MacRoeerrs, D.T. 1987.Reported Texas endemics. Castanea 52:72. 1218 BRIT.ORG/SIDA 22(2) MacRoserts, D.T. 1989. A documented checklist and atlas of the vascular flora of Louisiana. Bull. Mus. Life Sci., Louisiana State University-Shreveport 7—9:1-756. MacRoserts, D.T.,B.R.MacRoserts, and M.H.MacRostrts. 1997.A floristic and ecological interpretation of the Freeman and Custis Red River Expedition of 1806. Bull. Mus. Life Sci., Louisiana State Univer- sity-Shreveport 12:1-26. MacRoserts, M.H. and B.R. MacRoseats. 1998. Quality change of habitat in northwestern Louisiana. Phytologia 84:297-303. MacRoseets, M.H., B.R. MacRoserts, B. Sorrie, and R. Evans. 2002. Endemism in the West Gulf Coastal Plain: importance of xeric habitats. Sida 20:767-779. MacRoberts, M.H., B. R. MacRoserts, and L.S. Jackson. 2003. Louisiana prairies. In: E. Peacock and T. Schauwecker, eds. Blackland Prairies of the Gulf Coastal Plain: Nature, Culture and Sustainability. Univ. Alabama Press, Tuscaloosa. Pp. 80-93. MacRoserts, M.H. and B.R. MacRoserts. 2003a.West Gulf Coastal Plain ecoregions. Sida 20:1245-1274. MacRoserts, M.H.and B.R.MacRoserts.2003b.The east-west transition of flora in Texas:a biogeographical analysis. Sida 20:1693-1700. MacRoserts, M.H. and B.R. MacRoserts. 2004a. West Gulf Coastal Plain prairies:a first approximation at a synthesis. Third Eastern Native Grass Symposium, Chapel Hill, North Carolina: 5-21. MacRoetrts, M.H.and B.R.MacRoserts. 2004b, About 450 miles up the Red River in a large prairie: Peter Custis’ surviving botanical information from the Red River expedition of 1806. Sida 21:963-975. MacRoserts, M.H. and B. R. MacRoserts. 2005. Reference conditions of the Red River floodplain and upland, Caddo Parish, Louisiana. Sida 21:1793-1806. MacRoserts, M.H. and B.R. MacRoserrs (in prep.). Vascular flora and community structure of the C. Bickham Dickson Park/Red River Education and Research Park, Caddo Parish, Louisiana, Red River floodplain. Marsico, T.D. 2005. The vascular flora of Montgomery County, Arkansas. Sida 21: 2389-2423. McBryoe, J.B. 1933. The vegetation and habitat factors of the Carrizo sands. Ecol. Monogr. 3:247-297. Mclnnis, N.C., LLM. Smith, and A.B. Pittman. 1993. Geocarpon minimum (Caryophyllaceae), new to Louisi- ana. Phytologia 75:159-162. McKinney, LE. 1992. A taxonomic revision of the acaulescent blue violets (Viola) of North America. Sida Bot. Misc. 7:1-60. McKinney, L.E. and N.H. Russet. 2002. Violaceae of the southeastern United States. Castanea 67: 369-379. McLaucutin, S.P.(in press). Tundra to Tropics: The floristic plant geography of North America. Sida, Bot Misc. Nesom, G.L.and L.E. Brown. 1998. Annotated checklist of the vascular plants of Walker, Montg y, and San Jacinto counties, east Texas. Phytologia 84: 107-153. Nesom, G.L. 2004a. New species of Gamochaeta (Asteraceae: Gnaphalieae) from the eastern United States and comments on similar species. Sida 21:717-741. Nesom, G.L. 2004b. New distribution records for Gamochaeta (Asterac phali in the United States. Sida 21:1175-1185. Neyianp, R. 2002. An update to the vascular flora of Calcasieu Parish, Louisiana. Sida 20:431-433. Oversy, RE. 1974. A preliminary survey of the vascular flora of Caddo Parish, Louisiana. M.S. Thesis, Northeast Louisiana University, Monroe. Reio, C.and P. Fauckner (in press). Loeflingia squarrosa (Caryophyllaceae): new to Louisiana. Phytologia SKFEN, J.N., P.D. Doerr, and D.H. VAN Lear. 1993. Oak-hickory-pine forests. In: WH. Martin, S.G. Boyce, and A.C. Echtermacht, eds. Biodiversity of the southeastern United States. John Wiley, New York. Pp. 1-33 MACROBERTS AND MACROBERTS CADDO PARISH, LOUISIANA 1219 Smith, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas. Privately published. Fayetteville, Arkansas. Situ, E.B. 1994. Keys to the flora of Arkansas. Univ. Arkansas Press, Fayetteville. Teacue, J.and T. Wenor 1994. Caddo and Bossier parishes, Louisiana, natural areas survey. The Nature Conservancy, Baton Rouge. THomas, R.D.and C.M. Aiten. 1993-1998. Atlas of the vascular flora of Louisiana.Louisiana Department of Wildlife and Fisheries, Baton Rogue. THomas, R.D.,L.R. RavMonp, B.R. MacRoserts, and M.H.MacRoserts (in press). Documented and annotated checklist of the vascular flora of the Walter B. Jacobs Memorial Nature Park, Caddo Parish, Louisi- ana. Proc. Louisiana Acad. Sci. THorne, RF. 1993.Phytogeography. In: Flora of North America Editorial Committee, eds. Fl. North Amer. 1:132-153.Oxford Univ. Press, New York and Oxford. Taiska, FJ. 1984. Role of wood debris in modifying channel geomorphology and riparian areas of a large lowland river under pristine conditions: a historical case study. Verh. Internat. Verein. Limnol. 22:1876-1892. Turner, B.L., H.NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Sida Bot. Misc. 24:1-888. 1220 BRIT.ORG/SIDA 22(2) Book REVIEW Doucias J. Kennet and Bruck WINTERHALDER. 20060. Behavioral Ecology and the Transition to Agriculture. (ISBN 0-520-24647-0, hbk.). University of California Press, Califor- nia/Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A. (Orders: www.ucpress.edu/, 609-883-1759, 609-883-7413 fax). $60.00, 407 pp., 3 b/ w photos, 31 line illustrations, 22 maps, 30 tables, 7" x 10". Human behavioral ecology (as in the Foreword) focuses on how and why people have chosen, in social and cul- most funda- “This volume examines “one ol the m tural contexts, to exp dloit partic ular resources. From the Preface: mental economic shifts in human history—the evolutionary transition from foraging to farming through pro- he nenap tel authors use a variety of cen- cesses ? pen ane mee sscuiiat iaetcac ane. the emergence A apiculate. T spersed cz tral ee betas citeuncee. risk minimization, antl costly signaling shieae y. Their euanibirens are novel in presenting regionally comprehensive case studies that address the transition to agriculture from a consistent conceptual framework informed by neo-Darwinian theory.” A collection of 14 technical but readable papers.— Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S Book Notice W. SHENG YI, PeTeR H. Raven, and H. Deyuan (editorial co-chairs). 2006. The Flora of China, Volume 22, Poaceae. (ISBN 1-930723-50-4, hbk.). Missouri Botanical Garden Press, P.O. Box 299, Saint Louis, MO 63166-0299, U.S.A. (Orders: 877-271-1930, http:// www.mbgpress.org) $140.00, 752 pp.,9" x 111/2" le family, the Poaceae, ene -volume work. : umerades sing ) ] 809 < eas 22 of ne E ne of China isthe l2th of a 25 tribes, 226 genera, and 1795 species, among to China.” FE xcept for the protracted list of Asian Bambuseae genera (34), me sity in genera berieene a and North America is remarkable. The Chinese grass flora also is close in size to the North American one (nor of Mexico), perhaps even 20% greater in number of species. The first of two aes of oe FNA treatment was Many recent cl ts of grass genera are incor- the FNA treatments.—Guy published in 2003; it will be published in early 2007. porated in the Chinese flora, presumably making it parallel and peers sith Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4068, U.S.A. = rt SIDA 22(2): 1220. 2006 CAREX STIPATA (CYPERACEAE), CHAMAESYCE GEYERI (EUPHORBIACEAE), EURY TAENIA TEXANA (APIACEAE), PEDIOMELUM ESCULENTUM (FABACEAE), AND TALINUM CALYCINUM (PORTULACACEAE) DELETED FROM THE LOUISIANA FLORA Barbara R. MacRoberts and Michael H. MacRoberts Bog Research, 740 Columbia, Shreveport, Louisiana 71104, U.S.A. Herbarium, Museum of Life Sciences, Louisiana State University-Shreveport Shreveport, Louisiana 71115, U.S.A. anc Red River Watershed Management Institute, Louisiana State University-Shreveport Shreveport, Louisiana 71115, U.S.A. In the course of developing a vouchered checklist of the vascular flora of Caddo Parish, Louisiana (MacRoberts & MacRoberts in prep.), we discovered six species reported for Caddo Parish for which we could find vouchers neither for the Parish nor the State. We already have reported on one of these: Agrimonia gryposepala Wallr. (MacRoberts & MacRoberts 2005). Here we report on five others. Until vouchers are found, these species should be deleted from the Louisiana flora. Carex stipata.—MacRoberts (1979, 1984, 1989), Thomas and Allen (1993-1998), Kartesz and Meacham (1999), NatureServe (2005), and U.S.D.A. (2005) include Carex stipata Muhl. ex Willd. in the Louisiana flora based on a specimen collected by D.T. MacRoberts in Caddo Parish (MacRoberts 1349 LSUS). We examined the specimen and found it to be a misidentified Carex vulpinoidea Michx. (Phil Hyatt also annotated it to C. vulpinoidea). The reported distribution of C. stipata is confused. It is not reported for Loui- siana, Mississippi, or Arkansas in the Flora of North America but it is reported for these states by Kartesz and Meacham (1999), NatureServe (2005), and U.S.D.A. (2005). Turner et al. (2003) do not report it for Texas. Chamaesyce geyeri.—MacRoberts (1979, 1984, 1989), Thomas and Allen (1993-1998), Kartesz and Meacham (1999), NatureServe (2005), and U.S.D.A. (2005) include Chamaesyce geyeri (Engelm.) Small in the Louisiana flora ae on several specimens collected by D.T. MacRoberts in Caddo Parish (MacRoberts 1791, 1940 LSUS, 1951 LSU). We examined these specimens and found them to be misidentified Chamaesyce cordifolia (EIL.) Small. Chamaesyce geyeri is a central plains species that apparently does not occur nearer to Louisiana than central Texas and central Oklahoma (Turner et al. 2003, Oklahoma Vas- cular Plant Database 2005). Eurytaenia texana.—Thieret (1971), MacRoberts (1979, 1984, 1989), Thomas and Allen (1993-1998), Kartesz and Meacham (1999), LSU (2005), NatureServe (2005), and US.D.A. (2005) include Eurytaenia texana Torr. & A. Gray in the Louisiana flora on the basis of many collections from Caddo, Winn, Webster, and DeSoto parishes (e.g., [hieret 32309 LAE Thomas 45467, 65016, 70521, 99542, 99703 NLU, Thompson 485 LTU, Dixon 1342 NLU). We examined these specimens and others labeled Eurytaenia texana from Louisiana in BRIT, NLU, LSU, LTU, and LAF and found all of them to be misidentified Polytaenia SIDA 22(2): ): 1221-1223. 2006. 1222 BRIT.ORG/SIDA 22(2) nuttallii DC. The distribution of Eurytaenia texana is central Texas, the Texas coastal prairie region, and western Oklahoma (Mathias & Constance 1961, Taylor & Taylor 1989, Hatch et al. 1990, Turner et al. 2003, Oklahoma Vascular Plant Database 2005, TAMU 2005). It is, however, reported from Jefferson Co., Texas, on the Louisiana Gulf border and thus could possibly be present in the Louisiana coastal prairie region (Mathias & Con- stance 1961, Turner et al. 2003). A reported specimen of Eurytaenia texana from Arkan- sas housed at NLU turned out to be a misidentified Thaspium barbinode (Michx.) Nutt. (Eric Sundell, pers. comm.). Pediomelum esculentum.—MacRoberts (1979, 1984, 1989), Thomas and Allen (1993- 1998), Kartesz and Meacham (1999) and U.S.D.A (2005) include Pediomelum esculentum (Pursh) Rydb. in the Louisiana flora based on a specimen collected by D.T. MacRoberts from Caddo Parish (MacRoberts 1170 LSUS). We examined the specimen and found it to be a misidentified Pediomelum hypogaeum (Nutt. ex Torr. & A. Gray) Rydb. var. subulatum (Bush) J. Grimes. Pediomelum esculentum apparently does not occur nearer to Louisiana than northern Arkansas and central Oklahoma (Oklahoma Vascular Plant Database 2005, US.D.A. 2005). Talinum calycinum.—MacRoberts (1979, 1984, 1989), Thomas and Allen (1993-1998), Kartesz and Meacham (1999), NatureServe (2005) and US.D.A (2005) include Talinum calycinum Engelm. in the Louisiana flora based on specimens collected by Thomas in Caddo and Vernon parishes (Thomas 34513, 105079 NLU). We examined all Louisiana T. calycinum specimens (MacRoberts & MacRoberts 1997) and found them to be misidentified I’ parviflorum Nutt. Talinum calycinum apparenty does not occur nearer to Louisiana than central Texas, Oklahoma, and central Arkansas (Turner et al. 2003, Oklahoma Vascular Plant Database 2005, U.S.D.A. 2005). ACKNOWLEDGMENTS Diane Ferguson (LSU), Kerry Heafner (NLU), Garrie Landry (LAF), Amanda Neill (BRIT), Eric Sundell (UAM), John Thieret (KNK), Milan Vavrick (LTU), and Gary Hanson and Patricia Peyton, Red River Watershed Management Institute, aided with this work. R. Dale Thomas kindly reviewed the paper. REFERENCES Harcn, S.L, KN. GANDHI, and L.E. Brown, 1990. Checklist of the vascular plants of Texas. Texas Agricul- tural Experiment Station, Texas A&M University, College Station. Karresz, J.T. and CA. Meacuam. 1999. Synthesis of North American flora. Version 1.0 North Carolina Botanical Garden, Chapel Hill. LSU 2005. www.herbarium.|su.edu MacRoserts, B.R.and M.H. MacRosrers. 2005. Agrimonia gryposepala (Rosaceae) deleted from the Loui- siana flora. Phytologia 87:129-131. MacRoserts, B.R. and M.H. MacRoseets. in prep. Vouchered checklist of the vascular flora of Caddo Parish, Louisiana, with notes on regional phytogeography and ecology. MacRoseets, D.T. 1979. Checklist of the plants of Caddo Parish, Louisiana. Bull. Mus. Life Sci. Louisiana State Univ. 1:1—54. MacRoserts, D.T. 1984. The vascular plants of Louisiana. An annotated checklist and bibliography of the vascular plants reported to grow without cultivation in Louisiana. Bull. Mus. Life Sci. Louisiana State Univ. 6:1-165. FLORA 1223 MacRoseats, D.T. 1989. A documented checklist and atlas of the vascular flora of Louisiana. Bull. Mus Life Sci. Louisiana State Univ. 8:257-536. MacRoserts, M.H.and B.R. MacRoserts. 1997. Talinum rugospermum Holz.,new to Louisiana with notes on terete-leaved Talinum in Louisiana. Phytologia 82:86-93. Martuias, M.E. and L. Constance.1961. Umbelliferae. Flora of Texas. Vol. 3. Part 5. Texas Research Founda- tion, Renner. NatureServe. 2005. www.natureserve.org OKLAHOMA VASCULAR PLANT Database. 2005. http://g botanical/ TAMU 2005. www.csdl.tamu edu/FLORA/tracy2/main| hel Tayior, R.J.and C.E.S. Tayior. 1989. An annotated list of the ferns, fern allies, gymnosperms and flower- ing plants of Oklahoma. Southeastern Oklahoma State University, Durant. Tuieret, JW. 1971. Additions to the Louisiana flora. Castanea 36:219-222. THomas, R.D.and C.M. Atten. 1993-1998. Atlas of the vascular flora of Louisiana. Louisiana Department of Wildlife and Fisheries, Baton Rouge. Turner, B.L., N. NicHots, G. Denny, and O. Doran. 2003. Atlas of the vascular plants of Texas. Sida Bot. Misc. 24:1-888. U.S.D.A. 2004. http://plants.usda.gov 1224 BRIT.ORG/SIDA 22(2) Book NOTICES Yonc No Ler. 2006. New Flora of Korea (two volume set). ISBN 89-09-11801-6, vol. 1: ISBN 89-09-11802-4, vol. 2, hbk.). Kyo-Hak Publishing Co., Ltd., 105-67, Gongdeok-dong, Mapo-gu, Seoul, Korea. (Orders: 02-7075-155-156, 02-7075-160 Fax, chief @k yohak.co.kr, www:kyohak.co.kr) $345.00 (postage (surface) included to U.S.A.), 976 pp., vol. 1; 888 pp., vol. 2, abundant color Maia Ox Ly 2 So many beautiful and detailed color photos (1 estimate 80+% of the species with a photo: match photo number yer)... but Lcan read the scientifien names, that’s all. Ferns, grasses, sedges, trees, daisies, and all with species num the rest—all illustrated in two big volumes (boxed set). It’s not cheap but what a value for botanists. From the ‘flyer’ 4157 taxa of native and naturalized plants of Korea; almost all plant pictures have been taken by the au- thor, Vol. | with 976 pages, Vol. I] with 888; scientific names, proper Korean names, English names, Chinese names and Japanese names have been used; habitats, distributions, flowering seasons, and chromosome numbers have been recorded; new species and rearranged species names have been arranged in an appendix pinay an in- dex of new taxa and new combinations by the author in various literature from 1993 to early 2006, joined by many new combinations proposed here de novo). Since synonyms are given for very few of the scientific names, the index to scientific names functions as a checklist for the whole flora —Guy Nesom, Botanical Research Insti- tute of Texas, 509 Pecan Street, Fort Worth, 1X 76102-4068, U.S.A. Davin Baxter (Photographs by Laurence Parent). 2002. Nature of the Forest: Temple-Inland’s Timberlands in the Twenty-first Century. (ISBN 009718675-0-X, hbk.). Temple-In- land Inc. No other information provided. It's not easy to be enthusiastic about books about timber companies, published by timber companies, because so much private forest land has been turned to monoculture, the diversity of all plants reduced literally to a few species. Practices of the USDA Forest Service commonly give the same result over large portions of public land. This book obviously is intended to put the best face possible on the Temple-Inland timber company—nicely written is surely one of the white Gwhiter?) hats ces . ba: lates in Hmber and full of beautiful pictures. But T management, as it apparently has set aside an unusual amount of land for preservation The book notes that “Temple maintains about a quarter of its land as natural forest;” .. some seventeen percent af its land is in SMZs [streamside ne ment zones|—340,000 acres. Plans are to level olf at eighteen to twenty percent of the company’s total acreage.” Italso maintains a number of “special places” conserved within its hold- ings in east Texas, Alabama, and Georgia, and in 2006 the Native Plant Society of Texas recognized Temple-In- ynatur Qa with an award for its contribution to native plant habitat preservation in East Texas. As noted in the book, “Temple-Inland is in the tree puSnieSS Bu une greater Temple-Inland company has lanc expertise in real estate ove opment As such, Temple is company land for its ‘highest and best use.’ Temple already owns the land, so would not be out the cost ay is cai — might ~~ at oni ly types of development.” The ultimate fate of TI holdings and all timber lands has recent unpredictable—Boise-Cascade sold all of its 2.3 million acres of forestland in 2004, and in 2005 ineeroationel Paper announced the sale of its 6.8 million acres of North American timberland, including many conservation areas (see Nature Conservancy, Autumn 2006). Only a small fraction of this land has been moved into some kind of conservation status. At least commercial timber land, even with its greatly reduced species diversity, can be coaxed back into a semi-natural condition, but other kinds of development (housing or commercial) are more nearly irrevocable.—Guy Nesom, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, 1X 76102-4068, U.S.A SIDA 22(2): 1224. 2006 NEW VASCULAR PLANT RECORDS FOR NEW MEXICO Ronald L. Hartman Brian Reif Rocky Mountain Herbarium 53 Chof Trail Department of Botany, Dept. 3165 Flagstaff, Arizona 86001, U.S.A. ae versity of pa ing Brianreif_2000@yahoo.com 000 E. University Ave. Laramie ‘ee 82071, U.S.A. Rhartman@uwyo.edu B.E. Nelson Brian Jacobs Rocky Mountain Herbarium ource Management mites ae Dept. 3165 na National Monument University of Wyoming National Park Service 1000 - eee ity Ave. HCR-1, Box 1, Suite 15 Laramie Wyoming 82071, U.S.A. Los Alamos, New Mexico 87544, U.S.A. Bnelsonn@uwyo.edu Brian_jacobs@nps.gov ABSTRACT Additions to the vascular flora of New Mexico are reported herein: Achnatherum nelsonii var. sada ie glandulosa, Cardamine cordifolia var. incana, Carex deweyana var. deweyana, Carex rosea, Cynos Draba cate Equisetum Xnelsonii, Erigeron nivalis, Geum triflorum var. triflorum, ene hallii judians nigra, Lactuca biennis, Lepidium ramosissimum var. bourgeauanum, Neoparrya lithophila, Penstemon glaber var. alpinus, Piptatheru gens, Potentilla fissa, Prunus persica, Ranunculus alismifolius var. montanus, Ranuncu- lus repens, Renibod yes Rudbec kia laciniata var. laciniata, and Syringa vulgaris. RESUMEN \chnath lsonii var. nelsonii, Bet ula -] Se aportan las adiciones siguientes a la flora vascular de Nuevo México: glandulosa, Cardamine cordifolia var. incana, Carex deweyana var. dewe yana, Carex rosea, Cyn Draba grayana, Equisetum Xnelsonii, Erigeron nivalis, Geum t triflorum var. triflorum, eee inal it; ue nigra, Lactuca biennis, Lepidium ramosissimum var. bourgeauanum, Neoparrya lithophila, Penstemon glaber var. alpinus, Piptatherum pungens, Potentilla fissa, Prunus persica, Ranunculus alismifolius var. montanus, Ranuncu- lus repens, Rorippa sylvestris, Rudbeckia laciniata var. laciniata, y Syringa vulgaris Past and recent inventories on the flora of north central New Mexico have resulted in 24 native or naturalized taxa new to the state or verification of reports. Much of this work is part of an ongoing inventory of the Rocky Mountains by the students and staff of the Rocky Mountain Herbarium (http://www.rmh.uwyo.edu). Thus far more than 44 inten- sive, large-scale projects have been completed. More specifically, these records are the re- sult of the following inventories: Philmont National Scout Ranch (Hartman 1973), Bandelier National Monument (Jacobs 1989), Valles Caldera National Preserve (Hartman & Nelson 2005), and the Santa Fe National Forest (Reif @ Hartman 2005; Reif 2006). An additional study, Carson National Forest and adjoining Bureau of Management lands, is in its second year (Larson, Hartman, and Nelson, unpub.). All specimens are deposited at the Rocky Mountain Herbarium (RM) or at the herbarium of Bandelier National Monu- ment (here designated BAND). Determination that a taxon is new to New Mexico is based on Allred (2005). In several instances a taxon included in this report has been attributed to New Mexico, but without documentation. SIDA 22(2): 1225- 1233. 2006 1226 BRIT.ORG/SIDA 22(2) APIACEAE Neoparrya lithophila Mathias—The type locality for this species was attributed to New Mexico (“on rocks, Huefano[Huerfano] Mts, New Mexico, Sept., 1867, CC. Parry 83”) but is now considered to be from Huerfano Co., Colorado (Weber 1958, Theobald et al. 1964). Hartman (in O'Kane et al. 1988) reported it from several sites as far as 51 mi to the south- west and 70-80 mi to the west of the type locality, previously the only known site for the species. Based on this report and subsequent inventory in San Isabel and Rio Grande Na- tional Forests and vicinity, it is now documented from more than 30 localities largely along the eastern margin of the San Juan Volcanic Area. Most recently, collections (RLH) have been made in the South Pifion Hills in the extreme southern San Luis Valley, imme- diately to the north of the locality reported below. Thus a taxon originally attributed erroneously to New Mexico, then considered endemic to Colorado, indeed occurs in New Mexico but only by one-half mile. — Voucher specimen: Taos Co.: Carson National Forest and Vicinity: southern extension se Pinon Hills into New Mexico, N36.9915 W105.7935, 7900 ft elev, rocky outcrop, 29 Jul 2005, Hartman 8159 { ASTERACEAE Erigeron nivalis Nutt.—The following specimen was included in Allred (2005) as E. acris Var. asteroides (Anderz. CX Besser) DC. due to a misidentification (Hartman 2003). In the United States, E. nivalis ranges from the Rocky Mountains to the West Coast excluding Arizona and Nevada. It was attributed to New Mexico by Nesom (2006) based on the Ludwig and Smith and the Reif collections cited below. Voucher specimens: Sandoval Co.: Baca Land and Cattle Company, Redondo Peak, N35.87 W 106.66, 10925 ft elev, talus slope, 1-9 Jul 1975, J. Ludwig and S. Smith 1368 (NMC): Valles Caldera National Preserve, Jemez Moun- tains: upper slopes on NE side of Redondo Peak, N35.8806 W106.5484, 10160-10360 ft elev, roadside along volca- nic outcrop, open slopes, and forested areas, 15 Jul 2002, Reif 2403 (NMC, RM); trip from 7 roads to “Redondito peak” and down the La Jara Creek to headquarters: 0.5-0.7 air mi ENE of 7 roads, N35.8923 W106,5536 to N35.8908 W 106.5480, 9730- i ft elev, spruce-tir forest and adjacent felsenmeer and g grown over 2-tracks, 18 Jul 2003, Hartman 77974 (RM Lactuca biennis a Fernald—USDA, NRCS (20006) maps this taxon for New Mexico but without voucher data and it is not included in Allred’s checklist (2005). Thus docu- mentation is provided below. Voucher specimens: Colfax Co.: Philmont Scout Ranch: Middle Fork Cimarroncito Creek, 1 mi W of Hunting Lodge, N36.4844 W105.0801, 8200 ft elev. exposed waste run along road, 7 Aug 1968, Hartman 2626 (RM). San- doval Co.: Rito de los Frijoles, N35.7793 W106.2719, Aug 1910, Robbins 8210 (RM). Bandelier National Monument: Frijoles Canyon, ca. 2 mi below Upper Crossing, N35.8044 W106.3301, 6500 ft elev, moist canyon bottom ans stream, 24 Jul 1988, B. Jacobs and E. Jacobs 4353 (BAND); 20 Jul 1941, Clark 9935 (BAND). San uel C N35.8050 W105.4407, 8000 ft elev, Aug 1902, Cockerell 11 (RM). Santa Fe National Forest and Vicinity: Sangre ae Cristo Mountains: Maestas Creek on NM hwy 391, 4.5 air mi we ot Rack ada, N35.8568 W105. ee 8500- 8680 ft elev, riparian and moist banks of creek, 15 Aug 2002, Reif 4 cos Wilderness Area, Holy Ghost Creek, 3 air mi NW of Terrero, N35.7856 W105.7048, 8100-8360 ft elev, eee e meadow in canyon bottom, 28 Jul 2002, Reif 2768 (RM) oie Canyon on NM hwy 123 branching W of the Pecos River, 6 air mi NNW of Pecos, N35.6712 W105.7412, 7760 ft elev, wet to dry meadow and riparian with canyon constricting to the W, 6 Aug 2002, Reif 3392 (RM) Rudbeckia laciniata L. var. laciniata— Variety ampla(A. Nelson) Crongq. is not distinguished [rom the typical variety in Allred (2005). The former is the common variety throughout the Rocky Mountains with var. laciniata entering the region from the Great Plains. USDA, NRCS (2006) indicates its occurrence in New Mexico but without documentation. Thus HARTMAN ET Al 1227 var. laciniata is vouchered below for the state and the reader is referred to Urbatsch and Cox (2006) for the discrimination of the two taxa. Voucher specimens: Colfax Co.: NM hwy 72, ll rd mi E i -25 (milepost 11), N36.9219 W104.2783, 8200 ft elev, roadside through wooded area, 16 Aug 1998, Hartm RM). San Miguel Co.: Santa Fe National Forest and Vicinity: Sangre de Cristo Mountains: NM hwy 263 atone Gallinas Creek, 16 air mi NW of Las Vegas, N35.7083 W105.4505, 7750 ft elev, canyon bottom roadside, riparian, and bluffs of volcanic rock in mixed conifer forest, 13 Aug 2002, Reif 3820 (RM). Santa Fe Co.: Santa Fe National Forest and Vicinity: Sangre de Cristo Mountains: Rio Frijoles Trail 154 just E of Santo Domingo de Cundiyo Land Grant, 8 air mi SE of Chimayo, N35.9380 W105.8733, 00 ft elev, riparian of river, and meadows, in canyon bottom with mixed conifer slopes, 8 Aug 2002, Reif 3421 (RM) BETULACEAE Betula glandulosa Michx.—Furlow (1997) shows this taxon extending down the Rocky Mountains to south-central Colorado. Its presence on the Valles Caldera National Pre- serve was first noted by Allen (2004) who found numerous stunted birch plants growing on Alamo Bog. This site represents a disjunction of 100 miles or more from the nearest known location in Colorado (http://cumuseum.colorado.edu). Heavy browsing by elk appears to be jeopardizing the population. Voucher specimens: Sandoval Co.: Valles Caldera National Preserve: Jemez Mountains: Alamo Canyon, 1-15 mi ae N35.9182 W106.6032, 8461ft elev, over 43 healthy but cropped shrubs to 2' tall along 1/4 mi of shore- south side, 2 Jun 2003, Hartman 77223 (RM); 8 Jun 2001, Jacobs 14 (BAND). BRASSICACEAE Cardamine cordifolia A. Gray var. incana A. Gray ex ME. Jones—This markedly pubes- cent variety is quite distinctive when compared to the essentially glabrous typical one (Rollins 1993; Welsh et al. 2003). Holmgren (2005) does not recognize var. incana but notes that the epithet is available for the pubescent form. Previously it was known from Colo- rado and Utah. Voucher specimen: Rio Arriba Co.: Santa Fe National Forest and Vicinity, Jemez Mountains, upper Cananones Creek and W slope, 11. 3 air mi S of Youngsville, N36.0304 W106.5422 to N36.0342 W106.5271, 9300-10000 ft elev, springs on trail in | Pp kbark fir forest with mature aspen, 25 Jun 2002, Reif 1197 (RM Draba grayana (Rydb.) CL. Hitchc—This taxon has been considered endemic to Colo- rado, occurring in its northern and central portions (Rollins 1993). With its discovery on Little Costilla Peak, it is now documented from New Mexico. Voucl olfax Co.: Sangre de Cristo M , Carson National Forest- Valle Vidal Unit, Little Costilla Peak, Beiec N36.8260 W105.2234 and N36.8336 W105.2228, 12,231-12,584 ft elev, alpine slopes, 16 Aug 2002, Hartman 76640 (RM). Determined by Ihsan Al-Shehbaz (MO) Lepidium ramosissimum A. Nelson var. bourgeauanum (Thell.) Rollins—In the western United States, this variety ranges from Montana south to Colorado and west, including Arizona (USDA, NRCS 2006). According to Rollins 993) “the varieties here recognized are weak at best. The consistently glabrous siliques of var. bourgeauanumare not matched by consistency in var. ramosissimum, where the siliques are mostly sparsely pubescent, but on occasion both pubescent-fruited and glabrous-fruited plants are found in the same population. The chromosome difference of 2n = 32 for var. bourgeauanum and 2n = 64 for var. ramosissimum. .. helps to bolster the treatment where the two varieties are recog- nized... .” Holmgren (2005) accepts the two varieties without discussion. We maintain the two taxa pending further study. 1228 BRIT.ORG/SIDA 22(2 Voucher specimens: Sandoval Co.: Valles Caldera National Preserve, Jemez Mountains, Jemez Falls Road to turn iwack to southwest corner of Preserve, N35.8277 W100.6170, 8000-8164 [t elev, ponde- rosa pine forest with rock outcrops and swales, 30 Jul 2001, ase 73133 (RM); just S of Sulphur Point, N35.9116 W 106.6213, rocky slopes in mixed conifers, 8981 [t elev, 17 Sep 2001, Hartman 74216 (RM); Alamo Canyon, south side over a mile stretch to near lower end, N35.9117 W106. 5002 to N35.9109 W106.5927, 8650-8960 ft elev, scat- tered at edge of wet area over lower half of boggy portion, 19 Jul 2003, Hartman 78125 with Coop (RM); Puerta de Trasquilar NW up end of Trasquilar, N35.9398 W106.4960 to N35.9423 W106.5002, 8925-9360 ft elev. south ex- posure with oak and aspen, 24 Jul 2003, Hartman 78203. Puerta de Trasquilar NW up end of Trasquilar, N35.9398 W106.4960 to N35.9423 W106.5002, 8925-9360 ft elev, south exposure with oak and aspen, 24 Jul 2003, Hart- man 78234,(RM); Valle Jaramillo, near “B” and “M” road intersection, N 35.9184 W106.4981 to N35.9133 W106.4912, 8705-8720 ft elev. grassy meadow with creek and mud areas on slope with upwelling of water, 24 Jul 2003, Hartman 78258 (RM); Valle Toledo, crossing of Pipeline Road and San Antonio Creek to 0.4 air mito NE then S$ to 0.3 air mi ae N35.9602 W106.4827 to N35.9613 W106.4736, 8610-8639 [t elev, wetlands along creek to 1] ~ south, two track and bus — Qo. slopes to N w vith barren hau and adjacent “bouncy” ground, 10 Aug 2003, Hartman 78348 (RM); on Fork ae River ae segment in Preserve ca. 1 air mi SW of El Cajete (center of caldera), N35.8277 W106.5751 to N35.8277 W106.5780, on 7990 ft elev. along river with blue spruce and white fir on slopes, willow and birch scattered on bank, Ll Aug 2003, Hartman 78425(RM); Banco Bonito, 0.8-0.9 air mi NE of southwest corner of Preserve, N35.8385 W106.6136 to N35.8367 W106.6078, 8250-8332 ft elev., roadside through ponderosa pine, 12 Aug 2003, Hartman 78449 (RM); Pajarito Mountain, ca. | air mi WNW of summit, N35.8897 W106.4172 to N35.8897 W106.4173, 8970-9050 ft elev. lower grassy slopes to scattered ponderosa pine, 21 Aug 2003, Hartman 78802 (RM); off of G Road just above F Road, ca. Lair mi N of Cerro Santa Rosa, just N of Valle San Antonio, N35.9743 W106.5050, 8560-8800 [t elev, logged ponderosa pine forest, 5 Aug 2003, Nelson 59659(RM): Jemez Mountains, from 3/4 mi E to N35.9775 W106.4827 in Rito de los Indios drainage, 8750-9000 ft elev, dis- turbed areas, 5 Aug 2003, Nelson 59700 (RM); from tributary of Rito de los Indios NW to summit of ridge, ca. 1 1/ 2-1 3/4 air mi N of north summit of Cerros de Trasquilar, N35.9921 W106.5020, 9100-9740 ft elev, open, grassy slopes with scattered ponderosa pine, 5 Aug 2003, Nelson 59757 (RM); along G Road ca. 1 3/4 mi N of F Road, ca. 2 1/2-2 3/4 air mi N of Cerro Santa Rosa, N35.9978 W106.5006, 9600-9800 . elev, meadow, 5 Aug 2003, Nelson 59831(RM); on the ridge W of Valle Grande, ca. 11/4-1 3/4 air mi N of Ranch Headquarters, N 35.8806 W106.5180, 8900-9600 ft elev, roadside through mixed conifer forest, 7 Aug 2003, Ne en 59942 (RM); ridge S and E of Valle Jaramillo, ca. 1 air mi NW of Cerro Pinon, N 35.8993 W106.5124, es ft Bie clearcut Douglas fir forest with mos stly | blue spruce white fir regeneration, LO Aug 2003, Nelson 60025 (RM): a g K Re yad at the northeast base of San Antonio Sane ain, N35.9449 W106.5925 to N35.9437 W106. 5089, 8000- 9100 ft elev, roz mae and adja- cent open spen forest, 1] Aug cose Nelson ep0ee (RM); along the northeast and north side of San Antonio Mountain to the west boundary, N35.94 to N35.9463 W106.6204, 900¢ ee along old logging road, mixed conifer/aspen forest, a ae 1 Aug 2003, Nelson 60118 (RM); on the east slope of Cerros del Abrigo above the Abrigo aa : 0. 3mi on either side, N35.9301 W106.4726, ee ft Bley, i 0314(R old logging roa spen forest, and open areas,13 Aug 2003,) on south side of Cerros del Abies along the Ahaee Trail for 0.3 mi on either side, N35.9232 W106.4768, 9400 ft elev along old logging road, roadcut, and adjacent southwestern white pine/white fir/Douglas fir forest. 13 Aug 2003. Nelson 60243 (RM); Santa Fe Co.: Santa Fe National Forest and Vicinity, Sangre de Cristo Mountains, Rio Frijoles Trail 154 just E of Santo ati de Cundiyo Land Grant, c air mi SE . Chimayo, N35.9380 W105.8733, 7000 ft elev, riparian of river, and meadows, in canyon bottom v r slopes, 8 Aug 2002, Reif 3448 (RM). Rorippa sylvestris (L.) Besser—Introduced throughout much of the United States, this mustard is said to occur in New Mexico: “reported questionably by Kartesz and Meacham (1999): awaits verification” Allred (2005). Voucher specimen: San Miguel Co.: Santa Fe National Forest and Vicinity, Sangre de Cristo Mountains, Anton Chico Grant, 5 air mi ENE of Villanueva, along Pecos River, 35.2365 105.2558, 5500 ft elev, river bottom, mostly juniper with Russian olive and narrowleat cottonwood, 6 Aug 2004, Reif 10067 (RM). CYPERACEAE Carex deweyana Schwein. var. deweyana—Naczi (2002) lists but does not map this taxon flor New Mexico, although it is mapped for the state in USDA, NRCS (2006). Interesting]y, it isnot included in Allred (2005). Regardless, the first two sources do not contain voucher data. Consequently, our records are provided below. HARTMAN ET Al 1229 Voucher specimens: Los Alamos Co.: Santa Fe National Forest and Vicinity, Jemez Mountains, lower Guaje yon and Cabra Canyon, 3 air mi N of Los Alamos, N35.9310 W106.3432 to N35.9064 W106.2862, 7000- a ft elev, riparian drainage through mixed conifer forest, into a burn past the canyon, and then a ponderosa pine forest before joining the road, 28 Jun 2002, Reif 1475 (RM). Santa Fe Co.: Santa Fe National Forest and Vicinity, Sangre de Cristo Mountains, Pecos Wilderness, Rio Merlo tal 155 acincent to Rio Medio, 6 air mi due W of Truchas Peak, N35.9630 W105.7520, 8410 ft elev, rip or standing water, 23 Jun 2003, Reif 5806 (RM). > of lower canyon slopes, banks of river (=) Carex rosea Schkuhr ex Willd. According to Ball (2002), this taxon ranges from Wyo- ming, the Great Plains and Texas to the east coast of North America. Voucher specimens: Sandoval Co.: Santa Fe National Forest and Vicinity, Jemez Mountains, Cochiti Canyon, 4.3 air mi SSE of Rabbit Mountain, N35.7794 W106.4366 to N35.7645 W106.4266, 7200-8280 ft elev, canyon bottom and riparian, 23 Jun 2003, Reif 1116 (RM). San Miguel Co.: Santa Fe National Forest and Sana Sangre de Cristo ee. Sebadilla Creek S of Flying Cloud Ranch, 3 air mi SSW of Barillas Peak, u ek 1/2 air mi, N35.5320 7740-7880 ft elev, narrow riparian corridor in canyon bottom, canyon al by mixed are forest, sean bluffs and hee constrictions, 9 Jun 2003, Reif 5070 (RM); El Porvenir Canyon on Trail 247, mi W of San Ignacio, N35.770 W105.448, 8200-8450 ft elev, canyon bottom of mostly spruce, and moist ie slope of mixed conifers along am granite bluffs constricting canyon, 6 Jul 2003, Reif 6183 (RM). EQUISETACEAE Equisetum Xnelsonii (A.A. Eaton) J.H. Schaffn.—This plant represents a spontaneous hybrid between E. laevigatum and E. variegatum. The former putative parent is frequent in the Jemez Mountains; E. variegatum is not documented from New Mexico but has been collected as far south as Montezuma County, Colorado, some 130 mi to the northwest. In the Rocky Mountains, the hybrid is known from Montana, Wyoming, Utah, and Colo- rado (USDA, NRCS 2006) Voucher specimen: Sandoval Co.: Bandelier National Monument, White Rock Canyon at mouth of Alamo Can- yon along banks of Rio Grande, N35.7146 W106.2907, 5300 ft elev, 23 Aug 1975, Foxx s.n.(B ) JUGLANDACEAE Juglans nigra L—Black walnut, a native of the eastern United States, has been cultivated in areas of the western United States. As described below, it is naturalized in NM. J Voucher specimens: Sandoval Co.: Bandelier National Monument, vicinity of P 5 in Canyon de los Frijoles, along Rito de los Frijoles near and below Tyuonyi Ruin, N35.7839 W106.2772, 6, 100 ft elev. escaped third peneration seceine meattcree in colconnond bosqt ue alae stream, os Sep ee Jacobs 5083 (BAND), second gen- historic Frey Abbott orchard, 4 May 2003, Jacobs 5084 (BAND), OLEACEAE Syringa vulgaris |.—This introduced lilac occurs over much of eastern United States and scattered states in the West. The latter includes Kansas, Colorado, and Utah (USDA, NRCS 2006). This collection appears to be an adventive representing the first New Mexico record. Voucher ae ae Santa Fe Co.: Santa Fe National Forest and Vicinity, Sangre de Cristo Mountains, Tesuque n Winsor Trail 254, 3 air mi SE of Tesuque, N35.745 W105.888 SW to N35.743 W105.896, 7320-7370 ft ae narrow canyon bottom and riparian with mature ponderosa pine, horsetail, cotton wood, alder, and willow, granitic soils, 12 Jun 2003, Reif 5252 (RM). POACEAE Achnatherum nelsonii Scribn. var. nelsonii— According to Barkworth et al. (1979), this grass ranges from southern Montana and central Idaho south through Nevada and Utah. In Utah it has been documented in the southeast corner, immediately adjacent to New Mexico. In addition, USDA, NRCS (2006) maps it for Colorado and Arizona. Verified by Mary E. Barkworth. 1230 BRIT.ORG/SIDA 22(2) Voucher specimens: Rio Arriba Co.: Santa Fe National Forest and Vicinity, Jemez Mountains, vicinity of Presa ing on southern edge of Chama River Canyon Wilderness, 8 air mi NE of Gallina, N36.3120 W106.7340, 8340 elev, a forest of mature ponderosa pine, with thickets of Gambels oak and ao open areas below, 19 Jun 2003, - if 5617 (RM). Sandoval Co.: Sierra Nacimiento, Joaquin Canyon, ca. 2 3/4-3 1/4 air mi W of Rio Guadalupe, N35.79 x W106.8 oh coe 8050 ft elev, open area along logging road and adjacent mixed conifer forest, 20 Jul 2004, b 621 Cynosurus echinatus L. This grass has been introduced in much of eastern United States, the West Coast, and Montana (USDA, NRCS 2006). This represents the first report of its establishment in New Mexico. Voucher specimen: Sandoval Co.: Bandelier National Monument, Capulin Canyon, ca. 0.25 mi S of Base Camp, near where trail to Stone Lions joins canyon trail, N35.7541 W106.3262, 6100 ft elev, dense stand along stream, 13 Jun 1988, B. Jacobs 4167 (BAND). Piptatherum pungens (Torr. ex Spreng.) Dorn—This species ranges across western Canada and south in the Black Hills of Wyoming and South Dakota to the central cordillera of Colorado (Kartesz and Meacham 1999, USDA, NRCS 2006). In Colorado it is documented from Douglas, Grand, and Saguache cos. (http://cumuseum.colorado.edu). The distance from the nearest locality in Saguache Co. is about 150 miles. a Voucher specimen: Sandoval Co.: Valles Caldera National Preserve, Jemez Mountains, Cerros del Trasquilar, main portion, N35,9648 W106.4918, 8537-8740 ft elev, roadside and coniferous forest, 26 Jun 2001, Hartman 71472 (RM). RANUNCULACEAE Ranunculus alismifolius Geyer ex Benth. var. montanus S. Watson—According to Whittemore (1997), this taxon is restricted to southwestern Colorado, Idaho, Nevada, Utah, and Wyoming. It is one of six varieties recognized by this author, all of which occur from the Rocky Mountains to the west coast of North America. These specimens document its occurrence in New Mexico. Voucher specimens: Rio — Co.: San Pedro Mountains, N 36.0667 W106.8433, 10200 ft elev. seep under aspen and spruce, 4 Jul 2001 Dorn 8829 (RM); Santa Fe National Forest and Vicinity, Jemez Mountains, Forest Road 70 and Forest Road 50, Px = as Trail, NW into San Pedro Parks Wilderness to Vacas Trail, Forest Road 51, N36.0421 W106.7924 to N36.0662 W106.8121, 9300-10135 ft elev, ponderosa pine and some spruce, 20 Jun 2002, Hartman 75015 (RM); Rito de Abiquiu NE of Cienega del Oso, 2.5 air mi SSE of Polvadera Peak, N36.0290 W106.3900 to N36.0360 W106.3820, 9400-10430 ft elev. open riparian bottom and steep shaded slope of corkbark fir/blue spruce with an understory of Paxistima, Trauttvetaria, and Vaccinium, : Jul 2003, Reif 6567 (RM); NM hwy 70, South San Pedro Parks Wilderness trailhead from parking lot N to San Gregorio Reser voir, 36.0273 N106.8469 to W 36.0407 106.8460, 9200-9400 ft elev, open wet meadows in mixed coniferous forest with aspen, 2 Jun 2002, Reif 45 (RM Ranunculus repens L.—This taxon is naturalized throughout much of the eastern and western portions of the United States (Whittemore 1997). The two collections cited be- low also appear to represent self-maintaining populations. Voucher specimens: Deng al Co.: Santa Fe National Forest and Vicinity, Jemez Mountains, 8 air mi SW of La Cueva, Forest Road 376, 2 air mi N of Porter, N35.8442 W106.7810, 7490 ft elev, stream and adjacent meadow, 23 Jun 2002, Hartman ety (RM). San Miguel Co.: Sangre de Cristo Mountains, Forest Road 60 along Sapello River, 2 air mi SW of Rociada. N35.8129 W105.4480, 8000 [t elev, canyon bottom peat area at duius a5 trailhead, including riparian of river and south-facing bank (below road) of mature aspet Aug 2003, Reif 8741(RM) ROSACEAE Geum triflorum Pursh var. trifloruam—Holmgren (1997) states that var. triflorum ranges south in the Rocky Mountains, where it more or less straddles the continental divide from Montana to northern New Mexico and includes some populations as far west as HARTMAN ET AL., VASCULAR PLANT RECORDS FOR NEW MEXICO 1231 northeastern Arizona. The following specimens document its occurrence in New Mexico. Variety ciliatum, the more western version of the species is listed for New Mexico by Al- lred (2005). Voucher specimens: Colfax Co.: Philmont Scout Ranch, Beaubien, N36.4237 W105.1033, 9400 - oe common on dry, grassy slopes, 27 Jun 1966, Hartman 117 (RM); 6 Jun 1968, Hartman 1819 (RM); Rio A on pauona! EOL Canyon Bancosca He air mi W of pace 36.9008 107.2237 to 36.9143 107.2690, Sue: me 7? se 1-juniper, 28 May 1987, Hartman 22504 with D’Alcamo et al. (RM). Sandoval Ce Jemez Mountains, Hack Fore ae River, ca. 0.5 mi above Las Conchas Campground, N35.8200 W106.5212, 8500 ft elev, 18 Jun 1986, Jacobs 006 with Jacobs (RM). Potentilla fissa Nutt.—This species is common throughout the Rocky Mountains and sev- eral adjacent states; likewise it is included for New Mexico (USDA, NRCA 2006). Allred (2005) indicates that it is reported for New Mexico but awaits verification. Voucher specimen: San Miguel Co.: Santa Fe National Forest and Vicinity, Sangre de Cristo Mountains, Hermit Peak, N35.7447 W105.4139, 10000-10212 ft elev, open grassy and rocky areas and rim rock, 18 Aug 2002, Hart- man 76734 (RM) Prunus persica (L.) Batsch—The introduced peach is known from much of the eastern United States and several of the states to the west, but it is not reported from Montana south to New Mexico (USDA, NRCS 2006). In both localities here listed, it was found grow- ing in sites devoid of human disturbance and thus appears to be adventive. Voucher specimens: Sandoval Co.: Valles Caldera National Preserve, Jemez Falls Road to turn south, N35.8277 W106.6170, 8000-8164 ft elev, ponderosa pine forest with rock outcrops and swales, 30 Jul 2001, Hartman 73177 (RM); Santa Fe National Forest and Vicinity, Jemez Mountains, Battleship Rock Trail 137 at McCauley Warm Springs, N35.8222 W106.6281, 7300-7400 ft elev. warm springs and riparian drainage in os pine forest, 1 Sep 2003, Reif 9380 (RM). SAXIFRAGACEAE Heuchera hallii A. Gray—This taxon was considered by Rosendahl et al. 1936) as distinc- tive. It has been considered to be endemic to Colorado; in the vicinity of Pikes Peak (Chaffee, Custer, Douglas, El Paso, Jefferson, and Park cos.). (http://cumuseum. colorado.edu). These reports represent a range extension of about 90 miles to the south. ucher — Colfax Co.: Philmont ee Ranch, Rayado Canyon, 0.5 mi W of Old Abreu Lodge, N36.3765 ae 0331, 7500 ft elev, sandy stream bank and rock Loe 15 Jul 1968, Hartman 2385 (RM); base of Crater Peak alon ngR yado River, ca. 2 air mi SE of one Camp, N36.3746 W105.0380, 8200-8800 ft elev, slope above river 10 Jul 1991, a 92a with Hartman (RM); 4 Aug 1991, a bry 92b(RM). SCROPHULARIACEAE Penstemon glaber Pursh var. alpinus (Torr.) A. Gray—This variety is the second for the species to be documented in New Mexico. Its previous known range was southeastern Wyoming to south-central Colorado (Great Plains Flora Association 1986, see key to va- rieties). Variety brandegeei (TS. Porter & Rydb.) C. Freeman had been the only taxon of this species documented for New Mexico. Voucher speci :Los Alamos Co.: Santa Fe National Forest and Vicinity, Jemez Mountains, Mitchell Trail 69, from Los Alamos, NW 2.1 air mi to Guaje Reservoir, N35.9310 W106.3432, 7500-8900 ft elev, incised gravel stre- ambed through a burn, climbing canyon walls of tuft and rhyolite, toa mixed conifer forest with patchy burns, 28 Jun 2002, Reif 1379 with Hartman and Jacobs (RM); Taos Co.: Carson National Forest and Vicinity, Sangre de Cristo Range, Valle Vidal Unit, ridgeline above corral from Forest Road 1950 toward Little Castilla Peak, N36.7843 W105.2330, 10,000-12,584 ft - spruce fir and bristlecone pine forest with meadow openings, rocky ridgeline, 30 Jul 2005, Larson 3373 (RM 1232 BRIT.ORG/SIDA 22(2) ACKNOWLEDGMENTS The authors thank the following individuals for verification of specimens: Mary Barkworth, Utah State University; Ihsan A. Al-Shehbaz, Missouri Botanical Garden. Also thanked are Kelly W. Allred, Guy L. Nesom, and Richard W. Spellenberg for their helpful comments on the manuscripts. We thank Jill Larson, M.S. student (RM), for allowing us to publish her record of Penstemon glaber var. alpinus. REFERENCES Aten, C.D. 2004. Ecological patterns and environmental change in the Bandelier landscape. In: T.A. Kohler, ed. Village Formation on the Pajarito Plateau, New Mexico: Archaeology of Bandelier Na- tional Monument. Univ. of New Mexico Press, Albuquerque. Pp. 19-68. Allred, K. 2005 (September). A working index of New Mexico vascular plant names. http:// web.nmsu.edu/~kallred/herbweb/ Batt, PW. 2002. Carex sect. Phaestoglochin. In: Flora of North America Editorial jeu iaa eds. 2002. idae (in part): Cyperaceae. Flora of North America north of Mexico.Vol.23.Mag Oxford Univ. Press, New York, New York. Barkwortn, M.E.,J.McNeit, and J. Maze. 1979. A taxonomic study of Stipa nelsonii (Poaceae) with a key distinguishing it from related taxa in western North America. Canad. J. Bot. 57:2539-2553. Cronauist, A. 1994.In:A.Cronquist, A.H. Holmgren, N.H. Holmgren, J.L.Reveal, and P.K. Holmgren. Inter- mountain Flora. Vascular plants of the Intermountain West, U.S.A. Vol. 5, Asterales. New York Bo- tanical Garden, Bronx, New York. Furtow, J.J. 1997. Betulaceae. In:Flora of North America Editorial Committee, eds. 1997. Flora of North America north of Mexico. Vol. 3. Magnoliophyta: Magnoliidae and Hamamelidae. Oxford Univ. Press, New York, New York. Great PLAINS Flora Association. 1986. Flora of the Great Plains. Univ. Press of Kansas, Lawrence, Kansas Hartman, R.L. 1973. Notes. New plant records for New Mexico. SouthW. Nat. 18:241-242. Hartman, R.L. 2003. Preliminary checklist of the vascular plants of Valles Caldera National Preserve, New Mexico. Distributed by author. Hartman, R.L. and B.E. Newson. 2005. General floristic survey of the Valles Caldera National Preserve, New Mexico: report on field seasons 2001, 2002, 2003, 2004.Report to the U.S. Geological Survey, Biological Resources Division. Houmaren, N.H. 1997.In: A. Cronquist, NH. Holmgren,and PK. Holmgren. Intermountain flora. Vascular plants of the Intermountain West, U.S.A. Vol. 3, Part A, Subclass Rosidae (except Fabales). New York Botanical Garden, Bronx, New York. Houmoren, N.H. 2005. In:N.H. Holmgren, PK. Holmgren, and A.Cronquist. Intermountain Flora. Vascular plants of the Intermountain West, U.S.A. Vol. 2, Part B, Subclass Dilleniidae. New York Botanical Garden, Bronx, New York. Jacoss, B.F. 1989. Flora of Bandelier National Monument. Final report: National Park Service. Kartesz, J.T. and C.A. Meacoam. 1999. Synthesis of the North American Flora, CD format, Version 1.0. North Carolina Botanical Garden, Chapel Hill, North Carolina. Naczi, R.F.C. 2002. Carex sect. Deweyanae. In: Flora of North America Editorial Committee, eds. 2002. Flora of North America north of Mexico.Vol.23.Magnoliophyta:C linidae (in part): Cyperaceae. Oxford Univ. Press, New York, New York. Nesom, G.L. 2006. Erigeron. In: Flora of North America Editorial Committee, eds. 2006. Vol. 20. Magnoliophyta: Asteridae, part 7: Asteraceae, part 2. Oxford Univ. Press, New York, New York. O'Kane, S.L., Jr, D-H. Witken, and R.L. Hartman. 1988. Noteworthy collections: Colorado. Madrono 35: 72-74. HARTMAN ET Al 1233 Reir, B.P. 2006. A vascular plant inventory of Santa Fe National Forest (including the Valles Caldera National Preserve) and vicinity, North-Central New Mexico. M.S.Thesis, Univ.of Wyoming, Laramie, Wyoming. Reir, B.and R.L. Hartuan. 2005.A floristics inventory of the vascular plants of Santa Fe National Forest and the surrounding hydrographic region, New Mexico. Final report: Santa Fe National Forest. Rottins, R.C. 1993. The Cruciferae of Continental North America: Systematics of the mustard family from the Arctic to Panama. Stanford Univ. Press, Stanford, California. RoseNDAHL, C.O., F.K. Butters, and O. Lakria. 1936. A monograph on the genus Heuchera. Minnesota Studies in Plant Science, Vol. Il. Univ.of Minnesota Press, Minneapolis THeosatp, W.L., C.C. Tsenc, and M.E. MatHias. 1963. A revision of Aletes and Neoparrya (Umbelliferae). Brittonia 16:296-315. Ureatscu, L.E. and PB. Cox. 2006. In: Flora of North America Editorial Committee, eds. 2006. Vol. 21. Magnoliophyta: Asteridae, part 8: Asteraceae, part 3. Oxford Univ. Press, New York, New York. USDA, NRCS. 2006.The PLANTS Database, 6 March 2006 (http://plants.usda.gov). National Plant Data Center, Baton Rouge, Louisiana 70874-4490. Weser, W.A. 1958. Rediscovery of the genus Neoparrya Mathias (Umbelliferae). Rhodora 60:265-271. Wetsu, S.L., N.D. Atwoopo, S. GoopricH, and L.C. Hicains (eds.). 2003. A Utah flora. Third edition, revised. Brigham Young Univ., Provo, Utah. Wuittemore, A.T. 1997. Ranunculus.\|n: Flora of North America Editorial Committee, eds. Flora of North America north of Mexico. Vol. 3. Magnoliophyta: Magnoliidae and Hamamelidae. Oxford Univ. Press, New York, New York. 1234 BRIT.ORG/SIDA 22(2 Books RECEIVED Joun F Leste and Brett A. SUMMFRELL, Photographs by SuZANnr BuLLock. 2000. The Fusarium Laboratory Manual. (ISBN 0-8138-1919-9, pbk.). Blackwell Publishing. 2121 State Ave., Ames, IA 50014-8300, U.S.A. (Orders: 515-292-0140, 515-292-3348 fax 1-800-862- 6657, wwwblackwellprofessional.com, orders@ames.blackwellpublishing.com,). $124.99, 388 pp., numerous b/w illustrations, 81/2" x 11" From the Publisher—‘For the first time in over 20 years, a comprehensive collection of photographs and descrip- tions of species in the fungal genus Fusarium is available. This laboratory manual provides an overview of the biology of Fusarium and the techniques involved in the isolation, identification and characterization of indi- rich they occur. lt is the first time that genetic, morphological and mo- vidual species and the populations in w pigs oy oaches have been incorporated into a volume devoted to Fusarium identification. The authors in- f cies, both new and old, and provide protocols for genetic, morphological and molecular ide ee techniques. “The Fusarium Laboratory Manual also inc thinking that has begun to inform the understanding of agriculturally important fungal pathogens. In addition to practical “how-to” protocols it also provides guidance in formulating questions and obtaining answers about udes some of the evolutionary biology and population genetics this very important group of fungi. The need for as many different techniques as possinlet to be saa in the ide n- 8 ) q tification and characterization process has never been greater. The than those in the genus Fusarium. This volume presents an eee to the seneris usarium, the toxins retest fungi produce and tl the diseases tl they can cause.” J Kris M. Havstab, LaurA EF HueNneke, and WILLIAM H. SCHLESINGER. (eds.). 2006. Structure and Function of a Chihuahuan Desert Ecosystem. The Jornada Basin Long-Term Eco- logical Research Site. (SBN 978-0-19-511776-9, hbk.). Oxford University Press, 2001 Evans Road, Cary, NC 27513, U.S.A. (Orders: 800-451-7556; http://www.oup.com). $74.50, 465 pp., b/w halftones, maps and line illustrations, 6 1/8" x 9 1/4". From the Publisher—“The Jornada Basin LTER is located in the Chihuahan Desert, the largest in North America. This region of south central New Mexico hasa history of nearly 100 years as the basis for scientific research. This work gives a thorough, encompassing review of the tremendous array of observations resulting from experi- ments conducted in this ecosystem. Beginning with thorough descriptions of the most salient features of the region, the book then reviews a wide range of archived and active data sets on a diversity of biotic and abiotic features. It next presents a syntheses of important topics including livestock grazing and remediation efforts. A concluding chapter provides a synthesis of the principles that have emerged from this body of work, and how these relate to the broader fields of ecology and natural resource management. It concludes with recommenda- tions for future research directions. The insightful views expressed in this volume should guide management of arid landscapes globally. This is the sixth volume in the Long Term Ecological Network Series. GALIUM TRICORNUTUM (RUBIACEAE) AND PARENTUCELLIA VISCOSA (SCROPHULARIACEAE) NEW TO OKLAHOMA Amy K.Buthod Bruce W. Hoagland Oklahoma Biological Survey Oklahoma Biological Survey and ie t of Geography University of Oklahoma University of Okla Norman, Oklahoma 73019, U.S.A. Norman, Oklahoma at és aon amybuthod@ou.edu ABSTRACT This paper reports the occurrence of two species previously unknown to the flora of Oklahoma. Galium tricornutum, a Eurasian introduction, was discovered in a cement drainage ditch in Hughes County in central Oklahoma. Parentucellia viscosa, an introduction from Europe, was discovered in disturbed grassland in McCurtain County in extreme southeastern Oklahoma. RESUMEN Este articulo cita la ocurrencia de dos — Le tnens de Seon Oclads de a me de ee oa tricornutum, una introd iatica, fue descubierta en una zanja d ado de Hughes en el centro de Oklahoma. nner ie viscosa (L.) Caruel, una introduccién de Europa, fue descubierta en un prado al | >| condado de McCurtain en el extremo sudeste de Oklahom | Galium tricornutum Dandy, in the Rubiaceae, is an int d annual herb native to Eurasia that is typically found in waste places and as a weed of cereal crops. It has been reported previously in Arksansas, California, Georgia, New York, Oregon, Pennsylvania, Rhode Is- land, South Carolina, Washington, Wisconsin, and Oregon (USDA-NRCS 2006). One plant was encountered growing in a crack in a cement drainage ditch in the Holdenville city park in central Oklahoma. The combination of recurved pedicels and a tuberculate fruit distinguish G. tricornutum from other ey Galium species in Oklahoma. Voucher specimen for Galium tricornutum: OKLAH Hughes Co.: Holdenville, at Stroup City Park, T7N R9E secl8, ll May 2006, Buthod, Hoagland, and pein a 7138(OKL »: Parentucellia viscosa (L.) Caruel, in the Scrc pl 1ulariaceae. is an introduced annual herb na- tive to Europe typically found on moist, grassy places and frequently in old meadows. It has been reported in Arkansas, California, Hawaii, Louisiana, Oregon, Texas, and Washing- ton (USDA-NRCS 2006). Only one plant was located in a disturbed tallgrass prairie site on limestone in southeastern Oklahoma. Associated species included Centaurea americana Nutt., Echinacea atrorubens, Sorghum halepense (L.) Pers., and Rudbeckia hirta L. Voucher specimen for Parentucellia viscosa: OKLAHOMA. McCurtain Co.: 1.9 mi NWof Idabel, T7S R23E SE/4 sec 25, 25 May 2006, Buthod, Hoagland, and Arbour AB-7137(OKL). ACKNOWLEDGMENTS We appreciate the field assistance of David Arbour and Priscilla Crawford and the com- ments of Susan Barber and Ron Tyrl. REFERENCE USDA-NRCS 2006. The PLANTS Database [online]. Available: http://plants.usda.gov. National Plant Data Center, Baton Rouge, LA 70874-4490 USA. (Accessed on 13 June 2006). SIDA 22(2): 1235. 2006 1236 BRIT.ORG/SIDA 22(2) ANNOUNCEMENTS 2006 Delzie Demaree Travel Award Recipients The 18th Annual Delzie Demaree Travel Award was presented at the 52nd Annual Sys- tematics Symposium (13-14 Oct 2006) at the Missouri Botanical Garden. Three students were presented the Travel Award: Alejandra Vasco, New York Botanical Garden and City University; Diana Hurlbut, Central Michigan University; and Michael Sundue, New York Botanical Garden and City University. The 2006 Travel Awards were underwritten by 1) Delzie Demaree Travel Award En- dowment, 2) Members of the Delzie Demaree Travel Award Committee. and 3) John Clay- ton Chapter of the Virginia Native Plant Society. Anyone interested in making a contribution to Delzie Demaree Endowment Fund, which supports the travel award, may make contributions by VISA or MasterCard or by a check, payable to Botanical Research Institute of Texas, to Barney Lipscomb, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. 1-817-332-7432; Email: barney@brit.org. Thank you. The 2007 Applications for the Delzie Demaree Travel Award Applications for the 2007 Delzie Demaree Travel Award should include a letter from the applicant telling how symposium attendance will benefit his/her graduate work and letter of recommendation sent by the major professor. Please send letters of application to: Dr. Donna M.E. Ware, P.O. Box 8795, Herbarium, Biology Department, The College of William and Mary, Williamsburg, VA 23185-8795, U.S.A. 1-757-221-2799; Email: ddmware@wm.edu. The period for receiving applications will end three weeks prior to the date of the symposium if a sufficient number of applications are in hand at that time. Anyone wishing to apply after that date should inquire whether applications are still being accepted before applying. The Systematics Symposium dates for 2007 are 12-13 October 2007. The Delzie Demaree Travel Award was established in 1988 and honors Delzie Demaree who attended 35 out of a possible 36 symposia before he died in 1987. Delzie Demaree was a frontier botanist, explorer, discoverer, and teacher. His teaching career asa botanist began in Arkansas at Hendrix College in 1922. He also taught botany at the University of Arkansas, Navajo Indian School, Yale School of Forestry, Arkansas AS&tM, and Arkansas State University at Jonesboro where he retired as professor emeritus in 1953. One of the things he enjoyed most as a botanist was assisting students with their field botany re- search. SIDA 22(2): 1236. 2006 CAPRARIA MEXICANA (SCROPHULARIACEAE) IN CAMERON COUNTY, TEXAS: REDISCOVERED IN THE UNITED STATES Alfred Richardson Ken King Department of Biological Sciences 1008 W. 5" Street The University of Texas at Brownsville Weslaco, Texas 78596, U.S.A. 80 Fort Brown Brownsville, Texas 78520, U.S.A. ABSTRACT Two new sites are reported for Capraria mexicana (Scrophulariaceae) in the United States. RESUMEN Se reportan dos nuevos sitios para Capraria mexicana (Scrophulariaceae) en los Estados Unidos. Capraria mexicana (Mexican Capraria) is an attractive shrub reaching 2 m in height (Fig. 1). The leaves are alternate, lanceolate, with serrate margins. The flowers emerge from the leaf axils. Corollas are regular, white, basally sympetalous, with lobes about 9 mm long. Everitt et al. (2002) provide description and a picture. Most reports of C. mexicana are from Tamaulipas, Mexico, with records also from Jalisco, Guerrero, and Tabasco. The southernmost collections are from Belize (Ideker 1996). apraria mexicana was first discovered in the United States by Joe Ideker in 1993. He reported at least 66 mature plants plus seedlings in three small stands located at Fal- con State Park near the International Falcon Reservoir in Starr County, Texas. (Ideker 1996). The plants were in a precarious location where they could be flooded with a rise in the reservoir, and they were located within a proposed drilling pad site. After communi- cation with Joe Ideker in 1993, this species was listed as Endangered in Texas by the Texas Organization for Endangered Species (TOES). It is not considered Endangered in Mexico. Our attempt to relocate the population on 25 March 2006 was unsuccessful. Benito Tre- vino, well known authority on plants of Starr County, reported having seen no Capraria plants when he had visited the site. In September of 2005, while at the Laguna Atascosa National Wildlife Refuge, Cam- eron County, Texas, we were showna single plant of Capraria mexicana by Ellie Thomp- son, volunteer manager of the butterfly garden there. She had watched it grow from a volunteer seedling to maturity. As no nearby members of the species were known, it was assumed to be an inadvertent introduction from Starr County nursery stock. In March of 2006, we observed a large population of Capraria mexicana in Brownsville, Cameron County, Texas, along Highway 511, 0.4 miles north of Old Port Isa- bel Road. The population continued for at least 0.2 miles. The population is estimated to comprise several hundred individuals ranging from 0.3 m to 1.2 m tall The plants were seen growing on both sides of the highway, on saline clay, with Opuntia engelmannit var. lindheimeri, Borrichia frutescens, Prosopis glandulosa var. glandulosa, P reptans, Maytenus phyllanthoides, Fleischmannia incarnata, Dichanthium sp, and Bothriochloa sp. With the presumed extirpation of the population at Falcon State Park, the Brownsville SIDA 22(2): 1237— 1238. 2006 1238 BRIT.ORG/SIDA 22(2) site is the only confirmed population of Capraria mexicana in the United States. Further surveys might reveal a population at Laguna Atascosa National Wildlife Refuge. The plants can be easily overlooked if not in flower, since the leaves greatly resemble those of Baccharis. Voucher specimen: TEXAS. Cameron Co.: Brownsville, Hwy 511, 0.4 mi N of Old Port Isabel Road, 16 Mar 2006, Richardson and King 3284 (BRIT, TEX). ACKNOWLEDGMENTS We thank Mike Heep for identification of the grasses, and an anonymous reviewer who provided useful and constructive comments. REFERENCES Everitt, J., D.L. Drawe, and R.I. Lonarp. 2002. Trees, shrubs & cacti of South Texas. Second Edition. Texas Tech University Press, Lubbock. IbeKer, J. 1996, Capraria mexicana (Scrophulariaceae), an endangered addition to the United States Flora.. Sida 17:523-526 TEXAS ORGANIZATION FOR ENDANGERED Species (TOES). 1993.Endangered, threatened and watch lists of Texas plants. TOES Publ. 9, a rev (and 1993 addenda). RANGE EXPANSION OF HEXALECTRIS GRANDIFLORA (ORCHIDACEAE) IN TEXAS el Brown-Marsden Anne B. Collins iolo partment a of Land, Air and Water Resources Dee of Dallas ls and Biogeachemistry 1845 F. Northgate Drive Fes of California at Davis Irving, Texas 75062, U.S.A. 09 Veihmeyer Hall Davis, California 95616, U.S.A. ABSTRACT Plants of Hexale liflora A. Richard & Galeotti (Orchidaceae) collected in Dallas County, Texas in 2005 and 2006 ey fie oe of this species northeast by more than 700 km and indicates greater diversity for the cas enus He ke in ee Coun than previously observed. These observations suggest either the absence of t data on species’ range in intervening counties, or the loss of populations between the north central Texas and western Tees locations, and argue for continued study of Hexalectris throughout the state. ABSTRACT Las plantas de mealecin isgra ayn Bs Richard & Sai (Orchidaceae) colectadas en el condado de Dallas, ana en 2) v2 | noreste mas de 700 km e indican mayor diversidad lectri | lado de Dallas de la q Estas observaciones l iaded fici del Ide est ie en los eondndcs afectados, o la pérdida de pencones entre el norte-centro de Texas y las local idades del oeste de Texas, y se aportan argumentos para continuar el estudio de Hexalectris en todo el estado. The myco-heterotrophic orchid genus Hexalectris Rafinesque is comprised of seven spe- cies—five that are found in the United States and two that are found only in Mexico (Gold- man et al. 2002). Four of the species found in Texas are H. grandiflora A. Richard & Galeotti, H. warnockii Ames & Correll, H. nitida L.O. Williams, and H. revoluta var. revoluta Correll (H. revoluta var. colemanii Catling) has recently been described from Arizona (Catling 2004). The fifth species found in Texas, H. spicata (Walter) Barnhart, includes two varieties: H. spicata var. spicata and H. spicata var. arizonica (S. Watson) Catling & Engel . Hexalectris spicata var. spicata. ranges widely from Arizona through the south- eastern US and north to Maryland, and H. spicata var. arizonica, revalidated in 1993, is found only in parts of Texas and Arizona (Catling & Engel 1993). Dallas County has recorded populations of H. spicata var. spicata, H. spicata var. arizonica, H. warnockii,and H. nitida (Diggs et al. 1999; Hatch et al. 1990). Dallas County populations of Hexalectris are found in areas of the White Rock Escarpment, a geologi- cal region of lower Cretaceous limestone that extends northeast from Dallas to the Okla- homa border, and southwest past Waco and Austin into the Edwards Plateau (Dallas Department of Urban Planning 1977). The escarpment geological zone, known for its dis- tinctive flora, is most evident in the southwestern part of the county, and it is in this area where most of the Dallas County plants of Hexalectris have been found (Collins et al. 2005). Studies in Dallas County examined habitat structure and soil association for Hexalectris. They indicate an association with oak litter (generally Quercus buckleyit or Q. shumardii) mixed with Juniperus spp. and distributed primarily on Dallas soil series Eddy-Brackett complex (8-20 % slope) (Collins et al. 2005). Ongoing work attempts to SIDA 22(2): 1239- 1244. 2006 1240 BRIT.ORG/SIDA 22 = 2) confirm this soil-orchid relationship through targeted searches in preserved habitat ar- eas with Eddy-Brackett soils that are well-drained and have sufficient slope. Intensive studies of Hexalectris distribution in 2005 and 2006 led to the discovery of H. grandiflora, a species not previously found in Dallas County. On 21 June 2005, dur- ing Hexalectris studies on the focal soil series,a cluster of H. grandiflora was identified at Camp Wisdom, a 344-acre tract of escarpment habitat in southwest Dallas County owned and operated by the Circle 10 Council of Boy Scouts of America as an educational and See site. One cluster of six aboveground stems was found during a comprehensive search of the location (Fig. Ll). The stems were erect, leafless, and slender, pinkish red at base, and becoming lighter toward the inflorescence. The dorsal sepals were oblong to elliptic; white to pale pink at base, and light pink distally. The lateral sepals were falcate, white to pale pink at base and light pink distally. The petals were recurved, white to pale pink at base and light pink distally. The column was pink with a white tip and the lip (labellum) was lobed and lamellate. The lateral lobes were deep pink, and the middle lobe was pink at the margin and white at the center. Stems were found in an area domi- nated by Quercus spp. and Juniperus spp., although the plants were found directly below arusty blackhaw viburnum (Viburnum rufidulum). Fifteen blooming stems were noted on 9 June 2006 at the same location, and were observed to be more robust than when first sighted in 2005. A second population of Hexalectris grandiflora was found at Cedar Ridge Preserve in 2006 approximately 5.5 km southwest of the 2005 sighting. A cluster of 9 stems was observed on 13 May 2006. Definitive identification of the species was possible on 2 June 2006 when the plants were blooming (Fig. 2). These Hexalectris stems were immediately adjacent to one of the preserve’s trails in an area dominated by Quercus spp. and Jun iperus spp, and with a somewhat open canopy due toa trail cut. Only half of the individuals survived to flowering, due to losses from trail use and inadvertent trampling of the small stems. Specimens (Fig. 3) from the Camp Wisdom and Cedar Ridge locations were depos- ited at the herbarium at the Botanical Research Institute of Texas in Fort Worth (BRIT). In addition, tissue from the plant stem and roots was sampled at both locations for ge- netic analysis as part of a study of phylogeny, evolution and mycorrhizal specificity in Hexalectris orchids (A. Kennedy, pers. comm.) Voucher specimens: Texas. Dallas Co.: Camp Wisdom (6400 Redbird Ln.), Escarpment habitat, Eddy-Brackett soil series, N 32 40.552’ @ 96 56.640, elev. 730 ft, 21 Jun 2005, M. Brown-Marsden and A. Collins s.n. (BRIT): Cedar Ridge Preserve, White Rock Escarpment, Dallas Park and Recrreation/Dallas County Open a property, N 32° 37.056 W 096’, elev. 711 ft, mixed oak juniper woods, ground cover primarily juniper and oak leaves, some bare ial rock, adjacent to trail, one of nine stems seen emerging on 13 Mz vy 2006, first blooms 2 Jun 2006, M. Brown-Marsden sn. (BRIT) In Texas, Hexalectrisgrandiflora has previously been documented only within the Davis and Chisos Mountains of extreme western Texas at elevations of 700-2500 m (Goldman et al. 2002). Confirmation of H. grandiflora in Dallas County marks both a significant disjunct in its distribution, and deviation from previously described habitat types. The plants found at the Camp Wisdom and Cedar Ridge sites were at elevations of 217 and 243 m, respectively. They were found in woodlands dominated by oak and juniper, but lacking the pinyon pine of the extreme west Texas part of its normal range. In spite of these deviations from the range and habitat of conspecific populations, they were still found on the same soil types as other members of the genus throughout Dallas County BROWN 1 AND COLLINS TEXAS 1241 lam ie Fic. 1, Photograph of Hexalectri liffora infl t Camp Wisdom, Dallas County, Texas on 21 June 2005. Photo by Randy Schoen. (Collins et al. 2005). Additionally, all observations of H. grandiflora have occurred within years of moderate to severe drought, and have been the driest years on record since 1963- 1964 (Drought Preparedness Council report, March 2006). The discovery of this significant disjunct in the range of Hexalectrisgrandiflora gives rise to larger questions with broader conservation implications. If this species is present in Dallas County but unknown in the area between Dallas and its previously known range, the gap may result from a lack of studies on Hexalectris ecology and distribution. In this event, it would be possible for this species to be found in one or more of the inter- vening counties through targeted studies such as those conducted in Dallas County. How- ever, it is also possible that the historic range of H. grandiflora included area between 1242 BRIT.ORG/SIDA 22(2) Fig, 2, Photograph of Hexalectris grandiflora inflorescence at Cedar Ridge Preserve, Dallas County, Texas on 2 June 2006. Photo by Puttaswamy Ravishankar. Dallas and West Texas, but this has since declined due to increased urbanization, and habitat loss and degradation. Determination of the ultimate causes and consequences of the recorded range disjunct may depend on continuing to answer fundamental ecologi- cal questions about this genus and factors influencing persistence of Hexalectris th rough- out its range. BROWN LLIN 1243 260 Kilometers Fic. 3. Map of historic locations of Hexalectris grandiflora in Texas (circles; after Catling 2004) and the new locations found in Dallas County (triangle). ACKNOWLEDGMENTS We thank Circle 10 Council Boy Scouts of America admini ion and staff for allowing us access to the Camp Wisdom site. In particular, we acknowledge the assistance of Randy Schoen during our initial censusing effort, and Phil Augsberger, camp ranger at Camp Wisdom. Jim Varnum, Stephanie Varnum, Rigel Rilling, and Jenny Krabacher provided assistance in fieldwork. Aaron Kennedy (Miami University-Ohio) assisted in collecting tissue for laboratory analysis. Photography assistance was provided by P. Ravishankar. REFERENCES Carunc, BM. 2004. A synopsis of the genus Hexalectris in the United States and a new variety of Hexalectris revoluta. Native Orchid Conf. J. 1(2):5-25. Carine, PM. and VS. Encet. 1993. Systematics and distribution of Hexalectris spicata var. arizonica (Orchidaceae). Lindleyana 8:119-125. 1244 BRIT.ORG/SIDA 22(2) Cottey, J. 2006. Statewide drought situation report. Report of the Drought Preparedness Council, March 6, 2006. CoLuns, A., J. Varnum, and M. Brown-Marspen. 2005. Soil and ecological features of Hexalectris (Orchidaceae) sites. Sida 21:1879-1891. DALLAS DEPARTMENT OF UrBAN PLANNING. 1977. The escarpment report. Department of Urban Planning of the City of Dallas, Texas. Diccs, G.M., Jr., B.L. Liescome, and R.J. O’KENNON. 1999, Shinners & Mahler's llustrated Flora of North Central Texas. Botanical Research Institute of Texas, Fort Worth. Gotoman, D.H., R.A. Coteman, L.K. Macrath, and PM. Cattinc. 2002. Hexalectris. In Flora of North America, Vol. 26. Oxford University Press, New York and Oxford. Pp. 603-607. Hatcu, S.L., K.N. GaNoni, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agric. Exp. Sta. Misc. Publ. 1655:232-235. Licaio, J.and A.O. Liccio. 1999, Wild Orchids of Texas. University of Texas Press, Austin, ERODIUM MALACOIDES (GERANIACEAE), NEW TO TEXAS David E.Lemke and Jed L.Aplaca Department of Biology Texas State University San Marcos, Texas 78666-4605, U.S.A. ABSTRACT Erodium malacoides (Geraniaceae) is reported as an addition to the flora of Texas. A description of the species and a key for identifying the Te pecies of Erodium are provided. RESUMEN Se cita Erodium | id i ) nueva adicion a la flora de Texas. Se aporta descripcion de la especie y una clave de las especies de Erodium de Texas. Erodium L'Hér. comprises ca. 60 species of annual or biennial, rosette-forming herbs wide- spread in temperate and subtropical regions of the Old World and New World (Mabberley 1997). Three species have previously been recorded from Texas: the native FE. texanum A. Gray, which is broadly distributed across central and southern portions of the state west to Utah and California, and the non-native species E. cicutarium (L.) LHér. and E. botrys (Cav. Bert. Jones et al. 1997). In Texas, E.cicutarium occurs ina variety of disturbed habi- tats in central, northern, and western regions of the state, while E. botrys has been col- lected from similar habitats in Bexar, Taylor, and Throckmorton counties (Correll & Johnston 1970; Turner et al. 2003). In the spring of 2005, the senior author discovered a small population of Erodium malacoides(L.) Willd, a species previously unknown from the state, established on a dis- turbed site in Hays County, Texas. By 2006 the population had increased in size to hun- dreds of individual plants and occupied a substantially larger area, indicating a poten- tial for naturalization. In the system proposed by Pysek et al. (2004), FE. malacoides would be categorized as a “casual alien” in Texas. Voucher specimens: U.S.A. TEXAS. Hays Co.: E side of northbound frontage road of Interstate Hwy. 35, City of San Marcos, N29° 53’ 38", W97° 54' 31", 02 May 2006, Aplaca 287 (SWT): W side of northbound frontage road of Interstate Hwy. 35, City of San Marcos, N29° 53' 39", W97° 54’ 31", 02 May 2006, Aplaca 288 (SWT), E side of northbound frontage road of Interstate Hwy. 35, City of San Marcos, N29° 53’ 45", W97° 54' 25", 02 May 2006, Aplaca 289 (SWT); We can only speculate as to how E. malacoides may have been introduced at this loca- tion. Several years ago, activity associated with the construction of a new entrance ramp onto the interstate highway disturbed portions of the area now occupied by the species. The seed mix used for revegetation of the highway right-of-way, soil brought in as fill material, or vehicular traffic from Mexico represent the most plausible seed sources. Erodium malacoides is native to southern Europe (Brandes 1995) but has become naturalized across the Mediterranean region (El Hadidi et al. 1984). It has been intro- duced as widely as Australia and New Zealand (Webb etal. 1988; Westbrooke & Florentine 2005), South America (Figueroa et al. 2004), and North America, where it has been re- ported from a number of Mexican states (Villasenor & Espinosa-Garcia 2004) and from SIDA 22(2): 1245- 1249. 2006 BRIT.ORG/SIDA 22 Fic. 1. Upper photo: Individual plant of Erodium malacoides from Hays Co., Texas, showing the cha istic simple, palmately lobed leaf and relatively small (ca. 10 mm diam.) flower. Lower phowo: Close-up of flower showing the glandular pubescence of the calyx and the broad, entire-margined staminodes. LEMKE AND APLACA 1247 California (Taylor 1993), New York (Mitchell & Tucker 1997) and Massachusetts (Magee & Ahles 1999) in the U.S.A. Plants of E. malacoides (Fig. 1) are annual herbs with decumbent to ascending, pu- berulent stems that are glandular at the nodes. The leaves are simple, 4-15 cm long in- cluding the petiole, which is generally longer than the blade. The leaf blades are ovate, crenate to shallowly palmately lobed, and puberulent. The small flowers are borne in umbellate clusters and have glandular-pubescent sepals 4-6 mm long, lavender to purple petals about as long as the sepals, five fertile stamens, and five broad, entire-margined staminodes. The fruit is a five-parted schizocarp that separates from the style column into single-seeded segments, each consisting of an indehiscent fusiform body witha more or less rounded, glandular apical pit subtended by a single furrow, and a persistent style segment that becomes tightly coiled when dry. The following key is provided to distin- guish among the four species of Erodium presently known from Texas. KEY TO THE SPECIES OF ERODIUM IN TEXAS 1. Leaf blades simple, often shallowly or deeply palmately lobed 2. Stems appressed-canescent; sepals 5-10 mm long, apically strigose; petals unequal, 7-12 m long; staminodes long-awned, conspicuously ciliate; fruit body 5-8 mm long, the pit ae elliptic and eglandular, furrows absent; style column 3-7 cm long E.texanum 2. Stems puberulent, the nodes glandular; sepals 4-6 mm long, glandular; petals equal, 4-6 mm long; staminodes awnless, entire-margined; fruit body 3-5 mm long, the pit + round, glandular, and usually subtended by one furrow; style column 2-3 cm long E. malacoides ]. Leaf blades deeply pinnately lobed to pinnately compound 3. Stem pubescence glandular; sepals 3—5 mm lonag;fruit body 4-7 mm long, the pit subtended by a single furrow or furrow absent; style column 2-5 cm long E. cicutarium 3. Stem pubescence eglandular; sepals 10-13 mm long; fruit body 8-11 mm long, the pit usu- ally subtended by two furrows; style column 5-12 cm long E. botrys REFERENCES Branpes, D. 1995. The flora of old town centres in Europe. In: Sukopp, H.,M. Numata and A. Huber, Urban ecology as the basis of urban planning. SPB Academic Publishing by, Amsterdam, The Netherlands. Pp. 49-58. Corrett, D.S.and M.C. JOHNSTON. 1970. Manual of the vascular plants of Texas. Texas Research Founda- tion, Renner. Et Haoiol, M.N., A.A. Faveo, and S.M. Et Nacaar. 1984. Systematic revision of Erodium (Geraniaceae) in Egypt. Pl. Syst. Evol. 144:307-314. FicueroA, J.A., S.A. Castro, PA. Marauet, and FM. Jaksic. 2004. Exotic plant invasions to the Mediterranean region of Chile: causes, history and impacts. Revista Chilena Hist. Nat. 77:465-483. Jones, S.D., J.K. Wiprr, and P.M. Montcomery. 1997. Vascular plants of Texas: a comprehensive checklist including synonymy, bibliography and index. Univ. of Texas Press, Austin. Massertey, DJ. 1997. The plant-book. A portable dictionary of the higher plants, 2nd ed. Cambridge Univ. Press, Cambridge. Macce, D.W.AND H.E. Antes. 1999. Flora of the Northeast.A manual of the vascular flora of New England and adjacent New York. Univ. of Massachusetts Press, Amherst. MitcHett, R.S.and G.C. Tucker. 1997. Revised checklist of New York State plants. Contributions to a flora of New York State, checklist IV. New York State Mus. Bull. 790:1-400. Pysek, P, D.M. RicHarpson, M. RemAnek, G.L. Weester, M. WILLIAMSON, and J. KirscHNer. 2004. Alien plants in checklists and floras: towards better communication between taxonomists and ecologists. Taxon 53:131-143 1248 BRIT.ORG/SIDA 22(2) Taytor, M.S. 1993. Erodium. In: Hickman, J.C.,ed. The Jepson manual: higher plants of California. Univ. of California Press, Berkeley. Pp. 672-673. Turner, B.L., H. NicHots, G. Denny, and O. Doron. 2003.Atlas of the vascular plants of Texas. Sida, Bot. Misc 24:1-888. Vittasenor, J.L. and FJ. Espinosa-Garcia. 2004. The alien flowering plants of Mexico. Diversity Distrib. 10:113-123 Wees, C.J., WAR. Sykes, and PJ. Garnock-Jones. 1988. Flora of New Zealand IV: naturalized dicots, gymno- sperms, ferns and fern allies. Manaaki Whenua Press, Lincoln, New Zealand. Wesrerookt, M.E. and S.K. Florentine. 2005.Rainfall-driven episodic flood events: are they a major factor in moulding New South Wales arid land vegetation patterns? Austral. Geogr. 36:171-181. NEW DISTRIBUTION RECORDS FOR EUPATORIEAE (ASTERACEAE) INTHE UNEEE DS TALES Guy L.Nesom Billie L. Turner Botanical Research Institute of Texas Plant Resources Center 509 Pecan Street University of Texas Fort Worth, Texas 76102-4060, U.S.A. Austin, oe 78713, U.S.A. ABSTRACT Several species of BupAtoneae a occur in neoutnenn Arizona and New Mexico at the northern end of their otherwise Mexican distribution: St m New Mexico by a single collection, the exact locality un- known and perhaps in medal advicent ve Ageratina (Eupatorium) thyrsiflora (known from Ari- zona by a single ae and Koanophyllon (Eupatorium) palmeri (previously confused with K. solidaginifolium). Re of K. palmeri or K. solidaginifolium from New Mexico have not been confirmed. Chromolaena(Eupator ) bigelovii, primarily a species of northeastern Mexico, is d d for the U.S.A. by three Texas collections—its occurrence was recorded in Arizona (cited from only the type collection), but that record apparently reflects a confusion of label data for the type, which almost certainly was collected along the Texas-Mexico border. Lectotypes are chosen for Eupatorium arborescens, Koanophyllon (Eupatorium) palmeri (var. palmeri), and Ageratina (Kyrstenia) thyrsiflora. RESUMEN Vanigs especies de Eupatorieae viven en el sur de Arizona y N a pe ee ‘ ray eee pe | ] de NI Mévi la recoleccién 7) cuya exacta localidad esd i | México), A ti E tori ) th ifl de Arizona con una sola leans y Koanophyllon (Eupatorium) palmeri Gone contindic con K. solidaginifolium). Las citas de K. palmeri o K. solidaginifolium de Nuevo México no han sido confirmadas. Chromolaena (Eupatorium) bigelovii, primariamente una especie del noreste de México, se documenta para los Estados Unidos mediante tres colecciones de Texas—su ocurrencia os citada en Arizona (timicamente don la coleccién del tipo), p sa cita refleja nfu e los datos de la etiqueta del tipo, que casi con seguridad fue colectado alo largo de la tronitera de Texas y México. Se eligen lectotipos de coauall arborescens, I } Eup ium) palm var. palmeri), y Ageratina (Kyrstenia) thyrsiflor Preparation of Eupatorieae treatments for Mexico and the Flora of North America (FNA) region has brought to light three species previously unreported for the United States. Both are primarily distributed in western Mexico and are newly recognized as occurring in Arizona and New Mexico. A third primarily Mexican species is rare in Texas; it previ- ously has been recorded as a member of the Arizona flora but, as discussed here, it does not occur in that state. Morphological descriptions of all four species are provided in the FNA treatments. 1. Stevia salicifolia This species apparently has not been included for the U.S.A. in any published floristic account, but it was recorded in a dissertation study (Grashoff 1972) from southwestern New Mexico by a single collection of E.A. Mearns, naturalist for the United States and Mexican Boundary Survey of 1892-1894. Grashoff cited the collection as from Grant County, but the San Luis Mountains are a range primarily of northern Sonora and Chi- huahua, Mexico, with an extension into Hidalgo County. Voucher specimen: U.S.A. New Mexico. [Hidalgo Co.]: canon east side of San Luis Mts., 11 Sep 1893, Mearns 2220 (US) SIDA 22(2): 1249- 1253. 2006 1250 BRIT.ORG/SIDA 22(2) In further refinement of the San Luis Mountains locality, Hubbard (1999) concluded that Penstemoncampanulatus, which had been considered to be represented in the U.S.A. flora only by a Mearns collection, was collected on the Mexican side of the border rather than in New Mexico (canon, east side San Luis Mts., 11 Sep 1893, Mearns 2222 - US). In view of this, the U.S.A. record documented here for Stevia salicifolia should be considered doubt- ful, even though it seems likely that the species has stepped across the border in one place or another. Geographic-ecological summary.—Flowering Jul-)Aug-Oct (in northern Mexico). Oak and oak-pine woodland, rocky sites, crevices, boulder pockets; ca. 1500-2650 m (data from northern Mexico). New Mexico; Mexico (Michoacan, Edo. Mexico, Puebla, and Ver- acruz northward to Sonora, Chihuahua, Coahuila, Nuevo Leon, Durango, Zacatecas, San Luis Potosi). The range of var. salicifolia also closely approaches Texas and Arizona (see map in Turner 1997). Stevia salicifoliais a variable species—many infraspecific taxa have been named but most were regarded by Grashoff as synonyms of var. salicifolia. Among the U.S.A. species of Stevia, S. salicifolia is distinct in its glabrous and shiny stem and leaf surfaces, which results from the viscid exudate of sunken glands; stems and leaves of the other U.S.A. species usually are sparsely pubescent and not ee Stevia salic ifolia Cav le 32. pl. 354.1797. “The tyre... came from plants cultivated at the Royal Botani- cal Garden, Madrid; Ortega states that the seed was sent hon Mexico by Sesse.” (Mc Vaugh 1984, p. 895). 2. Ageratina thyrsiflora This species is native primarily to the states of Sonora, Chihuahua, Sinaloa, Durango, and Jalisco in western Mexico (Mc Vaugh 1984; Turner 1997). It is known from the United States only by a single collection near the international border. U.S.A. Arizona. [Santa Cruz Co.|: near Nogales, 15 Sep 1929, Harrison & Kearney 6039 (ARIZ fide Phil Jenkins, LL). Geographic-ecological summary.—Flowering Sep-Nov. Rocky sites, oak woodland; 1000- 2200 m (data from Mexican collections at TEX-LL and published notes of McVaugh 1984). Arizona; Mexico (Sonora, Chihuahua, Sinaloa, Durango, Jalisco). Ageratina thyrsiflora is recognized by its strict, unbranched or few-branched stems with alternate, densely arranged leaves, small heads densely clustered in a single, termi- nal aggregate, prominently orange-veined corollas and phyllaries, and close cauline and petiolar puberulence of minute, sharply upwardly bent hairs. Morphology is relatively uniform over the geographic range, except in two collections from Sonora (Muller 3655, LL) and Chihuahua (Gentry et al. 18026, ARIZ, LL), where stipitate glands are mixed with the eglandular puberulence on lower stems and petioles. One collection from Jalisco (Mc Vaugh 21771, LL) also shows a tendency to produce stipitate glands. Ageratina thyrsiflora (E. Greene) R. King & H. Robinson, Phytologia 19:227.1970. Kyrstenia hyrsiflora E. Greene, Leaf. Bot. Observ. Crit. 1:9. 1903. Eupatorium thyrsiflorum (E. Greene) B.L. Robin- son, Proc. Amer. Acad. Arts 43:36. 1907. Type: MEXICO. CHIHUAHUA: Southwestern Chihuahua, Aug 1885, i, ae 275 (LECTOTYPE selected here: US, internet image!). In the protologue, Greene cited the following: Chihuahua, Mexico, chiefly southward in the State; aes oy Palmer, P ae und E.A. Goldman, and lways distributed for E. occidentale var. Arizonicum. t GH was cited by Mc Vaugh (1984) as “isotype” of K. thyrsiflora (near C ce 1 Oct 1885, CG. Pringle 613), but there is no duplicate of this at NDG (fide Barbara Hellenthal) or at US. Greene perhaps studied the Palmer specimen at US before his time at the Smithsonian as an associate in botany (in 1904-1909); he did not annotate ses ees howe, n though he did make handwritten notations on U sheets of K sheet K. laeta, K ne ee and kK. rufa (fide Barbara Hellenthal, pers. comm), A 1251 Eupatorium thyrsiflorum var. holoclerum B.L. Robinson, Proc. Amer. Acad. Arts 43:36. 1907. Type: MEXICO. DURANGO: City of Durango and vicinity, Apr-Nov 1896, E. Palmer 755 (HOLOTYPE: GH; IsoTYPEs: MO|, US internet image!). 3. Koanophyllon palmeri Koanophyllon (Eupatorium) palmeri is native primarily to western Mexico (McVaugh 1984; Turner 1997) and is known from the U.S.A. by collections from south-central Ari- zona. A similar taxon (Eupatorium palmeri var. tonsum B.L. Robinson) occurs in near- coastal localities from Michoacan to Colima, Jalisco, Nayarit, and Sinaloa. The Arizona plants are var. palmeri( hoccurs southward through Sinaloa, Chihuahua, and Sonora to Durango and Jalisco). Eupatorium arborescens (as cited below) from Chihuahua is a synonym of K. palmeri var. palmeri. Vouchers for records of Koanophyllon palmeri in southwestern New Mexico (Hidalgo and Grant counties—as mapped by Martin and Hutchins 1980) have not been located in New Mexico herbaria J. Mygatt, UNM, pers. comm. and R. Spellenberg, NMC, pers. comm.). The basis for the Hidalgo County record may bea citation by Wooton and Standley (1915, p.647) of a Charles Wright collection “Guadalupe Pass, Wright 1146,” [4-5 October, 1851). According to Wooton and Standley (1915, p. 647), “Guadalupe Pass is on the southern boundary of the State, and Wright’s specimens may have come from either Mexico or New Mexico.” It seems reasonable to maintain K. palmeri asa member of the New Mexico flora, at least tentatively, especially in view of the map points shown by Martin and Hutchins (1980). U.S.A. Arizona. Cochise Co.: Guadaloupe Mts., Guadaloupe Canyon, 13 Oct 1946, Darrow et al. 3562 (LL). Pima o.: W side of Mt. Baboquivari, ees 1944, Gould ce 2670 (LL); Baboquivari Mts., 25 Sep, 1927, Harrison 4754 (LL); Baboquivari Mts., Sep 193] 932 (LL); Sta. Catalina Mts., Sabino Canyon, 27 Sep 1934, deed Peebles 10257 (LL); Babocunee Mts., 30 Sep 1934, Kearney & Peebles 10373 (LL); Baboquivari Canyon, 1] Oct 1 Peebles et al. 413 (LL); Tucson Mts., Hugh Norris Trail, 7 Nov 1976, Urry 840 (TEX), Waterman Mountains, NE = of road to Silver Hill Mine, 5 Dec 2001, Van Devender 2001-1014 (TEX). Santa Cruz Co.: Coronado Natl. Forest, just to the E end of Forest Rd 217, near the Dos Amigos Mine, E of of Old Glory Canyon and Warsaw Canyon, 31 Aug 2001, Goldman 2108 (BRIT); Peck Canyon, Atascosa Ranch, 18 Nov 1981, Van Devender s.n. (TEX): ene Canna, lmi NW of Bartlett Mts., 29 Oct 1981, Van Devender & Toolin s.n. (TEX). Geographic-ecological summary.—Flowering Sep-Nov(-Dec). Shaded rocks along streams, crevices, often in oak woodland; 850-1500 m. (data from Arizona collections). ie eae palmeri (A. Gray) R. King & H. Robinson, Phytologia 22:150. 1971. Eupato- d m palmeri A. Gray, Proc. Amer. Acad. Arts 21: 383.1886. Gray did not cite a aaa collection but note “type locality, ‘shady places high up in mountains above Batopilas.” Type: MEXICO. [CHIHUAHUAl: South- western Chihuahua, August to N ber 1885, Palmer 144 (LECTOTYPE ee ere: GH}; ISOLECTOTYPE: US internet image!). Ape collection, E. Palmer 263 (GH)), is similarly labeled. Both GH sheets were eee oa ie as “Eupatorium palmeri n. sp.” McVaugh’s account (1956) of Palmer’s seh indi- the vicinity of Batopilas were Aad in August through early October 1 Eupatorium arborescens M.E. Jones, Contr. West. Bot. 12:43. 1908. Type: MEXICO. CHIHUAHUA: Sierra Madre Mts., Guayanopa Canon, 3600 feet alt., in the Tropical Life Zone, 24 Sep 1903, M.E. Jones s.n. (LECTOTYPE pace ene ne ea hoe 1799, phOscops: ISOLECTOTYPE: RSA-POM 41800, photocopy!). Hi | g label oa and probably are ua: of Ene same Gels ec- tion; 41799 b gl I h: 41800 | } Each sheet is labeled apparently that of f Philli ip Munz, fide Michael Denslow at RSA-POM) “Part of type, Eupatorium ar horescens fos ’ Leaf position, vestiture and involucral features are those of K. palmeri, although the acuminate leaf apices are more similar to those of K. solidaginifolium. a: Plants of Koanophyllon palmeri in Arizona have been previously identified as K. solidaginifolium (A. Gray) King & H. Robinson (= Eupatorium solidaginifolium A. Gray, 1252 BRIT.ORG/SIDA 22(2) see citation below). As interpreted here, however, and in Turner (1997) and Blake (1924), K. solidaginifolium occurs in north-central Mexico (eastern Chihuahua, Coahuila, north- eastern Durango, northern Zacatecas) and into the trans-Pecos area of Texas, east of the range of K. palmeri. Their ranges approach each other in central Chihuahua, but the two apparently are allopatric. The following contrast separates the two species in the U.S.A. and northwestern Mexico. Leaves all opposite, blades apically acute, upper surfaces sparsely strigose to weakly hispidulous; involucres (3—)3.5-4 mm long; phyllaries herbaceous to the margins, at least the outer sald U lent, all usually weakly ciliate, outermost ovate-elliptic to obovate nophyllon palmeri Leaves usually subopposite to alternate on distal third of stem, blades apically long-acur serine upper surfaces glabrous; involucres 4.5—5.5 mm long; phyllaries with narrow but distinct t bas line-translucent margins, glabrous and eciliate, outermost narrowly lanceolate. phyllon side aici Eupatorium ue ium A. Gray, Smithsonian Contr. Knowl. 3(5) [Plant. Wright. 1]: ou paloie ye -“Collected in Expedition from Western Texas to El Paso, New Mexico, May-October, 1849, by Charles Gey [as on label], Wright 256 (HOLOTYPE: GH, photocopy! lsoTYPE: GH, photocopy!). Gray’s citation in the protologue was “Mountains between the Limpia and the Rio Grande, New Mexico.” Ac- ield notes (fide Walter Kittredge, GH), this locality cording to LM. Johnston’s commentary on Wright's fi matches Wright ’s field number of 256, as recorded on the type labels. L ee creek and Limpia Canyon are in heen day Jeff Davis Co., Texas, and it is probable that Wright's collection was made there (recent t the species in Jeff Davis Co. and Presidio Co.), proba bly in late August (as inferred from Wooton 1906). The specimens were annotated as ‘holotype’ and ‘isotype’ by D.E. Boufford in 1981. 4. Chromolaena bigelovii Chromolaena bigelovii has been included in various summaries of the Arizona flora (e.g, Kearney & Peebles 1951; Lehr 1978) on the basis of the protologue, which cited “On the Gila, Sonora; Parry.” Kearney and Peebles (1951, p. 845) observed that the species is “ap- parently known in Arizona only from the type collection on the Gila River (Bigelow),” and the present authors have located no further records of the species from Arizona. In contrast, plants matching the type of C. bigelovii (below) have been collected at numer- ous localities in northeastern Mexico (Coahuila, Nuevo Leon, San Luis Potosi—Turner 1997) and at two sites in Texas (mapped in Turner et al. 2003; documented here). Eupatorium madrense (as cited below) from Nuevo Leon is a synonym of Eupatorium bigelovii. U.S.A. Texas. Brewster Co.: Big Bend National Park, Basin of Chisos Mt. base of Baldy Peak, 6 Sep 1950, Warnock 9587 (SRSC); Big Bend National Park, Sierra Quemada, Claro 2 Spring, 29 Oct 2000, Bartel 211 (SRSC). Uvalde Co.: on chalk bluffs at Park Chalk Bluff along the Nueces River, ca. 15 mi NW of Uvalde, 12 Dec 1967, Correll 35433 (LL). Geographic-ecological summary.—Flowering Oct-Dec. Dry limestone hills in oak wood- lands, talus: ca. 1OOO-1800 m. (data primarily from Texas, Coahuila, and Nuevo Leén). Chromolaena bigelovii (A. Gray) R. King & i Robinson, Phytologia 20:208. 1970. Eupato- rium bigelovit A. Gray in Torrey, Rep. | 1s. & Mex. gund. Survey, Bot. 75. 1859. Type: U.S.A. [locality uncer- tain but probably along the Texas-Coahuila oie 1848-1855], J.M. Bigelow 12 (probable HOLOTYPE: GH, photocopy!). The protologue by Gray cited “On the Gila, Sonora; Parry.” Blake (1924) and Kearney and Peebles (1951) noted that the ely sou be Pi aced in Arizona than nian oo labels, however, ona collection that i Is reasonabl i g “E. bigelovii y n. sp.” and “E. bigelovii Gray” and do not have locality information. On the original label, “Mex. Bound. Survey No. 12,” a note by Gray says “Bigelow—locality not recorded”; another note (B.L. Robinson’s hand- writing) reads that “Identical material in Paris Herb. and Berlin Herb. bears Mex. Bound. Surv, label num- bered (in pencil) 471.” A Parry collection that would represent a potential type for Eupatorium bigelovii has not been located at GI NE IER THE U.S.A. 1253 The species represented by the Bigelow collection is the one that has been collected nu- merous times in northeastern Mexico and immediately adjacent Texas. The collection almost certainly was made along the Texas-Mexico boundary, in the known range of the species, in the region covered by the United States and Mexican Boundary Survey of 1848- 1855, for which Bigelow worked as a naturalist. Numerous collections by Bigelow from southwestern Texas are specifically cited in the Survey's “Botany” report. In the tentative interpretation here, a mistake in manuscript preparation apparently led to a substitu- tion both for the correct collector (Parry instead of Bigelow) and correct locality (On the Gila, Sonora, instead of ‘along the Texas border’). Gray’s choice of epithet supports this interpretation. Eupatorium madrense S. Watson, Proc. Amer. Acad. Arts 26:137. 1891. Typr: Mexico. Nuevo Leon: Sierra Madre near Monterey, 7 Jun 1888, C.G. Pringle 2201 (HOLOTYPE: GH photocopy!). ACKNOWLEDGMENTS We are grateful to Steve Boyd and Michael Denslow (RSA-POM) for information and pho- tocopies of Eupatorium arborescens, Phil Jenkins and Michael Chamberland (ARIZ) for checking potential occurrences of Chromolaena bigelovii at ARIZ, Phil Jenkins for com- ments on Ageratina thyrsiflora, Jane Mygatt (UNM) and Rich Spellenberg (NMC) for pro- viding information on New Mexico collections, Barbara Hellenthal for checking the po- tential occurrence of collections at NDG, Harold Robinson (US) for review comments, and Emily Wood and Walter Kittredge (GH) for providing information and photocopies of various GH collections. REFERENCES Bake, S.F. 1924. Asteraceae. In:P.C. Standley, Trees and shrubs of Mexico.Contr, U.S. Natl. Herb.23(4):1401- 1641. Grastorr, J.L. 1972.A systematic study of the North and Central American species of Stevia.Ph.D.diss., Univ. of Texas, Austin. Husearo, J.P. 1999. Penstemon pulchellus Lindl. [= P campanulatus (Cav.) Willd.]:A specious Mena or New Mexico's flora. New Mexico Botanist,|ssue No.11. Accessed June 2005. Lene, JH. 1978. A catalogue of the flora of Arizona. Desert Botanical Garden, Phoenix, Arizona. McVaucu, R.1956. Edward Palmer: plant explorer of the American West. University of Oklahoma Press, Norman. McVauch, R. 1984. Compositae. Flora Novo-Galiciana 12:1-1157. Martin, W.C. and C.R. Hutcuins. 1980. A flora of New Mexico. Vols.1 and 2. J. Cramer, Vaduz. Turner, B.L. 1997. The Comps of Mexico. A systematic account of the family Asteraceae. Volume 1. Eupatorieae. Phytologia Mem. 11:i-iv; 1-272. Turner, B.L., H. NicHols, G.C. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas.Vol. 1, Dicots; Vol. 2, Monocots, ferns, Sida, Bot. Misc. 24. Wooton, E.O. 1906. Southwestern localities visited by Charles Wright. Bull. Torrey Bot. Club 33:561- Wooton, E.O. and PS. STanpLey. 1915. Flora of New Mexico. Contr. U.S. Natl. Herb. 19:1-794. 1254 BRIT.ORG/SIDA 22(2) Book REVIEWS Bruct A. Bot. 2004. Hawai’i’s Native Plants. (ISBN 1-56647-066-6, pbk.). Mutual Publish- ing, 1215 Center Street, Suite 210, Honolulu, H1 96816, U.S.A. (Orders: 808-732-1709, 808-734-4094 fax, mutual@mutualpublishing.com, www.mutualpublishing.com). $22.95, 224 pp., approx. 200 color photographs, 6" x 9" Dr. Bruce A. Bohm’s Hawai’i’s Native Plants presents a historical view of botany on the Hawaiian islands and is quite pleasant to read. I must begin by saying that Hawaii’ Native Plants is nota field guide for quick identifica tion of plants. Rather, it carefully introduces the geologic, ecological, evolutionary and cultural history of each major taxonomic group, often in too much detail for the lay reader. This book is not organized like a traditional field guide; species information is presented in narrative form with headers of both Genus and Hawaiian names introducing each group. Excellent color photographs throughout the book provide identifying characteristics of species, though not all species are a gaa That being said, this book would be an excellent reference / companion for a “quick-identification” field guide of Hawaiian plants. is work begins with a chapter (The islands)on the geologic, ecological and cultural history of the Hawai- ian islands It gives a brief introduction to each island, eens deseri ested of vegetation patterns as well as brief historical backgrounds. This first chapter reads almost like a travel gui Hawai'i, but is a good introduction to the processes and historical events that have helped shape she island s current flora and fauna. Bohm focuses on plants that are easily accessible to the amateur naturalist/tourist but does include some more challenging-to- lind species. His stated audience is persons traveling to Hawai’; his goal is for the reader to become appreciative of Hawai’'s native flora and informed of the current threats to Hawai’i’s biodiversity. However, the average tourist (especially if he/she has no background in biology or geology) may find this book a challenging read because of the detail in which Bohm discusses biological and geological processes ative Hawatian Plants, the second chapter, is the one that provides the taxon-specific descriptions anc introduction to the major groups of native Hawaiian plants; thus, it makes up the majority of the book. As men tioned hee ai Plants are pateedneee uInE neading> of the Genus ane raya an TAMIeIe), Plants are first de- scribed fori history f most groups are discussed. For most ai the taxa, these short literature explorations may le sad the lay reader to loose interest in the technical information presented (ie. phylogenetic analysis or molecular genetics techniques). 2 conservation status, native cultural uses, factors that have affected the colonization, distribution, and populati species (e.g. major storms, volcanic activity, human activity) are a great addition to this book, as this nloumation is rarely included to this extent in traditional field guides. The end of this chapter contains a small section en- titled Ferns and Fern Allies; it is short and limited to relatively easy to find species. The author suggests Daniel — Hawat'i’s Ferns and Fern Allies (2003) tor a more soln iE enSING guide to Sa erouD e third chapter, Alien Invasion, discusses the extent t I native f] | a Th he author includes this chapter in hopes that the reader will gain an apeeeaaen for the “full impact” invasive species can have on native biota. He introduces invasive plants by taxonomic group and discusses how each group became introduced to the islands, focusing on species considered to be invasive as opposed to natural- ized or introduced species. There is also memes of the ecological CHATS of invaders that have made es so o successful on awe iand of s that are being try to eradicate) invasives. final chapter (There Is No Future In Extinction) begins with a discussion af extinction in general, con- ete species loss and biological characteristics sist may make certain taxa more prone to extinction. In this chapter Bohm gives the reader an abundance of literary references on extinction for further exploration. He includes a brief presentation of global hotspots of plant diversity and endemism, then discusses threats specific to Hawaiian flora and describes specific conservation efforts to preserve or restore native populations of Hawai- ian plants. Finally, the author provides a short appendix at the end of the book that explains methods commonly sed by taxonomists to identify and study eaeee history of species, including molecular phylogenetics ages that have allowed for better identification of plants. any or avid natural historians with some bac nee knowledge of geology, evolution and ecology, this book isa Stee addition to the botanical literature on Hawai'i. Professors teaching courses on Hawaiian natural history or botany would find this a useful volume; it is also quite affordable for students. Hawaii's Native Plants would best be read in ae entirety before a trip to Bona ito eee an unde standing of the basic lationship between plant taxaa logy of plant groups. TI ily find certain species (often roadside access), and would be a usef ful tool for planning outings on trips.—J. Hayley Gillespie, University of Texas at Austin, Integrative Biology, Austin, TX 78712, U.S.A, hayleygillespie@mail.utexas.edu SIDA 22(2): 1254. 2006 BOOK REVIEWS 1255 MELINDA A. ZEDER, DANIAL G. BRADLEY, EvE EMSHWILLER, and Bruce D. Smitu (eds.) 2006. Docu- nting Domestication: New Genetic and Archaeological Paradigms. (ISBN 978-0- 520-24638-6, hbk.). University of California Press. Berkeley, Los Angeles, London. (Orders: California Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, New Jer- sey 08618; Tel.: 609-883-1759; Fax: 609.883-7413) $70.00, 361 pp., b/w figures, b/w photos, tables. Apoues ae studies were well underway during the last century, Documenting Domestication: Ww haeological Paradigms definitely sets a new standard for scientific investigations, integra- tions, and interpretations for the 21st Century. Certainly, rapidly advancing electronic technologies for entering, storing, and retrieving data rapidly and accurately have ele the necessary tools to enable and facilitate a much larger body of assemblages and studies. More- research and interpretation and to utilize materials fro uch earns developments as high-power scanning electron microscopy and techniques “for the re- over, $ t macro- and micro-fossils” have greatly expanded and enhanced co identification of eee ae and genetic studies. mposium presented at the annual meetings of the American Association for the Advancement of Sci- ence (2001) featured four speakers, each addressing progress and future concerns and challenges in pene spe- cific studies “on the domestication of plants and animals from archaeological and genetic perspectives.” That kers are editors as well as contributors. omestication: Bringing Together Plants, Animals, provided the impetus for this volume, and three of those spea e overarching introductory chapter, “Documenting D Archaeology, and Genetics,” is written by the editors. It provides an in- de pe overview of earlier work from the mid 19th Century, beginning with perceptions from Darwin and through the present, with its rapidly expanding resources and technologies. Discussions of changing approaches and understandings of the scientific disciplines herald fresh new ways of ae and interpreting data, not only from newly collected evidence but from renewe tion sting data The book is designed ona ee matrix” (table 1.1). Research for domestication of plants and animals is documented in special sections based on archaeological studies and on Boe sas bach oe is respon- the genetic sible for one of tl gy of plant d documentation of plant domestication; Melinda Zeder, annie of animal domestication: and Dan Bradley, the genetics of plant domesticati ion ditor has iew chapter preced les in th egory. This is very helpful for the reader, and it pemmits rapid identi ication sof chapters ely 2 be relevant t to one’s own interest or research. 1 provide informa- The chapters tion that previously has not been easily available The ue mmterest will oe be drawn first by those that most oe oe interest pteee and os Re me, aus I enyea the entire volun TI of fi | visual percep- tion of data and eee are Grice additions. ther sad lor us volume? is the wide diversity of geographic areas covered by the researchers. The range of d hand South America, Eurasia, and the Middle East. That breadth of research enhances the ee of key animal and plant coverage and is eee helpful in analyzing the data. (To date, most studies have been centered on are: often very small areas; coma een wath aero iaeee from eis sites can be very distorted. It is not easy t formulate I the retrieved data were based on the same or femmes procedures and markers.) Interestingly, the new methods, tools, data retrieval, ease o to new information being immediately available from remote study areas have were workers to re-think older, accepted interpretations and to construct solid new paradigms ew questions; me eae and new pecan of evidence from ancient archaeological as- 1 | A data, ac 2SS to 7 inaccessible nd access semblages, « to phytoliths in plants or study studies. call of these things enable researchers to study op evidence again and refine or rane the interpretations. ning and execution of this Vohaniee are bom peas te and professional. The use of Vincent van Gogh's “The See for the cover is particularly p Scientists and historians usually concur that determining events in the history of mankind. This volume is ing.—Helen Jeude, Botanical Research Institute of Texas, Fort Worth, Texas 76102-4060, U.S.A. rr L d tication of pants and animals is one of the major stand- SIDA 22(2): 1255. 2006 1256 BRIT.ORG/SIDA 22(2) GFORGE Y ATSKIEVYCH. 2006. Steyermark’s Flora of Missouri Volume II- Revised edition. (ISBN 1-930723-49-0, hbk.). Missouri Botanical Garden Press, P.O. Box 299, Saint Louis, MO 63166-0299, U.S.A. (Orders: http://www bgpress.info/). $48.00, 1181 pp., b/w line drawings, maps, 7 3/8" X 97/8". fai yanyone interested in the Missouri F 4 ora Project, George Yatskievych’s revision of the original Flora of Missouri i Julian Steyermark was looked forward to with great anticipation. Volume 1, published in 1991, covered pteri- dophytes, gymnosperms, and monocots. It was so well written, organized, illustrated and thorough, that Mis- souri botanists anxiously awaited Volume 2. Published in June 2006, Volume 2 covers the first part of the dicots alphabetically from the Acanthaceae through two subfamilies of the legumes or Fabaceae (Caesalpinoideae and Mimosoideae). Volume 2 is so well done that the author has even surpassed the eae of Volume | and stu- dents of Missouri's flora have been greatly rewarded for their patience. Each species account includes a detailed description that is the most thorough I have ever observed in any North American floristic manual. Also pro- vided are blooming periods, county dot range maps, characteristic habitats, relative abundance and illustrations for the overwhelming majority of species. Most taxonomic treatments follow the findings outlined in most re- cently published literature, Sly for the most part, the rearrangement of genera within the Asteraceae that has been adopted in inuals (e.g., Flora of North America). Even without the seegone ae 2 isa monumental enue Nonet heless, the illustrations are so detailed and so complete that the 193 plates are the most accurate repres “treated species | have seen in any flora. One aspect | particularly like is the enlarged detail of the fruit illustrated ete is easy to recognize, especially for genera and species that are often difficult to visualize in sufficient detail to distinguish between closely related taxa (e.g,, species of Vernonia, plates 299 and 300; and species of Cuscuta, plates 364 and 365). Another highlight is that the Brassicaceae includes | ve lor both { Novwersand ED 651-659). Anyone who has ever tried to key out material in this family will fi ] a eS y helpful. Overall, the keys and the glossary on pages 1085-1102 covers terms tha at may be unfamiliar to some readers me 2 is such a quality compilation of information that it is hard to find fault. Nonetheless, Gee area few minor points that should be brought to light. The most obvious oversight is that the numerous genera and species within the Asteraceae are arranged by Tribe. Although this alignment is helpful in simplifying keying exercises (pages 167-171), it is extremely frustrating for readers who do not know what tribes various genera are ascribed, especially given the recent taxonomic changes within the oles To locate a eae genus within the Asteraceae, the reader must locate it by thumbing through 42 pages of index (pages | 139-1181). A potential solu- tion would have been to list all genera within each tribe near the - ginning of the discussion on the Asteraceae. Another shortcoming of the book is that the treatment does not cover the last subfamily of the Fabaceae (ie., Faboideace ) which includes most of the species of legumes documented for a state, Obvionsy ay writer of a } | } book this size establish a cut off somewhere, but it would have | family to b function wel covered together Alsou 1 undoubtedly a selfish and unrealistic expectation of his reviewer, it would be lave been helpful to include a key that included all dicot families, especially given the length of time that is likely required to complete the last treatment. With any book of such magnitude, minor errors are sure to be inherent but only a few have been identified to date. The couplets separating Helenium autumnale from H. virginicum in the key on page 490 were inadvertently transposed. Helenium virginicum should be 3a, not 3b. On plate 386 (page 1067), the fruiting branch illustrated for Senna obtusifolia should be labeled 10e and the fruiting branch for S.occidentalis should be labeled as 10f. In conclusion, Ste yermark’s Flora of Missouri- Volume 2 is nothing less than a botanical masterpiece in print and it clearly warrants a spot on the shelf of every botanist in Missouri. Because its application and coverage in- ludes many species that have been documented for many states in the Midwest and eastern Great Plains, it would make an excellent reference book for anyone studying plants in the eastern United States.—Paul M. McKenzie, U.S. Fish and Wildlife Service, 101 Park DeVille Dr; Suite A, Columbia, Missouri 65203, U.S.A., Email: paul_ mckenzie@fwsygov — SIDA 22(2): 1256. 2006 REPORT AND UPDATE 2006 SIDA, CONTRIBUTIONS TO BOTANY Source of current research in classical and modern botany for readers throughout the world. systematic ...bringing out the best in botanical science for plant conservation and education To all authors, we thank you for choosing Sida, Contributions to Botany as your forum to disseminate information and knowledge gained by scientific inquiry. Volume 22, com- plete in two numbers, will be the last volume of the journal title. Volume 22 has 1274 pages, 169 authors, 105 published contributions, and includes 97 new names and new combinations. Published papers are available online in Adobe Acrobat format (PDF files); the PDF files are true representations of the hard copy of Sipa . A word of thanks to the 298 individuals acknowledged on pages 1258-1259, who generously supported Sip through their time and expertise in reviewing 226 manuscripts submitted for volume 22. All manuscript submissions are peer-reviewed by distinguished reviewers; your support is deeply appreciated. If by chance you reviewed a manuscript and your name was left out, the error rests solely on the shoulders of this editor and | truly apologize. Please bring any omissions to my attention. Below are the dates of publication for the two issues of volume 22, subscriptions. We also bring to your attention (see bottom of this page) the Mary M. Hennen Scientific Pub- lications Endowment, established to provide support to the future of BRIT’s scientific publi- cation program (Sida, Contributions to Botany and Sida, Botanical Miscellany). The index to volume 22 (2006) follows. Guidelines to contributors are available online at http:// www.britorg/sida/SubmitPaperhtm. We thank all authors, reviewers, subscribers (individuals, institutions, organizations), and readers for your continued interest and support. It is our plan to continue bringing you the best sources of current research in classical and modern systematics with your continued support. Wishing you the best in 2006 and beyond.—Barney Lipscomb (BRIT), Editor: John W. Thierett (NKU), Associate Editor; and Félix Llamas (LEB), Contributing Spanish Editor. Dates of publication 22(1), pp. 1-846: 11 August 2006 22(2), pp. 847-1274: 23 November 2006 Mary M. Hennen Scientific Publications Endowment Established.—In honor of his wile, Mary, BRIT research associate, professor emeritus of botany at Purdue University, the world’s pre-eminent authority on the systematics and biogeography of the rust fungi, Joe Hennen has established the Mary M. Hennen Scientific Publications Endowment to pro- vide support to the future of BRIT’s scientific publication program. Joe and Mary (life- science librarian at Purdue University) have long recognized that publication of scien- tific botanical information is an integral part of conserving our natural heritage. Every gift to the endowment will make a difference for the future of BRIT scientific publications. There are many different ways to make a gift; use a credit card, give a check, stock, pledge, bequest, trust, memorials, employer/employee matching contributions, SIDA 22(2): 1257. 2006 1258 BRIT.ORG/SIDA 22(2) installments, etc. The BRIT Press seeks your support to ensure innovation and excellence in preparation, manufacture, and distribution of botanical research and scientific dis- coveries for the twenty-first century. For more information about the BRIT Press or any- one interested in making a contribution to the Mary M. Hennen Scientific Publications Endowment—which supports Sida and Sida, Bot. Misc.—contact Barney Lipscomb at the Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, Texas 76102-4060, US.A., 817-332-7432; barney @brit org; sida@britorg. REVIEWERS: VOLUME 22 (2006) 298 reviewers: several individuals reviewed more than one manuscript Thank you for supporting Sida, Contributions to Botany, Acedo, Carmen Allred, Kel ly W. Anderson, Loran C. Andreas, Barbara K. Arnold, A. E Arriagada, Jorge E. Austin, Daniel F Ayala-Silva, Tomas — izabeth Ayers, Tina J. Bailey, Jr, Claude J. Baird, Gary L. Baldwin, Bruce G. Ball, Peter W. Ballard, Jr, Harvey E. Ballenger, Julie Barber, Susan C. Barkworth, Mary E. Barlow-lrick, Patricia Barrie, Fred R Baskin, Carol C. Bz S > or) Brown, Larry E. Brunell, Mark S. Buck, William R Burnham, Robyn Burns, Jean SIDA 22(2): 1258. 2006 Campbell, Julian Cardellina, John Carlquist, Sherwin Carlsward, Barbara S. Carranza, Eleazar Carstens, Kenneth C. Carter, Richard Chambers, Kenton L. Chinnappa, C Christensen, a Ib Church, Sheri A Clark, Lynn G. Columbus, J. Travis Consaul, Laurie | Cruden, Robert W. Davidse, Gerrit Davila, Patricia D. Delprete, Piero G. Denslow, Michael W. Dorr, Laurence J, Dowling, R.M. Duvall, Jams Eggli, Urs Elisens, Wayne J. Endress, Mary E. Essig, Frederick B. Evans, Dan K Ezcurra, Cecilia Fantz, Paul R. Farmer, Susan B. Ferguson, C -aroly nj. Fernandez-Concha, Ger- man Carnevali Fishbein, Mark Ford, Bruce A. Ford-Werntz, Donna Freeman, Crai Freire, Susana Edith Freire-Fierro, Alina Frye, Christopher Fryxell, Paul A. Galloway, Cynthia M. Gandhi, Kanchi N. George, Robert German Ramirez, Maria T. Ghatoor, Abdul Gomez-Sanchez, Maricela Gonzalez-Elizondo, M. Socorro Gralt, Pamela Grant, Verne Hall, David W. Hallgren, Steven Hansen, Bruce F Hat e en Reuse L. > Hatch, Stephan L. Haynes, Robert R. Hennen, Joe Henrickson, James Herrera Arrieta, Yolanda Hess, Bill Hill, Steven R. Hoch, Pete Hoggard, Gloria Hoggard, Ronald K. Holmes, Walter C. Holst, Bruce K. Horne, Francis R. Howcrott, N.HS. Humber, Richard A. Hyatt, Philip Janovec, John Jarvis, Charlie Johns, Bob Johnson, Dale E. Johnson, George P. Jones, Thomas H. Judd, Walter S. Kaul, Robert B. Kawasaki, Lucia comes Sterling C. Kennedy, Robin C. Kephart, Susan R. Khan, Zaheer Kirkbride, Jr, Joseph H. Knapp, Sandra Kral, Robert Krapovickas, Antonio Kumar, Muktesh Kumar, Muktesh Lammers, Thomas G. t, Eric Landrum, Leslie R. Lee, Linda Lemke, David E. Liogier, Main Lipkin, Robert Liston, Aaron Livshultz, Tatyana Llamas, Félix Lohmann, Lucia G. Lonard, Robert I. Lowry IL, Porter P. MacDougal, John Macklin, James A. REVIEWERS Magill, Robert E. Magrath, Lawrence K. Malmquis oe le Washington Martin, David L Martinz Diaz, Mahinda M Teresa Mellichamp, T. Lawrence Middelton, David Mill, Robert Molpus, Jane Moore, Gerry Morefield, James D. Moreno, Nancy P. Mueller, Jochen Murray, David F. Murrell, Zack Naczi, Robert EC. Nasir, syed Mahmood Ne : Nepokroeff, Molly Nesom, Guy Nevling, Lorin L Nicolson, Dan H. Nordman, Carl Norris, Dan SIDA 22(2): 1259. 2006 Noyes, Richard D. Oberlies, Nicholas H. Olmstead, Richard G. Orzell, Steve L Palmer, Michael W. Paratley, Robert Peck. lal pceom, Paul M poner ie Kristen E. Powell, A. Michael Pratt, Donald B. Pringle, James 5. Pruski, John F Pyne, Milo Qiu, Yin-Long Rabeler, Richard Robertson, Kenneth R. Rock, Janet H. Rohrer, Joseph R. Rossman, Amy Rumely, John I. Rutishauser, R. Ryburn, Adam K. Saar, Dayle E. Salazar, Jackeline Sanchez - Pino, Ivonne Sander ae var E Schultz, Joanna Schuyler, aed aes Semple, ee Shaw, Joe Sheviak, les Shulkina, Tatyana Siddiqi, Akma Simpson, Beryl B. Smith, Latimore Smith, Marian Smith, S. Galen Soltis, Douglas E. Steury, Mr. Brent Stevens, G.N. Stewart, Scott Syring, John 1259 Taylor, Charlotte M Taylor, David D. Telfair IT, Ray C. Thiede, Joachim Umber, Ray Van Horn, Gene S. Vander pool Staria Scott Wagner, Warren L. Wallnofer, Bruno Ware, Stewart Weakley, Alan S. Webster, Grad Weigend, Maximilian Wilken, Dieter H. Wilkie, Peter Williams, Justin Wilson, Barbara L. Wilson, Carol A. Wipff, Joseph K. Witsell, Theo Wunderlin, Richard P. Yang, TY. Aleck Yatskievych, George A. Yokota, Masatsugu udio, Sergio Zarucchi, James L. Zmarzty, S Zona, Scott A. 1260 BRIT.ORG/SIDA 22(2) INDEX TO 103 TITLES WITH 148 AuTHoRS: VOLUME 22 (2006) We know you have a choice. Thank you for choosing and supporting Sida, Contributions to Botany. A demographic and ecological analysis of Geocarpon minimum (Caryophyllaceae): a federally threatened species in southwest Mis- souri by Tim SmitH and JoserH S. ELy—22(2):1145 A new combination in Polytrichastrum (Bryophyta: Polytrichaceae) by G.L. Smit Mer- Ri —22(1):547 A new combination in Tetraphis (Bryophyta: Tetraphidaceae) by JupitH A. Harpe. —22(1): 549 A new combination in the Bromus catharticus complex (Poaceae: Bromeae sect. Cerato- chloa) by Ana Maria PLANCHUELO—22(1):555 A new hybrid genus and 11 new combina- tions in North American grasses by Mary E. BARKWORTH—22(1):495 A new name for Sida, Contributions to Botany (1962-2006) by Barney Liescomsa—22(2):847 A new narrowly endemic species of Clematis (Ranunculaceae: subgenus Viorna) from northeastern Texas by Dwayne Estes —22(1):65 Anew species of Boltonia (Asteraceae) from the Ridge and Valley physiographic province, U.S.A. by JOHN F. Townsenp and Vesna KARAMAN- CASTR —22(2):873 A new species of Dodecatheon (Primulaceae) from the northern coast range of Oregon and Washington by Kenton L. CHAmBeERS—22(1):461 A new species of Eugenia (Myrtaceae) from the Monteverde region, Costa Rica by Frep R. Barrit-—22(2):1071 Anew species of Pediomelum (Fabaceae) from the lower Piedmont Plateau of Georgia and South Carolina by James R. ALUSON, MICHAEL WAYNE Morris, and AsHtey N. EcAN—22(1):227 Anew species of Stenanthium (Melanthiaceae) from Tennessee, U.S.A. by B. Eucene Worrorp— 22(1):447 Anew species of waterlily (Nymphaea minuta: Nymphaeaceae) from Madagascar by KennetTH LANDON, RicHarD A. Eowarps, and P. Ivan Nozaic— 22(2):887 SIDA 22(2): 1260. 2006 A new species of witch-hazel (Hamamelis: Hamamelidaceae) apparently endemic to southern Mississippi by S.W. Leonarpb— 22(2):849 A quantitative study of styles and achenes of terminal and basal flowers — of Schoenoplectus hallii (Cyperaceae), a rare plant species of transient wetland habitats by Marian SMITH, SARA AMMANN, Nancy Parker, and Paice Meriier-CHERRY—22(2):1159 A vascular plant inventory of Starkey Wilderness Preserve, Pasco County, Florida by Emity Fercu- SON and RicHarb P. WuNDERLIN—-22(1):635 Amelichloa: a new genus in the Stipeae (Poaceae) by Mirta O. ArriaAca and Mary E. BaRKWORTH—22(1):145 An updated, annotated vascular flora of Caddo Parish, Louisiana, with notes on regional phy- togeography and ecology by Barsara R. MacRoserts and MicHaet H. MaAcRoserts —22(1):119] Annotated checklist of the vascular flora of the Big Sandy Creek Unit, Big Thicket National Pre- serve, Texas by Larry E. Brown, Barsara R MacRoserts, MicHAEL H. MacRoserts, PAUL A. Harcomee, Warren W. Pruess, |. SANDRA Etsik, and STANLEY D. JONES —22(1):705 Annotated checklist of the vascular flora of the Lance Rosier Unit, of the Big Thicket National Preserve, Hardin County, Texas by Larry E. Brown, BarsarA R. MacRoserts, MICHAEL H. MacRoserts, Paut A. Harcomee, Warren W.Pruess, |. SANDRA Etsik, and Stanvey D. Jones —22(2):1175 Big-fruited buckthorn, Sideroxylon macro- carpum (Sapotaceae),a long-forgotten Geor- gia endemic by James R. ALuison—22(1):243 Bromus ayacuchensis (Poaceae: Pooideae: Bromeae),a new species from Peru, with a key to Bromus in Peru by Jerrery M. Saareta, Paul M. Peterson, and Nancy F, Rerutio-Ropricuez— 22(2):915 INDEX Capraria mexicana (Scrophularia¢ eae) in Cam- eron County, Texas: rediscovered in the United States by AtFred RICHARDSON and Ken Kinc— 22(2):1235 Carex bicknellii (Cyperaceae) new to Arkansas by Pau M. McKenzie, C. THeo WirseLt, and Joe Woo eriGHT—22(1):801 Carex reznicekii, a new widespread species of Carex section Acrocystis (Cyperaceae) from eastern North America by Davin A. WerieR— 22(2):1049 Carex stipata (Cyperaceae), Chamaesyce gereyi (Euphorbiaceae), Eurytaenia texana (Apiaceae), Pediomelum esculentum (Fabaceae), and Talinum calycinum (Portulacaceae) deleted from the Louisiana flora by Barsara R. MacRoserts and Michael H. MacRoserts—22(2):1 221 Cayratia japonica (Vitaceae) new to North Carolina and an updated key to the genera of Vitaceae in the Carolinas by ALEXANDER KriNcs and Roseet J. RICHARDSON—22(1):813 Chromosome numbers for western and arctic North American species of Antennaria (Asteraceae: Gnaphalieae) by Jerry G CHMIELEWSKI—2.2(1):561 Clematis morefieldii (Ranunculaceae) new to Tennessee by Dwayne Estes and Curls FLeminc— 22(1):821 Comments on the Gerbera-complex (Asteraceae: Mutisieae) by Hans V. HANsen— 22(1):539 Crataegus series Bracteatae and Triflorae (Ro- saceae) by J.B. PHipes, R.LaNce, and K.A. Dvorsky 22(2):1009 Crataegus series Parvifoliae and its putative hybrids in the southeastern United States by J.B. Puipes and K. Dvorsky-—22(1):423 Dendrophorbium restingae (Asteraceae: Senecioneae), a new species from Sao Paulo, Brazil by Aristonio M. Tees, Jimi N. NakAJima, and Joao R. STEHMANN—22(1):123 Drepanostachyum falcatum var. seng- teeanum: identity and origins (Poaceae: SIDA 22(2): 1261. 2006 1261 Bambusoideae) by C.M.A. STAPLETON— 22(2):1081 Ericameria nauseosa subsp. ammophila (Asteraceae), a new rabbitbrush from the San Luis Valley of Colorado by Loran C. ANDERSON 22(2):866 Eriogonum callistum (Polygonaceae), a new species from the Tehachapi Mountains of Cali- fornia by James L. ReveAaL —22(2):857 Erodium malacoides (Geraniaceae), new to Texas by Davio E. Lemke and Jeo L. APLAcA— 22(2):1 245 Estudios en las Apocynaceae neotropicales XXIV: Bahiella (Apocynoideae: Echiteae), un desapercibido nuevo género endémico de Bahia, Brasil by J. Francisco Morates—22(1):333 Estudios en las Apocynaceae neotropicales XXV: novedades y nuevos reportes en las Apocynaceae (Apocynoideae: Rauvolfioideae) de Venezuela by J. Francisco MorAtes— 22(1):355 Evidence for hybridization between two sympa- tric violet species, Viola grahamii and V. hookeriana (Violaceae), in central Mexico by Aurea C. Cortts-Patomec and Harvey E. BALLARD, Jp,—22(2):1119 Exotic plant introduction in Kansas, two new species by Iracee BARNARO—22(1):777 Floristic diversity and composition of terra firme and seasonally inundated palm swamp for- ests in the Palma Real Watershed in lower Madre de Dios, Peru by FerNANbo H. CorNeJO VALVERDE, JOHN P JANovec, and MatHias W. TosLer— 22(1):615 Four novelties and a lectotypification in Matelea (Apocynaceae: Asclepiadoideae) from His- paniola by ALexaNDeR KRINGS—22(2):941 Fungi associated with Borrichia frutescens (Asteraceae): insect galls and endophytes by Diane Te StRAKE, Amy Hapbock Keacy, and Peter D. STILING = 22(1):755 Galium tricornutum (Rubiaceae) and Parentu- cellia viscosa (Scrophulariaceae) new to Okla- homa by Amy K.ButHoo and Bruce W. HoaclAND— 22(2):1235 1262 Gratiola brevifolia (Plantaginaceae) new to the flora of Delaware, the Delmarva Peninsula, and the Mid-Atlantic by Westey M. Knapp and Dwayni Estes —22(1):825 Green fringed orchid (Platanthera lacera, Orchidaceae) in southern Louisiana by Cuaries ALLEN, SARA THAMES, HOWARD ANDERSON III, BILL NEw- FON II], RHONDA Hampton, and Georce FisHeR— 22(1):805 Hedyotis australis (Rubiaceae) new to Missouri and Florida and related species in the south- central United States by Watter H. Lewis— 22(1):831 Herbivory of feral goats on Espiritu Santo Island, Gulf of California, Mexico by Jose Luis Leon-pe LA Luz and Reymunno Domincuez-CADENA— 22(2):1135 High levels of seed inviability among seven populations of the endangered Short's gold- enrod (Solidago shortii: Asteraceae) by Patrick J. Catie, Ropert F.C. Naczi, CHRISTINA SHACKLEFORD, aNd Jessica CaiceDO—22(1):735 Hill cane (Arundinaria appalachiana), a new species of bamboo (Poaceae: Bambusoideae) from the southern Appalachian Mountains by J.K. Tripcett, A.S. Weaxtey, and L.G. Ciark— 22(1):79 John W. Thieret (1926-2005) by Rateu L. THOMPSON—2.2(1):3 John W.Thieret, a student's perspective (1967- 2005) by Rosert R. HAyNeS—22(1):25 John W. Thieret, associate editor of Sida (1972- 2005) by Barney Lipscomp—22(1):55 John W. Thieret, colleague and editor friend (1969-2005) by Wittiam F.MaHter—22(1):51 John W. Thieret, Dedication—22(1):1 John W.Thieret, the curator by Rosert F.C.Naczi— 22(1):21 John W.Thieret, valuable botanical friend (1965— 2005) by Ronato L. Stuckey —22(1):33 More American black sapotes: new Diospyros (Ebenaceae) for Mexico and Central America by Mircuett C. Provance and Anorew C. SANDERS— 22(1):279 SIDA 22(2): 1262. 2006 BRIT.ORG/SIDA 22(2) Morphometric analysis of an Amelanchier (Ro- saceae: Maloideae) complex on the Delmarva Peninsula (Delaware, Maryland and eastern Virginia) resolves the taxonomic identities of Amelanchier obovalis and A. canadensis by CHRISTOPHER T. Frye —2.2(2):1027 Naturalization and extirpation of water hyacinth (Eichhornia crassipes, Pontederiaceae) in southwestern Arkansas, U.S.A. by RENN TUMLISON and Brett Serviss—22(1):749 New and disjunct records of Eleocharis liesneri (Cyperaceae) from South America by Davio J. RoseN—22(1):781 New and noteworthy collections for Arkansas by James H. Peck and Brett E. Serviss —22(1):817 New and significant records of vascular plants for Florida and for Collier County and Lee County, Florida by Grorae J. Witber and MartHa R. McComas—22(1):787 New combinations in the genus Cymopterus (Apiaceae) of the southwestern United States by Ronato L. HarTMAN—22(2):955 New distribution records for Eupatorieae (Asteraceae) in the United States by Guy L. Nesom and Bittie L. Turner —22(2):1 249 New species of Myrtaceae from Ecuador and Peru by Bruce K. Hoist and Maria LUCIA KAWASAKI— 22(2):931 New taxa and combinations in cultivated bam- boos (Poaceae: Bambusoideae) by C.M.A. STAPLETON—2.2(1):331 New vascular plant records for New Mexico by RONALD L. HARTMAN, BriAN Rel, B.E. Nevson, and BriAN JACOBS—=2.2(2):1225 Notes on southwestern Moraceae by ALAN T. WHITTEMORE—2.2(1):769 Notes on the life and work of James Brigham McFarlin, Florida botanist by Tom Patmer— 22(1):608 Notes on types in Apocynaceae: Asclepia- doideae in Cuban herbaria and four lectotypi- fications in West Indian Gonolo-binae by ALEXANDER KRINGS and Paut R. Fantz—22(1):533 Novedades en Gentianaceae para América del INDEX Sur by Eva M. FivippA and Gtoria E. BARBOZA— 22(1):129 Poa matri-occidentalis (Poaceae: Pooideae: Poeae:Poinae),a new species from Mexico by Pau M. Peterson, Rosert J. SorENG, aNd YOLANDA Herrera ARRIETA —22(2):904 Psidium cauliflorum (Myrtaceae), a new spe- cies from Bahia, Brazil by Leste R. LanbruM and Marcos SopraL—22(2):927 Quadruple, triple, double, and simple pappi in the goldenasters, subtribe Chrysopsidinae (Asteraceae: Astereae) by JoHn C. SemPLe— 22(1):503 Range expansion of Hexalectris grandiflora (Orchidaceae) in Texas by MarcaretT BROWN- Marsden and Anne B. CoLLins—22(2):1239 Rejection of lectotypification of Aster amellus (Asteraceae: Astereae) and selection of a new lectotype by JoHN C. Semele —22(2):1087 Review of Crataegus series Pulcherrimae (Ro- saceae) by J.B. PHipes, RJ. O’Kennon, and KA. Dvorsky —22(2):973 Revision of Bauhinia subgenus Bauhinia sec- tion Amaria (Cercideae: Caesalpinioideae: Fabaceae) by RicHarp P. WuNDERLIN—22(1):97 Root-shoot anatomy and post-harvest vegeta- tive clonal development in Lophophora williamsii (Cactaceae: Cacteae): implica- tions for conservation by Martin Terry and JAMES D. MausetH—22(1):565 Sexual and apomictic prairie fleabane (Erigeron strigosus) in Texas: geographic analysis and a new combination (Erigeron strigosus var. traversii, Asteraceae) by RicHarb D. Noyes, Helo! GERLING, and CarLA VANDERVOORT—22(1):265 Seymeria falcata (Scrophulariaceae), a new record for Texas and the United States by JOSELYN FENSTERMACHER——22(1):8 11 Six new combinations in Dodecatheon (Primulaceae) by James L. ReveAt —22(2):863 Six new species of Sphagnum (Bryophyta: Sphagnaceae) from North America by RicHarD E. ANoRUS—2.2(2):959 SIDA 22(2): 1263. 2006 1263 Taxonomic overview of the Heterotheca villosa complex (Asteraceae: Astereae) by Guy L. Nesomg—22(1):367 Taxonomic review of Symphyotrichum patens (Asteraceae: Astereae) by Guy L. Nesom— 22(2):1075 Taxonomy and conservation of medicinal plants in canal-irrigated areas of Punjab, Pakistan by KHALID Faroog Aksar and MoHAMMAD ATHAR— 22(1):593 Taxonomy of Cuscuta gronovii and Cuscuta umbrosa (Convolvulaceae) by MiHAl Coste, Guy L. Nesom, and Sasa STEFANOVIC—22(1):197 Taxonomy of Lantana sect. Lantana (Verbenaceae): |. correct application of Lan- tana camara and associated names by Roctr W. SANDERS —22(1):381 Taxonomy of North American species of Oldenlandia (Rubiaceae) by Eowarn E. Terret and Harotp Rosinson—22(1):305 Taxonomy of the Cuscuta indecora (Convo- lvulaceae) complex in North America by Munat Coste, Guy L.Nesom, and Sasa STEFANOVIC— 22(1):209 Taxonomy of the Cuscuta pentagona complex (Convolvulaceae) in North America by Mina\ Costea, Guy L. Nesom, and Sasa STEFANOVIC— 22(1):151 Taxonomy of the Cuscuta salina-californica complex (Convolvulaceae) by Mixai Costes, Guy L. Nesom, and Sasa SterANOVIC—22(1):177 The lectotypification and 19" century history of Croton alabamensis (Euphorbiaceae s.s.) by KENNETH J. WurDACK—2.2(1):469 The north-south transition of flora across Arkan- sas:a preliminary phytogeographical analysis by MicHaeL H. MacRoserts and Bargara R. MacRoserts—22(1):725 Thomas Walter Typification Project, |. Observa- tions on the John Fraser folio by Daniet B. Ward—22(2):1111 Trichome morphology in selected Mexican red oak species (Quercus section Lobatae) by M. Lucia VAZQUEZ —22(2):1091 Two new combinations in Peyritschia (Poaceae: wa 1264 Pooideae: Aveninae) by Victor L. Finot, Paut M. PETERSON, aNd FERNANDO O. ZULOAGA—2.2(2):895 Two new species of Elymus (Poaceae) in the southern U.S.A. and other notes on North American Elymus species by Jutian J.N. CamPBetL—22(1):485 Type localities of vascular plants first described from Ohio: supplement by James S. Princie— 22(1):765 BRIT.ORG/SIDA 22(2) Validation of the name Dahlia sublignosa (Asteraceae) by Dayie E. Saar and Paut D. SORENSEN=2.2(1):545 Vascular floras of Sonoita Creek State Natural Area and San Rafael State Park: Arizona’s first natural-area parks by Steven P. McLauGHLiN— 22(1):661 Zeuxine pantlingjii, sp. nov. (Orchidaceae), a new species from India by Avistiek BHATTACHARIEE and H.J. CHOWDHERY—22(2):935 INDEX OF 148 AUTHORS: VOLUME 22 (2006) Thank you for choosing Sida, Contributions to Botany. Akbar, Khalid Farooq—22(1):593 Allen, Charles —22(1):805 Allison, James R.—22(1):227, 243 Ammann, Sara—22(2):1 159 Anderson Ill, Howard—22(1):805 Anderson, Loran C.—22(2):866 Andrus, Richard E.—22(2):959 Aplaca, Jed L—22(2):1 245 Arriaga, Mirta O.—22(1):145 22(1):593 Athar \Mohammad Ballard, Jr, Harvey E.—22(2):1119 Barboza, Gloria E—22(1):129 Barkworth, Mary E.—22(1):145, 495 Barnard, lralee—22(1):777 Barrie, Fred R.—22(2):1071 Bhattacharjee, Avishek—22(2):935 Brown, Larry E—22(1):705; 22(2):1175 Brown-Marsden, Margaret—22(2):1239 Buthod, Amy K.—22(2):1235 Caicedo, Jessica—22(1):735 Calie, Patrick J—22(1):735 Campbell, Julian J.N.—22(1):485 Chambers, Kenton L.—22(1):461 Chmielewski, Jerry G.—22(1):561 Chowdhery, H.J—22(2):935 Clark, L.G—22(1):79 Collins, Anne B.—22(2):1239 Cornejo Valverde, Fernando H.—22(1):615 Cortés-Palomec, Aurea C—22(2):1119 Costea, Mihai—22(1):151, 177, 197, 209 sees SIDA 22(2): 1264. 2006 Dominguez-Cadena, Reymund 22(2): 1135 Dvorsky, KA.—22(1):423; 22(2):973, 1009 Edwards, Richard A—22(2):887 Egan, Ashley N.—22(1):227 Elsik, |. Sandra—22(1):705; 22(2):1175 Ely, Joseph S$ —22(2):1145 Estes, Dwayne—22(1):65, 821,825 Fantz, Paul R—22(1):533 Fenstermacher, Joselyn—22(1):811 Ferguson, Emily—22(1):635 Filippa, Eva M.—22(1):129 Finot, Victor L—22(2):895 Fisher, George—22(1):805 Fleming, Chris—22(1):821 Frye, Christopher T.—22(2):1027 Gerling, Heidi —22(1):265 Hampton, Rhonda—22(1):805 Hansen, Hans V.—22(1):539 Harcombe, Paul A.—22(1):705;22(2):1175 Harpel, Judith A—22(1):549 Hartman, Ronald L.—22(2):955, 1225 Haynes, Robert R—22(1):25 Herrera Arrieta, Yolanda—22(2):904 Hoagland, Bruce W.—22(2):1 235 Holst, Bruce K.—22(2):931 Jacobs, Brian—22(2):1225 Janovec, John P-—22(1):615 Jones, Stanley D.—22(1):705; 22(2):1175 2873 — Karaman-Castro, Vesna—22(2 INDEX Kawasaki, Maria Lucia—22(2):931 Keagy, Amy Haddock—22(1):755 King, Ken—22(2):1239 Knapp, Wesley M.—22(1):825 Krings, Alexander—22(1):533, 813; 22(2):941 Lance, R.—22(2):1009 Landon, Kenneth—22(2):887 Landrum, Leslie R—22(2):927 Lemke, David E.—22(2):1245 Leonard, S.W.—22(2):849 Lewis, Walter H—22(1):831 Lipscomb, Barney—22(1):55; 22(2):847 Luis Leén-de la Luz, José—22(2):1135 MacRoberts, Barbara R.—22(1):/05, 725; 22(2):1175, 1191, 1221 MacRoberts, Michael H.—22(1):705, 725; 22(2):1175,1191,1221 Mahler, William F—22(1):51 Mauseth, James D.—22(1):565 McCombs, Martha R.—22(1):787 McKenzie, Paul M.—22(1):801 McLaughlin, Steven P-—22(1):661 Merrill, L. Smith—22(1):547 Mettler-Cherry, Paige—22(2):1159 Morales, J. Francisco—22(1):333, 355 Morris, Michael Wayne —22(1):227 Naczi, Robert F.C.—22(1):21, 735 Nakajima, Jimi N.—22(1):123 Nelson, B.E—22(2):1225 Nesom, Guy L.—22(1):151, 177, 197, 209, 367; 22(2):1075, 1249 Newton Ill, Bill —22(1):805 Noyes, Richard D.—22(1):265 Nozaic, P. lvan—22(2):887 O’'Kennon, R.J.—22(2):973 Palmer, Tom—22(1):608 Parker, Nancy—22(2):1 159 Peck, James H—22(1):817 Peterson, Paul M.—22(2):895, 904,915 Phipps, J.B. —22(1):423; 22(2):973, 1009 Planchuela, Ana Maria—22(1):555 Pringle, James S—22(1):765 SIDA 22(2): 1265. 2006 1265 Provance, Mitchell C.—22(1):279 Pruess, Warren W.—22(1):705;22(2):1175 Refulio-Rodriguez, Nancy F—22(2):915 Reif, Brian —22(2):1225 Reveal, James L.—22(2):857, 863 Richardson, Alfred —22(2):1 239 Richardson, Robert J.—22(1):813 Robinson, Harold —22(1):305 Rosen, David J.—22(1):781 Saar, Dayle E—22(1):545 Saarela, Jeffery M—22(2):915 Sanders, Andrew C.—22(1):279 Sanders, Roger W.—22(1):381 Semple, Jonn C.—22(1):503; 22(2):1087 Serviss, Brett E-—22(1):749, 817 Shackleford, Christina—22(1):735 Smith, Marian—22(2):1 159 Smith, Tim—22(2):1145 Sobral, Marcos—22(2):927 Soreng, Robert J.—22(2):904 Se@rensen, Paul D.—22(1):545 Stapleton, C.M.A.—22(1):331;22(2):1081 STEFANOVIC, SASA—2.2(1):151, 177, 197, 209 Stehmann, Jodo R.—22(1):1 23 Stiling, Peter D—22(1):755 Stuckey, Ronald L—22(1):33 Te Strake, Diane —22(1):755 Teles, AristOnio M.—22(1):123 Terrell, Edward E.—22(1):305 Terry, Martin—22(1):565 Thames, Sara—22(1):805 Thompson, Ralph L—22(1):3 Tobler, Mathias Wi—22(1):615 Townsend, John F—22(2):873 Triplett, J.K—22(1):79 Tumlison, Renn—22(1):749 Turner, Billie L—22(2):1 249 Vandervoort, Carla—22(1):265 Vazquez, M._Lucfa—22(2):1091 Ward, Daniel B.—22(2):1111 Weakley, A.'S.—22(1):79 Werier, David A.—22(2):1049 Whittemore, Alan T.—22(1):769 1266 Wilder, George J.—22(1):/87 Witsell, C. Theo—22(1):801 Wofford, B.Eugene—22(1):447 BRIT.ORG/SIDA 22(2) Woolbright, Joe—22(1):801 Wunderlin, Richard P—22(1):97, 635 Wurdack, Kenneth J—22(1):469 Zuloaga, Fernando O.—22(2):895 BOTANICAL NAMES AND SuBJECT INDEX: VOLUME 22 (2006) New names (97) in bold face Achnatherum arnowiae—22(1):496 Acrocystis—22(2):1049 Ageratina thyrsiflora—22(2):1250 Alternaria (fungus) —22(1):755 Amaranthus blitum (Arkansas record)— 22(1):817 Amblysperma spathulata—22(1):539 Amelanchier canadensis—22(2):1027 obovalis—22(2):1027 Amelichloa—22(1):146 ambigua—22(1):147 brachychaeta—22(1):147 brevipes—22(1):147 caudata—22(1):148 clandestina—22(1):148 hi Wi = —22(1):496 Antennaria—22(1):561 Apiaceae—22(2):955 Apocynaceae—22(1):333, 355, 533:22(2):941 Apocynoideae—22(1):333, 355 Appalachian Mountains—22(1):79, 873 Arizona—22(1):661 ArkansaS—22(1):725, 817, 749; 22(2):801 Arundinaria—22(1):79 appalachiana—22(1):88 gigantea—22(1):79 falcata var. glomerata—22(2):1083 Asclepiadoideae—22(1):533, 941 Asphondylia borrichiae (midge) —22(1):755 Aspidosperma rigidum—22(1):363 schultesii—22(1):364 Aster amellus—22(2):1087 Asteraceae —22(1):367, 503, 539,545, 735, 755, 22(2):866, 1075, 1087, 1249 Aveninae—22(2):895 SIDA 22(2): 1266. 2006 Bahia, Brasil—22(1):333, 927 Bahiella—22(1):342 blanchetii—22(1):342 infundibuliflora—22(1):345 Bamboos—22(1):331 Bambusoideae—22(1):79, 331, 1081 Bauhinia —22(1):97 amatlana—22(1):99 andrieuxiim22(1):101 arborea—22(1):102 augusti—22(1):104 ayabacensis—22(1):105 beguinotii—22(1):107 var. gorgonae—22(1):108 brachycalyx—22(1):107 Cookii—22(1):110 geniculata—22(1):111 haughtii—22(1):113 petiolata—22(1):114 var. caudigera—22(1):115 picta—22(1):116 section Amaria—22(1):97 seleriana—22(1):117 seminarioi—22(1):118 Series Stenanthae—22(1):1 20 stenantha—22(1):121 weberbaueri—22(1):120 Big Sandy Creek Unit (Big Thicket, Texas) — 22(1):705 Big Ticket National Preserve (Texas) —22(1):705, 22(2):1175 Big-Fruited Buckthorn—22(1):243 Black Sapotes—22(1):277 Boltonia asteroids var. asteroides—22(2):884 caroliniana—22(2):884 montana—22(2):874 INDEX Borinda angustissima—22(1):332 contracta—22(1):332 nujiangensis—22(1):332 utilis —22(1):332 Borrichia frutescens—22(1):755 Bradburia—22(1):503 Brazil—22(1):123 (Sao Paulo), 333 Bromeae—22(1):555,915 Bromus—22(1):555 ayacuchensis—22(2):919, 922 berteroanus—22(2):922 catharticus—22(2):923 var. elata—22(1):556 cebadilla—22(2):922 coloratus—22(2):923 flexuosus—22(2):923 lanatus—22(2):923 modestus—22(2):923 pitensis—22(2):923 segetum—22(2):923 Striatus—22(2):922 villosissimus—22(2):923 Bryophyta—22(1):547, 549, 959 Cactaceae—22(1):565 Cacteae—22(1):565 Caddo Parish (Louisiana) —22(2):1193 Caesalpinoideae—22(1):97 California—22(2):85 7 Calyptranthes manuensis—22(2):93 1 Cameron County (Texas) —22(2):1239 Capraria mexicana—22(2):1 239 Carex bicknellii—22(1):801 gigantea X Carex lupuliformis—22(1):787 reznicekii—22(2):1050 stipata—22(2):1 221 Carolinas—22(1):813 Caryophyllaceae—22(2):1 145 Cayratia japonica—22(1):813 Central AmMerica—22(1):277 Ceratochloa—22(1):555 Cercideae—22(1):97 Chamaesyce geyeri—22(2):1 221 SIDA 22(2): 1267. 2006 1267 hypercifolia (Arkansas record) —22(1):817 opthalmica (Arkansas record) —22(1):817 Chaptalia hintonii—22(1):539 Chelonanthus viridiflorus—22(1):137 Chromolaena bigelovii—22(2):1 252 Chromosome numbers—22(1):561 Chrysopsidinae—22(1):503 Chrysopsis—22(1):503 Cicendia quandrangularis—22(1):1 33 Clematis—22(1):65 addisonii—22(1):65 carrizoensis—22(1):67 glaucophylla—22(1):65 morefieldiim22(1):821 texensis—22(1):65 versicolor—22(1):65 Collier County (Florida) —22(1):787 Convolvulaceae—22(1):151,177, 197, 209 Costa Rica—22(2):1071 Crataegus ashei—22(2):1012 austromontana—22(2):1016 brittonii—22(1):423 eximia—22(2):973 gilva—22(2):973 harbisonii—22(2):1012 mendosa—22(2):973 opima—22(2):973 pallens—22(2):973 pinetorum—22(2):973 pulcherrima—22(2):973 sargentii—22(2):973 series Bracteatae—22(2):1009 series Parvifoliae—22(1):423 series Pulcherrimae—22(2):973 series Triflorae—22(2):1009 tecta—22(2):973 triflora—22(2):1018 uniflora—22(1):423 X vailiae—22(1):423 venusta—22(2):973 Croptilon—22(1):503 Croton alabamensis—22(1):469 Cuban Herbaria—22(1):533 Curtia tenuifolia—22(1):133 Cuscuta—22(1):151 1268 californica—22(1):1 77 campestris—22(1):15] coryli—22(1):209 glabrior—22(1):151 gronovii—22(1):197 harperi—22(1):151 howelliana—22(1):177 indecora—22(1):209 var. attenuata—22(1):216 jepsonli—22(1):209 occidentalis —22(1):177 pentagona—22(1):151 plattensis—22(1):151 polygonorum—22(1):151 runyonii—22(1):151] Salina—22(1):177 sandwichiana—22(1):151 Subinclusa—22(1):1 77 Suksdorfii—22(1):1 77 umbrosa—22(1):197 warneri—22(1):209 Cymopterus breviradiatus—22(2):955 davidsonii—22(2):956 glomeratus var. fendleri—22(2):956 sessiliflorus—22(2):956 Cyperaceae—22(1):/81,801,22(2): 1049 Cyperus nigromarginata—22(2):1049 umbellata—22(2):1049 Dahlia—22(1):545 sublignosa—22(1):545 Delaware—22(1):825; 22(2):1027 Delmarva Peninsula—22(1):825;22(2):1027 Dendrophorbium paranense—22(1):127 restingae—22(1):123 Diospyros conzattii—22(1):287 costaricensis—22(1):291 gomeziorum—22(1):282 pergamentacea—22(1):297 riojae—22(1):279 tuxtlensis—22(1):295 SIDA 22(2): 1268. 2006 BRIT.ORG/SIDA 22(2) Dodecatheon—22(1):461; 22(2):863 austrofrigidum—22(1):462 clevelandii var. gracile—22(2):863 var. insulare—22(2):863 var. patulum—22(2):863 frigidum—22(1):461 pulchellum var, Macrocarpum—22(2):863 var. Shoshonense—22(2):864 utahense—22(2):864 Drepanostachyum falcatum—22(2):1084 var. sengteeanum—22(1):332; — 22(2):1081 Ebenaceae—22(1):277 Ecuador—22(2):931 Eichhornia crassipes—22(1):749 Eleocharis liesneri—22(1):781 x Elyleymus hultenii—22(1):496 mossii—22(1):497 ontariensis—22(1):497 Elymus—22(1):485 xX cayouetteorum—22(1):498 churchii—22(1):486 hystrix—22(1):489 lanceolatus subsp. riparius—22(1):498 pringlei—22(1):485 texensis—22(1):488 Endophytes—22(1):755 Ericameria nauseosa subsp. ammophila— 22(2):868 Erigeron strigosus—22(1):265 var. traversii—22(1):273 Eriogonum callistum—22(2):857 Sect. Lanocephala—22(2):857 Erodium malacoides—22(2):1 245 Espiritu Santo Island—22(2):1135 Eugenia haberi—22(2):1071 yasuniana—22(2):934 Eupatorieae—22(2):1249 Eupatorium madrense—22(2):1253 INDEX Euphorbia graminea (Arkansas record)— 22(1):817 Euphorbiaceae—22(1):469 Eurytaenia texana—22(2):1221 Exotic Plant Introductions (Kansas) —22(1):777 Fabaceae —22(1):97 Fargesia apicirubens—22(1):331 Feral Goats—22(2):1135 Ficus palmata—22(1):/769 pseudos-carica—22(1):769 Florida—22(1):635, 787 Botanist—22(1):607 Fraser, John (Folio) —22(2):1111 Galium tricornutum (Oklahoma record)— 22(2):1235 Gentiana prostrata—22(1):133 Gentianaceae—22(1):129 Gentianella cosmantha—22(1):135 fabrisii—22(1):130 magellanica—22(1):135 multicaulis—22(1):136 ottonis—22(1):136 Geocarpon minimum—22(2):1145 Georgia—22(1):227, 243 Geraniaceae—22(2):1245 Gerbera—22(1):539 Goldenasters—22(1):503 Gonolobinae—22(1):533 Gratiola brevifolia—22(1):825 Green Fringed Orchid—22(1):805 Gulf of California, Mexico—22(2):1135 _ Hamamelidaceae—22(2):849 Hamamelis—22(2):849 macrophylla—22(2):855 ovalis—22(2):850, 854 vernalis—22(2):854 virginiana—22(2):854 Hardin County (Texas) —22(2):1175 Hedyotis australis—22(1):831 corymbosa (Arkansas record) —22(1):81 7 SIDA 22(2): 1269. 2006 1269 Heterotheca—22(1):503 depressa—22(1):379 horrida—22(1):378 polothrix—22(1):373 polothrix—22(1):379 sierrablancensis—22(1):374, 379 stenophylla—22(1):377 villosa—22(1):367 var. angustifolia—22(1):376 Hexalectris grandiflora—22(2):1 239 Hill Cane—22(1):79 Hispaniola—22(2):941 Jatropha cinerea—22(2):1142 India—22(2):935 Indocalamus hamadae—22(1):332 Insect Galls—22(1):/55 Ipomoea quamoclit (Arkansas record)— 22(1):81/7 Kansas—22(1):777 Koanophyllon palmeri—22(2):1251, 1252 solidaginifolium—22(2):1252 Lance Rosier Unit (Big Thicket, Texas)— 22(2):1175 Lantana—22(1):381 camara—22(1):381 subsp. aculeata—22(1):394 nivea subsp. mutabilis—22(1):395 strigocamara—22(1):392 Lee County (Florida) —22(1):787 Leibnitzia—22(1):539 Lespedeza thunbergii—22(1):/77 xLeydeum littorale—22(1):498 Leymus californicus—22(1):498 Lilaeopsis schaffneriana ssp. recurva—22(1):661] Lobatae (Quercus) —22(2):1091 Lophophora williamsii—22(1):565 Louisiana—22(1):805; 22(2):1193, 122] Madagascar—22(2):887 Madre de Dios, Peru—22(1):615 Mahler, Wm. F—22(2):847 Maloideae—22(2):1027 Mandevilla angustata—22(1):356 1270 boliviensis—22(1):359 tristis—22(1):359 vanheurckii—22(1):362 Maryland—22(2):1027 Matelea—22(2):941 corynephora—22(2):942 phainops—22(2):948 rhynchocephala—22(2):949 torulosa—22(2):951 McFarlin, James Brigham—22(1):607 Medicinal Plants —22(1):593 Melanthiaceae—22(1):447 Mexican Red Oaks—22(2):1091 Mexico—22(1):277; 22(2):904, 1119 (central) Mississippi—22(2):849 Missouri (record) —22(1):831; 22(2):1145 (southwest) Monteverde Region (Costa Rica) —22(2):1071 Moraceae—22(1):769 Morus nigra—22(1):/69 Mutisieae—22(1):539 Myagrum perfoliatum—22(1):777 Myrtaceae —22(2):927,931, 1071 New Mexico (records) —22(2):1225 North America Grasses—22(1):495 North Carolina—22(1):813 North-South Transition of Flora—22(1):725 Noticastrum—22(1):503 Nymphaea minuta—22(2):887 Nymphaeaceae—22(2):887 Odontadenia killipii—22(1):362 Ohio—22(1):765 Oldenlandia—22(1):305 boscii—22(1):305 corymbosa—22(1):305 drymarioides—22(1):305 lancifolia—22(1):305 microtheca—22(1):305 ovata—22(1):305 pringlei—22(1):305 salzmannii—22(1):305 uniflora—22(1):305 Opuntia tapona—22(2):1139 Orchidaceae—22(1):805, 935 SIDA 22(2): 1270. 2006 BRIT.ORG/SIDA 22(2) Oregon—22(1):461 Osbertia—22(1):503 Pakistan —22(1):593 Palm swamp forests—22(1):615 Palma Real River—22(1):615 Pantling, Robert—22(2):935, 939 (portrait) Parentucellia viscosa (Oklahoma record)— 2212 )51 35 Parietaria floridana 22(1):817 Pasco County, Florida—22(1):635 x Pascoleymus—22(1):499 bowdenii —22(1):499 Pediomelum—22(1):227 esculentum—22(2):1222 piedmontanum—22(1):229 Pennellia robinsonii—22(1):661 Peru—22(1):615;22(2):915, 931 Peyritschia conferta—22(2):902 deyeuxioides—22(2):902 howellii—22(2):897; 902 humilis—22(2):902 koelerioides—22(2):901 pinetorum—22(2):899; 901 pringlei—22(2):902 Phacelia sonoitensis—22(1):661 Piedmont Physiographic Province —22(1):227 Plantaginaceae—22(1):825 Platanthera lacera—22(1):805 Poa annua—22(2):912 bigelovii—22(2):912 compressa—22(2):913 fendleriana ssp. albescens—22(2):913 ssp. fendleriana—22(2):913 infirma—22(2):912 matri-occidentalis—22(2):906 subsp. mohinorensis—22(2):911 mulleri—22(2):91 3 orizabensis—22(2):91 2 pratensis—22(2):913 strictiramea—22(2):913 Poaceae—22(1):79, 145, 485, 495, 555: 22(2):1081, 895, 904,915 (Arkansas record)— INDEX Poeae—22(2):904 Poinae—22(2):904 Polygonaceae—22(2):857 Polytrichaceae—22(1):547 Polytrichastrum alpinum var. sylvaticum— 22(1):547 Pontederiaceae—22(1):749 Pooideae—22(2):895,915 Prairie Fleabane—22(1):265 Primulaceae—22(1):461, 863 Pseudoroegneria spicata f.inermis—22(1):499 f. pubescens—22(1):499 Psidium cauliflorum—22(2):927 Punjab, Pakistan —22(1):593 Quercus —22(2):1091 Rabbitbrush—22(2):866 Ranunculaceae —22(1):65, 821 Rhynchosia minima—22(1):661 Ridge and Valley Physiographic Province— 22(2):873 Rosaceae—22(2):973, 1009, 1027 Rubiaceae —22(1):831 San Luis Valley (Colorado) —22(2):866 San Rafael State Park (Arizona) —22(1):661 Sapotes, Black—22(1):277 Schoenoplectus hallii—22(2):1159 Schultesia guianensis—22(1):140 Scrophulariaceae—22(1):3811 Senecioneae—22(1):1 23 Seymeria falcata—22(1):811 Shinners, Lloyd H—22(2):847 Short’s Goldenrod—22(1):735 Sibara virginica—22(1):661 Sida, Contributions to Botany —22(2):847 Sideroxylon—22(1):243 lanuginosa—22(1):243 macrocarpum—22(1):245 Solidago shortii—22(1):735 Sonoita Creek State Natural Area (Arizona)— 22(1):661 South America—22(1):781 South Carolina—22(1):227 Southwestern Arkansas—22(1):749 SIDA 22(2): 1271. 2006 1271 Southwestern U.S.A—22(1):769, 955 Sphagnaceae—22(2):959 Sphagnum beothuk—22(2):966 bergianum—22(2):964 kenaiense—22(2):961 mcqueenii—22(2):959 sitchense—22(2):969 talbotianum—22(2):970 Starkey Wilderness Preserve—22(1):635 Stenanthium diffusum—22(1):450 Stevia salicifolla—22(2):1 249 Stipeae—22(1):145 Symphyotrichum patens—22(2):1075 var. gracile—22(2):1076 var, patens—22(2):1076 var, patentissinum—22(2):1075 Talinum calycinum—22(2):1 222 Tehachapi Mountains—22(2):857 Tennessee—22(1):447, 821 Tetraphidaceae—22(1):549 Tetraphis pellucida—22(1):551 var. trachypoda—22(1):551 Texas—22(1):65 (northeastern), 265, 705, 811; 22(2)31 175, 1239, 1259, 1245 Thieret, John W—22(1):1,3, 21,25, 33,51,55 Tomentaurum—22(1):503 Trichocline—22(1):539 hieracioides—22(1):539 Trichome morphology—22(2):1091 Type Localities of Ohio Plants—22(1):765 United Status (record) —22(1):811 Venezuela—22(1):355 Verbenaceae—22(1):381 Viola grahamii—22(2):1119 hookeriana—22(2):1119 Violaceae—22(2):1119 Viorna (subgenus)—22(1):65 Virginia (Eastern) —22(2):1027 Vitaceae—22(1):813 Walter, Thomas—22(2):1111 Washington—22(1):461 Water Hyacinth—22(1):749 — 1272 Waterlily—22(2):887 West Indies —22(1):533 Witch-Hazel—22(2):849 BRIT.ORG/SIDA 22(2) Zeuxine goodyeroides—22(2):935 pantlingii—22(2):935 Zygostigma australe—22(1):1 36 97 New NAMES AND NEw COMBINATIONS: Votume 22 (2006) Achnatherum arnowiae (S.L. Welsh & N.D. At- wood) Barkworth, comb. nov.—22(1):496 Amelichloa Arriaga & Barkworth, gen. nov.— 22(1):146 Amelichloa ambigua (Speg.) Arriaga & Barkworth, comb. nov.—22(1):147 Amelichloa brachychaeta (Godr.) Arriaga & Barkworth, comb. nov.—22(1):147 Amelichloa brevipes (£. Desv.) Arriaga & Barkworth, comb. nov.—22(1):147 Amelichloa caudata (Trin.) Arriaga & Barkworth, comb. nov.—22(1):148 Amelichloa clandestina (Hack.) Arriaga & Barkworth, comb. nov.—22(1):148 Ammophila breviligulata subsp. champlainensis (F. Seym.) Walker, Paris & Barrington ex Barkworth, comb. nov.— 22(1):496 Arundinaria appalachiana Triplett, Weakley & L.G. Clark, sp. Nov.—22(1):88 Bahiella J.F Morales, gen. nov.—22(1):342 Bahiella blanchetii (A. DC.) J.- Morales, comb. nov.—22(1):342 Bahiella infundibuliflo 22(1):345 Bauhinia amatlana Wunderlin, sp. nov.— 22(1):99 Bauhinia arborea Wunderlin, sp. nov.— 22(1):102 Bauhinia ayabacensis Wunderlin, sp. nov.— 22(1):105 Bauhinia geniculata Wunderlin, sp. nov.— 22(1):1 11 Bauhinia petiolata var. caudigera (S.F Blake) Wunderlin, comb. nov.—22(1):115 Boltonia montana JF Townsend & V. Karaman- Castro, Sp. NOV.—22(2):874 ).F. Morales, sp.nov,.— SIDA 22(2): 1272. 2006 Borinda angustissima (1.P.Yi) Stapleton, comb. NOV.—22(1):332 Borinda contracta (T.P. Yi) Stapleton, comb. NOV.—22(1):332 Borinda nujiangensis (Hsueh & C.M. Hui) Stapleton, comb. nov.—22(1):332 Borinda utilis (TP. Yi) Stapleton, comb. nov.— 22(1):332 Bromus ayacuchensis Saarela & P.M. Peterson, Sp.nov.—22(2):919 Bromus catharticus var. elata (£. Desv.) juelo, comb. nov.—22(1):556 — Plane Calyptranthes manuensis 8. Holst & ML. Kawasaki, Sp NOv.—22(2):93 | Carex reznicekii Werier, 5p. nov.—22(2):1050 Clematis carrizoensis D. Estes, sp. nov.— 22(1):67 Cuscuta indecora var. attenuata (Waterfall) Costea, comb. & stat. nov.—2.2(1):216 Cymopterus breviradiatus (W.L. Theob. & C.C. Tseng) R.L. Hartm., comb. & stat. nov.— 22(2):955 Cymopterus davidsonii (J.M.Coult.& Rose Hartm., comb. nov.—22(2):956 Cymopterus glomeratus var.fendleri (A. Gray) R.L. Hartm., comb. & stat. nov.—22(2):956 Cymopterus sessiliflorus (W.L. Theob. & C.C. Tseng) R.L. Hartm., comb. nov.—22(2):956 Dahlia sublignosa (P.D.Sarensen) D.E.Saar & PD. S@rensen, comb. & stat. nov.—22(1):545 Dendrophorbium restingae A Jeles, J. N.Nakaj. & Stehmann, sp. nov.—22(1):1 23 Diospyros costaricensis M.C. Provance & A.C. Sanders, sp. nov.—22(1):291 Diospyros gomeziorum M.C. Provance & AC. Sanders, sp. nov.—22(1):282 — R.L. INDEX Diospyros tuxtlensis M.C. Provance & A.C. Sand- ers, SO. NOV.—22(1):295 Dodecatheon austrofrigidum K.L. Chambers, sp. NOV.—22(1):462 Dodecatheon clevelandii var. gracile (Greene) Reveal, comb. nov.—22(2):863 Dodecatheon clevelandii var. insulare (HJ. Thomps.) Reveal, comb. nov.—22(2):863 Dodecatheon clevelandii var. patulum (Kuntze) Reveal, comb. nov.—22(2):863 Dodecatheon pulchellum var. macrocarpum (A. Gray) Reveal, comb. nov.—22(2):863 Dodecatheon pulchellum var. shoshonense (A. Nelson) Reveal, comb. nov.—22(2):864 Dodecatheon utahense (N.H. Holmgren) Re- veal, stat. Nov.—2.2(2):864 Drepanostachyum falcatum sengteeanum Stapleton, var. nov.— 22(1):332 Val. xX Elyleymus hultenii (Melderis ex Hulten) Barkworth, comb. nov.—22(1):496 Elyleymus mossii (Lepage) Barkworth, comb nov.—22(1):497 Elyleymus ontariensis (Lepage) Barkworth, comb. nov.—22(1):497 Elymus Xcayouetteorum (Boivin) Barkworth, comb. nov.—22(1):498 Elymus churchii J.J.N. Camp., sp. nov.— 22(1):486 Elymus lanceolatus subsp. riparius (Scribn. & J.G. Smith) Barkworth, comb. & stat nov.— 22(1):498 Elymus texensis J.J.N. Camp., sp. nov.— 22(1):488 Ericameria nauseosa subsp. ammophila L.C. Anderson, subsp. nov.—22(2):868 Erigeron strigosus var. traversii (Shinners Noyes, comb. & stat nov.—22(1):273 Eriogonum callistum Reveal, sp. nov.— 22(2):857 Eriogonum Sect. Lanocephala Reveal, sect. Nov.—22(2):857 Eugenia haberi Barrie, sp. nov.—22(2):1071 pale’, SIDA 22(2): 1273. 2006 1273 Eugenia yasuniana B. Holst & M.L. Kawasaki, sp. nov.—22(2):934 Fargesia apicirubens Stapleton, sp. nov.— 22(1):331 Gentianella fabrisii Filippa et Barboza, sp. nov.—22(1):130 Hamamelis ovalis S.W. Leonard, sp. nov.— 22(2):850 Heterotheca polothrix Nesom, nom. et stat. nov.—22(1):373 Heterotheca sierrablancensis (Semple) Nesom, comb. & stat. nov.—22(1):374 Indocalamus hamadae (Hatus.) Stapleton, comb.nov.—22(1):332 Lantana camara subsp. aculeata (L.) R.\W.Sand- ers, comb. & stat. nNOV.—22(1):394 Lantana nivea subsp. mutabilis (WJ. Hook.) RW. Sanders, comb. & nov.—22(1):395 Lantana strigocamara R.W. Sanders, sp. nov.— 22(1):392 xX Leydeum littorale (H.J. Hodgs.& W.W. Mitch.) Barkworth, comb. nov.—22(1):498 Leymus californicus (Bol. ex Thurber) Barkworth, comb. nov.—22(1):498 Mandevilla angustata (Steyerm.) J.F. Morales, comb. & stat. nov. —22(1):356 Mandevilla tristis J.F. Morales, sp. nov.— 22(1):359 Matelea corynephora Krings, sp. nov.— 22(2):942 Matelea phainops Krings, sp. nov—22(2):948 Matelea rhynchocephala Krings, sp. nov.— 22(2):949 Matelea torulosa Krings, sp. nov.—22(2):951 Nymphaea minuta K. Landon, R.A. Edwards & P|. Nozaic, sp. nOV.—22(2):887 x Pascoleymus Barkworth, gen. hybr. nov.— 22(1):499 x Pascoleymus bowdenii (Boivin) Barkworth, comb. nov.—22(1):499 Pediomelum piedmontanum JR. Allison, M.W. Morris & A.N. Egan, sp. nov.—22(1):229 1274 Peyritschia howellii (Hitchc.) Finot & PM.Peter- son, comb. nov.—2.2(2):897 Peyritschia pinetorum (Swallen) Finot & P.M. Peterson, comb, nov.—22(2):899 Poa matri-occidentalis P.M.Peterson & Soreng, Sp. NOV.—22(2):906 Poa matri-occidentalis subsp. mohinorensis soreng & PM. Peterson, subsp. nov.— 22(2):911 Polytrichastrum alpinum var. sylvaticum (Menzies) G.L. Merrill, comb. nov.—22(1):547 Pseudoroegneria spicata f. inermis (Scribn. & J.G. Sm.) Barkworth, comb. nov. & stat. nov.— 22(1):499 ~ AL ta f kK (Cl Barkworth, comb. nov. & stat.nov.—22(1):499 Psidium cauliflorum Landrum & Sobral, sp. nov.—22(2):927 Psidium cauliflorum Landrum & Sobral, sp. NOV.—22(2):931 SIDA 22(2): 1274. 2006 BRIT.ORG/SIDA 22(2) Sideroxylon macrocarpum (Nutt.) J.B. Allison, comb. nov.—22(1):245 Sphagnum beothuk Andrus, sp. nov.— 22(2):966 Sphagnum bergianum Andrus, sp. nov.— 22(2):964 Sphagnum kenaiense Andrus, sp. nov.— 22(2):961 Sphagnum mcqueenii Andrus, sp. nov.— 22(2):959 Sphagnum sitchense Andrus, sp. nov.— 22(2):969 Sphagnum talbotianum Andrus, sp. nov.— 22(2):970 Stenanthium diffusum Wofford, sp. nov.— 22(1):450 Tetraphis pellucida var trachypoda (Kindb. ex Paris) Harpel, comb. nov.—22(1):551 Zeuxine pantlingii Av. Bhattacharjee & HJ. Chowdhery, sp. nov.—22(2):935 i A olume | of the Illustrated Flora of East Texas is the first fully Ne flora for East Texas, a species-rich area that contains roughly two-thirds of all the plant species known for Texas. The book covers all the native and naturalized ferns and similar plants, gymnosperms, and monocotyledons (1,060 species) known to occur in East Texas. Published ape) Botanical Research Institute of Texas clo WMusriated ts plofas ro oje Ilak . €7; RIT Texas (BRIT and the Austin College Center for Environmental Studies. www.easttexasflora.org 2 vy" x10 ve 1,614 pages 1,200+ bist Soules 1,000+4 Sida, Botanical Miscellany 126 Illustrated Flora issn 0833-1475 of East Texas isbn 1-889878-12-X, hbk. George Diggs, Barney Lipscomb, May 2006 Monique Reed, Robert J. O’Kennon Big “D" to the Big Thicket there is a land ee interwoven with groves of timber, stately pines, hardwoods, wed) ROnEYsuCNG a hid: d, d: €20905 please inquire) Available from Botanical Research Institute of Texas Press 509 Pecan Street md VAL, LL 761 4060, U.S.A | E-mail: dies Fax: 1-81 Bar aie aoe Pee nC ie sear 1 EA 9 BRIT TS photographic guide to the Wild Flowers of WILD FLOWERS OF MOMBACHO Mombacho is color-coded and arranged by Family NICARAGUA within the color sections. It covers flowering plants found on the whole of the Volcano; which includes both tropical cloud forest and dry, deciduous forest, at lower altitudes. The book covers 200 plants, each of which has two or three photographs and a short description in both English and Spanish giving common name(s); a short description of the plant; height; flowering period; habitat and any known use of the plant. The vast majority of plants are likely to be found in similar habitats throughout tropical Ss HELEN PACKERING America and nearly one third are pan tropical, making this book of interest to a wide audience throughout the tropics. WILD FLOWERS OF MOMBACHO ICARAGUA Sida, Bot. Misc. No 28, 2006 lca issn 0823-1475 FLORES SILVESTRES DEL MOMBACHO isbn 1-889878-14-6 BY HELEN PICKERING 6.5" x 8.5", x + 217pp. over 500 photographs ) 5 + p&p" SA $3 “International: $7 BI Texas residents add $1.24 sales tax PRESS UISIANA TEXAS. GEORGIA NORTH. \N AHOMA. VIRGINIA OREGON. NEBRASKA ALABAMA NEW MEXICO ea ITZERLAND “ze YA JORDON ‘en : XACRUZ ME NOY ALGO GUADALAJARA od COAHUILA YUCATAN Sih