NUMBER 1, 24 JULY 2008 JOURNAL OF THE BOTANICAL RESEARCH INSTITUTE OF TEXAS J. Bot. Res. Inst. Texas ISSN 1934-5259 VOLUME 2 NUMBER 1 24 JULY 2008 COPYRIGHT 2008 Botanical Ge Institute of Texas 509 Pecan Street Fort Worth, pan 76102-4060, USA = SEH EDITOR Barney L Lipscomb Botanical Research Institute of Texas 509 Pecan Street HISTORY AND DEDICATION 1962—Lloyd H. Shinners (left), a member of the Fort Worth, Texas 76102-4060, USA Southern Methodist University 817-332-7432; 817-332- dis tax (SMU) faculty and a prolific Electronic mail: ! g; jbrit@brit researcher and writer, published the first issues of Sida, Contributions to Botany (now J. Bot. Res. Inst. Texas) CONTRIBUTING SPANISH EDITOR Félix Llamas 1971—William F. Mahler (right), professor of ^ Dpto. de Botánica, Facultad de Biologia botany at SMU and director emeritus of BRIT, ^ Universidad de Léon inherited editorship and copyright. ` E-2471 Léon, SPAIN 1993— BRIT becomes publisher/copyright holder. SUBSCRIPTION PRICES (2008) $41. Personal (Individual/Family) $90. USA (Institutional) MISSION AND GOALS ` 5100. Outside USA (Institutional) The BRIT Press seeks innovation and besse in 2007 —First issue of J. Bot. Res. Inst. Texas. SUBSCRIPTIONS/BACK VOLUMES J. Bot. Res. Inst. Texas and Sida, Contr. Bot. Publications Assistant preparation, manufacture of botanical research and scientific Ee for the twenty-first century. The BRIT Press—bringing out the best in botanical science for plant conservation and education. Botanical Research Institute of Texas 509 Pecan Street Fort Worth, Texas 76102-4060, USA DIRECTION AND COVERAGE Electronic mail: orders@brit.org The BRIT Press considers original research papers COMPOSITION concerned with classical and modern systematic rhorngraphics, Plano, Texas; rlrchorn@verizon.net botany, sensu lato, for Re in J. Bot. Res. Inst. Texas. P INTI submissions are peer- reviewed. tinitad Creat £f NTING Prepress production and printing in the America by Millet the Printer, Dallas, Texas www.millettheprinter.com Guidelines for submissions are available from the BRIT Press website, http://www.britpress.org. BIBLIOGRAPHICAL COVER ILLUSTRATION Gemen Tu for the | : Du a | TS PA Electronically tinted botanical illustration | f Texas is of Liatris aestivalis originally used on BRIT's atu liver Ja! y HAUT 2001. Summer a MERE mid O J. Bot. Res. Inst. Texas following the principles P.H. (informally JBRIT). International Standard Serial No. (ISSN 1934-5259) Hol c ) Jul-Aug(-Sep) FREQUENCY OF PUBLICATION Oklahoma and Texas. J. Bot. Res. Inst. Texas is published semiannually ^ Sida 19:768. 2001. (summer/fall) as one volume Botanical illustration by by the Botanical Research Institute of Texas. Linny Heagy ©2001 TABLE OF CONTENTS SYSTEMATICS Xyris panacea (Xyridaceae)—a new yellow-eyed grass from the Florida panhandle Loran C. ANDERSON AND ROBERT KRAL Molecular analysis of Solidaster cv. Lemore, a hybrid goldenrod (Asteraceae) EDWARD E. SCHILLING, JAMES B. BECK, PATRICK J. CALIE, AND RANDALL L. SMALL Novae Gesneriaceae neotropicarum XV: Kohleria hypertrichosa, a new species of Gesneriaceae from northwestern Ecuador JOHN LirTNER CLARK AND LAURENCE E. SKOG Two new species of Guadua (Bambusoideae: Guaduinae) from Colombia and Bolivia XIMENA LONDONO AND ENEIDA ZURITA Tetrazygia paralongicollis (Miconieae: Melastomataceae), a new species from the Sierra de Baoruco and Sierra Martin Garcia, Dominican Republi WALTER S.. JUDD, GRETCHEN M. IoNTA, TEODORO CLASE, AND J. DAN defe ¿JR Una nueva especie de Cuatresia (Solanaceae) de Costa Rica y Panamá D. ARMANDO SOTO AND A.K. RO Two new species of Baccharis sect. Caulopterae (Asteraceae: Astereae) from southern Brazil ANGELO ALBERTO SCHNEIDER AND Ia Jop BOLDRINI Two new Cotoneasters etuer Rosaceae) from Yunnan Province, China JEANETTE FRYER AND BERTIL HYL Meliosma antioquiensis: una nueva Sabiaceae de Colombia XAVIER CORNEJO Anisocapparis y Monilicarpa: dos nuevos géneros de Capparaceae de América del Sur XAVIER CORNEJO, HuGH H. ILTIS, AND A. SPENCER TOMB A revision of Colicodendron (Capparaceae) XAVIER CORNEJO AND HuGH H. ILTIS Revision of Gonolobus s.s. (Apocynaceae: Asclepiadoideae) in the West Indies ALEXANDER KRINGS Index of names and types in West Indian Gonolobinae (Apocynaceae: Asclepiadoideae), including fourteen new lectotypifications, one neotypification, a new name, and a new combination ALEXANDER KRINGS Molecular phylogenetic analysis of the American Stipeae o resolves Jarava sensu lato polyphyletic: evidence for a new genus, Pappos KONSTANTYN ROMASCHENKO, PAUL M. PETERSON, ROBERT J. e NURIA GARCIA-JACAS, OKSANA FUTORNA, AND ÁLFONSO SUSANNA Infrageneric classification and nomenclatural notes for Pycnanthemum (Lamiaceae) KENTON L. CHAMBERS AND HENRIETTA L. CHAMBERS Nomenclatural notes in North American Potentilla (Rosaceae) BARBARA ERTTER A synopsis of the Mexican and Central American species of Vaccinium (Ericaceae) RosERT L. WILBUR AND JAMES L. LUTEYN Re-discovery and typification of Thibaudia laurifolia, Macleania insignis, and M. coccinea (Ericaceae: Vaccinieae), an adventurous history James L. LUTEYN Revision of Siphonandra (Ericaceae: Vaccinieae), a genus endemic to Peru and Bolivia James L. LurEYN AND EDGARDO M. ORTIZ 139 165 193 207 243 Folia taxonomica 4. Conspectus of Myriopus (Heliotropiaceae: Boraginales) in the Guiana Shield CHRISTIAN FEUILLET Folia taxonomica 5. A new name for Passiflora heterophylla (Passifloraceae) from Cuba and Haiti CHRISTIAN FEUILLET AND JOHN M. MACDOUGAL Folia taxonomica 6. Two new species of Besleria (Gesneriaceae) from the Venezuelan Guayana CHRISTIAN FEUILLET Folia taxonomica 7. Two new species and a new section in Episcia (Gesneriaceae) from the Venezuelan Guayana CHRISTIAN FEUILLET Folia taxonomica 8. Passiflora tecta (Passifloraceae), a new species in subgenus Passiflora from the Guianas CHRISTIAN FEUILLET A synopsis of Pachyphyllum (Orchidaceae) Eric A. CHRISTENSON Lectotypification and notes on Baccharis riograndensis (Asteraceae: Astereae) Gustavo HEIDEN AND ANGELO A. SCHNEIDER New species of Myrtaceae from Ecuador Bruce K. Hoist AND MARIA Lucia KAWASAKI Clarification of Borreria gymnocephala, Diodia gymnocephala, Diodia schumannii, Borreria flavovirens, and Spermacoce schumannii (Rubiaceae PIERO G. DELPRETE AND JOSEPH H. KIRKBRIDE, Jr. Aristidae eludendae II: a re-evaluation of the Aristida gibbosa complex (Poaceae: Aristideae), including A. marginalis, A. orizabensis, and A. sorzogonensis RoBERT T. STRAHAN AND KELLY W. ALLRED Taxonomy of Bromus (Poaceae: Pooideae: Bromeae) sections Bromopsis, Bromus, and Genea in British Columbia, Canada ` JEFFERY M. SAARELA Munrochloa, a new genus (Poaceae: Bambusoideae) with a new combination from India MukrESH KUMAR AND M. REMESH Two new species of Cyperaceae from Peninsular India M.A. Wapoop KHAN AND P. LAKSHMINARASIMHAN Lyonothamneae, a new tribe in the Rosaceae (Rosales) Luc BROUILLET The taxonomy of North American loti (Fabaceae: Loteae): new names in Acmispon and Hosackia Luc BROUILLET Micranthes nelsoniana var. porsildiana (Saxifragaceae), the proper name at the varietal level Luc BrROUILLET Nomenclatural changes in Nemacladus (Campanulaceae) ANCY R. Morin Phylogenetic analysis of North American plums (Prunus sect. Prunocerasus: Rosaceae) based on nuclear LEAFY and s6pdh sequences JoserH R. ROHRER, MEGAN A. O'BRIEN, AND JULIE A. ANDERSON Recognition of three taxa of eastern North American “Waldsteinia” and their appropriate names when incorporated into Geum (Colurieae: Rosaceae) ALAN S. WEAKLEY AND KANCHI N. GANDHI The genus Abronia (Nyctaginaceae) in Colorado, with notes on Abronia bolackii in New Mexico JENNIFER ACKERFIELD AND WILLIAM F. JENNINGS 291 305 323 373 379 385 387 395 397 401 414 Further transfers of glandular-pubescent species from Chenopodium subg. Ambrosia to Dysphania (Chenopodiaceae) SERGEI L. MOSYAKIN AND STEVEN E. CLEMANTS New combinations in the Panarctic vascular plant flora REIDAR ELVEN AND David F. MURRAY Heuchera woodsiaphila (Saxifragaceae), a new species from the Capitan Mountains of New Mexico PATRICK J. ALEXANDER Three new species of Ixora (Rubiaceae) from the state of Tocantins, Brazil PIERO G. DELPRETE Lasiambix dominicensis gen. and sp. nov., a eudicot flower in Dominican amber showing affinities with Fabaceae subfamily Caesalpinioideae GEORGE O. POINAR, JR., KENTON L. CHAMBERS, AND ALEX E. BROWN Kruschke names in North American Crataegus (Rosaceae): a correction and clarifications J.B. Puiprs Thomas Walter Typification Project, V: neotypes and epitypes for 63 Walter names of genera D through Z DANIEL B. WARD CHROMOSOME NUMBERS Chromosome numbers for (Asteraceae: Gnaphalieae) JERRY G. CHMIELEWSKI — North American species of Antennaria Chromosome number of Thevetia ahouai (Apocynaceae: Rauvolfoidae: Plumerieae) with discussion on the generic boundaries of Thevetia JUSTIN K. WILLIAMS AND JULIA K. STUTZMAN ANATOMY AND MORPHOLOGY Anatomía foliar de algunas gramíneas alpinas y subalpinas del Eje Volcánico Transversal, México MARICELA GÓMEZ-SANCHEZ AND KATHIA GEORGINA TÉLLEZ-PIMIENTA FLORISTICS, ECOLOGY, AND CONSERVATION A novel design for a light weight and durable field press CHARLES T. BRYSON AND RICHARD CARTER Structure and diversity of a riparian forest at Kaieteur National Park, Guyana CAROL L. KELLOFF Rediscovery of Mirabilis hintoniorum (Nyctaginaceae), a striking four-o'clock endemic to the Sierra de Coalcomán, Michoacán, México MARK FISHBEIN AND VICTOR W. STEINMANN Floristic composition of seasonally dry tropical forest fragments in Central Bahia, northeastern Brazil DOMINGOS Benício OLIVEIRA SILVA CARDOSO AND LUCIANO PAGANUCCI DE QUEIROZ Diversity and floristic composition of the vascular plants in the forest fragment in southeastern Rio de Janeiro, Brazil REGINA H.P. ANDREATA, HAROLDO C. DE LIMA, ANGELA S. FONSECA Vaz, JOSÉ FERNANDO A. BAUMGRATZ, AND SHEILA R. PROFICE La familia Nymphaeaceae en el estado de Nuevo León, México CARLOS VELAZCO-Macías, RAHIM FOROUGHBAKHCH POURNAVAB, MARCO A ALVARADO VAZQUEZ, Y GLAFIRO J. ALANIS FLORES 425 433 473 475 575 593 Refugio de fitodiversidad en la ciudad de México, el caso de la cuenca del río Magdalena V. ÁVILA-AKERBERG, B. GONZALEZ-HIDALGO, M. Nava-Lórez, Y L. ALMEIDA-LENERO 605 Praxelis clematidea (Asteraceae), a genus and species new for the Flora of North America J. RICHARD ABBOTT, C. LEANN WHITE, AND S. BARRY DAVIS 621 Exotic species of Celtis (Cannabaceae) in the Flora of North America ALAN T. WHITTEMORE Luziola subintegra (Poaceae: Oryzeae), new to Florida and the United States Jonn M. KUNZER AND MICHAEL J. BODLE 633 New and noteworthy records of several non-native vascular plant species in Arkansas Brett E. SERVISS AND JAMES H. PECK 637 Noteworthy vascular plant collections from northwest Louisiana CHRISTOPHER S. REID, PATRICIA L. FAULKNER, BARBARA R. MACROBERTS, AND MICHAEL H. MACROBERTS 643 Silene flos-cuculi ssp. flos-cuculi (Caryophyllaceae) and Euphorbia peplus (Euphorbiaceae), new to North Carolina DERICK B. POINDEXTER Annotated checklist of the vascular flora of the Beech Creek Unit of the Big Thicket National Preserve, Tyler County, Texas Larry E. Brown, BARBARA R. MACROBERTS, MICHAEL H. MACROBERTS, PAUL A. HARCOMBE, WARREN W. PRUESS, I. SANDRA ELSIK, AND SUZANNE BIRMINGHAM WALKER 651 Early historical references to the Big Thicket, Texas MicHAEL H MACROBERTS AND BARBARA R. MACROBERTS 661 The Big Thicket of Texas as floristically unique habitat MiCHAEL H. MACROBERTS AND BARBARA R. MACROBERTS 665 Seed germination response of Zizania texana (Poaceae: Oryzae) to soil inundation M.L. ALEXANDER Predicting Viola guadalupensis (Violaceae) habitat in the Guadalupe Mountains using GIS: evidence of a new isolated population Timothy C. MULLET, FRED ARMSTRONG, BENJAMIN ZANK, AND CHRISTOPHER M. RITZI 677 Annotated vascular flora of the Dead Horse Mountains, Big Bend National Park, Texas, with notes on local vegetation communities and regional floristic relationships JoseLYN FENSTERMACHER, A. MICHAEL POWELL, JOE SIROTNAK, AND MARTIN TERRY 685 Distribution and taxonomy of Symphyotrichum sericeum and S. pratense (Asteraceae: Ástereae) RoNALD L. Jones, C. THEO WITSELL, AND Guy L. NESOM 731 Ranunculus ficaria (Ranunculaceae), naturalized in Texas Guy L. NESOM 741 Paspalum vaginatum (Poaceae), a new threat to wetland diversity in southern California RICHARD E. RIEFNER, JR. AND J. TRAVIS COLUMBUS 743 Book Reviews AND Notices 6, 52, 94, 164, 200, 206, 242, 262, 266, 274, 284, 290, 296, 304, 424, 432, 454, 472, 474, 494, 516, 546, 574, 604, 620, 642, 648, 664, 672, 740, 760 INDEX to new names and new binati in J. Bot. Res. Inst. Texas 2(1), 2008 Achillea alpina subsp. multiflora (Hook.) D.E Murray & Elven, comb. et stat. nov.—442 cmispon americanus var. helleri (Britton) SC EE nov.—388 Acmispon argophyllus (A. Gray) Brouillet, comb. n Acmispon argophyllus var. adsurgens (Dunkle) Brouillet, comb. nov.—388 Acmispon argophyllus var. argenteus (Dunkle) Brouillet, comb. nov.—388 Acmispon argophyllus var. fremontii (A. Gray) Brouillet, comb. nov.—389 Acmispon argyraeus var. multicaulis (Ottley) Brouillet, comb. nov.—389 Acmispon argyraeus var. notitius (Isely) Brouillet, comb. nov.— Acmispon argyreus (Greene) Brouillet, comb. nov.—389 Acmispon cytisoides (Benth.) Brouillet, comb. nov.—389 Acmispon dendroideus (Greene) Brouillet, comb. nov.—389 Acmispon dendroideus var. traskiae (Noddin) Brouillet, comb. nov.—389 Acmispon dendroideus var. veatchii (Greene) Brouillet, comb. nov.—389 Acmispon glabrus (Vogel) Brouillet, comb. nov.— Acmispon glabrus var. brevialatus (Ottley) Brouillet, comb. nov.—390 Acmispon grandiflorus (Benth.) Brouillet, comb. nov.—390 Acmispon grandiflorus var. macranthus (Greene) Brouillet, comb. nov.—390 Acmispon greenei (Wooton & Standl.) Brouillet, comb. nov. Acmispon haydonii (Orcutt) Brouillet, comb. nov.—3 Acmispon heermannii (Greene) Brouillet, comb. nov.—390 Acmispon heermannii var. orbicularis (A. idt raios comb. nov.—390 Acmispon intricatus (Eastw.) Brouillet, comb. n 390 Acmispon junceus (Benth.) Brouillet, comb. n 1 Acmispon junceus var. biolettii (Greene) Brouillet, comb. nov.—391 Acmispon maritimus var. brevivexillus (Ottley) Brouillet, comb. nov.—391 Acmispon mearnsii (Britt.) Brouillet, comb. nov.—391 Acmispon mearnsii var. equisolensis (J.L. Anderson) Brouillet, SE nov.—391 Acmispon micranthus (Nutt. ex Torr. & A. Gray) A comb. n 1 Acmispon nevadensis (S. Watson) Brouillet, comb. nov.—391 Acmispon nevadensis var. davidsonii (Greene) roule comb. nov.—391 Acmispon niveus (S. Watson) Brouillet, comb. nov.—3 Acmispon nudatus (Watson) Brouillet, comb. MK Acmispon oroboides (Kunth) Brouillet, comb. nov.—391 Acmispon procumbens (Greene) Brouillet, comb. nov. —392 Acmispon procumbens var. jepsonii (Ottley) Brouillet, comb. nov.—392 Acmispon prostratus (Nutt. ex Torr. & A. Gray) Brouillet, comb. nov.—392 Acmispon rigidus (Benth.) Brouillet, comb. nov.—392 Acmispon strigosus (Nutt. ex Torr. & A. Gray) Brouillet, comb. nov.—392 Acmispon utahensis (Ottley) Brouillet, comb. nov.—392 Acmispon wrightii (A. Gray) Brouillet, comb. nov.—392 Anisocapparis X. Cornejo & H.H. Iltis, gen. nov. —62 Anisocapparis speciosa (Griseb.) X. Cornejo & H.H. Iltis, comb. nov.— Arabidopsis petraea subsp. septentrionalis (N. Busch) Elven & D.F Murray, comb. nov.—438 Arabidopsis petraea subsp. umbrosa (Turcz. ex Steud.) Elven & D.F Murray, comb. nov.—439 Aristida sorzogonensis f. orizabensis (E. Fourn.) Strahan & Allred, comb. nov.—322 Artemisia kruhsiana subsp. alaskana (Rydb.) D.E Murray & Elven, comb. et stat. nov.—443 Baccharis apicifoliosa A.A. Schneid. & Boldrini, sp. nov.—45 Baccharis flexuosiramosa A A Schneid. & Boldrini, sp. nov.—48 Besleria neblinae Feuillet, sp. nov.—269 Besleria yatuana Feuillet, sp. nov.— Bistorta elliptica (Willd. ex Spreng.) V.V. Petrovsky, D.E Murray & Elven, comb. nov.—435 Calamagrostis purpurascens subsp. laricina (Louis- EHE SEH comb. et stat. nov.—434 Calyptranthes websteri B. Holst & M.L. Kawasaki, sp. nov. sn Colicodendron bahianum X. Cornejo & H.H. Iltis, sp. nov— Colicodendron valerabellum H.H. Iltis, T. Ruiz & G.S. Bong sp. nov.—82 Coptidium x spitsbergense (Hadac) Elven, comb. nov.—437 Cotoneaster floridus J. Fryer & B. Hylmó, sp. nov.—55 Cotoneaster qungbixiensis J. Fryer & B. Hylmó, sp. nov.—53 Cotoneaster series Sterniani J. Fryer & B. Hylmo, ser. nov.—53 Crataegus mollis var. incisifolia Kruschke, var. nov.—473 Crataegus schuettei var. gigantea Kruschke, var. nov.—473 Cuatresia amistadensis D.A. Soto & A.K. Monro, sp. nov. —41 Cyperus karthikeyanii Wad. Khan Sr Lakshmin., sp. nov.—379 Dryas punctata subsp. hookeriana (Juz.) Jurtz., comb. nov.—441 Dysphania andicola (Phil.) Mosyakin & Clemants, comb. nov.—4 Dysphania bonariensis (Hook. f.) Mosyakin & Clemants, mm nov.—428 Dysphania burkartii (Aellen) Mosyakin & Clemants, comb. nov.—428 Dysphania congolana (Hauman) Mosyakin & Clemants, comb. nov.—429 Dysphania dissecta (Moq.) Mosyakin & Clemants, comb. nov.—429 Dysphania dunosa (L.E. Simón) Mosyakin & Clemants, Ges nov.—428 Dysphania mandonii (S. Watson) Mosyakin & Clemants, comb. nov.—429 Dysphania melanocarpa (J.M. Black) Mosyakin & Clemants, comb. nov.—427 Dysphania minuata (Aellen) Mosyakin & Clemants, comb. nov.—429 Dysphania nepalensis (Colla) Mosyakin & Clemants, comb. nov.—428 Dysphania oblanceolata (Speg.) Mosyakin & Clemants, comb. nov.—428 Dysphania procera (Hochst. ex Moq.) Mosyakin & Clemants, comb. nov.—429 Dysphania pusilla (Hook. f.) Mosyakin & Clemants, comb. nov.—427 y . nov. Dysphania stellata (Standley) Mosyakin & Clemants, comb. nov.—429 Dysphania tomentosa (Thouars) Mosyakin & Clemants, comb. nov.—428 Dysphania truncata (PG. Wilson) Mosyakin & Clemants, comb. nov.—427 Dysphania venturii (Aellen) ee & Clemants, comb. nov.—428 Episcia duidae Feuillet, sp. no Episcia rubra Feuillet, sp. n » Episcia sect. SEET Se Ene ) Feuillet, sect. nov.— Equisetum arvense subsp. alpestre (Wahlenb.) Schónswetter & Elven, comb. et stat. nov.—433 Eugenia concava B. Holst & M.L. Kawasaki, sp. nov.— Eugenia grossa B. Holst & M.L. Kawasaki, sp. nov.—299 Fimbristylis naikii Wad. Khan & Lakshmin., sp. nov. —381 Gentianopsis barbata subsp. raupii (A.E. Porsild) Elven, comb. nov.—442 Geum donianum (Tratt.) Weakley & Gandhi, comb. nov.—417 Guadua chaparensis Londoño & SCH sp. nov.—31 Guadua incana Londoño, sp. nov.— Hedysarum boreale subsp. mees (Turcz.) D.E Murray & Elven, comb. et stat. nov.—441 Heuchera woodsiaphila PJ. Alexander, sp. nov.—448 Hosackia crassifolia var. otayensis (Moran ex Isely) Brouillet, e nov.—388 Hosackia oblongifolia var. cuprea (Greene) Brouillet, comb. n 388 Hosackia stipularis var. ottleyi = Brouillet, comb. eee a Ixora araguaiensis Delprete, sp. no 56 Ixora congestiflora Delprete, sp. nov. 456 Ixora irwinii Delprete, sp. nov.—459 Koeleria pyramidata subsp. seminuda (Trautv.) Elven, comb. nov.—434 Kohleria hypertrichosa J.L. Clark & L.E. Skog, sp. nov. — Lasiambix Poinar, Chambers & Brown, gen. nov.—464 Lasiambix dominicensis Poinar, Chambers & Brown, sp. nov. —464 Limnorchis aquilonis (Sheviak) Rebrist. & Elven, comb. nov. — 434 Limnorchis huronensis (Nutt.) Rebrist. & Elven, comb. nov.—434 Matelea costata var. goodfriendii (Proctor) Krings, comb. nov.—150 Matelea dictyopetala (Urb. & Ekman) Krings, comb. nov.—130 Matelea proctori Krings, nom. nov.—151 Matelea pubescens (Griseb.) Krings, iua nov. - 31 Meliosma antioquiensis X. Cornejo, sp. n Micranthes hieraciifolia subsp. longifolia med & Irmsch.) Elven & D.F Murray, comb. nov.—439 Micranthes lyallii subsp. hultenii (Calder & Savile) Elven & D.E Murray, comb. nov.—440 Micranthes merkii (Fisch. ex Sternb.) Elven & D.E Murray, comb. nov.—440 Micratithes nelsoniana subsp. aestivalis (Fisch. & CA Mey.) Elven € D.F Murray, comb. nov.—440 Micranthes nelsoniana subsp. insularis (Hultén) Elven & D.E Murray, comb. nov.—440 Micranthes nelsoniana var. porsildiana (Calder & Savile) Gornall & H. Ohba, comb. nov.—395 Micranthes porsildiana (Calder & Savile) Elven & D.F Murray, comb. nov.—440 Micranthes redofskyi (Adams) Elven & D.E Murray, comb. nov.—441 Monilicarpa X. Cornejo & H.H. Iltis, gen. nov.—-67 Monilicarpa brasiliana (Banks ex DC.), X. Cornejo & H.H. Iltis, comb. nov —71 Monilicarpa tenuisiliqua (Jacq.) X. Cornejo & H.H. Iltis, comb. nov.—70 Munrochloa M. Kumar & Remesh, gen. nov.— Munrochloa ritchiei (Munro) M. Kumar & Remesh, comb. nov.—376 Myrcia subcordifolia B. Holst & M.L. Kawasaki, sp. nov.—303 Myriopus candidulus (Miers) Feuillet, comb. nov.—264 Myriopus maculatus (Jacq.) Feuillet, comb. pi i Myriopus paniculatus (Chamisso) Feuillet, comb. no Myriopus paniculatus var. spigeliiflorus (A. DC.) Bei ‘comb nov.—264 Nemacladus australis (Munz) Morin, comb. et stat. nov.— Nemacladus calcaratus Morin, sp. nov.— Nemacladus californicus (A. Gray) Morin, comb. nov.—397 Nemacladus orientalis (McVaugh) Morin, comb. et stat. nov.—398 Nemacladus secundiflorus var. robbinsii Morin, var. nov.—399 Nemacladus tenuis (McVaugh) Morin, comb. et stat. nov.—398 Nemacladus tenuis var. aliformis Morin, var. nov.—398 Orthaea laurifolia (M. Martens € Galeotti) Luteyn, comb. nov.—244 Pachyphyllum sect. Capitulum E.A. Christ. sect. nov.—288 Pachyphyllum sect. Orchidotypum (Kraenzl.) E.A. Christ., stat. nov.—287 Packera hyperborealis subsp. Pan e (Jurtz., Korobkov & VW Petrovsky) Jurtz., Korobkov & VV. Petrovsky, comb. nov.— Papaver labradoricum CA T & Elven, comb. et stat. nov.—438 Pappostipa (Speg.) Romaschenko, PM. Peterson & oleo comb. et stat. nov—181 Pappostipa ameghinoi (Speg.) Romaschenko, comb. nov. 4 Pappostipa ameghinoi var. digona (Parodi) ponte semen comb. nov.—184 Pappostipa ameghinoi var. precordillerana (FA. Roig) Romaschenko, comb. nov.—184 Pappostipa atacamensis (Parodi) Romaschenko, comb. nov.— Pappostipa barrancaensis (EA. Roig) Romaschenko, comb. nov.—184 Pappostipa braun-blanquetii (EA. Roig) Romaschenko, comb. nov.—183 Pappostipa chrysophylla (E. Desv.) Romaschenko, comb. nov.—185 Pappostipa chrysophylla var. cordillerarum (Parodi) Romaschenko, comb. nov.—185 Pappostipa chrysophylla var. crispula (Kuntze) Romaschenko, comb. nov.—185 Pappostipa chrysophylla f minuta (FA. Roig) Romaschenko, comb. nov.—185 Pappostipa chrysophylla f. modica (EA. Roig) Romaschenko, comb. nov.—185 Pappostipa sect. Chrysovaginatae Romaschenko, sect. nov.— Pappostipa chubutensis (Speg.) Romaschenko, comb. nov.— Pappostipa chubutensis var. hirsutissima (EA. Roig) Romaschenko, comb. nov. —185 Pappostipa frigida (Phil.) Romaschenko, comb. nov.—185 Pappostipa frigida var. parvispicula (Parodi) Romaschenko, comb. nov.—185 Pappostipa hieronymusii (Pilg.) Romaschenko, comb. nov.—183 Pappostipa humilis (Cav.) Romaschenko, comb. nov.— Pappostipa humilis var. decrescens (Kuntze) Romaschenko, comb. nov.—185 Pappostipa humilis var. ruiziana (Parodi) Romaschenko, comb. nov.—185 Pappostipa ibarii (Phil.) Romaschenko, comb. nov.—1 Pappostipa ibarii var. anomala (Parodi) Romaschenko, comb. nov.—184 Pappostipa ibarii f pallescens (Parodi) Romaschenko, comb. nov.—184 Pappostipa maeviae (EA. Roig) Romaschenko, comb. nov.—183 Pappostipa major (Speg.) Romaschenko, comb. et stat. nov.—184 Pappostipa malalhuensis (EA. Roig) Romaschenko, SEHR nov.—183 Pappostipa nana (Speg.) Romaschenko, comb. n 1 Pappostipa nicorae (FA. Roig) Romaschenko, s id 83 Pappostipa sect. Pappostipa, sect. nov. —182 Pappostipa parodiana (FA. Roig) Romaschenko, comb. nov.—183 Pappostipa semperiana (FA. Roig) Romaschenko, comb. nov.—183 Pappostipa sorianoi (Parodi) Romaschenko, comb. nov.—184 Pappostipa speciosa (Trin. & Rupr.) Romaschenko, comb. nov.—182 Pappostipa speciosa f. abscondita (FA. Roig) Romaschenko, comb. nov —182 Pappostipa speciosa var. atuelensis (EA. Roig) Romaschenko, comb. nov.—182 Pappostipa speciosa var. manqueclensis (EA. Roig) Romaschenko, comb. nov.—182 Pappostipa speciosa var. media (Torres) Romaschenko, comb. nov.—182 Pappostipa speciosa var. parva (EA. Roig) Romaschenko, comb. nov.—182 Pappostipa vaginata (Phil.) Romaschenko, comb. nov.—182 Pappostipa vaginata var. argyroidea (EA. Roig) Romaschenko, comb. nov.—183 Pappostipa vaginata f. contracta (EA. Roig) Romaschenko, comb. nov.—183 Pappostipa vaginata var. dilatata (EA. Roig) Romaschenko, comb. nov.—183 Pappostipa vaginata f. inmersa (EA. Roig) Romaschenko, comb. nov.—183 Pappostipa vaginata f. laevis (FA. Roig) Romaschenko, comb. nov.—183 Pappostipa vatroensis (EA. Roig) Romaschenko, comb. nov.—184 Passiflora insueta Feuillet & MacDougal, nom. nov.— Passiflora tecta Feuillet, sp. nov.— Potentilla anserina subsp. yukonensis (Hultén) Soják ex Elven & D.F Murray, comb. nov. —441 Potentilla jepsonii Ertter, nom. nov.— Puccinellia phryganodes subsp. neoarctica (Á. Lóve & D. Lóve) Elven, comb. et stat. nov.—435 Puccinellia phryganodes subsp. sibirica (Hadac & Á. Lóve) Elven, comb. nov.—435 Pycnanthemum sect. Californicae K.L. Chambers & H.L. Chambers, sect. nov.—195 Ranunculus subborealis subsp. pumilus (Wahlenb.) Elven, comb. nov.—437 Ranunculus subborealis subsp. villosus (Drabble) Elven, comb. et stat. nov.—438 Rhododendron tomentosum subsp. decumbens Elven & D.F Murray, comb. nov —441 Rosaceae tribus Lyonothamneae Brouillet, tribus nov.—385 Scorzoneroides autumnalis subsp. pratensis (Hornem.) Elven, comb. nov.— Silene involucrata subsp. furcata (Raf.) VV. Petrovsky & Elven, comb. et stat. nov.—436 Silene soczavana var. macrosperma (A.E. Porsild) V.V. Petrovsky, D.E Murray & Elven, comb. et stat. nov.—436 ov. Silene villosula (Trautv.) VV. Petrovsky & Elven, comb. nov.—437 Silene violascens (Tolm.) V.V. Petrovsky & Elven, comb. nov.— 437 Siphonandra nervosa Luteyn & E.M. Ortiz, sp. nov.—25 Siphonandra santa-barbarense Luteyn & E.M. Ortiz, sp. nov.— Solidago multiradiata subsp. arctica (DC.) Korobkov & Elven, comb. et stat. nov.—443 Stuckenia filiformis subsp. borealis (Raf.) Tzvelev & Elven, comb. nov.—433 Tetrazygia paralongicollis Judd, lonta, Clase & Skean, sp. uc Vaccinium beamanianum Wilbur € Luteyn, nom. nov. —2 Vaccinium campanense Wilbur & Luteyn, sp. nov —234 Vaccinium chihuahuense Wilbur & Luteyn, sp. nov.— Vaccinium uliginosum subsp. vulcanorum (Kom.) Alsos & Elven, comb. et stat. nov.—442 Vahlodea latifolia subsp. paramushirensis (Kudó) Elven, comb. nov.—435 Xyris panacea L.C. Anderson & Kral, sp. nov.— XYRIS PANACEA (XYRIDACEAE)—A NEW YELLOW-EYED GRASS FROM THE FLORIDA PANHANDLE Loran C. Anderson Robert Kral Professor Emeritus, B e of Biological Science Professor Emeritus, Vanderbilt University lorida State University 1425 Pine Circle, ea Ge 32306-4370, U.S.A. Cairo, Georgia 31728, USA ABSTRACT Á new species of apud de Brees (Xyris pane dip BEE ponds in the Big Bend area of the Florida panhandle (USA) is technically described, RESUMEN Án intensive plant survey by the senior author of the St. Marks National Wildlife Refuge included discovery of an unusual Xyris that in normal years would probably have been accessible only by boat, but the drought for the spring months of 2007 was the most severe on record for the Florida panhandle (USA). This enabled collecting the plants on foot—albeit I was up to my armpits in water on one occasion because at some sites the new Xyris grows on floating islands. The species description follows the format and terminology of Kral (2000) Xyris panacea L.C. aida » Kral, EDEN (Figs. 1-2). Tyre: U.S.A. Froripa. Nao Coz 2 Mate ee IT Refuge, water of "droughted" y and Nymph dorata SE edge of P Field, ca. 1.7 air mi SW of RN Lat. 30.00742 N, bons 84.42203 W 23 Aug 2007, 1 C And 23,436 ( : FSU; isotypes: MO, NY, VDB). Planta robusta, perennis, caespitosa, 7—10(-13) a ee incrassati, varie Pic (basibus in substratio profundo elongati, ramificantes ascendentibus) cum (frequente) tibus. Folia pricinpalia rigid, disticha, anguste Habellate PEU ca. Der REECH cm longa, pou ee longiora; laminae planae vel leveter tortae, 3-58) mm latae, compres- sae, 1-2.5-plo vag , integrae, | pups RO dud contracti, incurato-acuti; li ae carinatae, ea ates bue eod : is, glabris, rufobrun laminae gradati gentibuis, infimis gradati t is, dilute | li SE usque ad 1 lat oa a basin convolutae | icarinatae, | 1 lae, tortae, levit lticostatae, laminis erectis, planis, usque ad 15 cm longis et 3 mm ue Scapis MM recti, ee versus in sectio transversali elliptici, vallis unicostati, ca. 1 mm crassi. E a ovoideae go) cylindricae, 2-4(-4.5) cm EE obtusae, multibracteatae, | I convexis, ecarinatis, lat t anguste obovatis. Bract teriles pl i , pari infima cari- , rotundatis. B fertilies ] tanguste ol tae, 7-8 mm longae, See, lamprol lae, areis dorsalis distinctis, lat I Sepala lateralia lil , anguste obl , 7-8 mm longa, pallide la; al inali sug li iliat pil iliata.I i petal late ol te, 5mm longae, l latae, denticulatae. Staminodia bibrachi brachiis longipenicillatis. Anth blongae, ca. 2 mm longae, filiis ca. 0 Sfm longe Capsulae anguste obovoideae, ca. 5 mm SE dorsalit Semi bcylindri gula psoidea, 0.7-0.8 mm longa, translucida, brunneola, longitudinae valde multicostata et subtiliter transversilineata. Robust, solitary to cespitose perennials 7-10(-13) dm long, the bases typically slightly bulbous, firm, buried in a mucky, often submersed, substratum, thus periodically producing (on larger plants) elongate ascending branches (internodes) and new “rosettes,” the older, more basal nodes producing dense mats of pale, spongy roots, whereas new nodes produce spreading-arching stolons tipped with new plants. Princi- pal leaves subdistichously in narrow fans, ca. 32-45(-50) cm long, longer than the scape sheaths, entire, multicostulate, green proximally shading to red, flat or slightly twisted, 3-5(-8) mm wide, 1—2.5 times shorter than the sheathes, gradually narrowed to incurved-acute tips; sheathes entire, narrowly convex at | Rot Rac [nct Tavac ATI, 1— 5. 2008 TEA! lexas 2(1) £T : Ae d i im % - + & te v P ge t: H n ai H Es 15 cm h laaf base, mid-sector, and anex. € Spike d tal sal ol Li J Il AA, E ` if E a - J \ Fertile bracts. a. Lat Fic. 1. A De > Fa 4: Is 4l L A L E Viet. EI | EL. LI LI T d [| H E * A | B e L . LI d i 41 I Im elei T "TI H ted | Ic. b. Distal views of a | At a a H ate senals, Y E , IR a a HI disposed style branches. very base, there laterally 1-11.5 cm wide, lustrous dark red aging red-brown or deep brown, distally keeled, gradually narrowing into blade, reddish shading to green, eligulate. Scape sheaths shorter than principal leaves, proximally convolute, sharply keeled, lust brown or red-brown, twisted, shallowly multicostate, distally opening to an erect, keeled blade. Scapes slender, erect, 60-115(-130) cm long, red-brown shading to green distally, oval or elliptic in cross-section and 1-1.2 mm wide with one firm, smooth costa. Spikes usually narrowly cylindric, also narrowly ovoid to narrowly ellipsoid, 2-4(-4.5) cm. long, obtuse, multi- bracteate, the bracts imbricate in high spiral, convex, ecarinate (except sometimes with carinate basal pair), broadly to narrowly obovate, entire and distinct dorsal areas. Sterile bracts several, shorter than the fertile and grading into them; fertile bracts 7-8 mm, lustrous dark to pale brown, in contrast to the subapical, broadly ovate to elliptic, gray-brown to dark green dorsal areas. Lateral sepals free, narrowly oblanceolate, /—8.5 mm long, inequilateral, the keel above middle (often) pilose, thence distally increasingly lacerocili- ate or pilose-ciliate, with tip acute and slightly to noticeably exserted. Petal blades broadly obovate, 8—9.5 mm long, 6-7 mm wide, apex shallowly rounded, denticulate. Staminodia 3-4.5 mm long, bibrachiate, branches elongate-penicillate. Anthers oblong, 2.5—3 mm long, on stout filaments 0.5—0.8 mm long. Styles 4.5—5 mm long, first ascending, then spreading horizontally between petals with age. Capsules narrowly obovoid, ca. 5 mm long, adaxial side plane, abaxial side convex; placentation 3-parietal. Seeds narrowly to broadly fusiform, 0.7-0.8 mm long, light brown, translucent, longitudinally strongly multiribbed with numerous finer, straight or diagonal connecting lines. Etymology.—The specific epithet commemorates its geographical area (no curative powers are known). [| [| E al n.a H in LI dd nd Common name. —*“St. Marks yellow-eyed grass” would be an appropriate common name Phenology.—Transplants of all relevant collections were potted in orginal substrate and cultivated in a flooded wheelbarrow in Anderson's backyard for observation. The new species blooms from late June to early October. Flower buds unfurl and petals are fully expanded around 11:30 am (EDT); in a close associ- ate, X. fimbriata Elliott, the flowers open around 1:00 pm and stay open about two hours, whereas those of X. panacea remain open until around 4:00 pm. Flowers of X. smallinana Nash (in this region) open around 5:00 pm, and their flowering does not overlap with the other species. Flowering observations among the transplants were confirmed in the field. Habitat.—The Refuge Comprehensive Conservation Plan (U.S. Fish and Wildlife Service 2006) lists ponds in the Panacea unit variously as “coastal depression ponds” or “basin lakes” or “flatwoods lakes.” The St. Marks yellow-eyed grass grows either in mucky, loosely matted sandy loam in shallow depression ponds in longleaf pinewoods (with stems bases submerged in 10 or more centimeters of water or in poorly congealed loam of floating islands in those depression ponds (with stem bases only slightly submerged). Some ponds are ringed with a thicket of Cyrilla racemiflora L., whereas others are bounded by Hypericum fasciculatum Lam. The new species was vouchered from five ponds in the area; it is presumably present in several other ponds that Anderson planned to visit, but his 2007 field season was terminated abruptly with a broken leg. Associated species —Frequently seen taxa (other than other Xyris) at one or more sites include: Bacopa caroliniana (Walter) B.L. Rob. , Burmannia biflora L., Cyrilla racemiflora, Decodon verticillatus (L.) Elliott, Drosera intermedia Hayne, Eleocharis baldwinii (Torr.) Chapm., E. elongata Chapm., E. equisetoides (Elliott) Torr., Eriocaulon compressum Lam., Fuirena breviseta did pns F. N Michx., o repens Nuttall, Hydrocotyle bonariensis Comm. ex Lam., H q th ( ) Dandy, Ludwigia alata Elliott, Lycopodiella appressa (Chapm.) Cranfill., Mayaca ee Aubl., Re laxum Shuttlew. ex Chapm., Nuphar advena (Aiton) W. Aiton ssp. orbiculata (Small) in m B iii pilis Sol., Nymphoides aquatica (J.F. Gmel.) Kuntze, Panicum verrucosum Muhl., Pontederia careyana Fernald, R. cephalantha A. Gray, R. tracyi Britton, Sagittaria inead L., Scleria — MERE Triadenum virginicum (L.) Raf., Utricularia floridana Nash, U. juncea Vahl, and U. purpurea Walter. Paratypes. Anderson 23,437 (AUA, FLAS, FSU, GA, USF, VDB, VSC), Anderson 23,438 (FLAS, FSU, GA, USCH, USF), R. Kral 98,248 B). Additional collections (all FLORIDA. Wakulla Co.: St. Marks National Wildlife Refuge). East Renfro Lake near type locality, Lat. 30.00754 N, Long. 84.42194 W., 7 Aug 2007, And 23396 (FSU, VDB); “Point 4 Pond” (0.4 mi S of Otter Lake Rd on Refuge Rd 319), Lat. 30.02258 N, Long. 84.40673 W, 4 Sep 2007, Anderson 23473 (FSU); “Point 8 Pond” (0.8 mi S of Otter Lake Rd on Refuge Rd 319), Lat.30.01715 N, Long. 84.40813 W, 4 Sep 2007, A ld eg d ap. penus side Rte 372 Dee Refuge Rd 334), Lat.30.02234 N, Long. 84.44504 W,11 Sep 2007, A dq th Lake, S of Refuge Rd 329, SW of Otter Lake, Lat.30.01434 N, Long. 84.42978 W, 25 Sep 2007, A And 23595 (ELAS, FSU, FIG, GA, MO, UNA, VSC). DISCUSSION The common associate Xyris in these ponds is X. fimbriata, a species of similar habit and pigmentation but with two scabrid scape costae (rather than a single, smooth one), with spikes mostly shorter and of broader outline, with lateral sepals strongly exserted, their keels densely fimbriate (Fig. 2). Also, a few diminutive flowering plants of X. jupicai L. Rich. were seen at the type locality. Ponds 3.8 km and 4.3 km NE of the new taxon’s range had the “X. panacea zone” replaced with X. smalliana exclusively, see Anderson 23456, 23457 (FSU, VDB). In the general area, though not an associate, is X. stricta Chapm., another tall species with similarly ascending, long-sheathed, slender-bladed leaves and similar pigmentation. This taxon also hasnarrowly ellipsoidal to subcylindric spikes with similar design of bracts. However, this taxon has scapes distally broader, flatter, actually 2-edged, its lateral sepals are broader, with broader, firmer, ciliolate keels whose tips are not exserted; its seeds are typically farinose. The St. Marks yellow-eyed grass is distinctive with a suite of vegetative and floral features. The dark red stem bases are somewhat bulbous and firm (being pale green and soft in X. fimbriata or pale green and A J Iu Il A H Ev. LEN E Al ri SA 1 Jl 5 F 1:7 4 -l ls + Ew mucilagionous in X. smalliana); larger plants have stout vertical, rhizome The flowering scapes are smooth with weakly developed ribs below the spikes. The spikes are cylindric, longer than those of any other taxa in North America (Kral 2000). Flowers open earlier in the day and are larger than those of other species (equally large petals are found in X. caroliniana Walter, but that species differs in habitat and very many E e pd M More attention ae to be given to style position amongst North American species; the | X. panacea may be exclusively unique. The known range of the new species is ida Sech to warrant designating it a threatened or endangered species. ACKNOWLEDGMENTS Valuable logistical support (i.e., maps, gate keys, vehicles) was provided by the St. Marks National Wildlife Refuge staff. Photographs of living material were taken by Ken Womble. Barry J. Conn, J.F. Hays III, and A.B. Thistle provided helpful suggestions on the manuscript. Funds from the Friends of the Robert K. Godfrey Herbarium (FSU) helped defray publication costs of this paper. REFERENCES KraL, H 2000. The Xyridaceae family. Flora N. Amer. 22:155-167. U.S. FisH AND WiLbure Service. 2006. St. Marks National Wildlife Refuge Comprehensive Conservation Plan. Atlanta, Georgia. BOOK REVIEW Harry S. Paris. The Drawings of Antoine Nicholas Duchesne for his Natural History of the Gourds. (ISBN 9782856536049, hbk.). Muséum National d'Histoire Naturelle, Publications Scientifiques, Dif- fusion 57 Rue Cuvier, F-75231 Paris Cedex 05, France. (Orders: www.mnhn.fr/publication, diff. pub mnhn.fr, 33 (0)1 40 79 48 d $257.00, 454 pp., 258 color plates, 17 1/2" x 12 3/4". AT T VTA 1707075 I AË (1747-1827) was born at ne Duchesne became a careful observer of nature under the tutelage of his scholarly and well-traveled father, and later ] (the secon d of the five de Jus- = [I1 x H Ae x Paris, in was da e o Linnaeus. Duchesne eo with Linnaeus ] s. Tho 4 e on botanical matters, particularly on his fi dh it was ird at «lI qo aj 1 H aut Ls lf. 1 1 + A f el 1 11 1 r PE rdg Reps O Cro “tar 1 1 ] +l 1 Cal 8 es WS? y) u :11 1 E | ] ELS ts T] 1 Linnaeus said, I g g oped that some botanists would de: pane: that they d each choose their pan a) ] ine it ttl ghly; in thi 14 1 1 Lë Gg 1 3 n liilaillitl 5 Is WWII ILL YY Duchesne’s first work, L’Histoi turelle des frasiers (Natural history of the strawberries), was mee in 1766 Wee ee was ony 19 years old. One product E this research EE his own | ical ill , in pencil and gh he | see them published in his lifetime). Duchesne was particularly i ted l l systems as they pertained to the creation of domesticated Mn and through cross- ane experiments he sans ed ae attempted to retrace "i arentage of) various le strawberry cultiv | what v tation of the doctrine of constanc zu immutability of speci intained by other MCN of his era. He called himself a cultivator botanist,” distinct from otl tanists who observed without After all, this was the Age of the Enlighten His next major work boused on mpini: squash, and EDS of ie genus E de Cua a arose in the New Midi din thus only Kee Se to p NER after 1492. 1 Du the grown YY VLA. i YY LA E Per a I «1 4 latah] ] 1 : ] 1 1 day jos of northern summers. Duchesne was is drawn t to Cucurbita E its s incredibly varied fruit GE E to the En and hand of a botanical artist), | ts, and a similar LA L + O 1 4 1 E ] E mt h q nis d Seer v & e AA [TA 1 DH re 1:71] 4 £41 1 TE «Lee a 231221 $ +51 +1 d re EL ke La r entire collection of illustrati f Cucurbita is published for the first time. The author Dr. Harry Paris, one of the foremost experts on ia is Gees for aly locating Duc! : ] illustrat the Library of Ges e uid Museum in iu e 1 A EJE T. 1 D + A Li 6 idi are ae e Geste S illustrations, o of those cultivars coma called BECH These E idide Ee a nr] ne Į Us are full-color watercolors (258 color plates). The primary focus is on the form and color pat- terns of fruits, and their textures, sia ing att to Wal d. f all kinds. Since this book is gigantic, many of these illustrations approach life-size. The works "e from hazy i isti g lingly photo-realistic pieces worthy of any Dutch Master's x Mi iau ER too are a number of D ] 's illustrati f Cucurbita fl d their developmental stages, and [ I f the vines, EE are more ee oe in nad RE 1 LO 1 "- ( to h; y of Duchesne y I . 8 and botanical significance of his work, as well as a chapter on the botany of tl Cucurbita pe à Ea e CN to scientific names. This is the sine qua non coffee table book fi un al E horticulture, or konica illustration, pary with the interpre- 1 4 1 H :11 4 +1 q ss PEE + tematists.— Aman eill, Director of the the Herbarium (BRIT-SMU- VDB), Botanical Puch Institute of Texas, 500 East 4th Street, Fort BEE Texas 76102-4025, U.S.A. J. Bot. Res. Inst. Texas 2(1): 6. 2008 MOLECULAR ANALYSIS OF SOLIDASTER CV. LEMORE, A HYBRID GOLDENROD (ASTERACEAE) nana E SEHR James B. Beck Depart FE y Biology Department of Biology University of Tennessee Washington University, Campus Box 1137 gie lene Sse hike. U. e i t. Louis, Missouri 63130, U author foi hilling@ lu) and Missouri Botanical Garden St. Louis, Missouri 6313 Curent aaa Department of Biology y, th Carolina 27708, U.S.A Patrick J. Calie Randall L. Small A, Sciences Depart t of Ecology and Evolutionary Biology Fastern Kentucky University University of Tennessee Richmond, Kentucky 40475, U.S.A Knoxville, Tennessee 37996, U.S.A. ABSTRACT ly : r 1 21 1 1TS and CTC 1 3-4 Ja Phi 1-45 I ir £ +1 » | OR EE | Cnlidaster CV. T emore 1 1 cds 1 1: 1, 1 1 2 gu 1e | Haras D + fil 1 1 1 f the g ic pl tof fits putative [ t the Upland White Aster, Said pt icoides. B fit perficial external alo: 2 H 2211 Ce P Ts | Aster, 1+1 C 1 +1 + CR 1 111 1 sified within salidas Phylog i is of the combined ITS S and ETS seq data sl 1 ti ] inclusion of S. Re _ Solidago, e on its a in a well supported clade ca d of al other sampled See of the g s. The IT S sequence a evidence of a hy "brid origin base base ss polymorphisms. C ing experi ] liff i idual ITS sequences, dentical to that obt d f SR 1 1 1 1 | TE E canade ensis Thuc DNA EEN E WH ] 1 Cl ] Lemore isa e brid Stee that involved a a Cross Kees 5. o ans i a The d further pu that Paus TE i Ct + x: re 1 1 1 + :11 11 nd [e] d us f lid [a] fad [e] I f I a L I I oO L r evolving markers. Key WORDS: Asteraceae, Astereae, Solidaster, Solidago, ITS, ETS, hybrid, phylogeny RESUMEN C "ES z -]- 0: 1 ] ] 1 JI "rte ere 1 1 1 1 s 1 1 1 ás a] Solidaster cv. Lemore, lti ] t tiemp yóq híbrido int genéri (Solid g Aster). Parte del análisi ae la evaluación e la Stee SC género de uno de los padres putativos, Solidago ptarmicoides. Debido al parecido externo de Actor ] ] J : f. t a 3.1 z 1 epe L sr L Kë L Solidago II xq. cz [=] Les J J LE ] 1.1 o 1.1 Tre ore PET d 1 14 fe 1 1 se 1 + i i ] yl + : J de S ptarmicoides en Solidag | z i Li Los cae de Sé secuencia Rene ITS de EE mostraron pruebas de un origen híbrido See en 4 presencia de varios una r A : SN 1 y EA J Vi Tn A 1 1 J J; : E | dE O canas Pori x ds E + Lei L 1.7 Ja TARTA anima Em 1 Gs t Lemore es una r1 Liked I ] te d t S ptarmicoides Y 5. canadensis. Los datos indi d lid ible, Futhami inifoli tal de Solidaster Fue también E r 7 e + ? Is matakhla la falta da ri AAA AA la secuencia de TTE cp ETE + E d Cuba DE pe ] + ] EI e dix ^ Ir Le a a] L J J 1 m z 2 5] [e] L Y Consideration of the role of hybridization in evolution has received new impetus SIM the availability of molecular genetic data (Arnold 2006). It has long been clear that intersy y is a widespread J. Bot. Res. Inst. Texas 2(1): 7 — 18. 2008 [| E ai n.a D in L | L'ALA ET TEA! phenomenon in plants, with numerous clearly documented examples (e.g., Grant 1980), but there are still questions regarding its evolutionary significance. Of perhaps greatest interest is hybridization between species of lineages that have diverged morphologically to the point of being recognized as distinct genera. This represents a potentially dramatic reversal of the process of evolution, which is normally considered to be divergent, and it also presents a challenge to systematic classification based on phylogeny. It is thus of considerable interest to examine closely possible cases of intergeneric hybridization to be able to assess how frequently this phenomenon occurs. A persistent myth is that intergeneric hybridization is common and perhaps even pervasive within Asteraceae. For example, Robinson (1983) and King and Robinson (1987) invoke intergeneric hybridiza- tion as an explanation for discordant distribution of morphological characters in Liabeae and Eupatorieae, respectively. A search of the literature, however, reveals few well documented cases of natural intergeneric hybridization in Asteraceae. Naturally occurring intergeneric hybrids have recently been documented by McKenzie et al. (2004) and Saito et al. (2006). In some cases, artificial intergeneric hybrids have been made successfully (e.g., Powell 1985; Carr 2003), and studies have found incongruence between nuclear and chloroplast markers suggestive of past wide hybridization (Schilling & Panero 1996; Fehrer et al. 2007). But documentation of intergeneric hybrids in most groups remains elusive. This prompted us to undertake a detailed study of a putative case of intergeneric hybridization within the Astereae. The name of the horticultural plant known as Solidaster cv. Lemore (Fig. 1) reflects its presumed hybrid origin as a cross between species of Solidago L. and Aster L. The plant is unknown in the wild, but appeared in European gard Iter the import of plants from North America (Nesom 1993). Its status as an intergeneric hybrid depends in part, however, on the classification of one of its putative parents, the Upland White Aster, which has been variously placed in Aster, Solidago, and Oligoneuron Small. The Upland White Aster, S. ptarmicoides (Torrey & A. Gray) B. Boivin, has long been a source of puzzlement for taxonomists. Its superficial outward appearance, especially its open corymbose and relatively large heads with white rays and disk flowers, seems clearly that of an aster, as traditionally conceived, and is reflected in its common name. Despite this, considerable evidence has PEEL a bal affinity to the goldenrods, including the observation of abundant natural hybrids with species go (Boivin 1972), considerations of phytogeography and technical features of morphology (Brouillet & apie 1981), and a chloroplast DNA restriction site study placing it within Solidago L. tl al et al e Nev ertheless, it is still widely referred to as an Aster s.l. Another, somewhat different int tion of the genus Oligoneuron as distinct from Solidago (e.g., Nesom 1993), which would include the Upland | White Aster as well as five other species. Although the second parent of Solidaster has generally | considered to be a Solidago, particularly S. canadensis (Brouillet & Semple 1981), Nesom (1993) suggested that based on morphology it was more likely to be Euthamia graminifolia (L.) Nutt. The taxonomic history of the Upland White Aster, Solidago ptarmicoides, has been reviewed by Brouillet and Semple (1981), and only a brief summary will be presented here. Not only the generic placement but also its species epithet has varied. The plant was originally described by Nuttallin Inula L., where it received the appropriate epithet alba for its white rays and disk HOMES Menem GE to either Aster or Solidago this epithet proved to be already occupied, and the alt its | to Achillea ptarmica L., a widely cultivated garden plant; Fernald 1950) v was BE ig Nees. When placed in still another genus, such as Oligoneuron (e.g., Nesom 1993), it reverts to the epithet album. Over its taxonomic history, the species has been placed in nine separate genera by various authors (Semple & Cook 2006). The taxonomic treatment of Solidago has been notoriously difficult (Semple & Cook 2006). Although there are a large number of e similar SE in eastern North America, no clearcut morphological apomorphy has been discovered that genus. Similarly, delimitation of infrageneric groups has been elusive as has aa and ion of individual species, many of which differ from one another by minute details of inflorescence form or pubescence. The situation is further compli- cated by the frequent occurrence of both interspecific hybridization and polyploidy. Some representatives Schillina et al.. Molecul lysis of Solidaster cv. Lemore 9 pa E d Fic. 1. Solidaster cv. Lemore, habit of the genus are quite common and widespread and are particularly significant as dominants during early successional stages of old fields, where several are components of a well studied ecological model system (e.g., Abrahamson et al. 2005). Additionally, Solidago taxa have become invasive in Europe (Weber 2001) and Asia (Dong et al. 2006), and are being used as models to understand the general evolution and ecology of invasive species (Van Kleunen & Schmidt 2003; Jakobs et al. 2004; Meyer et al. 2005). The abundance and ecological importance of Solidago therefore make a sound taxonomy highly desirable. A number of studies have applied molecular data to the resolution of systematic problems in Astereae (Suh & Simpson 1990; Morgan & Simpson 1992; Lane et al. 1996; Zhang 1996; Noyes & Rieseberg 1999; Roberts & Urbatsch 2003; Urbatsch et al. 2003; Beck et al. 2004), so a considerable body of molecular data has begun to accumulate for the tribe. Other than a chloroplast DNA restriction site analysis (Zhang 1996; Semple et al. 1999), there has been no focus as yet for DNA sequence analysis on Solidago, although a few sequences are available from other work. As part of an initial survey for suitable molecular markers, we analyzed ITS and ETS sequence variation for several samples of Solidago that represented the major subunits of the genus, as delimited by Semple and Cook (2006). Although the lack of suitable variation in these markers led us to abandon any attempt to make a complete survey of the genus, one of the species that we sampled was S. ptarmicoides, and sufficient data have been acquired to allow for an assessment of its generic placement, as well as to clarify the parentage of its hybrid offspring, Solidaster cv. Lemore, which we report here. MATERIALS AND METHODS Plant Material A sample of Solidaster cv. Lemore (Fig. 1) was grown from material obtained commercially (Bluestone Perennials, Ohio, USA). Samples of Solidago (Appendix) were either collected in the field or obtained from herbarium specimens. Sampling was designed to include at least one member of each of the currently rec- ognized sections and subsections of the genus (Semple & Cook 2006). To evaluate intraspecific variability, multiple accessions of four species (S. canadensis, S. riddellii, S. ptarmicoides, and S. nitida) as well as an additional sample of Brintonia discoidea were analyzed. Note that we have elected to follow the most recent treatment of Solidago (Semple & Cook 2006), which deviates from nomenclature used in our earlier paper 10 | tl | ical | f Texas 2(1) (Beck et al. 2004) by including Oligoneuron within Solidago but recognizing Brintonia parate from it; the classification of related genera also follows the Flora of North America treatments. Molecular Methods Preparations of total DNA made from fresh (0.5-2.0 g) leaves generally followed the procedure of Doyle and Doyle (1987), and those from herbarium material (ca. 0.1 g) were performed with the Dneasy Plant Minikit (Qiagen, Valencia CA). The crude DNA extracts of some samples required further purification using the Wizard Kit protocol (Promega, Madison, Wisconsin). Methods for DNA amplification and sequencing were as described in Schmidt and Schilling (2000). Amplification and sequencing reactions for the IT'S region were both performed using primers “ITS-4” and “ITS-5” (White et al. 1990). Amplification and sequencing reactions for the ETS region were performed using primers Ast-1 (Markos & Baldwin 2001) and 18-S-ETS (Baldwin & Markos 1998). All PCR products were checked by agarose gel electrophoresis. Sequencing was done at the University of Tennessee Automated Sequencing Facility, utilizing the ABI Prism Dye Terminator Cycle Sequencing reaction kit on an ABI 373 or ABI 3100 DNA sequencer (Perkin-Elmer Inc. Foster City, CA). The initial sequence data text files were edited following comparison with the same data displayed in four-color electropherograms before further analysis. Sequence alignment was performed using the Clustal X (version 1.6) program (Thompson et al. 1994 RIDE of the ITS region for the sample of Solidaster cv. Lemore was undertaken to confirm ITS poly- | from direct sequencing. The purified PCR products were ligated into pGEM-T (Promega, Madison: Wisconsin) according to the manufacturers instructions. Competent Topl0 Fº (Invitrogen, San Diego, California) cells were transformed via electroporation and the resulting colonies were screened for plasmas d inserts by PCR using the original amplification primers. Plasmids were isolated from single g an alkaline lysis/PEG precipitation protocol (Sambrook et al. 1989). Sequences were obtained for ten independent clones. Data for additional ITS and ETS sequences for Solidago were obtained from GenBank, as well as for all of the samples placed in the same clade as Solidago s.s. in Beck et al. (2004; “Clade III”). Two data sets were analyzed, the first of which utilized ITS sequences from a broad sampling of Solidago, with samples of Brintonia discoidea as outgroups. The second utilized both ITS and ETS data with a broader sampling of related genera (Appendix) to assess the placement of Solidago ptarmicoides and of Oligoneuron relative to other Solidago, and samples of Sericocarpus tortifolius and Cuniculotinus gramineus were added as outgroups for this analysis. ENEE E were analyzed using both maximum parsimony and Bayesian approaches. Parsimony analys ted using PAUP* 4.0b10 (Swofford 2003), with gaps treated as missing data, using a heuristic ac with 1,000 random addition replicates and with TBR branch swapping. Bootstrap analysis (Felsenstein 1985) was performed with 10,000 replicates using the FASTSTEP search option. Bayesian analysis was implemented in MRBAYES 3.0B4 (Huelsenbeck & Ronquist 2001) run for ten million generations with four separate chains and trees saved every 100 generations. The number of trees to discard as “burn-in” was assessed by plotting likelihoods of trees sampled throughout the run and discarding all trees prior to the stable likelihood plateau (in both analyses the first 10% were discarded). An appropriate maximum likelihood model of sequence evolution (GTR+I+G; General Time Reversible model with a proportion of invariant sites and gamma distributed rates) for the Bayesian analysis was chosen for both analyses using Modeltest (Posada & Crandall 1998). RESULTS Newly obtained ITS sequences for Solidaster cv. Lemore and for Solidago were consistent in length with previous reports for the genus. In particular, there was extraordinarily little variation in length among any of the sampled members of the genus; eae Sequence lengths varied from 627-629 bp. Insertion of single 1 bp indels ired for five samples (tl 1 two deletions) to produce a completely aligned matrix of length 631 bp for Ee entire ITS region. Similarly, there was little sequence length varia- tion in allied genera, with individual sequences varying from 627-630 bp, and the aligned matrix was of Schilling et al., Molecul lysis of Solidast Lemore 11 bi a 634. Tasse was also ony limited Palo variation between species of Solidago for the ITS region; airwise comparisons revealed sequence divergence f generally less than 1%, with total differences between samples of 1- 10 bp. Samples of Bier senem: were generally, although not always, more divergent, with pairwise divergence values of 1-3% and total differences of 7-19 bp. The sample of Sericocarpus tortifolius differed from all Solidago samples for at least 15 bp, but was still relatively similar, with overall divergence values of 2—396; the sample of Cuniculotinus gramineus was somewhat more divergent, differing by 23—29 bp (496 overall divergence) from samples of Solidago. In the five species (S. canadensis, S. riddellii, S. nitida, S. ptarmicoides, and Brintonia discoidea) where multiple accessions were examined there was at most one bp difference among samples and no evidence of nonmonophyly. The ITS sequence for Solidaster cv. Lemore was very similar to other Solidago sequences, but displayed bp polymorphisms (Fig. 2) at the following positions: 18 (A/G), 106 (C/A); 514 (C/T), 611 (G/T), 612 (C/T), 613 (A/G). Visual inspection revealed that this pattern of bp polymorphisms would fit exactly with a com- bination of the ITS sequences characteristic of S. ptarmicoides and S. canadensis (Fig 2.) Cloned ITS repeats from Solidaster cv. Lemore exhibited ITS sequences that matched closely either those of S. ptarmicoides or those of S. canadensis. Phylogenetic analysis in comparison to a broad sampling of Solidago showed that consensus sequences from the clones clustered with some ESTE DEEN ie those Eon S. pone D D D D TI J is į e and S. Roques id 3), on E pr g polymorj 1 from direct f a combination of tl f th ies. The ITS sequence of uland graminifolia, proposed by Nesom (1993) as the second potential parent of Solidaster, has been shown to be relatively divergent from those of Solidago (Noyes and Rieseberg 1999), and its reported ITS sequence (GenBank AF046982) differs from that of S. ptarmicoides by 4 indels as well as a minimum of 32 bp. Because the ITS sequence of Solidaster cv. Lemore did not show any evidence of indel polymorphisms, and only a few bp polymorphisms, Euthamia graminifolia is clearly not one of its parents. Approximately 450 bp of the ETS region were amplified for each sample of Solidago with the primer pair 18S-ETS and Ast-1. Removing regions of poor sequence and uncertain alignment produced a matrix with a total length of 444 bp, including 18 bp of 18S coding sequence. As with ITS, length variation among sequences was minimal among Solidago samples: only a single 1 bp deletion in S. patula, and only a single 1 bp insertion in Stenotus and a 3 bp deletion in both outgroup taxa, Cuniculotinus and Sericocarpus, were required for complete alignment of this portion of the ETS region. Levels of sequence divergence for ETS were similar to those for ITS, with pairwise divergence values among Solidago samples of 0-1% (0-4 bp) and among all samples reaching a maximum of 4% (17 bp). The ETS sequence of Solidaster cv. Lemore was basically identical to that of S. ptarmicoides, although each sequence ex hibited a single, non-informative bp polymorphism (position 124 in Solidaster, 36 in S. tadas both A/G). Other than the polymorphic positions, the ETS sequence of S. canadensis differed from those of S. ptarmicoides at only a single position, and there was no evidence of polymorphism in the sequence of Solidaster cv. Lemore at this position. For phylogenetic analysis to assess the placement of S. ptarmicoides relative to Solidago, the ITS and ETS sequence data were analyzed together, producing a combined matrix of 1078 bp in which there were 46 potentially parsimony-informative characters and 85 additional variable but parsimony-uninformative characters. The results of phylogenetic analyses (Fig. 4) reflected the overall low levels of divergence in providing a poorly Ee tree with ne dni > ver of EEN for many branches. The results of the parsimony and Bayesian h the Bayesian tree was more resolved. A monophyletic group m to clade HI af Beck et al. (2004) was defined relative to Cuniculotinus gramineus and Sericocarpus tortifolius with a posterior probability of 1.00 (Bayesian) and 100% (bootstrap), respectively. At the next level of branching there was a large polytomy in the strict consensus tree, within which there was a strongly supported clade formed by the three species of Chrysothamnus, C. scopulorum, C. stylosus, and C. viscidiflorus (1.00; 99%), a variably supported clade with Petradoria and Stenotus (0.97; 57%), and a strongly supported clade formed by Solidago, Chrysoma, and Brintonia (1.00; 76%). Within the last clade, there was a Chrysoma + Brintonia clade (0.79; <50%) that was sister to a Solidago s.s. clade (1.00; 69%). Within the Solidago s.s. clade, there was littl lution in the tree, with the only bootstrap y yy S. ptarmicoides | C ^ ^ CG C G T TG T CAT G A À T CAA! S. canadensis [cAAcCGCGT T GG T GT G AA T CA | l 240 250 Solidaster CAACGCGTTGGTGTG AA TI CA A bw 2 its putative parents, 5. canadensis and $. ptarmicoides. support greater than 80% for multispecific clades going to a strongly supported S. canadensis + S. gigantea clade (1.00; 94%); weaker support was provided to clades consisting of S. rigida + S. ohiensis (0.99; 64%), S. caesia + S. riddellii (0.62; 64%), and S. arguta + S. patula (0.50; <50%). The three samples of S. ptarmicoides were placed in a large polytomy with other members of the genus (Fig. 4). Results of the Bayesian analysis also provided weak to strong support for ci ps Geet less than 50% bootstrap support, Ge Petradoria/Stenotus with Solidago/Chry 'B Oreochrysum 4 Tonestus (0.51), with sequential addition e Lorandersonia (0.52), and Eastwoodia (0.76), and this clade combined with the three species of Chrysothamnus (0.60). These results are pletely congruent with those presented by Urbatsch et al. (2003) in placing Oligoneuron within Solidago as well as showing Chrysoma as a near outgroup to Solidago relative to Sericocarpus. They also support the revised classification of Chrysothamnus and related genera presented by Urbatsch et al. (2005). clades not present in the strict DISCUSSION Molecular data showed that Solidaster cv. Lemore is not an intergeneric hybrid by providing evidence not only to identify its parents (Figs. 2, 3) but to show clearly that both are accurately placed as members of Solidago (Figs. 3, 4). The combination of Solidago ptarmicoides and S. canadensis as its progenitors fits well with what is known of the origin of Solidaster, and has been proposed previously (e.g., Brouillet & Semple 1981). It has been clear based on morphology that S. ptarmicoides was one of the parents, and S. canadensis was one of the few goldenrod species naturalized near the nursery where Solidaster was first discovered (Nesom 1993). Although Solidaster does not show complete additivity in morphological features relative to S. ptarmicoides and S. canadensis (Nesom 1993), there are now numerous well documented cases of hybrids failing to exhibit one or more seemingly characteristic traits of their progenitors (Rieseberg 1995). Note that these results apply specifically to the cultivar Lemore, and other material marketed as “Solidaster” (or in florists shops as “Aster”), which may exhibit somewhat different morphologies, may not have a common origin. Laureto and Barkmann (2005, pers. comm.) present molecular-based evidence for the hybrid origin of another species, S. houghtonii Torr. & A. Gray ex A. Gray, that involves members of sect. Ptarmicoidei (S. riddellii) and sect. Triplinerve (S. gigantea), and similarly does not show additivity in morphology. Ge of Euthamia graminifolia as a potential parent of Solidaster also refut ther potential interg ization hypothesis (Nesom 1993). Schilling etal 1 M Solidaster Copy 1 «dm 0.73/53 S. ptarmicoides 1* S. ptarmicoides 2* S. ptarmicoides 3* Solidaster Copy 2 u 0.86 / 64 S. canadensis 1 1.00/94 S. canadensis 2 S. canadensis 3 S. gigantea —Ó 2 D Fit speciosa . sempervirens shortii rigida* arguta caesia 1.00 / 99 patula E -/ 59 virgaurea ———— E decurrens nitida 1* nitida 2* nitida 3* ohiensis* riddellii 1* riddellii 2* riddellii 3* simplex o flexicaulis S. juncea S. petiolaris S. nemoralis Brintonia discoidea 1 Brintonia discoidea 2 F 2 Phyl 4’ I A wf ITC £ Callas 1 f 1 leat A P E CAIS £ AC das " LI IG | nnl d 1 CL H J T Earr sa” I PE A H H la tal afr n o1.DI n 00V ahta L| i lysis. using Brintonia eter RN F AA J f 3 . £ el LR + i € 4 [| £ n » Il End, fig 4 all rans) al L L L as outaroup a a | ia a E ki Lu -f Ptarmicoidea (Oligoneuron). 1.00/76 0.94/- 1.00/100 Al h m 1.00/94 0.99/64 0.50/- c g S ES 1.00/98 0.73/- 1.00/98 l 1.00/100 0.97/57 1.00/99 0.51/- Solidago canadensis Solidago gigantea Solidago fistulosa Solidago sempervirens Solidago shortii Solidago ngida* Solidago ohiensis* Solidago arguta Solidago patula Solidago caesia Solidago riddellii 1* Solidago riddeilii 2* Solidago nddeliii 3* Solidago virgaurea Solidago nitida 1* Solidago nitida 2* Solidago nitida 3* Solidago ptarmicoides 1* Solidago ptamicoides 2* Solidago ptamicoides 3* Brintonia discoidea 1 Brintonia discoidea 2 Chrysoma pauciflosculosa Petradoria pumila Stenotus acaulis Columbiadona hallii Oreochrysum parryi Tonestus pygmaeus Lorandersonia linifolia Eastwoodia elegans Chrysothamnus viscidiflorus Chrysothamnus scopulorum Chrysothamnus stylosus Cuniculotinus gramineus Sericocarpus tortifolius land, C1—0.69, RI=0.85, obtained f a EITE AUTE Jak Shawn le A LL dis rea Li LEE j frre hai + Schilling et al Molecul lysis of Solidast Lemore 15 - E TS The major conclusion from phylogenetic analysis of ITS = ETS sequence data is that there is strong support for the inclusion of S. ptarmicoides within Solidago. PI learly placed it in the clade EE to Solidago s.s s Fig, 4), ine it Seneca in lle sequence (ITS and ETS combined) from other lit ff (vs. S. shortii). There wa sequence data to segregate Oligoneuron, presea in the data set by five of its six species dude S. ET as a distinct genus; the combined ITS and ETS data in fact placed one of the Oligoneuron species, S. riddellii, with S. caesia from sect. Solidago, albeit with only weak support (Fig. 3, 4; see asterisks). The sequence data did, however, provide weak support for restricting the limits of Solidago to exclude Brintonia and Chrysoma, although the group formed by inclusion of these genera with Solidago would be monophyletic. These results Cd mirror those presented by ara Kee reproduced in Semple et al. 1999) dran on pepe NA restriction site data, and ] tional support for the most recent and treatment of the abovementioned taxa (Semple & Cook 2006). A notable aspect of the ITS and ETS sequence data was the striking lack of divergence among mem- bers of Solidago. By contrast, ITS divergence in Eupatorium L. another genus of Asteraceae with a similar geographic range and occurrence in old field habitats, is 2-6% between species (12-43 bp differences; Schmidt & Schilling 2000; Schilling et al. 2007); Eupatorium is further differentiated from Eutrochium Raf., which are widely treated as congeners, by two indels (loss of a 13 bp region in Eupatorium and addition of a 3 bp region in Eutrochium) in addition to a minimum of 896 (33—61 bp) divergence. It should be noted that a previous chloroplast RFLP analysis (Zhang 1996) reports a higher level of variation within Solidago, and the resulting phylogeny strongly supports certain previously hypothesized groups. These contrasting levels of phylogenetic signal are best explained by the relative amount of DNA sequence scored in each study. Extrapolating from the values reported in Lane et al. (1996), approximately 2.5—3 kb of chloroplast sequence was assessed for variation in the Zhang study, compared to the approximately 1.1 kb ITS/ETS dataset analyzed here. In addition, previous work in Asteraceae indicates that chloroplast RFLP datasets can be more informative relative to those from ITS sequence (Morgan 1997). The most obvious possible source of the lack of ITS divergence in Solidago is that species level divergence is very recent. Other possible contributing factors include relatively large effective population sizes and the potential for homogenization through widespread interspecific hybridization followed by concerted evolution. Hybridization is tradition- ally viewed as widespread within Solidago (Cronquist 1980), and homogenization via concerted evolution is potentially a rapid process DIE o (Franzke & Mummenhoff 1999). The relative lack of differentiation in ITS seq y distantly related congeners, which because of geographical considerations are unlikely to be part di a Geen hybridization network, suggests that the primary cause is recency of divergence. This suggests that the striking ecological differences between Solidago species may have arisen during relatively short periods of time and indicates that resolving the phylogeny of Solidago will be a major challenge that will require markers evolving more rapidly than ITS and ETS. APPENDIX Ce of Solidago lat enera pled for molecular analyses, with collector and herbarium or literature refer- nce and GenBank ERC (ITS, ETS). Solidago L. Solidaster cv. Lemore, Schilling 07-06 (TENN), EU125353, EU125374. Sect. Solidago, Subsect. Argutae, 5. arguta Aiton, Beck481 (MO), EU125354, EU125375; S. patula Muhl., Beck 482 (MO), EU125355, EU125376. Subsect. Glomeruliferae, 5. caesia L., Beck 483 (MO), EU125356, EU125377; S. flexicaulis L,Kress et al. 2005, DQ005979. Subsect. Humiles, S. simplex Kunth, Kress et al. 2005, DQ005982. Subsect. Juncea, S. juncea Aiton, Kress et al. 2005, DQ005981. Subsect. Maritimae, 5. sempervirens 4, Urbatsch et al. 2003, AF477668, dps Subsect. Nemorales, S. nemoralis Aiton, Estes d d EU125357. Sub- sect. Solidago, 5. virgaurea L., Dinies n. (MO), EU125358, EU125378; S. decurrens Lour. EF103140. Subsect. Squarrosae, S. speciosa Nutt., p 07-05 (TENN), EU125359. Subsect. Thyrsiflorae, S. petiolaris Aiton, Noyes & Rieseberg 1999, AFO46968. Subsect. Triplinerve, S. canadensis L., Urbatsch et al. 2003, AF477665, AF477729; Laferriere 3564 (TENN), EU125360, — Weldon I 7/22/81 (TENN), EU125361; S. gigantea Aiton, o (MO), EU125362, EU125379; S. shortii Torr. & A. SE Beck etal 2004, ib [submit]. Subsect. pao fistulosa Mill, Urbatsch E al. acd AF477666, b bin ser t. ay) B. Boivin O), EU125363, EU125380 16 t i titute of Texas 2(1) (MO), EU125364, EU125381; And MO221, EU125365, EU) 25382; S. nitida Torrey & A. Gray, Thomas 141724 (MO), EU125366, EUT25383; Thomas 97470 (MO), EU125367, EU125384; Thomas 138143, EU125368, EU125385; S. ohiensis Riddell, Kral 48497 (MO), EU125369, EU125386; S. riddellii Frank, Smith 3617 (MO), EU125370, EU125387; Vogt: »06 (MO), E e Luges s.n. (MO), EU125372, EU1 25389; S. rigida L., Beck et al. 2004, AY523851, EU125390. B (E Greene, ROBOTS E Urbatsch 2003, AY 170930, AY! 69727; Anderson 20021 ipis: EUIS, vq uh dcum: Nutt C Urbatsch et al. 2003, ME AF47 Cl pul (M.E. Jones) Urbatsch, R.P. Roberts & Neubig, Roberts & Urbatsch 2003, AY 170956, erc e SE (East ) Urbatscl RP en & Neubig, Roberts & Urbatsch 2003, AY170973, pe iscidifl rts & Urbatsch 2003, AY1 70947, AY169744. o o a E, c hallii (&. Gray) G.L. Nesom, Roberts e Urbatsch ei AE. a a rd Urb ibig, C. gramineus (H. M. Hall) Urbatsch, R.P u 70936, AY] pul Eastwoodia ro E. elegans Brandegee, Roberts €: Urbatsch 2003, AY! 70949, AY169746. ust iue. Urbatsch, P dad rts & Neubig, L. linifolia (Greene) Miedo R.P. Roberts & Neubig, Roberts & Urbatsch 2003, AY170936, AY169737 Ib., O. parryi Rydb., Roberts 8 atsch 2003, AY 170958, AY169755. Petradoria Greene, P pumila Greene, Roberts & Urbatsch 2003, AY! ie AY 169756. Seri lees, S. tortifolius Nees, Urbatsch et al. 2003, AF477664, AF477728. St lutt., S. acaulis Nutt., Roberts €: Urbatsch 2003, AY1 70960, AY169757. Tonestus A. Nelson, 7. pygmaeus À. Nelson, Roberts & Urbatsch 2003, AY1 70972, AY169769. ACKNOWLEDGMENTS The authors thank G. Beattie, P. Heise, and J. Miller for technical support, and G. Nesom and J. Semple for helpful comments on the manuscript. Financial support from the L.R. Hesler Fund of the Department of Botany, University of Tennessee, and NIH award #P20 RR164841, is gratefully acknowledged. REFERENCES ABRAHAMSON, W.G. K.D. DosLey, H.R. 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NOVAE GESNERIACEAE NEOTROPICARUM XV: KOHLERIA HYPERTRICHOSA, A NEW SPECIES OF GESNERIACEAE FROM NORTHWESTERN ECUADOR John Littner Clark Laurence E. Skog Department of Biological Sciences Botany Department, MRC-166 Box 870345 Smithsonian Institution, PO Box 37012 The University of Alabama National Museum of Natural History Tuscaloosa, Alabama 35487-0345. U.S.A. Washington, DC 20013-7012. U.S.A. skogl@si.edu ABSTRACT A new species of Kohleria (Gesneriaceae, tribe Gloxinieae) is described from the Andean cloud forests of the Carchi and Esmeraldas = DH el 4 ke 1 1 [PRI 1 J 41 1 1 = "TL 1 1-1 T 1 pe we | £ 1.5 ws Lm | + Se dl E 1 E T zl T le JA e| P 1 Axa Iff Kc uk] D 1 Ae? zl [. 41 Í L^-Lkl1 t RESUMEN Cc J 1 : 1 Tah] H Feo cl pase | roy z : ^ A 1 1 17 ] T2 q 1 s : a ST BS d y E ld el noroeste de Ecuador, en donde es una especie endémica abundante localmente. Los largos tricomas lanosos junto con la dehiscencia del fruto por medio de una hendidura dorsal y el hábito epifítico diferencian a Kohleria hypertrichosa de otras especies de Kohleria. Key Wonps: Kohleria, Capanea, Monopyle, Ecuador, Gesneriaceae The most recent treatment of the genus Kohleria Regel is that by Kvist and Skog (1992), who recognized 17 species. More recently, Roalson et al. (20052) transferred the two species of the genus Capanea Decaisne ex Planchon to Kohleria making it currently a genus of 19 recognized species. The revised circumscription of Kohleria to include Capanea (Roalson et al. 2005a) is based primarily on molecular data from the nuclear ribosomal DNA internal transcribed spacer region, the chloroplast DNA trnL intron and trnL-trnF intergenic spacer region (Roalson et al. 2005b). Traditional Kohleria (e.g., the recent monograph by Kvist and Skog, 1992) would be paraphyletic with the exclusion of Capanea. Thus, many of the features that differentiated Capanea are autapomorphic for a well-supported clade nesting in Kohleria (Roalson et al. 2005b). The new species described here has a history of being collected in the Carchi province where it is locally abundant in cloud forests. It was initially assumed to be a new species of Capanea by Hans Wiehler when it was collected and appeared in a photograph in the horticultural journal The Gloxinian (McDowell 1995). Images of K. hypertrichosa also appeared in the Gesneriad Journal as a new species of Capanea in a report of a collecting expedition to Ecuador (Dunn 1997). The species was assumed to be a member of this formerly recognized pane ARN e Ge woolly pubescence the shape of the corolla, and its epiphytic habit. Vegetatively Kol y] pr ilar to dad Lip because of the SHORE mi d eis leaves +] ] CT Ah] the eneripe nth X and dorsiventral shoots. I ol is also abundant (cf., field ci É Clark 6298). The two species often grow E by-side and e them apart vegetatively is challenging. The species described here is remarkable in that it has been collected and specimens annotate d as be- longing to Monopyle or Capanea, but not Kohleria. The correct placement of this new species has perplexed many Gesneriaceae workers because of its habit (facultative epiphyte instead of the more typical terrestrial habit in Kohl ena Gees E pune on the outer surface of the corolla Ee typical of Capanea); and . of Monopyle) The key sy y tor recognizing traditional str ongl Kohleria is a ERU dehiscing by a single dorsal slit from apex to base (Biss 1D and 2G). The presence of this type of fruit in this species was unknown until it was observed and photographed in 2001 (cf., field collec- tion J.L. Clark 6288). Many other collectors (e.g., Tirado and Hoover) had documented this new species in J. Bot. Res. Inst. Texas 2(1): 19 — 24. 2008 [| [| £ al Dos A In LI POP ET */41 20 flower, but not in fruit, thus making it a difficult species to place without molecular data. The presence of a bilobed stigma is a SR ee that K. LL adn shares SH most dani adi Á EE stigma is present in À y in the two traditionally g d species of Capanea that were eg — in Kohleria o et P 20053). Another morphological feature of many species in Kohleria, which is also present in K. hypertrichosa, is the presence of a ventricose apical pouch on the lower side of the corolla. The ventricose pouch in K. hyper- trichosa is sometimes obscured a d: and on pressed herbarium collections (e.g., the pouch was not EES in ds photograpl t e L. Clark & R.W. Dunn 2406 and J.L. Clark et al. 2408) Th t of K. hypertrichosa is al ly supported by molecular data from the nuclear ribosom al DNA internal t ] region, the ala DNA trnL intron and trnL-trnF intergenic spacer region (Roalson et al. 2005b). One of t] that appeared in the pl y published by Roalson et al. is a tissue sample from silica Ln dried leaves of a paratype of K. eri hasi (cf., field collection J.L. Clark & R.W. Dunn 2446). The sp llection is noted in Roalson et al. (2005b) as, “Kohleria sp. nov. 2446” where it was shown as being strongly supported as the sister taxon to Kohleria villosa (Fritsch) Wiehler. Kohleria hypertrichosa J.L. Clark & L E Skog, SP. nov. (Figs. 1-2). Tee: ECUADOR, Carchi: Cantón Espejo, Par- roquia Guatal, Mirador de las Gol olondrinas), trail from Santa Rosa (El R ) to the refugio at El Corazon, 00?49'46"N, 78°07'03"W, 1600-2000 m, 5 Jul 2003, JL. Clark, S.G. Clark, E. Folleco & B. Syka 8450 (HoLorveE: US; isoTYPES: AAU, BRIT, C, CAS, COL, E, E GH, K, MO, NY, QCA, QCNE, SEL, UNA, US, W). A A et à D 1 t 1 Lara 4$ halb "ael PAn 4 ` ; lh d rrr + r E D f is bivalvati dd liter 1 itudinaliter unifissis, distincta. Facultative epiphytic climber; rarely terrestrial, stems dorsiventral to erect, frequently branched, to 2 m long, subwoody to herbaceous, terete, glabrescent below, densely pilose to sericeous above. Leaves opposite, unequal in a pair; larger leaf with petioles terete, 3-10 mm PAR green, RÀ sericeous, blade EE ceous when dry, elliptic to slightly falcate, 6-14 x 1.5—4 cm, base acute an trical, apex attenuate, margin serrate, adaxially pale green, sparingly to densely pilose especially on Hs) ais green to reddish-green, uniformly pilose; smaller leaf with petioles sessile to 3 mm long, green, densely sericeous, aue subcoriaceous when dry, broadly ovate (rarely narrowly ovate), 1-2.5 x 1-1.5 cm, base and gin serrate adaxially pale green, sparingly to densely pilose (especially on veins), abaxially seen to reddi en uniformly pilose. Inflorescence epedunculate, reduced cyme, appearing fasciculate, with 1 (rarely 2) flower per node at or near stem apices; bracteoles absent; pedicels longer than the petiole, 1.5-3 cm long, densely sericeous. Fl hic, not inate; calyx lobes 5, ventral lobe free, lateral and dorsal lobes basally connate for 3-5 mm, SEET erect at anthesis, reflexed in fruit, equal, ovate, 1.5-2.0 x 0.5-1.0 cm, apex acute, margin nearly entire with 13 pairs of serrations, bright red, outside densely sericeous, dd: pasen sericeous; EE e in niet 2.5-3.0 cm long; base 1 cm in diameter, mpliate on , throat slightly constricted, EE laterally — a to 1 cm wide Ge — outside yellowish-white, eegen with dense villous inside mostly yellow with red spots, mouth with glandular trichomes, lobes glabrous, red, subequal — to 3 mm long, to 5 mm wide, rounded, entire; nectary of five separate mem M distributed around ovary, each 1.0-1.5 mm high and 0.5-1.5 mm wide at base, glabrous; stamens 4, didynamous, included; filaments 1.6 cm long, adnate to the base of the corolla tube, pilose; anthers longer than broad, ca. 2 mm long, ca. 1.5 mm wide, dehiscing by longitudinal slits; staminode not observed; ovary inferior, villous, 2-5 x 3—6 mm, style 9-12 mm long, sericeous, stigma included, forked. Fruit an oblong capsule, 20-30 mm long, diameter 4—8 mm, dehiscing by a single dorsal slit from apex to base; seeds numerous, subglobose, irregularly striate, 0.4 x 0.3 mm, dark brown. Phenology.—Flowering in January, April, May, November, and December. The only collections with fruits are from May, July, and December. Distribution. —Kohleria hypertrichosa is known from northwestern Ecuadorian cloud forests on the western Andean slopes of the Carchi and Esmeraldas provinces from 1100 to 2000 meters. It is locally Clark and Skog, A new species of Kohleria from northwestern Ecuador E Fic. 1, Kohleria hypertrichosa J.L. Clark & LE Skog. A. Front view of corolla. B. Immature flower. C. Front view of capsule. D. Fruit dehiscing by a single dorsal slit. E. Lateral view of flower. (A from J.L. Clark et al. 6359; B & E from J.L. Clark et al. 8462; C & D from the holotype, J.L. Clark et al. 8450) UN É pe RES d d | M = ed == M NS Tak SD A SS rs «y! JI SUN GN A el l P | PS | all De 3 Vais Sat asf i 11 Clark 2, | E Clan À Hahit D GR 2 ££ rr H En n d LI d et al. 6359). ds. G. Mature fruit. H. Seeds. (A-H from LL. Clark Clark and Skog, A pecies of Kohleria f ti tern Ecuad 23 abundant sometimes covering trees and trailsides, especially in the Mirador de las Golondrinas, a private reserve managed by Fundación Golondrinas and the Gualpi area near the Reserva Awá (collections exist both inside and outside the Territorio Awá). No collections of Kohleria hypertrichosa have been made outside of the Carchi and Esmeraldas provinces, but it is likely to also occur in the adjacent Department of Nariño, Colombia where limited recent botanical exploration has taken place. Etymology.—The epithet hypertrichosa is derived from the congenital condition of generalized “Hyper- trichosis,” the medical term referring to a condition of excessive growth of body geg in humans. The condi- tion is commonly known as “Werewolf syndrome” which comes from a mythological werewolf of which the person is completely covered in hair or fur. Although not Ge PINE in ión: the plant species is known among horticulturalists as “Chewbacca,” for the famous hairy biped character (^Wookie") from the popular cult phenomenon 1 1977 movie Star Wars IV: A New Mad d E | T Jla eaha PARATYPES. ECUADOR del Corazón, 00%49'N, ii ed W, SS 20001 m, BAN 1996 JL L. Ge P. Sabbe Gë W. Dam id (MO, QCNE, SRP, US); Cantón Espejo, , trail SW of La Cal I ine, 00 Mi dp Hee 2000 m, ial 1996 ,J.L. Clark & RW. Dunn 2446 (MO, QCNE, SRP, US); C lor d of Las Juntas and La Cabaña del Corazón, 00°49'N, reor w, 1400- 2000. m, Er ER 1996 J L Mic & R.W. Dunn 2466 (MO, OCNE, SRP, US); Cantón Tulcan, Parroquia Chical, path from th “Gualpi” (near the border e the Reserva did ek iios f bi 1200—1700 m, 6 Dec 2001 (2. J.L. Clark & O. an 6288 8 (MO, QCNE, Sach Kee Sie ly as “Crystal” via Ri (ca. 6- GER SW B Chical), 0095349" 'N, 78?34"W, 1200- ii»: m, Roc 2001 om Ge Ji. Eu O. ac al e d E E Ed MO, NY, QCA, QCNE, US); Cantón Espejo, P fi zon towards La Cortadera (2 km NE of o 00°49) 46"N, 78207 03"W, a m, dg ir d Ge J L. Clark & E. Folleco m (AAU, CAS, COL, E, F, K, MO, NY, QCNE, SEL, US, UNA); f 78°8'W, 1890 m, 28 Nov 1987 (fD, W.S. Hoover & S. Wormley 1906 (MO): embankments along Río Verde, from point at on trail = Rafael's Mountain Finca crosses river, 1.5 km, 00°52'N, 78°8'W, 1890 m, 29 Nov 1987 (fl), W.S. Hoover 1918 (MO); ridge to NE of Rafael Quind's mountain finca, 0°52'N, 78°8'W, 2000 m, 29 Nov 1987 (fl), W.S. Hoover 2046 (MO); Gualpi Chico, Awá encampment, trail on reservation border going iiid: Ge N, 78°16'W, 1330 m, 15 e 1988 (fD, W.S. Hoover, P. Gelpi, R.A. Lorentzen & A. Arguello 2463 (MO, 1 El 0°58'N, 78°16'W, 1330 m, 20 Jan 1988 (1D), W.S. Hoover, A. Arguello, P. Gelpi n RA. Greg 2832 (MO, US) trail to Pailon on scam DERE Gualpi Chico area of Awd, 00°58'N, 78°16'W, 1350-1400 m, 21 Jan 1988 (fD, W.S. Hoover, A. Arguello, P. Gelpi & R.A. Lorentzen 3618 (MO, US); Cantón Tulcan, Parroquia Tobar Donoso, sector El Baboso, Reserva Indígena Awa, 00º53'N, 78°20'W, 1600 m, 3 Oct 1991 (fl), G. Tipaz, D. Rubio & M. Taicuz 267 (SEL, US); Cantón Espejo, Parroquia Tobar Donoso, Reserva Indígena Awá, centro Baboso, 00°53'N, 78°25'W, 1800 m, 17-27 Aug 1992 (fl), G. Tipaz, M. Tirado, C. Aulestia, N. Gale & P. Ortíz 1800 (US); Cantón Mira, El Carmen, Cerro Golondrinas, 00°50'N, eebe, 2000-2400 m, 18-25 Aug 1994 (fD, M. Tirado, P. Fuentes, R. Zurita & L. Chamorro 1286 (MO, QCNE, US); Espejo, R d , El Corazón, sendero a Río El Corazón, 00°50'N, 78°08'W, 2010 m, 24 Jan 2004 (fl), H. Vargas et al. 4406 (MO, US); E idas: C San Lorenzo, Parroquia Alto Tambo, mature forest 4-8 km W of El Cristal, 00°50'16"N 078?31'04"W, 1500-1650 m, 27 May 2008 (fr), J.L. Clark, J. Melton III & O. Solarte 10310 (CAS, E, F, K, MO, NY, QCNE, SEL, UNA, US, W); Cantón San Lorenzo, Parroquia Alto Tambo, mature forest 4—8 km W of El Cristal, 00°50'16"N 078°31'04"W, 1500-1650 m, 27 May 2008 (fl), J.L. Clark, J. Melton III & O. Solarte 10311 (QCNE, SEL, UNA, US); area of Cristal, above Lita, 1100-1150 m, 24 Apr 1995 (fD, H. Wiehler et al. 9574 (SEL, US). ACKNOWLEDGMENTS Support for this project for the first author was provided by the Elvin McDonald Research Endowment Fund of The Gesneriad Society, the National Science Foundation (DEB 0206512), the Explorers Club Washington Group, and a diu s EE Study and Research Abroad Scholarship. We are grateful to Alice Tangerini for preparing illustrations of the n cies and Harold Robinson f Isti g with the Latin diagnosis. Herbaria MO, QCNE, and SEL are Mice due for making their collections available to us. We also acknowledge Eric Roalson and Christian Feuillet for their careful reviews and comments on the manuscript. REFERENCES Dunn, R.W. 1997. A bright and shining star. Gesneriad J. 8(3—4):4—9. McDoweLL, M. 1995. ;Dondé Está Lita? Gloxinian 45(4):16-22. Kvist, L.P. and LE Skoc. 1992. Revision of Kohleria (G Í ). Smithsonian Contr. Bot. 79:1-83. 24 laries in the Gloxinieae RoaLson, E.H., J.K. Boacan, and L.E. Skos. 2005a. Reorganization of tribal and generic (Gesneriaceae: Gesnerioideae) and the description of a new tribe in the Gesnerioideae, Sphaerorrhizeae Selbyana 25:225-238. RoaLson, E.H., J.K. BOGGAN, L.E. Sec, and E.A. Zimmer. 2005b. Untangling Gloxinieae (G | ). 1. Phylogenetic patterns and generic boundaries inferred from nuclear, chloroplast, a morphological cladistic datasets Taxon 54:389-410. TWO NEW SPECIES OF GUADUA (BAMBUSOIDEAE: GUADUINAE) FROM COLOMBIA AND BOLIVIA Ximena Londoño Eneida Zurita Instituto Vallecaucano de Herbario Nacional Forestal Investigaciones Cientificas “Martin Cárdenas” A 11574, Cali, COLOMBIA aa Bod ximelon@telesat.com.co eneidaz@gmail.com ABSTRACT Guadua Hana a G. Ge two ne species of woody bamboo from South America, are described and illustrated. Guadua incana, fi 1 in the foothills of tl t ide of the Cordillera de los Andes, and G chapa- rensis, from DESTA occurs in "Be FM lowlands in the District of Chapare. Based on morphological evidence, G. incana and G. to Guadua weberbaueri, which is lectotypified here. We discuss other related species and provide a y the faleu Regi to them. RESUMEN Se describen e ilustran d peci para Sur América papi Md G. chaparensis. Guadua incana del suroriente de Colombia, crece en el pie d te del tiente oriental d e los Andes, y G. chaparensis de Bolivia, se localiza en el Distrito Biogeográfico Amazónico del Chapare. Con base en evidencias morfológicas, G. incana Gs G. MU CAD se relacionan entre si y también con Guadua weberbaueri, la cual se lectotipifica aquí. Además, se hace otras especies afines y se incluye clave para identificación. During a 1987 field trip to Caquetá and Putumayo, Colombia (Colciencias-Inciva Project No. 2108-07-009-85), sterile specimens of a new species of Guadua (Londoño & Quintero 144 & 214) were collected. A living rhizome was brought to the bamboo germplasm bank of the Juan Maria Cespedes Botanical Garden in Tuluá, Valle del Cauca, Colombia, and after 14 years in cultivation (1987—2001), accessions Londoño 144 & 214 flowered and made possible their identification. Because of the absence of flowering material in the majority of woody bamboo herbarium specimens (in part, a result of their long flowering cycles), several species of bamboo are described from vegetative material alone, e.g., Eremocaulon setosum Londoño & L.G. Clark, Chusquea riosaltensis L.G. Clark and Ch. caparaoensis L.G. Clark (Londoño & Clark 2002a; Clark 1992). Vegetative characters such as culm leaves, buds, branches and rhizomes have been utilized to delimit many new species (McClure 1966; Judziewicz et al. 1999; Soderstrom & Young 1983). We waited 14 years for the opportunity to describe G. incana with flowering material, but for the new species from Bolivia, G. chaparensis Londoño & Zurita, we decided to describe it on the basis of sterile material alone, in order to provide a name for this distinctive species. In addition to the vegetative and reproductive characters used to delimit G. incana, the genetic AFLP analysis done by Marulanda et al. (2002) reported a clear genetic differentiation between G. incana and G. angustifolia (accessions, varieties and biotypes), and also between G. incana and Guadua uncinata Londoño & L.G. Clark (2002b), a species that is sympatric with G. incana. This genetic result helped corroborate the diagnostic morphological features that identify Guadua incana as a new species. Guadua chaparensis was collected by the second author in central Bolivia. It occurs not only in the type locality (Comunidad Israel) but in various communities such as Capinota, Hermanos Ledesma, Agua Rica, 24 de Septiembre, Monte Sinaí, Villa Jerusalen, Villa Imperial, Valle de Sajta, Valle Ivirza, Villa Nueva, Alto San Pablo, Tarija and Puerto Aroma, all of them in the Chapare area. The two new pecies, G. incana and G chaparensis, are clearlv ref to Guadua | 1on their] ked rhizomes, white bands on the culms, thorny branches, enla culm leaves, and, in the case of E incana, pseudospikelet structure with winged-keeled palea (Judziewicz et al. 1999). Both species are closely related, | Rot Res inet Tayac 2(1): y UA 34. 2008 26 I Ze Dos a In LI Eine £'T, ^n BIT) grow in lowland tropical rain forests, and appear to be endemic to the localities in which they were collected. Both are also utilized by local communities for corrals, fences, water conduction and light rural construction. Guadua weberbaueri, the taxon most closely allied to the two new species, was described by Pilger (1905), who selected as the holotype number 4562 collected by the German botanist August Weberbauer (1871-1948) in Moyobamba, Perú. As with many other collections by Weberbauer, the type chosen by Pilger and deposited in Berlin (B) was destroyed during World War II. In 1991, the first author visited the herbarium of the Universidad Nacional Agraria, La Molina (MOL) in Lima, Peru, and found a complete du- plicate of Weberbauer 4562. This isotype, which we have selected to serve as the lectotype, includes a foliage leaf complement and one inflorescence branch. Morphological study reveals that Guadua weberbaueri Pilger, G. ol usd Sege and e sarcocarpa Londoño & Peterson (1991) share several features with each other and the two species, forming the basis for a group within Guadua, here recognized informally as the G. weberbaueri eos (Table) The following combination of characters delimits this group: a) culms erect at the base with the apical portion leaning on or pendent from trees; b) internodes hollow, with walls up to 1.5 cm thick, elongated up to 90 cm; c) culm leaves with persistent blades, these 1/4 to 1/5 as long as the sheaths; d) culm leaves with canescent inner ligules; e) synflorescences terminating in leafy or leafless branches with capitate coflorescences in the earliest state of development; f) pseudospikelets with wide-winged palea keels (0.8 to 2.5 mm); and g) ovaries fusoid in shape. These five species, Guadua weberbaueri, G. sarcocarpa, G. incana, G. chaparensis and G. tagoara, grow in very humid, lowland forests and they have in common the presence of water inside of the hollow in- ternodes. Herbarium labels from Guadua sarcocarpa (Calderon & Soderstrom 2348-US#2810196 and Smith 5275-US#3080541), the description for G. tagoara in Londoño and Clark (2002b), and Louton et al. (1996) for G. weberbaueri reported the presence of internodes filled with water. This internal water phenomenon seems to be associated with very high relative humidity and a high water table. Why water accumulates inside the internodes is still uncertain. It may occur after strong changes of temperature that break tissues allowing water to leak into and accumulate in the lumen. A key to the species of the G. weberbaueri group based on vegetative characters is presented here. KEY TO SPECIES OF THE GUADUA WEBERBAUERI GROUP 1. Culml | ally F | | let ly with difficulty from tl f th j | li t the junction of the sheath and blade: 2. Foliage leaf sheaths abaxially pubescent, bearing fimbriae at the summit; folage leaf blades abaxially pilose and inconspicuously tessellate; inner ligules pubescent; pseudopetioles abaxially setose-pilose; culm leaf auricles absent G. weberbaueri 2. Foliage leaf sheaths abaxially glabrous with oral setae and fimbriae at the summit; foliage leaf blades aaa lis dd a and conspicuously tessellate; inner ligules glabrous; f auricles dBSent or present G. sarcocarpa L Culm leaves Tn densely pubescent to canescent, ti letached from ti f the g aig tly tthe Junon of the sheath ai blade. 3 Cul SR | ing heee, 6 | -foliage leaf blad glabrous o on bot surf. | abaxially strongly! late; | lopetiole abaxi ially gla! ; Brazil G. tagoara 3. Cul hal ; ee | n m 4. Foliage E sheaths glabrous fimbriae al uE summit fimbriae 4-5 mm long; inner x figules glabrous; leaf I liy glabrous; Colombia . incana 4. Foliage id sheaths subglabrous is fimbriate at summit; inner ligules puberulous; leaf m abaxially g | G. chaparensis Guadua incana Londoño, sp. nov. (Fig. 1). Tre: COLOMBIA. Caquetá: Km 26.7 via Florencia-Guadalupe, Quebrada La Rebolcosa, vertiente oriental de la Cordillera Oriental, 750 m, cultivated at the Botanical Garden Juan María Céspedes, 2 Jun 2001 (fl), X. Londoño 972 (poLorvre: COLI, isorvres: CUVC, TULY, ISC, US Bambusa lignosa, spinosa. Rhizoma sympodiale, pachymorphum. Culmi 10-15-18) m alti, 7-9(-12) cm diam.; internodia 20-)30-65 cm longa, cava. Folia culmorum leviter coriacea, canescentia, deciduas, vagina et lamina conspicue distinctae in ju- fire E: E en m pa * pn Em A e Et, Se e bru gU pu remi E IAE (A E m L—.5 IET Rn UA oe E a en MÀ é SÉ ~ h CE uA + ET. s e samt e iu. ` ` cmm EE = : EE Wf De E eg NS "o Eus die BIN "m i EN TÀ x dun má a Fic. 1. Guadua incana. A. Culm leaf, sdayial viaw R m H E | | K .L LI L| L [| L all “a adavial «urfara Ü mr." | gl a - ed wë we area, abaxial view D | t of leafv fl i is EP losnikelet sl i | t , a sterile LI lemma, fertile florets, it inal rudi t fl ECH Prophyll deg view ano apex H- I Basal gemmiparous bracts, abaxial view and ap J- K. Lemma, abaxial view and apex. EN Palea, baxi 0-Q. Ca aryopsis with a stylar column; Sr | view (0). (A-C, london 144, TULV; D-Q, Londoño 972 TULV). || E sl n.a H Inm LI d. da adis £T. TA Vi 28 ventute; vagina (19-)24-33 cm longa; lamina (3.535-10 cm longa, persistens, erecta, triangularis. Ramificatio intravaginalis. Folia ramorum cujusquisque complementi 5-8(-10); vagina subglabra, fimbriata, margine imbricata, conspicue e ciliata; pseudo- petiolus 2-4 mm longus; lamina (3.5211-13C-16) cm longa, (0.7-)1-2(-2.2) cm lata, lineari-lanceolata. Synflorescentia 1-4(-6 coflorescentiis, 1-5 multifloris pseudospiculis munitis. Pseudospiculae 2-4 cm longae, 0.4-0.7 mm lata; lemma 9-11.5 mm longum, 6-8 mm latum; palea 9-11 mm longa, 4-5 mm lata. Ovarium fusiforme, (1.522-3 mm longum, 0.4-0.5 mm latum. Woody, thorny bamboo. Rhizomes pachymorph. Culms 10-15(-18) m tall, 7-9(-12) cm in diam., erect at base, arching apically, whitish-green when young; internodes (20-)30-55(-65) cm long, cylindrical, hollow, some filled with water, the wall 1-1.5 cm thick, the surface whitish-green pubescent when young, becoming evidently whiter below the nodal line, covered by very appressed, white, soft, slender, retrorse hairs; nodes solitary, the nodal line horizontal, with a supranodal band 1-1.7 cm wide of appressed, white, soft, slender hairs, and an infranodal band 1.5-2 cm wide of long, soft, slender, white, appressed hairs, the two bands unambiguously evident, the supranodal ridge slightly pronounced, the canescence from the infranodal band extending 2/3 along the internode; bud single, triangular, the shoulders of the prophyll ciliate. Culm leaves (22230—44 cm long, 29-44 cm wide at the base, coriaceous, deciduous, triangular, blade and sheath conspicuously distinguished by color and pubescence when young; sheaths (19-)24-33 x 29-44 cm, whitish-green with yellowish spots when young, becoming whitish-brown to stramineous, abaxially canescent-hispid, densely covered by two different types of hairs (1) short, matted, white and wavy hairs, looking like wool and easily detached, and (2) straight, stiff, sharp, amber hairs, up to 3 mm long, adaxially glabrous and shiny, the margins papery, the overlapping one ciliate, the cilia transparent, up to 2.5 mm long; inner ligules 0.5-1.0 mm long, truncate, straight to slightly curled in the middle, extending completely from margin to margin, adaxially glabrous, abaxially canescent, densely ciliate on the margin, the cilia up to 0.6 mm long, whitish in color and conspicuous; blades (3.5-)5-10 x 7-12 cm, 1/4—1/5 as long as the sheath, triangular, erect, persistent, inflated, green to reddish-green when young then brown, abaxially glabrous at the central inflated part, sparsely hispid and pubescent through the apex and margin- ally, adaxially densely pubescent between the nerves, covered by two types of hairs (1) firm, stiff, transpar- ent hairs up to 2.5 mm long, and (2) short, soft, loose hairs up to 0.5 mm long, the margins papery, shiny and smooth at the middle and upper portion, basally ciliate, wrinkled at the junction with the sheath and bearing conspicuous fimbriae and cilia, the cilia up to 2.5 mm long, hyaline, rubbing off at the middle and upper portion, the fimbriae 3-5 mm long, ivory, wavy to curled, basally scabrid, apically smooth, the apex strongly mucronate and usually split in two parts, the mucro up to 1 mm long. Branching intravaginal, consisting of one main branch and 1 to 2 secondary branches, the branches manifestly armed, thorns 1 to 3(-4) per node, the central one dominant, bigger and straight to slightly curved, the other two slightly curved to curved. Foliage leaves 5 to 8(-10) per complement; sheaths abaxially pubescent to glabrous, usually with a patch of villous hairs on one side of the midnerve at the upper portion, green when young then stramineous, the overlapping margin conspicuously ciliate, the underlapping one minutely cililate to smooth and papery, bearing a few fimbriae at the summit; fimbriae 4—5 mm long, ivory, basally straight and scabrid, apically wavy to curled and smooth; inner ligules 0.2-0.3 mm long, short, brown, puberulous, ciliolate to smooth at the margin; outer ligules 0.3-0.4 mm long, stramineous, glabrous, shiny, the margin ciliolate to smooth; pseudopetioles 2—4 mm long, adaxially da toward one margin, otherwise gla- brous, abaxially glabrous, pulvinate, the pulvinous viilous, st to brown; blades (3.5-)11-13(16) x (0.721-2C2.2) cm, LW = 5-141, 13-15 nerved, linear-lanceolate, adaxially glabrous to subglabrous, with scattered, strigose, transparent hairs up to 1.2 mm long, with 4—6 raised scabrid submarginal nerves on one side, becoming basally evidently scabrous by one side, abaxially glabrous to subglabrous, with scattered strigose, transparent hairs up to 1 mm long, papillose and tessellate, the midnerve and primary nerves prominent and yellow in the middle and lower portion, one EEN EE the other scabridulous, the apex acuminate, with a mucro 2-3 mm long. Synfl ting usually leafy branches of all orders, polytelic, consisting of 1-4(-6) MN with 1-3C5) riuiletflowered pseudospikelets; main axes basally glabrous, apically pilose; subtending bracts varying in size and shape through the main axis, from fully developed and similar to a small foliage leaf to triangular with a reduced apiculus, the blades Taste 1. C f veg hologi f Guadua i , G. chaparensis, G. tagoara, G. weberbaueri, and G. sarcocarpa CHARACTER incana chaparensis tagoara weberbaueri sarcocarpa Culm habit erect basally, erect basally, erect basally, erect basally, erect i arching apically arching apically ranching arching apically ` branchin and reaching and cn over trees over trees Culm size length (m) 10-15(-18) 18-25 10-15-20) 8-15(-20) 10-20(-30) diameter (cm) 7~9(-12) (3.5-)7-12 5-10 4—8(-12) -10 Internode size length (crn) 20-65 22-70 25-80 25-70 25—90 wall thickness (cm) 1-1.5 1-1.5 0.5-1 1-1.5 1 Culm leaf auricles absent absent absent absent absent/present Culm leaf inner ligule size imm 5-1 — — - abaxialindument ^ canescent canescent canescent canescent canescent Foliage leaf sheath indument subglabrous, with subglabrous with subglabrous with subglabrous glabrous a patch of hairs a patch of hairs a patch of hairs at the apex at the apex at the apex margin conspicuously conspicuously ciliate ciliate smooth and ciliate ciliate summit fimbriate glabrous fimbriate, rarely fimbriate oral setae or with auricles fimbriae Pseudopetiole length (mm) 2 — 4—10(-15) 5-7 (4-)6-10 abaxialindument ^ glabrous glabrous glabrous setose-pilose glabrous Foliage leaf blade, glabrous to glabrous glabrous pilose irregularly abaxial indument subglabrous subglabrous to glabrous Pseudospikelet 2-4 x 0.4-0.7 not seen 0.5-4 x 0.2-0.6 0.5-2.5 x : 2-8. > x (0.2-) size (cm) 0.2-0.7 — Lemma, abaxial subglabrous not seen usually glabrous pubescent n surface Palea sulcus re not seen oe pubescent pubescent pu width (mm) not seen -1.2 8- (1-)1.8-2.5 Fruit type n — not seen de caryopsis dry caryopsis fleshy caryopsis a Colombia olivia Brazil Peru Peru, Bolivia, distribution razil, Colombia Ecuador, Guyana, Surinam & Venezuela when present deciduous, the sheaths adaxially pubescent and bearing fimbriae at the summit, the overlap- ping margin conspicuously ciliate. Pseudospikelets 2-4 x 0.4-0.7 cm, linear-lanceolate, straight to slightly curved, green when young becoming stramineous, consisting of a subtending bract, a prophyll, 1-4(—5) gemmiparous bracts, 1-2 sterile lemmas, 5- 2 functional Get terminating in a rudimentary anthecium; prophylls 4 x 2 mm, with the 2 keels winged, t ] , abaxially BABE i in sericeous at the tip and strigillose along the keels, the enfoldi lal d keels, with the margins ciliolate, the wings 0.2-0. e mm da peca on both sides, conspicuously ciate on the margin, the cilia up to 0.5 mm long; gemmiparous bracts 1—4(—5), 5-9 x 3-6 mm, 7-11-nerved, 30 ovate to ovate-lanceolate, deciduous, adaxially shortly pubescent toward the upper third, otherwise glabrous and shiny, abaxially Py pubescent, with the midnerve EE Ge thg apezi mucronate, Se mucro 0.5-0.7 mm long, the ly ciliat te, except basally; sterile lemmas E d p 8x a Gm. 10- ies -nerved, ovate- med stramineous, ;, slightly lighter in color than the lemmas, mucronate, the mucro 0.5 mm long, abaxially glabrescent, adaxially pubescent on the upper 1⁄2, being densely pubescent toward the apex, the overlapping margin minutely ciliolate, the underlapping one smooth, enclosing a rudimentary palea or not; rachilla segments 3-4.5 mm long, shortly pubescent except at the most lower portion, with a rim of hairs at the uppermost portion, disarticulating below the attac hment of each lemma and falling attached to the floret. Fertile florets 5—7, 10—11 x 2-4 mm, with the winged palea keels exceeding the lemma margins, the lemma exceeding the palea apically only by the mucro; lemmas 9-11.5 x 6-8 mm, 16-18-nerved, ovate-lanceolate, green when young with purplish margins, becoming stramineous, abaxially glabrous, except with a patch of very short hairs dorsally, and a purplish-tinged to darker color line along the margins, adaxially pubescent on the upper half otherwise glabrous, the apex mucronate, the mucro 0.5-1 mm long, the margins smooth, papery; paleas 9-11 x 4-5 mm, stramineous, the apex acute, the sulcus 2 mm wide, 3—4-nerved, strigillose, covered by transparent, short, appressed hairs, the enfolding margins 2-3-nerved, glabrous, the margins smooth, the keels winged, the wings 1-1.2 mm wide, wider apically than basally, 2-nerved, purplish to stramineous, abaxially glabrous and shiny, adaxially strigillose along the sulcus and apically, otherwise glabrous and shiny, not prolonged at the apex, the margins ciliolate on the upper half, basally sparsely ciliolate. Lodicules 3, 2.5—5 x 0./-1.8 mm, 8—12-nerved basally, membranous, acute, the upper half pul and thinner, covered by hyaline prickles, basally glabrous, thickened, nerved and darker, tl ly ciliolate at the tip, the anterior pair slightly asymmetrical, the posterior one symmetrical, narrower than the anterior pair. Stamens 6, the anthers 4—5.2 x 0.1-0.5 mm, brown, basally sagittate, apically apiculate, the filament free, up to 2 mm long. Ovary (1.5323 x 0.4-0.5 mm, fusoid, the basal half receptacle-like, 0.7-1 x 0.5 mm, glabrous, brown, the upper half cone-shaped, 12 x 1 mm, densely antrorse-hispidulous, stramineous; style 2-2.5 mm long, densely antrorse-hispidulous, darker than the ovary, stigmas 3, plumose, ca. 5 mm long, Fruit 7-9 x 3 mm, an asymmetric fusoid caryopsis, with a persistent hispidulous style base at the apex, shiny and glabrescent immediately below the style, the basal two-thirds minutely strigillose, opaque, glaucous; embryo 1.5 x 1 mm, circular, lateral at the base of the caryopsis. Etymology.—Ihe specific epithet refers to the evidently canescent abaxial surfaces of the culm leaf sheath when young Distribution and habitat. —This species occurs in southern Colombia, in the departments of Caqueta and Putumayo, in the foothills of the eastern side of the Cordillera Oriental de los Andes, at elevations between 280—1200 m. It grows in the interior of very humid mountain forests, along creéks, but does not form dominant clumps and is generally uncommon. Common name.—Guadúa (with accent). Uses.—Farm fences and light rural infrastructure. Phenology and Fruiting.—The flowering behavior of bamboos is still unexplained and mysterious. The factors that switch a bamboo plant from a vegetative to a flowering state are not fully understood (Janzen 1976; Judziewicz et al. 1999). Guadua incana can flower continuously for at least 2-3 years and without dying. A flowering culm in this species is leafy and the production of fruit and seedlings is uncommon. As in G. weberbaueri, only a few culms from the whole clump flower at any given time. Despite this research, it is still impossible to establish its regular flowering cycle, but we know that it takes at least more than 14 years. The BEE of mature fruit in G. incana is scarce. Only one fruit was found in a functional i from tl iddle part of t] lospikelet. This floret falls along with the rachilla, lemma and } to the base as in other ree species, e.g., G. chacoensis (Rojas) Londoño & Peterson (Londoño & Peterson 1992). The dorsal surface of the fruit has a conspicuous linear hilum, purplish in color, extending down the | | Z I ta Al . Er | £ fe Lo E A 31 length of the fruit and the ventral surface has a prominent embryotegium. The embryo is 1/5-1/6 of the length of the whole fruit, and is located at the base of the ventral surface. The indument of the fruit is very peculiar, with the upper 1/9th e MESE the fruit is strigillose and covered with a whitish waxy coating. Pseudospikelet.—The p tisa phy for Guadua + Eremocaulon within the subtribe Guaduinae (Ruiz et al., in tess) and among estaria woody bamboos is also found only in the genera Elytrostachys McClure, Amta Soderstrom & Londoño, and Atractantha McClure, all of them in the sub- tribe Arthrostylidiinae. The term is used to describe synflorescences of woody bamboos that rebranch to produce successive orders of spikelets (McClure 1966; Judziewicz et al. 1999; Young & Judd 1992, Bamboo Phylogeny Working Group 2005). According to Londoño and Clark (2002b) the following structures are all homologous at some level: subtending bracts, gemmiparous bracts, glumes, and both sterile and fertile lemmas. However, this is not the case with the prophyll, which is homologous to the palea (Stapleton 1997; Bamboo Phylogeny Working Group 2005). In G. incana, the most proximal bract to the prophyll gives the impression of being the prophyll itself, because the midnerve is similarly keeled and conspicuously asym- metrical, the apex bifurcate and shortly mucronate, and always fertile, enclosing a bud (Fig. 1: HD The prophyll when present is conspicuously ciliate but frequently the hairs rub off in situ. Affinities —Guadua incana is most similar to G. weberbaueri Pilger and G. sarcocarpa Londoño & Peter- son. The three species share the following characters: a) triangular culm leaves with the margins slightly discontinuous at the junction with the blade; b) culm leaf blades 1/4 to 1/5 as long as the sheaths, adaxially pubescent and with a strong mucro at the apex; c) adaxial surfaces of the lemmas See on their upper 1] 1 "mi OH "pm E ingin halves; d) paleas with pubescent ; e) conspicuous winged | an acute apex; f) lodicules strigilllose in the upper third; and g) ovaries and styles antrorse-hispidulous. However, G. incana differs from G. weberbaueri and G. sarcocarpa in having (1) blade and sheath of the culm leaves conspicuously distinguished by color and pubescence (vs. inconspicuously distinguished by color and pubescence); (2) subglabrous adaxial surfaces of the leaf blades (vs. entirely glabrous); and (3) strigillose sulcus of the palea (vs. pubescent) (Table 1). Guadua incana differs from G. sarcocarpa in its (a) culms erect at base, arching apically (vs. erect at base, then branching above and reaching over trees for support); (b) white-greenish and hairy culm leaves when young, (c) conspicuously ciliate overlapping margin of the foliage leaf sheaths, (d) shorter and narrower pseudospikelets, (e) the winged palea keels marginally ciliolate, and (f) dry caryopsis fruit. It differs from G. weberbaueri in its (a) longer pseudospikelets, (b) smooth and papery margins of the lemma, (c) abaxially BEE put basally SEHEN, ciliolate Map ps iet Ge x heath margins, (b) similar pseudospikelet size, (c) gemmiparous bracts with the miden strongly ed (d) the lemma margins exceeded by the winged-keels of the palea, (e) a palea with two conspicuous winged-keels and a pubescent sulcus between nerves; and (f) a fusoid antrorse-hispidulous ovary (Table 1). According to the molecular analysis done by Marulanda et al. (2002), G. incana shows a wide genetic distance in relation to G. angustifolia and greater genetic similarity with G. amplexifolia which shares a canes- cent indument on the abaxial surfaces of the culm leaf sheaths, the inner ligule extending from margin to margin and the strong mucro at the apex of the culm leaves; DNA isolation for G. weberbaueri, G. sarcocarpa and G. tagora, the three closely allied species, was not successful in the above mentioned research. Amana EE E Geen E Km 26.7 vía EM AE re La Pune Di vertiente E , 750 m, 10 Feb 19 itero ied, No. XL114, 2 Jun 2004 (£D, Londoño 982 GE TULV, uS Km 22 vía Florencia- E 6701 m, 9 Feb 1987, EE €» Quintero 122 (TULV). Putumayo: Mpio. Mocoa, Vereda Suiza, carretera Mocoa-Pasto, después de La Tebaida, a orilla de Q. Suiza, 1050 m, 6 Mar 1987, Londoño & A 214 (COL, TULV, US); cultivated Jardín Botánico Juan Maria Céspedes, Plot £3 collection No. XL214, 2 Jun 2001 (fl), Lond 73 EENS US). Guadua chaparensis Londoño & Zurita, Sp. nov. (Fig. 2). Tyre: BOLIVIA. Cocuasamba: Prov. Carrasco, Distrito Chapare, localidad Israel, orilla del Rio Sajta, 270 m, 17° 12' 33" S, 64º 49' 47" W 8 Aug 2004, E. Zurita & J. Huaranca EZ 302 (HOLOTYPE: BOLV; isotypes: BOL, COL, TULY, US). [| [| E s n.a H Im LI PE ET “ia 32 ICAOA £11] Ic 3 e J L H A ein ` E J Sne R AMA 1 1 Sal, L L p PN | H [| L L [| Jal £ a Li E] P Thornv branch. D. Foli leaf liaul abaxial view. E. Culm seament. with tl t tl les (Zurita 302. BOLV) d bi kl Li ki 7 V y, J Bambusa lignosa, spinosa. Rhi ta pachy pha. Culmi 18-25 m longi (3. nd cm in d internodia a cm EE Folia culmorum 29-57 cm SE 35-54 lata ad basim, coriacea, decidua, triangularia, (vagina albido-bru ersus lamina ); vagina 23-50 cm longa; isa (44-)9-15 cm loni a, 14—17 fo) | 6B cm lata, triangularis, erecta, viridis ES i-rubescentem, inflata in j Folia ramorum Use ibd od 5-8(-10); . 1 EM 1.1 “4 Mas 1 1 : e 3 fimbri llae; 1 lopetiolus 2-4 longus; lamina (5-}15-23(-25)c imos (Q. 621. 3-2 Gina: bale glabra de Papos: et tessellata, api inato. Synfl ti visae. Caryopsis non visa. Woody, thorny bamboo. Rhizomes pachymorph. Culms (15-)18-25 m tall, (3.5-)7-12 cm in diam ., erect at base, arching from the middle to the apical portion, whitish-green when young, dark green at maturity; internodes 22—50(-70) cm long, cylindrical, hollow, some filled with water, the wall 1-1.5 cm thick, densely pubescent, with white hairs when young, glabrous and smooth when mature; nodes solitary, the nodal line horizontal, with a pubescent supranodal band 1.45-2.2 cm inn of dense, Bde soft, appressed short hairs, and an infranodal band 1-2 cm wide of white, appressed hairs, t I 1; bud single, triangular. Culm leaves 29-57 cm long, 35-54 cm wide at the base, coriaceous, Ea triangular, I Jaú E A AI H £r J £ ral L: pa se 33 blade and sheath conspicuously distinguished by color and pubescence when young; sheaths 23-50 cm long, whitish-green and maculate with red when young then becoming cream-brownish to stramineous, abaxially sericeous becoming glabrous, covered when young by a single type of hairs, appressed, transpar- ent or silky hairs up to 1 mm long, looking like wool, easily detached from the surfaces, adaxially glabrous and shiny, the margins smooth and papery; inner ligules 0.5-1 mm long, straight to slightly curved at the middle, extending completely from margin to margin, adaxially glabrous and shiny, abaxially canescent, the margin densely ciliate, the cilia up to 0.6 mm long, whitish or grey in color; blades (4-)9-15 cm long, 14—17 cm wide, 1/5 as long as the sheath, broadly EH erect, pee inflated at the central part, green to reddish-green, abaxially glabrescent, less 1 ifferent in color, adaxially densely hispid between the nerves, covered by transparent, non-appressed, stiff hairs, up to 2.5 mm long, stramineous or brownish when young becoming gray, glabrescent toward the margins, the margins papery, fimbriae to smooth, the apex mucronate, the mucro 1.2-1.5 mm long. Branching intravaginal, consisting of one main branch and 1 to 6 secondary branches, the branches manifestly armed, thorns (123-4 per node, curved or recurved. Foliage leaves 5—8(-10) per complement; sheaths subglabrous, with a patch of hyaline, villous hairs along the midnerve near the apex, the summit glabrous, the overlapping margin conspicuously ciliate, the underlapping one ciliolate to smooth and papery; inner ligules 0.3-0.5 mm long, puberulous, the margin minutely ciliolate to smooth; outer ligules 0.2-0.3 mm long, glabrous and shiny, stramineous, the margin smooth; pseudopetioles 2-4 mm long, adaxially scabridulous, abaxially glabrous and pulvinate, the pulvinus yellowish, shiny; blades (5-)15-23(-25) cm long, (0.6-)1.3-2 cm wide, LW = 8-13:1, 11-14-nerved, linear-lanceolate, adaxially usually glabrous, rarely bearing sparse, transparent, strigose hairs up to 1 mm long, with 3-5 raised scabrid submarginal nerves on one side, abaxially glabrous, densely papillose and tessellate, the midrib and primary nerves prominent and yellow at the middle and lower part, one margin scabrous, the other scabridulous, the apex acuminate with a mucro 1.5-2.5 mm long. Synflorescences not seen. Feann not seen. Etymology.—Th itl to the Amazon biogeographical district of the Chapare River, Cochabamba, Bolivia where this species is found. Distribution and habitat. —Known only from the department of Cochabamba, Carrasco, Chapare, below 900 m of elevation, with a total annual precipitation estimated to be 6000 mm and a median temperature of 25°C. It occurs along rivers and creeks, such as the Sajta and Ivirza, and is associated with Hura crepitans L., Guadua sarcocarpa Londoño & Peterson, Mendoncia aspera Nees, Cecropia sp., and Costus sp. Common name.—Tacuara hembra or tacuara. Uses.—Guadua chaparensis culms are used by local communities to make fences, walls, water contain- ers, and for conducting water. Affinities —Guadua ch j t closely resembles G. tagoara (Nees) Kunth and G. incana. They share a culm that is whitish-green ¡nen young to dark green at maturity; er and SE nua 20 to 80 cm long with walls less than 15 mm thick; coriaceous culm leaves wit ight to slightly curved at the middle, abaxially with a canescent indument, looking like wool and easily dead ed with the margins densely ciliate; a eae ee page as leaves in complements of 5 to 8(-10); foliage leat sheaths bearing a patch of I the apex; pulvinate pseudopetioles that are abaxially glabrous; and a prominent and vello midnerve and primary nerves on the abaxial surfaces of the foliage leaf blades (Table 1). Although they share several characters, Guadua chaparensis has a combination of vegetative features that distinguishes it from G. incana and G. tagoara even in the absence of inflorescences. The most evident ones are (a) the sericeous abaxial surface of the culm leaf sheaths, covered by one type of hair (vs. pubes- cent, covered by two or three different types of hair and forming a hispid surface); and (b) the absence of fimbriae at the summit of the foliage leaf sheath (vs. present) (Table 1). With respect to G. weberbaueri and G. sarcocarpa, G. chaparensis also shares several vegetative characters listed in Table 1. Additional specimens examined. BOLIVIA. Cochabamba: Prov. Carrasco, localidad Israel, 270 m, 17? 14' 56" S, 64? 51' 84" W, 270 m, 5 May 2004, Zurita & Baldelomar EZ176 (BOL, CUVC); al borde del camino, 270 m, 17? 12' 20" S, 64° 49' 36" W, 270 m, 4 May 2005, 34 t tani itute of Texas 2( Zurita & Soto EZ320 (BOL, COL); Israel, 270 m, 22 Nov 2005, Zurita & Soto EZ324 (BOL, TULV); Proyecto Valle del Sacta, km 240 en la carretera Santa Cruz-Villa Tunari, 64° 46' W, 17? 0' S, 290 m, 12-14 Jul 1989 (2), D. N. Smith, V. Garcia, M. Buddensiek, J. Leon & C. Negrete 13720 (MO). Guadua weberbaueri Pilger, Feddes. Repert. Spec. Nov. Regni Veg. 1:152. 1905; Bambusa weberbaueri (Pilger) Mc- Clure, Smithsonian Contr. Bot. 9:68. 1973. Type: PERU. San Martin: MODE emia ‘marona,” Weberbauer 4562 (HOLOTYPE: H 1 ] LADO 3 , here designated: MO g Etymology.—The "m epithet refers to the German botanist and expedition leader August Weberbauer (1871-1948), who devoted his life to studying the flora of Peru. ACKNOWLEDGMENTS The first author thanks Colciencias and INCIVA for supporting the fieldwork (Grant No. 2108-07-009-85). Special thanks are given to Daniel Debouck and Alba Marina Torres for their support and to herbarium facilities in the International Center of Tropical Agriculture (CIAT) in Palmira, Colombia, and to the curators and staff of the US herbarium at the Smithsonian Institution, Washington, DC. Special thanks go to Lynn G. Clark for imp d ipt with her comments and to Tarciso Filgueiras for the Latin diagnoses. The illustrations were FN by Jesús Salcedo and the final art (scale and letters) by Alice R. Tangerini. The second author thanks ASDI-SAREC and the Institute for Research in Architecture-IIA, project Bolbambu. Special thanks to Jose Luis Reque, and to the herbarium BOLV — Biodiversity and Genetic Center. Final preparation of the manuscript was supported by National Science Foundation grant DEB-0515712 to L.G. Clark. REFERENCES Bamsoo PHYLOGENY WORKING GROUP. 2005. www.eeob.iastate.edu/research/bamboo/index.html CLARK, L.G. 1992. Chusquea sect. Swallenochloa (Poaceae: Bambusoidea) and allies in Brazil. Brittonia 44:387—422. JANSEN, D.H. 1976. Why bamboos wait so long to flower? Ann Rev UEM & t 7 ÓN SC Jubziewcz, E L.G. CLARK, X. Lonpoño, and MJ, STERN. 1999. A Press, Washington 8 London. LonboKo, X. and L.G. CLARK. 2002a. A revision of the Brazilian bamboo genus É lon (Poaceae: Bambuseae: Guaduinae). Syst. Bot. 27:703-721. Lonpoño, X. and L.G. Clark. 2002b. Three new taxa of Guadua (Poaceae: Bambusoideae) from South America. Novon 12:64-76. LONDONO, X. and P. Pererson. 1992. Guad is (P Bam ideae), its taxonomic identity, morphol- ogy and relationships. Novon 2: i -47. Lonboño, X. and P. Peterson. 1991. G pa (P Baml ), a new Amazonian bamboo with fleshy fruits. Syst. Bot. 16:630—638. Louton,J., J. GELHAUS, and R. BoucHARo. 1996. The aquatic macrofauna of water-filled bamboo (P Bambusoid Guadua) internodes in a Peruvian lowland tropical forest. Biotropica 28:228- did MARULANDA, M.L. P. MÁRQUEZ, and X. LoNpoRo. 2002. AFLP's analysis of Guad lia (P Bambusoideae) in Colombia with emphasis in the Coffee Region. Bamboo Sci. Culture 16(1):32-42. McClure, EA. 1966. The bamboos — A fresh perspective. Cambridge. Harvard Univ. Press. Ruiz-SANCHEZ, E, V. Sosa, and MT. Mena. In press. Phylogenetic position of the American bamboo Otatea (Poaceae: Bambusoideae: Bambuseae) based on coDNA and morphological data. Syst. Bot. Soperstrom, T.R. and S.M. YOUNG. 1983. A guide to collecting bamboos. Ann. Missouri Bot. Gard. 70:128-136. STAPLETON, C MA 1997. The morphology of woody bamboos. In: GP Chapman, ed. The bamboos. Academic Press, London. Pp. 251-267. Young, S.M. and WS. Jupo. 1992. Systematics of the Guadua angustifolia complex (Poaceae: Bambusoideae). Ann. Missouri Bot. Gard. 79:737—769. TETRAZYGIA PARALONGICOLLIS (MICONIEAE: MELASTOMATACEAB), A NEW SPECIES FROM THE SIERRA DE BAORUCO AND SIERRA MARTIN GARCIA, DOMINICAN REPUBLIC Walter S. Judd & Gretchen M. lonta E od Clase Department of Botany lardín Bot ional Dr. Rafael Ma. Moscoso 220 Bartram Hall . Box 118526 Santo Domingo, REPÚBLICA eli University of Florida ted claseghotmail.co NA Horida 32611, U.S.A. wjudd@botany.ufl.edu J, Dan Skean, Jr. ir ent of Biology A College Albion, See 49224, USA. dskean@albion.edu ABSTRACT l e TT yc 16; Maria Circa P alus DAR Republic, is described TL: A o E r " 1 : r r 720-860 m, EP p 1 Los I 1 1 lia: T. longicollis and T. elaeagnoides. RESUMEN Se describe En poa una nueva especie de la adds E Baoruco y de la Sierra Martin Garcia de la Repüblica Do- los 720—860 m de altitud, y se compara con sus minican presuntos parientes más cercanos: T. EH y E ehemals Key Woros: Hispaniola, Dominican Republic, Melastomataceae, Miconieae, Tetrazygia During the course fheldworl lucted i ti ith the study of the systematics of Miconieae (Melas- + ua herbarium material of a distinctive species was collected in the Sierra Martin Garcia (by W.S. Judd, in 1992 and 2006) and Sierra de Baoruco (by T. Clase, in 2001). It is now evident that these collections represent an undescribed species of Tetrazygia (Miconieae, Melastomataceae), which appears to be closely related to T. longicollis Urb. & Cogn. and T. elaeagnoides (Sw.) DC. These plants are described here, provided with the name T. paralongicollis, and compared morphologically with the above mentioned species. Recent molecular-based phylogenetic analyses (F. Michelangeli, R. Goldenberg, W. Judd, and others; unpublished data) indicate that Tetrazygia, a genus of ca. 20 species (Judd & Skean 1991; Liogier 2000), is not monophyletic as traditionally circumscribed. The species of Tetrazygia Rich. are intermixed in prelimi- nary cladograms with members of SE other pe of Miconieae, especially Pachyanthus A. Rich. and Calycogonium DC., two genera that y Antillean in distribution, asis Tetrazygia. Clarification of generic limits within this sidade of the EEN i.e., an Antillean complex (see Michelangeli et al. 2004, 2008) is beyond the scope of this paper, but we note fat T. elaeagnoides, which has 4-merous flowers (as does T. longicollis and the species described herein), in preliminary cladistic analyses does not appear to be closely related to the species with 5- or 6-merous flowers, such as T. bicolor (Mill.) Cogn., T. coriacea Urb., and T. lanceolata Urb. Tetrazygia paralongicollis Judd, lonta, Clase & Skean, sp. nov. (Fig. 1). Tree: DOMINICAN REPUBLIC. PEDERNALES n cs da Ral 1601 Cees Ta El DAS 1 e 1 oA mare y | jc A Eus Ix E lentalis. Lat 71° 37' 63" W, Long. 18? 6' 7" N, 720 m, 19 Jul 2001 (fl, fr), T. Clase & P Delprete 3035 (HoLoTYrE: JBSD; isotypes: FLAS, NY, S). J. Bot. Res. Inst. Texas 2(1): 35 — 40. 2008 36 Jo tani itute of Texas 2(1) 1 1 ae 1 : : “rr - 4 1 14] : : trir] d d 31 471.1 &r Coon J etellatic (sre PRIMA veins a ES E : : D f. . D 0 Species haec ab Tet WIE I ULL LIC. Del L L i" D D ca. 1.2 mm longis (vs. 1-5 mm longis). Shrub or small tree to 6 m tall, with gray, horizontally furrowed bark; lateral | hes arising from the main trunk « horizontally, then arching upward. Indumentum of See pele Aerrugineous, nearly sessile to long-stalked, globular-stellate hairs, and often also simple, el 0.6 to 1.4 mm long, these sometimes with minute antrorse bristles. Young twigs 2— 5 mm wide, slightly quadrangular, with the interpetiolar face (in relation to the distally adjacent node) slightly concave, the adjacent sides slightly convex, becoming terete with age, with moderate to dense globular-stellate hairs and often sparse simple, elongate multiseriate hairs, many breaking off with age; internodes 1.1-6.1 cm long (to 8.5 cm in vigor- ously growing vegetative shoots). Leaves with petiole 6-9.4 mm long, (but to 28 mm in leaves of vigorously growing vegetative shoots), with dense globular-stellate hairs; blade 2.5—6.8(-20.4) cm long, 0.6-2.2(-7.7) cm wide, 3.1—4.2 (2-3.5 in leaves of vigorously growing sucker-shoots) times longer than wide, ovate to el- liptic, flat, coriaceous, the apex acute to acuminate, with the tip of the leaf with upturned margins, forming a mucro 0.5—1.1(-6) mm long with adaxial pocket, the base narrowly acute to cuneate or slightly cordate, the margin plane to slightly revolute, nearly entire, i.e., with only a very few teeth along distal portion of blade, and these at most 0.1 mm long, to + clearly serrulate or ciliate-serrulate, with teeth 0.1-0.3 mm long, sometimes associated with a short multiseriate hair, + evenly spaced along margin; venation acrodromous, basal to suprabasal, with prominent midvein and 4 secondary veins (6 in leaves of rapidly growing vegeta- tive shoots), 2 conspicuous secondary veins positioned 1.7—4(-16 in sucker-shoot leaves) mm from margin, and 2 inconspicuous secondary veins closer to margin (but sucker-shoot leaves with an additional pair of inconspicuous, + intramarginal, secondary veins), numerous percurrent tertiary veins oriented subper- pendicular to midvein, the tertiary veins occasionally separated by composite-intertertiary veins (but such veins more common on leaves of sucker-shoots), the higher-order veins reticulate; adaxial surface green, initially with numerous globular-stellate hairs, but quickly glabrescent, the midvein and major secondary veins moderately impressed, minor secondary and tertiary veins slightly impressed to flat, and higher order veins flat, the surface appearing minutely papillose after drying ES to ene of numerous subglobose subcuticular druse crystals; abaxial surface light E with d , globular-stellate hairs obscuring the epidermal surface, and sometimes elongate —€—— Gerten some stellate hairs on the major veins dans ferrugineous, the midvein — raised, j ] lerately raised, minor secondary veins, tertiary veins, and some composite-intertertiary veins slightly raised, and higher- order veins flat (or very slightly raised in some leaves of sucker-shoots). Inflorescences 3- to 12-flowered pyramidal cymes of 1 to 2 branch-pairs, 4—7.5 cm long, 1.7—5 cm in diameter; proximal segment of lowermost inflorescence branches 1.4-2.5 cm long, distal internodes of inflorescence branches increasingly shorter, ultimate branches 1.1-1.9 cm long, and flowers appearing in 1- to 3-flowered dichasia, with dense globular- stellate hairs and sparse elongate multiseriate hairs; peduncle 0.5-2.1 cm long, with similar indumentum; proximal inflorescence branch associated with pair of persistent leaflike bracts, 2.5-4.3 cm long, 0.7-1.2 cm wide, similar in form to vegetative leaves, other inflorescence branches ted with pair of caducous bracts (and not present on specimens examined). Flowers perfect, zygomorphic (due to androecium form), with pedicel 1.4-1.5 mm long, the indumentum similar to that of inflorescence branches. Free portion of hypanthium slightly constricted above ovary, flaring and funnelform distal to the constriction, 2.9-3.8 mm long, outer surface with dense to moderate globular-stellate hairs and sparse elongate multiseriate hairs, the inner surface slightly 16-ridged, glabrous except for occasional minute-globular hairs on the ridges. External calyx lobes 4, ca. 1.2 mm long, ca. 0.8 mm wide, narrowly triangular, with acute apex, and terete in cross section, with dense globular-stellate hairs; internal calyx lobes 4, 0.6-0.8 mm long, 2.9-3.4 mm wide, broadly triangular, apex rounded, green to red-tinged, with dense globular-stellate hairs abaxially, glabrous adaxially, the margin membranaceous, minutely erose; calyx tube ca. 1.5 mm long. Petals 4, im- bricate in bud, + asymmetrical, ovate-obovate, 5.2-6.1 mm long, 4.3-4.9 mm wide, glabrous, white; apex rounded; margin entire. Stamens 8, anther elongate-ovate, 4.6-5.2 mm long, glabrous, pale yellow, very Judd et al., A ies of Tet iaf the Domini R | bli 37 AA A SS NN. AA ON Ay oH ES S PF Ny, ne REY ez D FH delo a lo E um 2 a d et Sg oH " E E 3 mm SE É a — Su Ne Y Ke pa A q* Fic. 1. A-I. Tet ygi | / gi llis. A. Dichasium, witl fl | B. Stamen C 0 * i f haxial leaf surf E Ovarv. in cross-section. E Petal. G. Berrv. H Habit |. Leaf. abaxial surface d d F except for “I” which is from Judd 8148 (FLAS)). : D. Stellate I I Eat Dos H In LI eh nd fT fai 1*7 38 slightly sagittate at base, fertile the entire length, and opening by a single dorso-apical pore, the filament terete, 5.8-6 mm long, glabrous, white. Ovary 4-loculate, ca. 4/5 inferior, ellipsoidal and slightly 4-lobed, 3.5-3.9 mm long, 2.1-2.5 mm wide, with cylindrical apical crown ca. 0.7 mm long encircling base of style, the crown slightly 8-ridged, with sparse stellate hairs and elongate-uniseriate hairs at apex; style ca. 15 mm long, terete, glabrous. Placentation axile with placentas inserted into locules with narrowed longitudinal placental stalk with flattened ellipsoid distal portion, T-shaped in cross-section; ovules numerous. Nearly mature berries 9-12 mm in diameter, globose-ellipsoid, with strongly constricted persistent hypanthium, green when immature, but probably turning purple-black at maturity, with sparse globular-stellate hairs. Seeds angular-obovoid, 0.7-0.9 mm long; testa + smooth. Distribution and habitat —Tetrazygia paralongicollis is restricted to the southern part of the Dominican Republic where it has been collected only in the Sierra de Baoruco and Sierra Martin Garcia (Fig. 2), from 720—860 m. In the Sierra de Baoruco it occurs in Pinus occidentalis Sw. forests, while in the Sierra Martin Garcia it grows in moist montane forest (on limestone). In the Sierra Martin Garcia, associated melastomes include Calycogonium hispidulum Cogn., Miconia laevigata (L.) DC., Sagraea fuertesii (Cogn.) Alain, Tetrazygia elaeagnoides (Sw.) DC., and T. longicollis Urb. & Cogn. Etymology.—Ihe specific epithet highlights the purported close relationship of this species with Tetra- zygia longicollis, especially as evidenced in the characteristics of its flowers and fruits. Vegetative anatomy.—Stem, leaf, and petiole anatomy were assessed in material of Tetrazygia paral- ongicollis, i.e., Judd 8148 (FLAS) and Clase & Delprete 3035 (FLAS), and leaf and petiole anatomy were as- sessed in material of T. longicollis, i.e., Judd 6656 (FLAS) and T. elaeagnoides, i.e., Judd 6553 (FLAS) using the phloroglucinol-hydrochloric acid technique outlined by Howard (1974). As is typical of Melastomataceae, me stems =o eS "eue a ring of xylem with phloem das both externally and internally of Miconieae, medullar y vascular bundles are presen t. The pith i 15 ligni- fed, and petivásculi fibers are lacking, but there are scattered lignified idioblasts in the inner portion of the cortex, which may have a protective function (because they surround the stele). Druses also are present in the cortex. The nodes are unilacunar/unifascicular, but the vascular bundle divides within the petiole base, forming several bundles in the petiole. The petioles (sectioned at their midpoint) of all three species exhibit a U-shaped pattern composed of 7 to 11 vascular bundles, with the individual bundles composed of xylem surrounded by phloem. The ground parenchyma is unlignified, but in T. paralongicollis there are a i Ge lignified idioblasts, and in T. viii Pad are numerous lignified idioblasts. The petioles of : ] lignified idioblasts, but | ant EC tal ee parena 15 wind packed with PR In contrast, the p hy of T. Petras and T. 1 rystals The leaves of all three species have dorsoventral blades with an epidermis, palisade and spongy —— the midvein has several vascular bundles, which are more or less arranged in a ring. Lignified idioblasts are present in the parenchyma associated with the midvein vascular bundles of T. paralongicollis and T. longicollis, while these cells are lacking in T. elaeagnoides. The abaxial leaf surface of all three species is covered with a thick layer of stellate hairs, which are variably lignified. As with the petiole, the parenchyma of the midvein of T. elaeagnoides contains very abundant druses. Druses are present in the lamina of all three species, with these iun placed below the epidermis in either the palisade or spongy mesophyll. Addit 1 collecti DOMINICAN REPUBLIC. Azua Prov.: Sierra Martin Garcia, Loma del Aguacate, hill d rid f ntain west of Barreras, ca. 850-860 m, See collecteg along trail from Barreras to El Copey, 17 May 1992 (sterile), Judd 6552 (FLAS, JBSD); Sierra Martin Garcia, hill SW of Barreras, along trail from Barrera toward El Copey (from N side of town, trail starts to NNW, cuts W, then SW, hens: then more or less W, with variance); near Barahona, 810-820 m. Lat. 18° 19' 4.4" N, Long. 70° 56! 47.2" W, Datum: WGS84, 6 Jun 2006 (sterile), Judd 8147 (FLAS, JBSD, MICH, MO, MSC, NY, S, US) DISCUSSION Tetrazygia paralongicollis is most similar to T. longicollis (incl. T. brevicollis Leonard). This is especially seen in the form of its flowers and fruits, i.e., 4-merous, with conical-terete external calyx lobes, broadly trian- gular internal calyx lobes, ovate-obovate, white petals, stamens with ovate-elongate anthers, an ellipsoidal Judd et al., A I i f Tet iaf the D 20°N SS 18" | Hispaniola /4"W 72° 70° Fic. 2. Distributi f Tet ovary (only slightly 4-lobed) and with the placentas inserted into locules, T-shaped in cross section, and globose-ellipsoid berries with strongly constricted hypanthium. However, it is consistently differentiated from T. longicollis by its globose-stellate hairs, i.e., with branches radiating in all directions from a globose central region (vs. flattened-stellate hairs, i.e., with branches more or less radiating outward from a central region), the tertiary veins usually connected by reticulate quat y veins, i.e., only occasionally separated by composite-intertertiary veins (vs. tertiary veins usually separated by composite-intertertiary veins, thus making the leaf “tertiary” veins appear to be closer together than they actually are), and external calyx lobes that are ca. 1.2 mm long (vs. variable in length, ca. 1-5 mm long). Tetrazygia paralongicollis is also quite similar to T. elaeagnoides, especially in the morphology of its stellate hairs and in the pattern of venation of its leaves. Tetrazygia paralongicollis is easily distinguished from this species by its hypanthia (and fruits), which are globose-ellipsoid and only very slightly 4-lobed when in flower, becoming unlobed in fruit (vs. subglobose and strongly 4-angled), placental form, i.e., the placenta extended into the locule and T-shaped in cross-section (vs. elliptic in cross-section), and the presence of lignified idioblasts in the parenchyma of the petioles (vs. lignified idioblasts lacking). In addition, many plants of T. elaeagnoides have shorter exter- nal calyx lobes, e.g., ca. 0.2-0.4 mm long, but in some plants Ge B BE as long as 1.6 mm. Plants of T. le, elon gate, multiseriate paralongicollis from the type locality differ from both of tl hairs intermixed with the globular-stellate hairs on their stems, abaxial leaf Seet kal margin (in associa- tion with the teeth), inflorescence axes, and hypanthium. The plants of the Sierra Martin Garcia, however, lack these hairs, but these plants (unfortunately, only collected in sterile condition) are considered within T. paralongicollis because their leaf shape, venation, and stellate-hair morphology closely match that of the type specimens. It is noteworthy that the leaves of the Sierra Martin Garcia population are frequently larger than those of either T. longicollis or T. elaeagnoides. This new species occurs with both T. longicollis and T. elaeagnoides in the Sierra Martin Garcia, but no intermediate plants were seen in that region. Only T. longi- collis, however, is known from the Sierra de Baoruco, in the vicinity of the type EES Tetrazygia paralongicollis, exhibiting a distinctive combination of | characters, satisfies the expectations of the morphological-phenetic (Judd 2007) and diagnostic (Wheeler & Platnick 2000) species concepts. The description of Tetrazygia paralongicollis brings the number of species of Tetrazygia known from 40 | PR | A a | Li sta fTexas 2( Hispaniola to eight (Liogier 2000), given that we consider T. urbaniana (Cogn.) Croizat ex Moscoso to be a synonym of the morphologically variable T. tuerckheimii (Cogn.) Ekman & Urb. All the other species of Tetrazygia show fairly broad geographical distributions on the island compared to T. paralongicollis. ACKNOWLEDGMENTS We thank Norris Williams, Keeper, and Kent D. Perkins, Collections Manager, of the University of Florida Herbarium (FLAS) in the Florida Museum of Natural History, for assistance in processing specimen loans. We are indebted to the director of JBSD, Daisy Castillo, for the loan of herbarium material. Special thanks are given to Milcíades Mejía, Francisco Jiménez, Brígido Peguero, and other herbarium personnel at the Jardín Botánico Nacional, Santo Domingo (JBSD) for their support of fieldwork in the Dominican Republic. We thank Frances Combs for her help with the illustration of this new species. We also thank J. Richard Abbott, Reuben E. Judd, and P. Delprete for their assistance in the field. This research was supported, in part, by NSF Grant DEB-0515636. Two anonymous reviewers provided helpful suggestions for improving the manuscript. REFERENCES Howanb, R.A. 1974. The stem-node-leaf continuum of the Dicotyledoneae. J. Arnold Arbor. 55:125-181. Jup, WS. 2007. Revision of Miconia sect. Chaenopleura (Miconieae, Melastomataceae) in the Greater Antilles. Syst. Bot. Monogr. 81:1-235. Jupp, WS. and J.D. Skean, JR. 1991. Taxonomic studies in the Miconieae (Melastomataceae). IV. Generic realign- ments among terminal-flowered taxa. Bull. Florida Mus. Nat. Hist, Biol, Sci. 36:25-84 Liocier, A.H. 2000. La flora de la Hispañola, Vol. 9. Melastomataceae. Jardín Botánico Nacional "Dr. Rafael Ma. Moscoso" & Instituto Tecnológico de Santo Domingo (INTEC). Santo Domingo, Dominican Republic. MICHELANGELI, FA., D.S. Penneys, J. Giza, D. Souris, M.H. Hiis, and J.D. Skean, Jr. 2004. A preliminary phylogeny of the tribe Miconieae (Melastomataceae) based on nrlTS sequence data and its implications on inflorescence position. Taxon 53:2/9—290. MICHELANGEL!, EA. W.S. Juop, D.S. Penneys, J.D. SKEAN, JR, E.R. Becquer, R. GOLDENBERG, and C.V. Martin. 2008. M Itiple e of dispersal and radiation of the tribe Miconieae (Melastomataceae) in the Caribbean. Bot. Rev. 74:53-77. WHEELER, Q.D. and N.I. PLatnick. 2000. The phylogenetic species concept (sensu Wheeler and Platnick). In: Q.D. Wheeler and R. Meier, eds. Species concepts and phylogenetic theory: a debate. Columbia University Press, New York. Pp. 55-69. + UNA NUEVA ESPECIE DE CUATRESIA (SOLANACEAE) DE COSTA RICA Y PANAMÁ D. Armando Soto A.K. Monro Instituto Nacional de Biodiversidad (INBio) The Natural History Museum (BM) Apdo. 22-3100, Santo Domingo de Heredia Cromwell Road London, SVV7 5B COSTA RICA UNITED KINGDOM asotoainbio.ac.cr a.monrognhm.ac.u ABSTRACT Cuatresia amistadensis, a new species from Costa Rica and Panamá is described and illustrated, and its taxonomic relationships are discussed. Key Wonps: Solanaceae, Cuatresia, Talamanca, Parque Internacional La Amistad, Costa Rica, Panamá. RESUMEN Cuatresi istadensi para Costa Rica y Panamá, es descrita e ilustrada, y se discuten sus relaciones taxonómicas. PALABRAS CLAVE: Solanaceae, Cuatresia, Talamanca, Parque Internacional La Amistad, Costa Rica, Panamá. La familia Solanaceae se distribuye en áreas templadas, tropicales y subtropicales, pero su mayor represen- tación se encuentra en el trópico, con ca. 95 géneros y 2200 especies (Nee 2004). El género Cuatresia Hunz. (Hunziker 1977) de origen Neotropical, posee 12 especies publicadas y 6 para Costa Rica (A.Soto en prep.) se distingue del género Witheringia L'Hér. por sus anteras ee y pe no ados (Hunziker 1987); un carácter importante de notar es que el género Cuatresia se distingue de otros g relacionados por su patrón de estivación induplicada (N. Sawyer, com. pers.). En el trabajo de campo para el proyecto 'Baseline tools for management in PN La Amistad (Costa Rica/Panama), realizado en el Parque Internacional La Amistad entre Costa Rica y Panamá, se recolectó la siguiente especie de Cuatresia, la cual es descrita a continuación. Cuatresia amistadensis D.A. Soto & A.K. Monro, sp. nov. (Fig. 1). Tro. COSTA RICA. Liston: Cantón de Talamanca, Parque Internacional La Amistad, camp 1, Río Lori, secondary forest adjacent to camp, 17 Feb 2007, A.K. Monro & D. Santamaría 5425 (moLoTIPO: INB, isoriros: BM, MO, PMA) Planta glabra foliis oblongo-ellipticis vel ellipticis, 12.5-21 cm longis et 5-10.5 cm latis, dimidiatis. Inflorescentia cymosa axillaris, pedunculis 3,8-12 cm. Corolla pentamera, campanulata, 7-20 mm longa, lobis oblongo-obtusis, 3-5 mm longis, filamentis 1 mm longis, et tubo corollae ad medium affixo. Gynoecium 6-10 mm. Fructus globosus, 8-12 mm longus. Arbustos t ionalmente epífitos, hasta 4 m de altura. Ramitas glabras, rojo palido—marrón, tallos principales con a engrosados, tallos con hojas con una constricción arriba del nudo, internudos 2,8—8 cm de largo, 2,5-6 cm de ancho. Hojas oblicuas y en pares desiguales, por lo general las hojas menores 4 a 8 veces más pequeñas que las mayores; hojas mayores oblongo elípticas a elípticas, 12,5-21 x 3,5-8.8 cm; hojas menores 0,9-65 cm x 0, ids cm de ancho; el margen poco ondulado y repando, de un color café claro a anarajado-amarillento est te en el envés hacia la vena central, glabras en ambas superficies y un poco escabrosas hacia los márgenes, con pequeñas escamas blanquecinas-cremosas, bases agudas, obtusas y asimétricas, ápice largamente acuminado, peciolos 3-9 mm de largo 1-1,5 mm de ancho. Inflorescencias péndulas de 9~20cm de largo con 2 a 19 flores en 2-3 dicasios, pedúnculos 3,8-12 cm, de menos de 1mm de diámetro, glabros y angulados. Flores con pedicelos de 8-16 mm, obcónicos, menos de 1 mm ancho proximalmente hasta 3 mm distalmente; cáliz glabro, apos de 2-3 mm de largo, erue acostillado donde cada costilla finaliza en un lóbul I ola verde lada, 7-20 mm de largo, 5-lobada en 1/2 a 2/3 de su extension, lobos oblongo- idos y cuculados, con venación evidente, 3-5 mm de largo, ciliolados en | é , más largos que el tubo, filamentos 1 mm de largo, fusionados a la corola por la mitad des su longitud, labras: anteras basifijas, 2 mm de largo, no apiculadas; ] Rot Ras Inet Tevac 2(1): AT — 44. 2008 - AN A lcm GK icm 1mm C Fic. 1. Ilustración de Cuatresia amistadensis. A. Rama florífera, B. Infrutescencia, C. Flor, D. Corola abierta, F. Antera, G. Fruto. pistilo 6-10 mm; estilo apical y estigma claviforme. Frutos 8-12 x 7,5-12 mm, redondeados, cáliz no acres- cente. Semillas 2,5-3 mm de diámetro, discoides, de color pardo claro con la superficie foveolada. Distribución, hábitat y ecología. —Bosques muy Húmedos y nubosos, bosques primarios y secundarios, principalmente en la vertiente Caribe, más escasa en el pacífico, al Sur de la Cordillera de Talamanca, en Costa Rica y Panamá, entre 980-2050 m. Cat ^J RA Il Cuaoro 1. Cuad parativo entre Cuat tad C. ph y Witheringia cuneata. Carácter Cuatresia amistadensis Cuatresia plowmanii Witheringia cuneata Pedúnculos Pubescencia en hojas e inf Pedicelos Cáliz floral Corola Estambres Rango de altitud 3,8-12 cm de largo, «1 mm de diámetro, angulados Hojas glabras con diminutas escamas blancas. Infl. glabras 8-16 mm muy delgado y difícil de distinguir del cáliz Hipocrateriforme Lobos 3-5 mm, divididos por » 1/3 de su largo, tubo excerto del cáliz por 2-3 mm Filamentos glabros, 1 mm, unidos al a por > Y de su largo, eras 2 mm i 2050 msnm 4-25 cm de largo, 0,75-1 mm de diámetro, cilíndricos Hojas glabras sin diminutas escamas blancas. Infl. glabras 5-8 mm, muy delgado y difícil de distinguir del cáliz Campanulado Lobos ca. 4 mm, dividido por < 1/3 de su largo, tubo excerto del cáliz por 7-10 mm Filamentos piliformes, 2-3 mm, unido al corola por < Y e su largo, anteras 2 m 0-2000 msnm 0,3-6 cm de largo, 1-2 mm de diámetro, cilíndricos puberulentas a pubescentes -6 mm, robusto y claramente definido del cáliz dla y truncado lobos 4-8 mm muy angostos, tubo Gebees inserto en el cáliz Filamentos pilosos, 1 mm, anteras 2,5-3 mm 100-1900 msnm Fenología.—Se ha recolectado con flores de Diciembre a Mayo, y con frutos de Febrero a Marzo. Etimología.—El epíteto de este taxón está dedicado al parque internacional La Amistad, ubicado entre Costa Rica y Panamá (PILA). Cuatresia amistadensis es fácilmente distinguible por poseer pedúnculos péndulos, muy largos y del- gados, hasta 10.5 cm, además con hojas dimidiadas secando de un color café claro. Esta especie se podría confundir con Witheringia cuneata (Standl.) Hunz., ya que comparten un rango de distribución geográfica similar y porque tienen pedúnculos alargados, pero en W. cuneata son más cortos y robustos, miden como máximo 6 cm. Las diferencias entre ambos taxones se mencionan en el Cuadro 1. Otra especie muy similar es Cuatresia plowmanii A.T. Hunziker, que se distribuye en el Sur de Colombia y Ecuador, el largo de los pedúnculos es similar pero los lobos de la corola en C. plowmanii son triangulares vs. oblongo—acutados, están divididos por < 1/3 del largo de la corola vs. divididos por > 1/3 del largo; además la inflorescencia es una cima unípara vs. compuesta por dicasios y las hojas presentan una venación terciaria muy continua y d otras diferencias entre estos taxones se anotan en el Cuadro 1. COSTA RICA.P C de Coto Brus, Z Protect Las Tablas, Cuenca Térraba Sierpe, fi La Neblina, finca boa 1122 (CR, INB, MO). I Cantón de Talamanca, Parque ] 9?21'25" N, 83°13'20" O, 1900 de Lucho. 9? 05'052' N, 82244733" W. 1850 m, 92041270" N, 82°44'174" O, 1750 m, 11 Mar 2004, (fl) , ridgetop, 9°04' 270 N, Pii 174" O, 1750 m, 13 Mar 2004, f ‘Falso a 9°09'54" N, l d between a M PR 825417" N, 82?45'03" O, 2050 m, 1 Mar 1997, B. G le Talamanca, Río Lori d ] Ujan 4s a San losé Cabé 70] (CR INB) PANAMÁ Dora del T I S Em nis m Yi m. 1993 (f) A. E 19 Mar 2004 (fl) E. Alfaro & A.K. Monro 5628 (BM, INB, PMA); La pat A.K Monro & E. Alfaro 4302 (BM, INB, MO, MEXU, PMA); La pata del Ced (£1) A.K Monro & E. Alfaro 4379 (BM, INB, MEXU, MO, PMA); Caribl g 82°40'45" O, 980 m, 25 Mar 2005, A.K Monro & S. Cafferty 4826 (BM, INB, MO, PMA). Chiri Chiriquí Grande, 4—4,5 km N of dam over Fortuna Lake, 8°43'N, 82?17' O, 1100-1135 m, 8 Mar 1985, TB. Croat & M.H. Grayum 60046 (BM, MO); 28 Dic 1995, E. Montenegro & B. Cuevas 1148 (BM); 28 Dic 1995, E. Montenegro & B. Cuevas 1185 (BM); SE slopes of Cerro Pate Macho, trail for Río Palo Alto, 4 km NE of Boquete, 1700—2100 m, 26 May 1981, K. Sytsma et al. 4858 (BM, MO) AGRADECIMIENTOS A la Iniciativa Darwin por la beca 415/027 que nos permitió el trabajo de campo. A Cyril H. Nelson en TEFH, por revisar y corregir la traducción latina de la sinopsis. A los curadores de los Herbarios MO, BM, PMA por permitir el acceso a sus colecciones. 44 [| [| £ al Dos H In LI rr fTexas 2(1 ) REFERENCIAS Hunziker, A.T. 1977.Estudios sobre Solanaceae VIII. Novedades varias sobre tribus, géneros, secciones y especies de Sud América. Kurtziana 10:7-50 Hunziker, A.T. 1987. Studies on Solanaceae. XXI, A preliminary synopsis of Cuatresia. Opera Bot, 92:/3-82. Nee, M. 2004. Solanaceae. In: Smith, N., S. Mori, A. Henderson, D. Stevenson, and S.V. Heald. 2004. Flowering plants of the Neotropics. Princeton University Press. The New York Botanical Garden. New Jersey. E.F.UU. TWO NEW SPECIES OF BACCHARIS SECT. CAULOPTERAE (ASTERACEAE: ASTEREAE) FROM SOUTHERN BRAZIL Angelo Alberto Schneider Ilsi lob Boldrini Geen Federal do Rio Grande do Sul Universidade Federal do Rio Grande do Sul e Pós-Graduacáo em Botánica rograma de Pós-Graduação em Botânica P Bento Goncalves, 9500, Porto Alegre Av. Bento Goncalves, 9500, Porto Alegre Rio Grande do Sul, 91501-970, BRASIL Rio Grande do Sul, 91501-970, BRASIL angeloschneidereyahoo.com.br ilsi.boldrinigufrgs.br ABSTRACT T peci f Baccharis L. sect Caulopterae DC. (Ast )f highlands of ] Brazil ted, Baccl foliosa A.A. Schneid. & Boldrini and Baccharis fl i A.A Schneid & Boldrini. The new species are del. illustrated es | d P ¿1 : +7 I D 1 : Dn) fl (I 3 DC and D d g Dal , respective ly RESUMEN Dos nuevas especies de Baccharis L. sect. Caulopterae DC. (Asteraceae) que vi n regiones de altitud del de Brasil se pr Baccharis NOIA A. Schneid. & Boldrini and Baccharis fl i A A Schneid. & Boldrini. Se deseen, ans 3i las similares, Baccharis milleflora (Less.) DC. y Baccharis organensis Baker, respectivamente. r The infrageneric classification of Baccharis was recently worked for Giuliano (2001, 2005) to Argentina, Giuliano and Nesom (2003), and by Müller (2006) to Bolivia. Müller used the name “Baccharis genistelloides Group” for the species belonging to Baccharis sect. Caulopterae, which consists of perennial herbs, subshrubs, and shrubs with 2-3- alate stems, normal to squamiform leaves, capitula sessile and arranged in spikes or vithout paleae, and achenes (5-20)-ribbed and glabrous or papillose. Section Camlonienies is ded. to South America. During a taxonomic revision of Baccharis sect. Caulopterae of southern of Brazil two new species were recognized with morphological characteristics that distinguish them from all other species. Baccharis apicifoliosa A.A. Schneid. & Boldrini, sp. nov. (Figs. 1-2). Te: BRAZIL. Rio GRANDE po Sur: Municipio de São Francisco de Paula, rodovia RS 20, km 97 (29° 26! 31.1"S, 50? 32! 35,2"W), 20 Nov 2007, A.A. Schneider 1542 (HOLOTYPE: ICN; isotypes: K, M ), alis Affinis B. milleflora sed alis angustioribus, foliis oblongis vel 1 et foliis granulosis albidis punctiformis differt Shrub, 0.8—1.5 m tall; shoots branched, terminating in a capitulescence; stems 3-winged, wings to 2-8 mm wide, slightly undulate, distinctly interrupted, grayish green, surface cover with whitish granules (stomata), indument of flagellate hairs. Leaves obovate to elliptic, sessile, seemingly 1-veined, covered with whitish granules, indument of flagellate hairs, larger leaves 0.5—1.5 cm long, 0.2-0.6 cm wide, apex obtuse to rounded, base cuneate, margins entire. Capitula sessile, in terminal short spikes 1-3 cm long, forming pyramidal panicles. Male capitula 3-5 mm long.; flowers 10-17; involucre 2-3 mm long, 2—2.5 wide, campanulate; phyllaries in 3—4 series, outermost phyllaries ovate, median phyllaries lanceolate, innermost phyllaries linear, 2.6—3.1 times as long as the outermost; all phyllaries with margins broadly scarious; apex obtuse fimbriate; corolla 3—4.3 mm long, tube 2-2.5 mm long, lobes 1-1.8 mm long, coiled at maturity; anthers with apical appendage 2.73.2 times as long as the filaments, anther apices acute; style slightly exceeding the corolla, with sweeping hairs, the apex nearly fully divided into lanceolate branches, achenes abortive, glabrous and reduced; pappus uniseriate, 3-4 mm long, bristles 15-20, apically broadened and scarcely barbellate. Female capitula 5-7 mm long; flowers 20-30; involucre 3-5.5 mm long, 2-3 mm wide, cylindrical; phyllaries in 3—4(—5) series, outermost and median phyllaries like those of male capitula, in- nermost phyllaries linear, 0.5-0.7 mm long., 0.1-0.2 mm wide; corolla 2.5-3 mm long, 0.1-0.2 mm wide, I Rot Rac Inct Tavac 2(1): 45 e 51. 2008 Fic. 1. Holot 2 0 tu 018 Y243H d ha AMI 1 V € 2 S H "d Pta di ae =] mm | Herb. Depto. Bet. ta E ERGS - Porto Ai gei | Nome bits Alt 505 12007 Dati: S0 a Habitat Arbusir ep carp ern Fam ¿Apia UNS 190258 | Nome pop. Col: ^ A Senner i547 Det: A A Schnee Loc Bras En e boro o ce Pauia Bir af «RU S E ES To sell Biriga Acervo: HOLO EH I rn » np (A.A. Schneider 1542, ICN). eic d I Boldrini, N e e ee 47 apt enn À f, T aif x. z iei a m rv $ au x LE M EU. A LM Ba ne z " ` "xd px S25 + F O " EP Pm Fa = LC z - uda ar E f Pd Fic 2. Bacchari icifoli A Flowering branch, famale plant BR leaf (A IB f P Wacum 802 PA CA) C. Female capitula D. Male ranitulum. E A A m | by Fal mi ri (C Female flower. H mL T | . £r | VEA IT (E from A eL FAFPFADAFTA! G E, G and H from A.A. Schneider 1542, ICN). filiform, apex short-ligulate with ligule shallowly 3—5-dentate, glabrous; style 5.2-6.1 mm long, branches 0.4—0.8 mm long; achenes 0.9-1.3 mm long, 0.3-0.5 mm IES nearly cy lindric, slightly bos later- ally, covered by low papillae; 57) -ribbed, ribs with p 3 mm long, bristles 20-25, persistent, basally fused. Etymology.—The specific epit] the shoots. {_pnoranhical Aictedhaitian and habitat e L r r EE t refers to the presence of conspicuous leaves only in the apical part of Restricted to highlands of south Brazil, in the states of Rio Grande do Sul and Santa Catarina. This species grows in grassland, with many individuals occurring abundantly near and in damp areas and often in secondary forests, at altitudes between 750-1500 m, often growing with B. milleflora, B. trimera (Less.) DC. and B. articulata (Lam.) Pers. PanArvPEs. BRAZIL. Rio G de do Sul: B I ,F da Fundo das Almas, 23 Nov 2001, R. Wasum 1262 (HUCS, PACA); Cambará J 48 t tani i Texas 2( do Sul, Feb 1948, B. Rambo 36246 darn GE ee a 20 Nov 1950, A. Sehnem 5050 (PACA); Sáo Francisco de Paula, rodovia RS 235, 12 Nov 2000, R. Wi Taquara, 26 Nov 2000, R. Wasum 773 (HUC S, PACA); Pró-Mata, 1 Oct 2001, M. Sobral et al. 9394 (ICN). Santa Catarina: Bot Retire: Campo dos Padres, 19 Dec 1948, R. Reitz 2619 (PACA). Urubici, Serra do Corvo Branco, 17 Oct 2006, A. A. Schneider 1358 (ICN). Baccharis apicifoliosa is similar to Baccharis milleflora (Less.) DC.; the two species are differentiated by con- trasts in the following key. 1. Species with conspicuous leaves, leaves oblong-lanceolate; surface of wings as with whitish granules; wings to 2-8 mm wide accharis apicifoliosa 1. Species with squamiform leaves; surface of wings without whitish granules; wings to E gra 20) wide Baccharis milleflora Baccharis flexuosiramosa A.A. Schneid. & Boldrini, sp. nov. (Fig. 3-4). Tyre: BRAZIL. Rio Granne DO Sur: Municipio de Cambará do Sul, Parque Nacional dos Aparados da Serra, Canyon Itaimbezinho, 7 Dec 2006, A.A. Schneider 1419 (HOLOTYPE: ICN; IsoTYPES: K, MO, RB) BMW. : “1 Al Bacchari g i Baker similis, 1 folii | ii paci (vs. vernicosis), ramis alatis (vs 1 Shrub, 0.8-2.5 m tall; shoots | hed, terminating in a capit , stems 3-winged, wings to 2-8 mm wide, plane, attenuate near to the nudes internodia long, pur green. Leaves oblanceolate-obovate or elliptic, sessile or with a short petiole to 4 mm long, seemingly 1-veined on the adaxial face and 3-veined on the abaxial face, covered with hair tufts, opaque, 1.2-3 cm long, 0.7-1.8 cm wide, apex obtuse to rounded, base attenuate, margins entire, l-nervate in adaxial face and 3-nervate in abaxial face. Capitula sessile, in terminal short spikes, 1-3 cm long, forming pyramidal panicles. Male capitula 5 mm long; flowers 20; involucre 4 mm long, 2-2.4 mm wide, en Rue in 4 4 series, outermost phyllaries ovate, median phyllaries See lanceolate, innermost phy nceolate or lanceolate, 2.8-3.2 times as long as the outermost; all phyllaries with 11 | te with Ambas hairs; corolla 3-3.5 mm long, tube 2-2.5 mm long, with i» hairs, lobes | mm dono coiled at maturity; anthers with apical appendage 3 times as long as the filaments, apices acute; style slightly exceeding the corolla, with sweeping hairs, the apex nearly fully divided into lanceolate branches, dilated towards apex, with few hairs, achenes abortive glabrous and reduced; pappus uniseriate, 3-3.5 mm long, bristles 15-20, apically broadened and scarcely barbellate. Female capitula 5-6 mm long; flowers 20-25; involucre 4.5—5.5 mm long, 2-2.5 mm wide, cylindrical; phyllaries in 3—4 series, outermost and median phyllaries like those of male capitula, innermost phyllaries lanceolate-linear; corolla 2.3-2.6 mm long, 0.1-0.2 mm wide, filiform, ligulate with dentate ligule, glabrous; style 3.1-4 mm long, with few hairs; achenes 1.5-1.8 mm long, 0.3-0.5 mm wide, nearly cylindric, slightly compressed laterally, covered with inconspicuous papillae; 6(-8)-ribbed, ribs with papillae more prominent; pappus uniseriate, 2.8-3.3 mm long, bristles 15-20, persistent, basally fused. Etymology.—The specific epithet refers to the flexibility of the shoots. Geographical distribution and habitat. —Restricted to the highlands of southern Brazil, in the states Rio Grande do Sul and Santa Catarina. This species grows on the borders of cold forests and on shores of rivers at altitudes between 900-1500 m. PARATYPES. BRAZIL. Rio G de do Sul: Municípi de Cambará do Sul, Parque Naci ld AT los dat , Canyon lt bezinho, 7 Dec 2006, 4.4. Schneider 1420 (ICN); M le São F le Paula, Distrito de Tainhas, Camping Passo da IIha, 29? 05' 09.2"5, 50? 21' 48.9"W, 900 m, 4 Dec 2006, AA. ORUM 1448 (ICN). Santa Catarina: Município de Urubici, S lo C B , 28° 03' 36.5"S , 49? 21' 48.1"W, 1400 m, 18 Oct 2006, A.A. Schneider 1348. Baccharis flexuosiramosa has 3-veined leaves and apparently is closely related to B. organensis; the two species differ by contrasts in the following key. 1. Shoots winged; leaves seemingly 1-veined on adaxial face, 1.2-3 cm long, 0.7-1.8 cm wide Baccharis flexuosiramosa 1. Shoots not winged; | 3-veined on adaxial face, 3.5-5.5(-6.5) cm long, 1.5-2.6 cm wide wide Baccharis organensis J Nome: Fa “Ties etre TU wo S P t £ i 0 N231I-OIHVUHUZJH 9 EA k, e JA ra ZC Pots pisado, A ES Is 1 D 1 4 8 E t3 ty v3 INL Lidl Hd b EA iy parador da Senta i Bes bz OS AR (Or Data: CG tS VOIR Habitat Fordo e Mara Biene Acer prod pon lr Fic. 3. Holoty; (A.A. Schneider 1419, ICN). 50 | | of the Botanical R h Institute of Texas 2(1) o 4 Zum 2 Ii no cp A gr. TE: a an TEE Ra caia aint c Bega e ix, r^ POT PPM ha « su MED “a RALE e ^ E Ada we Hi et. ae MUI TURA Se A pa dio 2 pd D E «i ets $ Kc q35- 7 D m, moy 5 E .. a. ee 2 E é pr Max a Le woe ue he H s e 4 - ` ». "S da do 1, e SS frg + E " ï r " 3E e ES. i AM ao d A den a T0. wre eee = Ben ie E ‘ = E EE et S TP a E t wo t D ah E Fi A Lei à 4 A 4 D + ei - - ra ae LET E "E he Seuss WE ER Fic, 4. Baccharis fl i A Flowering branch, female plant B. Leaf C. Part off | ikes. D. Female capitula E Phyllaries of | itul ter t ). F. Male flower. G. Female flower. H. Acl ined by SEM (A Cand G-H H from A.A. Schneider 1419, ICN). ACKNOWLEDGMENTS The authors thanks Rafael Trevisan and E dee EN ee L. Nesom and Jochen Müller for the 13 reviews and suggestions, Jair G. Kray for , Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for the al support of our cies REFERENCES Guuiano, D.A. 2001. Clasificación infragenérica de las especies argentinas de Baccharis (Asteraceae, Astereae). Darwiniana 39:131-154 GIULIANO, D.A. 2005. New infragenera in Baccharis (Asteraceae, Astereae). Novon 15:535-541. f. * | IDA: AI H £n | 5 E al n 2] 51 GIULIANO, D.A. and G.L. Nesom 2003. A new section of Baccharis (Asteraceae: Astereae), and notes on allied taxa. Sida 20:1481-1484, Overa, A.S., L.P. DEBLE, AA SCHNEIDER, and J.N.C. Marchiori. 2006. Checklist do género Baccharis L. (Asteraceae- Astereae) para o Brasil. Balduinia 9:17-27. MULLER, J. 2006. Systematics of Baccharis (Compositae-Astereae) in Bolivia, including an overview of the genus. Syst. Bot. Monogr. 76:1—339. BOOK REVIEW lan R. Hatt, GORDON T. BROWN, AND ALESSANDRA ZAMBONELLI. 2007. Taming the Truffle: The History, Lore, and Science of the Ultimate Mushroom. (ISBN 978-0881928600, hbk.). Timber Press, Inc., 133 S.W. Second Avenue, Suite 450, Portland, Oregon 97204-3527, U.S.A. (Orders: www.timberpress.com, mail? timberpress.com, 1-503-227-2878, 1-800-327-5680, 1-503-227-3070 fax). $29.95, 304 pp., 9" x 6". Contents: Preface Acknowledgments 1. From the Past Comes the Present.—Regions of Renown; La Grande Mystique; The Golden Age of Truffles; Magical Powers and Me- dicinal Virtues 2. Science to the Rescue.—Mycorrhiza: An Intimate Relationship; Types of Mycorrhizas; Structure of Ectomycorrhizas, Underground Warfare; The Collapse of pe encon New PS 3. Identifying Truffle Species. , Nomenclature, and T: y; Identifying Species of Truffles; Périgord Black Truffle; ee or Summer inci Winter Truffle; Bagnoli Truffle or Truffe Mésentérique; Smooth Black Truffle; Asiatic Truffle; Italian White Ti oloure ies; North OEA Pale- Coloured Ti Truffles; dis Truffle: Desert Truffles; Inedible and Poisonous “Truffles; Other Species of Truffle; False Truffles; Ee Truffle Species from Their M izas; Identifying Truffl g Molecular Techniques 4. The Habitats of Some Commercial Truffles.— Périgord eu Truffle; Burgundy or Summer Truffle; Italian White Truffle; Bianchetto; Desert Truffles This was a delightful book covering most of ything you ever wanted to know about truffles. For instance, I learned that during medieval times, truffles garded as witches spit—quit Il from th rin É the gods during the time ee Roman rule! Dr. Hall tnl] Lost TL we (egal I ] JA 11 1 £T CELLS tio [ar] -rl 1 : 1 1 “41 FÉ. 3 asf PRE 1 lies f, fei £11 a E 1 1 soil dod "i temperature, as Ge sun/shade conditione: and tn trees iid which. Ss aus il bes: Since carmen are and growing of truffles. ME : £m 1 a | | 1 A NT E : : E lr n led KA P" L i [= PES n LI to home to train dnd truffle EH tl ion that truffle pigs may be less A tog p their found truff! tl TÉ 1 a 64 cr] E. 1: " "mi f: " rr] 71 1 e [tf] dogs. If y j I gg are in season, and [ ide at least F hj b, “Rain in July, truffles for Christmas.” Tal : Fg A | 1 Te 1 1 1 H 1 7 HËNN Le JA H I thoroughly enjoy y yp gs. That PE eid have a wonderful r1 EP 1: 1 1 1 1 ti Kay Yount O L ia E fuerte, Tarrant County Master Gardener and BRIT Volunteer, Fort Worth, Texas, U. e A. J. Bot. Res. Inst. Texas 2(1): 52. 2008 TWO NEW COTONEASTERS (COTONEASTER: ROSACEAE) FROM YUNNAN PROVINCE, CHINA Jeanette Fryer Bertil Hylmót Cornhill Cottage, Honeycritch Lane Bjuv, SWEDEN Froxfield, Petersfield, Hampshire U32 1BE ENGLAND ABSTRACT Two new cotoneasters are described from Yunnan Province, dude Cotoneaster qungbixiensis is disti df the similar C , less rugose, decid llipti ly Both species are in the hem described series Sterniani, e differs nen series Franchetioides in having semi-evergreen to E foliage that is green not y its less shiny and less rugose a and na no) didi leay a A DEE fet RESUMEN So d da e gt incia de Yunnan, ee t ter qungbixi i disti de C. sternianus por sus hoj d ás delgadas, menos rugosas, que son te elípti poco ao das Están en la Serie Sterniani d d? EE EENG TE | 1 L: 4 7 E 3 E d : | verde grisaceas, y el color de las anteras mucho más padoe t ter floridus de la Serie Dielsiani parable de C. dielsi I , brillantes y menos rugosas, inf] flores, y frutos obovoides. In EE of a DE of the genus Cotoneaster two new species and one new series have come to light, all in the subgenu Cotoneaster, and are described | t] for use in the upcom- ing treatment of the genus in the Flora of North America. Cotoneaster series Sterniani J. Fryer & B. Hylmó, ser. nov. Wee: Cotoneaster sternianus (Turrill) Boom, Jaarb. Ned. Dendrol Ver. 20:81. 1957 Affinis Cot ies F hetioides Flinck & B. Hylmö sed folii I i , SUT trovi , antheris albis, pyrenis 3, —4, —vel 5 differt. Shrubs, small t lium sized, often densely branched. Leaves deciduous or semi-evergreen, subcoriaceous or coriaceous, lower surface tomentose, upper surface rugose. Inflorescences 1-20-flowered; the fertile shoots 10-70 mm long. Flowers with hypanthium tomentose-pilose. Anthers mostly white. Fruit orange- red. Pyrenes (2—)3—4(—5). Fruiting mid to late season. Cotoneaster series Sterniani includes five species: Cotoneaster elegans (Rehder & E.H. Wilson) Flinck & B. Hylmó; C. induratus J. Fryer € B. Hylmó; C. insculptus Diels; C. qungbixiensis J. Fryer & B. Hylmó; and C. sternianus. Members of the series Sterniani are distributed in Asia from China (Yunnan, Xizang (Tibet), Sichuan) to Myanmar. Species belonging to series Sterniani are closely related to those of series Franchetioides, which differ in having evergreen leaves, with the upper leaf surface grayish-green, anthers mauve, pink, or purple, and pyrenes 2 or 3 per fruit. Cotoneaster qungbixiensis J. Fryer & B. Hylmó, sp. nov. (Fig. 1). Tyee: CHINA. Yunnan: Dali Xian, near Dali, eastern slope of Cangshan, Qung Bi Xi, 15 Sep 1987, K. Brickell & A. Leslie 12455 (HoLoTyrE: WSY; isotypes: E, KUN). Affinis Cotoneaster sternianus nian B d foliis decid ] hart l teril gusto-ovatis vel angusto-ellit is t tis et venis 2—4 bini dier, Shrubs, 23 m tall. Branches erect and arched, spreading; branchlets distichous o piraled, reddish-brown, initially strigose-pilose. Leaves deciduous or semi-evergreen, chartaceous, on sterile shoots narrowly ovate or narrowly elliptic, 28-42 x 13-21 mm, base cuneate or obtuse, apex typically acute-truncate, or rarely J. Bot. Res. Inst, Texas 2(1): 53 — 56. 2008 Fic. 1. Cotoneaster qungbixiensis. A. Flowering shoot. B, C. Fruiting ` Fic. 2. Cotoneaster floridus. A. Flowering shoots. B, C. Fruiting shoots. S acute, lower surface greyish tomentose, upper surface slightly rugose, dark green, shiny, initially pilose, lateral veins 2—4 impressed pairs; petiole 3-4 mm long, often red, strigose-pilose. Inflorescences compact, 3-9(-15) flowered; fertile shoots 30-50 mm long with 4-6 leaves; pedicels 1-3 mm, strigose- pilose. Flow- ers (including hypanthium) 7-8 mm long; hypanthium infundibulate, silky tomentose; sepals cuspidate or acuminate, tomentose. Corolla open; petals erect-incurved, pink with dark red base, apex yellowish to off- white. Stamens 20; filaments dark red; anthers white. Fruit orange-red, slightly shiny, globose or obovoid, 8-10 mm, pilose, calyx lobes infolded, appressed and forming a flat apex, tomentose. Pyrenes 2-34 (ca. 10%-80%-10%), rarely 5, persistent style on abaxial surface, 2/3 from base. E | Hyl ó, T ras j £ PL 55 Growing | g batcl f seeds y ield ifi plants, suggesting it is an icti id ,like many species of Cotoneaster in the subsets Cotoneaster (Hjelmqvist 1962; Battish: et al. 2001) and within other genera of the subfamily Maloidea (Campbell & Dickinson 1990; Campbell et al. 1991). The chromosomes of cultivated material in England were recently counted by both John Bailey and Hugh McAllister, confirming O. aida isa dad 2n = 68 Otl 1: CHINA. Yunnan: Dali, E sl [C M i ktol Peak, 4 Nov 1996, J. Fryer JFWY-063 (GB); N between Zhongdian and Dali, Beahan Chiang, 22 Sep 1994, Alpine Garden Society China Expedition 1114 (E); Yunnan, Dali, on Cangshan, above a small temple, Nov 1985, C. Wingfield 772 (seed accession). SWEDEN: "e Ge grown from seed from type collection, Hylmó accession 2087, 11 Jul 1995, B. Hylmó s.n. (WTU). ENGLAND. H Clanfield, cultivated, — Gardens, seed source Wingfield 772, 29 Jun 2001, J. Fryer 758 (WTU); Hampshire, Sir Harold Hillier Coen cultivated, Hillier cession 1994.1175*A, seed source AGS China Expedition No. 2490, 2000, J. Joyce 293 (HILL), J. Joyce 13 (HILL). U.S.A. ae ae King Co.: Seattle, Washington Park Arboretum, doa accession 189-60-A, 11 Oct 2000, P.F. Zika 15550 (WTU), same shrub, 24 Jun 2006, P.F. Zika 22816 (MO, GH, NY, WTU, WTUH); same site, accession 189-60-B, 14 Dec 2006, P.F. Zika 22901 (WTU, WTUH): same site, accession 162-57-A, 24 Jun 2006, P.F. o 22815 (GH, OSC, WTU, WTUH). CANADA. BririsH COLUMBIA: Saanich, Rithets 0 Jun 2006, P.F. Zika 22807 (CAN, DAO, GH, NY, UBC, WTU), same site, 21 Oct 2007, PF Zika 23 o CAS, DAO, GH, UBC, Uc, Y, WTU). E e Phenology and ecology.—Flowering June to July; fruiting October to November. Natural populations in Yun- nan Province, China, are in full sun on rocky slopes over calcareous bedrock. In cultivation, Cotoneaster qungbixiensis is hardy to -18°C, and is an attractive shrub with good autumn fruit color. Cotoneaster qungbixiensis is named for the type locality on Cangshan Mountain in Yunnan. It is closely related to C. sternianus, which differs in having leaves that are evergreen, thicker and coriaceous, broadly elliptic to broadly ovate with apex acute or acuminate and lateral veins in 4—6 pairs. SERIES DIELSIANI G. Klotz, Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss. Reihe 21:991. 1972 Cotoneaster floridus J. Fryer & B. Hylmó, sp. nov. (Fig. 2). Tres: CHINA. Yunnan: Muli, Washin area, near lamasery, 5 ct 1937, TT Yi 14430 (uororveE: E; ISOTYPE: A, BM). Affinis Cotoneaster dielsianus E. Pritz. sed foliis chartaceis ad subcoriaceis, supra planities obscurus, inflorescentiis 2—5-floris, fructu oblongo-obovoidus differt. Shrubs, 1.5-2 m tall. Branches erect and arched; branchlets distichous, maroon and minutely verruculose, initially pilose-strigose. Leaves deciduous, chartaceous or subcoriaceous, on sterile shoots elliptic, often broadly so, 15-25 x 11-25 mm, base cuneate or obtuse, apex acuminate or acute, lower surface whitish tomentose-pilose, upper surface dark green, dull, strigose, veins 3(4) slightly impressed pairs; petiole 3—4 mm long, strigose. Inflorescences 2—5 flowered; fertile shoots 20-40 mm long, with mostly 4 leaves; pedicels 2-6 mm long, strigose. Flowers (including hypanthium) 5-6 mm long. Hypanthium turbinate, densely strigose; sepals cuspidate or acuminate, densely strigose, membranous, reddish-brown, and glabrous near margin, margin villous. Corolla closed; petals erect-incurved, pink with dark red base and white and pink border. Stamens 14-17, filaments dark red or red with pink apex; anthers white. Fruit rich red, intensely shiny, narrowly obovoid, 10-11 mm long, sparsely pilose, calyx lobes infolded, appressed, densely pilose. Pyrenes 2—3—4 (ca. 1096—6096—3096), persistent style on abaxial surface, 2/3 from base. Growing large batches of seed yields uniform plants, suggesting this is another apomictic species, but the chromosomes have not been studied to confirm this. Other specimens examined: CHINA. YUNNAN: Chungtien [Zhongdian], Pica, 30 Oct 1937, T.T. Yü 13905 (A, BM, E). ENGLAND. Hamp- SHIRE: Sir Harold Hillier Gardens, cultivated, Hillier accession 1992.0451*B, 2001, J. Joyce 329 (HILL). Cotoneaster floridus is named for its florid red ins Es is closely related to S E E. Pritz ex Diels which differs in its thicker (cori eaves with PI gntiy rug and shiny, inflorescences 3-7(-10) flowered, and in its globose a Phenology and ecology.—Flowering June; fruiting September to October. Little is] bout its habitat or associates where it is native in China, in Yunnan and Sichuan Provinces. In cultivation it is hardy to -21°C. Cotoneaster floridus is frequently found in cultivation as C. rubens hort. non W.W. Sm., a misnomer under which it was distributed as seed from the Royal Horticultural Society Wisley Gardens in England. It can also be found under the name e dielsianus var. rubens hort. Unfortunately, Professor Yü's collection bers have | confused in cultivation, hence various species of Cotoneaster can sometimes be found SE a single collection number. ACKNOWLEDGMENTS We thank the herbaria cited for loans and access to specimens, we are grateful to John Bailey and Hugh McAllister for chromosome counts , and extend ppreciation to Ken Chambers, Allen Coombes, Randall Hitchin, John Jewsbury, Peter Hylmó, and Peter Zika for research assistance. We also appreciate the helpful comments of David Bouford (GH) and an anonymous reviewer. REFERENCES Barts, IN. B. Hyimô, and H. Nysom. 2001. RAPD analysis of interspecific relationships in presumably apomictic Cotoneaster species. Euphytica 120:273-280. CAMPBELL, C.S. and T.A. Dickinson. 1990. Apomixis, patterns of morphological variation and species concepts in subfam. Maloideae (Rosaceae). Syst. Bot. 15:124-135. CampseLL, C.S., CW. Greene, and T.A. Dickinson. 1991. Reproductive biology in subfam. Maloideae (Rosaceae). Syst Bot. 16:333-349. Hue. vovsr, H. 1962. The embryo sac development of some Cotoneaster species. Bot. Not. 115:208-256. MELIOSMA ANTIOQUIENSIS: UNA NUEVA SABIACEAE DE COLOMBIA Xavier Cornejo The New York Botanical Garden 200" St. and Kazimiroff Ave. SOn New York 1 0438- 5126 a S. id Ej id A - +? — Soc RESUMEN Cag -7 : | EVI (Calz a ATA Aca : Colombia Est I IF 1 ? E E F1 . - 1.4 Iw es S | 1 i J: L 3 Iu fai h] lad ] t 1 Ee en las ramas, pétalos internos muy pequeños, bífidos y estilos muy duédoso o ausentes. PALABRAS CLAVE: Sabiaceae, Meliosma, pétalos bífidos, endémica, Colombia ABSTRACT f Meli (Sabi ) is described f. he Depart f Anti dee Ti is characterized by having large PA leaf blades with attenuate, decurrent base, paniculate infl ] gI fl , that are laxly arranged, very small, bifid inner petals, and very short or absent stv vies Key W Sabi , Meliosma antioquiensis, bifid petals, endemic, Antioquia, Colombia INTRODUCCIÓN Meliosma Blume (Sabiaceae) es un género de árboles y arbustos nativos de Asia y América tropical. En el Neotrópico comprende ca. 70 especies, y se encuentra distribuido desde México hasta Brasil, Bolivia y las Indias Occidentales (Gentry 2001; Morales 2003; Arbeláez 2004; Cornejo, unpubl.). En América del Sur Meliosma se encuentra concentrado al norte de los Andes (Gentry 1980, 1986, 1992, 2001; Cornejo 2006), pero el conocimiento taxonómico de muchas de las especies andinas de este género es aún limitado debido a la falta o insuficiencia de especímenes con flores en los herbarios. En la mona grata de Meliosma para Co- oia Geen & EE SE se trataron 12 especies, seis de | f nuevas para la ciencia Post han sido descritas para la flora de este país (Cuatrecasas & Idrobo 1988; Cuatrecasas 1988; Idrobo 1988). Al revisar el material colombiano, encontré la siguiente novedad. Meliosma antioquiensis X. Cornejo, sp. nov. (Figs. 1, 2). Tiro: COLOMBIA. Antioquia: Mun. Urrao, Parque Nacional Natural Las Orquídeas, margen derecha del Río Calles, en el filo NW de la Cabaña de Calles, bp-PM, Parcela G, subparcela G-13, 06°32'N 76°19'W, 1450 m, 14 Oct 1993 (fl), A. Cogollo, A. Duque, E Giraldo, W. Rodríguez & J. Vélez 6887 (HoLotipo: NY-937622; isóTIPO: MO-5010854). L T e Speciei Meliosma ee Urb. similis, a qua differt foli tiolis | ioribus (1,5-3,5 cm nec 0,3-1,1 cm), floris pedicellatis (0,7—1 mm nec ca. P bifid (nec sessilis), 0,1 mm, nec E 0,8-1 mm). Árbol 5 m de alto. Ramas jóvenes densamente estrigosas. Hojas espiraladas; pecíolos 1,5-3,5 cm (incl. pul- vínulo 1-1,5 cm) x 4-6 mm, densamente estrigosos; lámina oblanceolada, firmemente cartácea, 40-60 x 15-20 cm, haz café PES al secar, x densamente estrigoso hacia la base del nervio medio, el resto de la lámina glabro, envés café l , estrigoso a lo largo del nervio medio, el resto de la lámina glabro, con numerosos e inconspicuos puntos elle de color rojizo-marrón; ápice + redondeado, base atenuada, decurrente y obtusa en el extremo basal, margen suavemente repando y remotamente dentado, con incon- spicuos y dispersos dientes dispuestos hacia la porción distal de la lámina, nervios secundarios ca. 20-30 pares, fuertemente prominentes en el envés, broquidódromos, nervios terciarios reticulados, prominentes en el envés. Inflorescencia una panícula axilar y subterminal, laxa, ca. 50-65 cm, con ramificaciones hasta J. Bot. Res. Inst, Texas 2(1): 57 — 60. 2008 copyright reserve COLUMBIA SADIARLEAZ SS doses ANVIOQUEMS (= 6.1 709 Tu, NEO, ue. ao. fintioguia: Urrao Municipio : dar OUR 3 Voresda Callen. P Mactroral | ure "ias DrguaideeeU. Margin art sel Ei Celli Ze Senn U. Tria NB ye la voa chm we ES Ek Parcela " athe tied... Ej Col#eetade con A Fique, T. Di w,. "c gues EK J. Veloz. Ge 32 'm Ye "t 6T gn &rbol. 3 o Fier Di ce con cé] verde. Ka WEEK A. Eeneilo wn ai, EDT Supported by National Ceographic Society JARDIN BOTANIC J. A. LDRIB'U (JAUM) MISEOLRO BOTANICAL GARDEN (MO) pn lis! T J A. Cog tal. 6887 (NY). Fic 1 TT NL Cornejo, U H M | | Y, sl? ol Fal LE 59 Së vow a oen mimm PAE ZX wë CN 1 mm weg At par A ZE e Ki Géi * » 5 DN 4 - a - rd v a”. Ca Mit dfe ^ B Fic. 2. Meli tí is X. Corne jo AO to estil tami átalo ext ición hacia | tesis. B Un pétalo interno. Dibujo del holótipo, A. Cogollo et al. 6887 (NY). de 4 to orden, densamente i idit pedunculo principal ca. 5 mm diám., ramas secundarias 5-20 cm, las de tercer y cuarto orden con fl laxamente dispuestas; brácteas florales ovadas a lanceoladas, 0,6—1,2 x 0,3-0,5 mm, ciliadas; pediceles 1,5-2,5 mm, mayormente ebracteolados. Sépalos 5, ovados a deltoides o suborbiculares, ca. 0,7-1 x 0,6-1 mm, verdes (en vivo), margen ciliado, el resto glabro. Pétalos externos 5, 1,2-1,8 x 1-2 mm, + suborbiculares, blancos (en vivo), glabros; pétalos internos 2, oblongoides, 0,7-1 mm, cuya longitud es aproximadamente la mitad de la de los estambres, adnados hacia la base de los filamentos, glabros, bífidos en el ápice. Estambres fértiles 2, 1-1,7 mm. Ovario subgloboso, ca. 0,5 mm, negro al , £labro; estilo au o muy corto, ca. 0,1 mm, estigma truncado, no diferenciado, con orificio estigmático a veces visible en forma de cruz. Infructescencias glabrescentes, frutos asimétricos obovados, 1,72 x 1,3-1,7 cm, 1-quillados, púrpura obscuro al madurar PARÁTIPO. COLOMBIA. Antioquia: Mun. Urrao, Vereda Calles, Parque Nacional Natural Las Orquídeas, margen derecha del Río Calles, en el filo NW de la Cabaña de Calles, bp-PM, Parcela G, subparcela G-22, 6º32'N 76°19’W, 1450 m, 19 Oct 1993 (fr), 4, Cogollo, A. Duque, F. Giraldo, W. Rodríguez & J. Vélez 7197 (MO). Etimologia. —El epíteto “antioquiensis” se refiere a la localidad del tipo. Distribución. — Conocida solamente de la localidad del tipo. ls MA MN es una s las especies ne este género que posee las hojas mas grandes en América del Sur. Vegeta! a M. donnel i, distribuida en Costa Rica y Nicaragua (Gentry 2001; wl Morales 2003), pero facilmente se diferencia de ésta por tener las hojas con los peciolos más largos (1,5-3,5 vs. 0,3-1,1 cm), las flores pediceladas están laxamente dispuestas a lo largo de las ramificaciones (vs. sésiles, densas), los pétalos internos son más pequeños (0,7—1 vs. ca. 1,8 mm), éstos poseen el ápice bífido (vs. en- tero) y el estilo si presente es muy corto (ca. 0,1 vs. 0,8-1 mm, linear). Por sus flores pediceladas con pétalos internos bífidos, distintivamente más pequeños que los estambres y estilo muy corto, Meliosma antioquiensis es parecida a la especie colombiana M. ellipticifolia Cuatrec. Sin embargo, esta última difiere por presentar láminas foliares mucho más pequeñas (7-12 x 3,5-5 cm), con menor número de nervios laterales (9-11), 60 j t tanical h Insti Texas 2( peciolos más cortos (0,6—1 cm), inflorescencia más corta (8—12 cm), y flores con sépalos más grandes (1,5—2 mm). Por sus grandes láminas foliares Meliosma antioquiensis es algo similar a M. itatiaiae Urb., restringida a las tierras bajas de Brasil, pero ésta presenta hojas con peciolos más cortos (ca. 1 cm), láminas con base redondeada hasta subauriculada y margen fuertemente aserrado, con curvos dientes corniculados e inflo- rescencias opuestas a las hojas, más densas, que portan flores sésiles. AGRADECIMIENTOS A Rosa Ortíz-Gentry (MO) y un revisor anónimo por sus constructivos comentarios del manuscrito y a Jim Solomon por coordinar el préstamo de las colecciones de MO. REFERENCIAS AnBtLÁEZ, A. 2004. Two new species of Meliosma (Sabiaceae) from Bolivia. Novon 14:12-16. Cornejo, X. 2006. Meliosma stellata, una nueva Sabiaceae de Ecuador. Novon 16: 328-330. CUATRECASAS, J. 1988. Miscellaneous notes on Neotropical Flora XVII. New species of Meliosma. Phytologia CUATRECASAS, J. & J.M. IDrOBO. 1955. El género Meliosma en Colombia. Caldasia 7:187-211. CUATRECASAS, J. & J.M. loroso. 1988. Tres nuevas especies de Meliosma Blume (Sabiaceae) de Colombia. Ernstia 49:8-14. Gentry, AH. 1980. Sabiaceae. In: R.E. Woodson, Jr, RW. Schery & W.G. D'Arcy, eds. Flora of Panama. Ann. Missouri Bot. Gard. 67:949-963. Gentry, A.H. 1986. New neotropical species of Meliosma (Sabiaceae). Ann. Missouri Bot. Gard. 73:820-824. Gentry, A.H. 1992. Four new species of Meliosma (Sabiaceae) from Peru. Novon 2:155-158. Gentry, A.H. 2001. Sabiaceae En: W.D. Stevens, C. Ulloa Ulloa, A. Pool y O. M. Montiel, eds). Flora de Nicaragua, 3:2303-2306. Missouri Botanical Garden Press, St. Louis Missouri. IbROBO, J.M. 1988. Una Sabiaceae nueva de la Flora de Mutis. Mutisia 72:1-3. MORALES, J.F. 2003. Sinopsis del género Meli (Sabiaceae ) en Costa Rica y Panamá, con tres nuevas especies. Sida 20:931-943. ANISOCAPPARIS Y MONILICARPA: DOS NUEVOS GÉNEROS DE CAPPARACEAE DE AMÉRICA DEL SUR Xavier Cornejo Hugh H litis The New York Botanical Garden Department of Botany h St. and Kazimiroff Avenue University of Wisconsin Bronx, New York 10458-5126, U.S.A. 430 Lincoln Drive Madison, Wisconsin 53706, U.S.A. swisecharter.net A. Spencer Tomb Division of Biolo Kansas State University Manhattan, Kansas 66506, U.S.A. mbeksu.edu RESUMEN Ts E ] : L 1 Lr io T a ER iustrad Ani is X Cornejo HH Iltis, o 3 e e Er ir ae tor; r1 41; sna] EE 153 lesu DEE lesde | f] ión, nectarios Ir E ^ B fi 1 P ERR PP " Mn eL A e] EE fuertemente UTE con un eee oe x compacto y otro menor muy di o ausente. Anisocapparis es un e Pu fof + A D a D oa: LN az a L gi a ayes F H PI I ( ) Corio & H.H dis. comb. nov. g g I p a & E Tab dé b e Las 1 12 Ba PE E ET a cp H.H. Htis I I , hasta de y I inflorescencias. f] Ali anal ño hi i ios floral ] discifi en cuvos E O L L l L i SÉ E H + L a AS f. 1 1 + 1 cry H 1 1 q 7 r fw A xu voran FL 1 fa | [ul A 1z 4 1 Beg) (A E ] E H 1.7 J 1 A EN :11 1: FALLOS Lal Lo E d 1 r DA | Je Lan A ge J 1 J dal LA ] A I ? E E I E r I Monilicarpa tenuisiliqua (Jacq.) X. Cornejo & H.H. Iltis, comb. nov., distribuid l norte de Colombia y Venezuela y Monilicarpa brasiliana (Banks ex DC), X. Cornejo & H.H. Iltis, comb. nov., disyunta al este de Brasil discuten | laci d | 5 zx — A 1 1 , ds en us eee : 1 mS m Pm 1 La aj 1 A E » t F 5 EF J t D : E PALABRAS CLAVE: Anisocapparis, Monilicarpa, Capparis, Capparidastrum, Capparaceae, new genera, Sudamérica ABSTRACT Texn nor Caith A : 4 - A a H HH 1 ius 1 : S : ES = ; i X. E H. Iltis, y g I vces, y ) g E ` - : P " H 1 Fara] which nave a Pup pae cotyledon and a a very reduced or even totally i dd minor cotyledon Anisocapparis is is a a monotypic genus, Argentina 1 e o D + EE speciosa (Griseb. ) X. HE & H. H. Tis, comb. SE proposed for a ver li ies herein transferred from Capparis s.l. second genu pror p ESA & H.H. Iltis. This is segregated a virtue s its Rasa ei n to Toad fib | ] ] flows: re nit a carincanalaimie renha that anntaime a cmall hem am D ^ a I 4 PEU ds if " aq ER A [Leib ] f. ] $ " TL Er) ` jua l+ " 1 ] y, e edges of whorls, solían grains with a striato-reticulat d if li torulose fruits of owly-attenuate ape peeing a ae row of 114 1d T1 le dedu ge a AR Ewech Ads f «I f. Lt W x DEI YA 3 XA É D Á L D L M Ls X. Cornejo & H.H. Iltis, comb. nov., f tl Colombia and A and Monilicarpa brasili (Bank DC.) x. Cornejo & H.H. Iltis, comb. nov., disjunct to E Brazil. TI lationshi di d, and a key to the American genera of Capparaceae with simple leaves, glabrous or with a simple bs is also provided. Key Wonps: Anisocapparis, Monilicarpa, Capparis, Capparidastrum, Capparaceae, new genera, South America Capparis L. s.s. es un pequeño género de distribución casi estrictamente Holárctica, centrado alrededor de su tipo Capparis spinosa L., la especie proveedora de las conocidas alcaparras y alcaparrones comestibles, ] Rot. Rec Inst Tava« 2(1): 61 — 74, 2008 62 J t tani tit Texas 2( que son los botones florales utilizados como condimento y entremés y los frutos respectivamente (Iltis & SOHO SEN Los EE s.s. son Ar bugt Usan mente po o pendulares que poseen un par de ilare itarias, zigomorfas, erectas, con el sépalo abaxial EEn (en forma de ad. eege con pétalos le: “androginóforo ausente, estambres en número de 50 hasta numerosos y nectarios florales con el ápice dirigido hacia el interior de la flor (Inocencio et al. 2006). Desde Linneo, la mayoría de las especies americanas de hojas simples de Capparaceae s.s. (excepto Morisonia L. y posteriormente Steriphoma Spreng. ) han sido incluidas y conocidas como M ao s.l., un ince extremadamente poménico y Deiere cuyos representantes americanos a tipo es molecularmente más cercano a Apophyllum F. Muell., Lit un género de dee enemies de Australia, antes que a Md Capparis del Nuevo Monde (Hall et al. 2002: 1839; Inocencio et al. 2006: 123). Basado en gicos, Hutchinson (1967: 307) concluyó que más de un género debía de ser reconocido en Capparis s.l., y propuso su reordenamiento genérico. 5in embargo, debido a la falta de un conocimiento satisfactorio y a las muy incompletas colecciones de las espe- cies de esta familia en América hace 40 afios atrás, el reordenamiento genérico de Capparis s.l. propuesto por Hutchinson prácticamente ha sido ignorado. En aiios recientes, al realizar los estudios de Capp para Flora? | y Flora de Ecuador (Iltis & Cornejo, ms. sometido; Cornejo & Iltis, en prep.), con base a las colecciones Gees relativa- mente más completas obtenidas en las pasadas décadas, así como meticulosas Io en pe ha llegado a ser claro que los Capparis s.l. de América son divisibles en varios géneros Cornejo & Iltis 2006, 20083, b, c; Iltis & Cornejo 2007, 2008). En los Capparis s.l. de América, el tipo de pubescencia es un caracter m útil, estable, bien definido y confiable (Iltis et al. Po Desde este pue Es vista, los Capparis sl dividen en dos grandes grupos, que están j t ados con el número cromosómico: 1) los de ec simple o glabros, y 2) los de pul ia peltada a estrellada, candelabra o dendroidea (Iltis & Cornejo, manuscrito sometido). Entre los Capparis s.l. de pubescencia simple o glabros están "v géneros Cynophalla (DC.) J. Presl y Capparidastrum Hutch. I pect le e ás un nuevo género de este grupo están en proceso (Cornejo & Iltis 2008b, C). Ene esta publicación se presentan dos nuevos géneros sudamericanos pertenecientes al grupo de las Capparaceae de hojas y pubescencias simples. Para obtener las imágenes de polen se tomaron prefloraciones justo antes de la antesis de los espe- címenes respectivos (Vogel 483, de Michel 156, Gentry et al. 10992, Santos et al. 2066, Mori & Kallunki 5379) que reposan en el herbario WIS y fueron enviadas a la Estación de Agricultura Experimental de Kansas (Kansas Agricultural Experiment Station). Éstas prefloraciones fueron suavizadas en KOH al 10 % y acetolizadas de acuerdo con Erdtman (1960). Las muestras de polen fueron dividas en dos grupos para ser estudiadas en el microscopio de luz (ML) y en el microscopio electrónico de barrido (SEM). Para ML, las muestras de polen fueron montadas en un gel de glicerina y examinadas con un microscopio óptico Zeiss Universal de campo brillante. Para SEM, las muestras de polen fueron suspendidas en agua destilada, pipeteadas sobre un portaobjetos y secadas al ambiente. Las muestras fueron rociadas y recubiertas con una película de oro- paladio y examinadas con un microscopio electrónico Eteck Autoscan SEM, en la Estación de Agricultura Experimental de Kansas. Las placas con los respectivos granos de polen y las imágenes SEM reposan en el herbario WIS de la Universidad de Wisconsin, en Madison l. nd X. pua € H.H. Iltis, gen. nov. (Figs. 1, 2, 4). Trus: Capparis speciosa Griseb. 5 impl l glab , folia simple, espiralata, sempervirens, i fl inalia et/vel solitaria et axilaria, flori lyx dialisepalis 2-seriatis, d ti E d nectariae 4- EE T duo EE duo minoribus, poll btili ticulatus, f D ind ^ b E 1 li ar] s JL DN ] , anisocotila, iled joril | bgloboso, to et albo: Árbustos o árboles, glabros o con pubescencia de delicados y frágiles pelos simples. Hojas simples, espirala- das, con peciolos cortos de similar longitud y articulados con las ramas, pulvínulo aparentemente ausente LR Af FAO N F be 1 Anisocapparis X C ornejo & H.H. ic A. R [| £z Y) B. Flor en antesis LI v hn € dem d d 14794, NY). Fic. 2. Anisocapparis X. Cornejo & H.H. Iltis, ivo: | inal i i tesis, escala 1 cm. Foto tomada en Bolivia, Dept. Santa Cruz, Prov. Caballero, 64°28'W 17º59.84'5, 1650 m, 8 Ago 2003, en los especímenes secos. Inflorescencias en racimos pate GE ido flor ada y nd ien distintivamente 2-seriado, decusado-imbricado y anisosépalo desde las pretioraciones Jóvenes, los dos sépalos externos de menos de la mitad del tamaño que ios dos sépalos internos. Een nectarias cuatro, puce las, dimórficas, escuamiformes y carnosas, éstas son una proyección de la capa carnosa del tejido nectario que cubre al receptáculo. Las escamas mayores son transversalmente SCH geg hasta semicirculares, siempre más anchas que altas y varían desde rectas hasta có ia el receptáculo, a manera de una concha. Corola de aestivación torsiva y/o imbricada, MEM 4, sésiles, insertos en el receptáculo. Estambres exsertos, ca. 20-50, insertos al mismo nivel sobre un grueso andróforo carnoso (en vivo), éste es plano y a veces algo expandido en el ápice, a manera de un torus (nectarífero?); los estambres internos son glabros, los estambres externos poseen filamentos abundantemente pubescentes en la base, con tricomas unicelulares, + complanados, + transparentes; anteras basifijas, granos de polen tricolpados, prolados, con una ornamentación finamente reticulada. Fruto un pepo pendular, globoso o subgloboso, con un pericarpo coriáceo, flexible y con una pulpa de color amarillo hasta anaranjado brillante en la madurez. Semillas uno hasta ocho, con testa dura, subleñosa, cubierta por una sarcotesta de abundantes pelos infiltrados, y embriones fuertemente anisocótilos de cotiledones separados, que poseen un cotiledón mayor compacto, subgloboso, de color blanco marfil y no clorofílico, glabro, y un cotiledón menor muy reducido o ausente SPranceschint id Tressens 2004: 214, Dë 6 [Capparis speciosal). ,restringido a Sudamérica en Bolivia, Brasil, Paraguay y Argentina. Éste se et y distingue de codos] los demás gé (Hutchinson 1967) tradicionalmente tratados en Capparis s.l. por presentar: 1) Escamas nectarias dimórficas (vs. nectarios [e.g., Cynophalla], escamas nectarias [Beautempsia] o apéndices nectarios [e.g., Capparicordis], siempre isomórficos); 2) semi- llas x subglobosas (vs. cocleadas o reniformes hasta elípticas); y, 3) embriones con cotiledones separados, Cornejo et al., N South A i fC 65 extremadamente anisocótilos, muy distintos en tamaño y forma entre sí, con un cotiledón mayor compacto, especializado para almacenar (Fig. 1D), y un cotiledón menor muy reducido o ausente, único en las Cap- paraceae del Nuevo Mundo. Éstos han sido discutidos en detalle e ilustrados por Franceschini & Tressens (2004 [ver Capparis speciosa]). Por la apariencia externa de sus frutos y el color blanco marfil de los embriones de sus semillas, Ani- socapparis superficialmente se asemeja a algunos miembros de Capparidastrum Hutch. (e.g., Caparidastrum petiolaris [Kunth] Hutch., comparar con Cornejo & Iltis 2005a: 158, fig. 2C [tratado como Capparis peti- olaris]). Pero además de los tres caracteres anteriormente mencionados, Anisocapparis también se diferencia de Capparidastrum por tener un cáliz notoriamente anisosépalo, con aestivación decusada-imbricada (vs. cáliz isosépalo, frecuentemente 1-seriado) y granos de polen con una ornamentación finamente reticulada (Fig. 4, vs. tectada-espinulosa [Fig. 7] o tectada-perforada). Otras diferencias menores de Anisocapparis en comparación con Capparidastrum son la presencia de láminas foliares de menores dimensiones, 3-8 cm (vs. usualmente 10-50 cm), y una densa Tue en la base de los filamentos de los estambres externos (vs. filamentos glabros), cuya función es desconocida. Bajo el estereoscopio se ha observado que ésta retiene granos de polen, por lo que se presume podría estar relacionada con la polinización. Entre las Capparaceae de América, el tipo de cáliz anisosépalo que presenta Anisocapparis, con los sépalos dimórficos dispuestos en dos series, decusados-imbricados desde la prefloración, cuyo par de sé- aie EE externos es pa menor dimensión en comparación con el par de sépalos internos, es ola Ha (Iltis & Cornejo 2005, tratado como Capparis subg. Cynophalla). Debido a esta similitud, la especie SH E EN (Capparis speciosa) fue originalmente descrita como perteneciente a las “Cynophalla” (Grisebach 1879: 18). Pero Anisocapparis fácilmente se diferencia de Cynophalla por la ausencia de las características glándulas nectarias extraflorales, dispuestas en las axilas de las ramas termi- nales e inflorescencias; también por la diferente estructura y disposición de los nectarios florales (éstos en Cynophalla son siempre E dci hasta un pee cóncavos y dispuestos + horizontalmente sobre el receptáculo), y por el polen de iculada (vs. tectado-espinuloso en Cynophalla). Además, los pepos globosos a AA con semillas subglobosas, D de cotiledones separados con un 1 cotiledón mayor muy desarrollado y compacto, est ma , de color blanco y no clorofílico de Anisocapparis, contrasta grandemente con los frutos capilares, nas, con semillas elipsoideas, isocótilas, de embriones de cotiledones delgados, convolutos, verdes y clorofílicos de Cynophalla (véase en Franceschini & Tressens 2004 [Capparis speciosa vs. C. flexuosa L. y C. retusa]; Iltis & Cornejo 2005). Etimología.—Aniso, el prefijo de su nombre, se refiere a los distintos tamaños de los sépalos, nectarios florales y cotiledones que caracterizan a este nuevo género. Anisocapparis speciosa (Griseb.) X. Cornejo & H.H. Iltis, comb. nov. Basonmo: C Griseb., Abh. Konigl Ges. Wiss. Göttingen 24:18. 1879. Non C iosa Moric., nomen nudum escrito en Blanchet 2564 (NY!), citado en Eichler in Martius, Flora Brasiliensis 13:272. 1865 [a Noir yco Martius). Non Capparis e. Miranda, Anal. Inst. Biol. Mex. 4:77. 1953, nom. illeg. (= M ale mollicella Standley). Capparis speciosa Griseb. var. normalis Kuntze, Rev. Gen. Pl. 3:7. 1898. Tiro: ARGENTINA. Gran Craco: Prov. Salta, Dept. Orán, prope Dragones (Fuerte Sarmiento), 13-18 Ago 1873 (fl), “Naranjillo,” PG. Lorentz 6 G. Hieronymus 601 Seege GOET!, designado en Gó 1953: 298; isóripos: B-10 02427341, K-220554!, NY-967902! [herb. O. Kuntze]). Capparis pruinosa Griseb., Abh. Kónigl. Ges. Wiss. Ud M 24:19. 1879. ile inde var. pruinosa Hassler, Repert. Spec. Nov. Regni Veg. 12:254. 1913. Tiro: ARGENTIN y Santa Rosa, Nov 1873, P G. Lorentz & G. Hieronymus 227 (Lecróriro, aquí designado: GOET!). BCE speciosa Griseb. var. ovata Kuntze, Rev. Gen. Pl. 3:7. 1898. Tiro: BOLIVIA: Sierra de Santa Cruz, 2000 m, sin fecha, O. Kuntze n. (LECTÓTIPO, aquí designado: NY 581261! [herb.O. Kuntze]). Dee speciosa Griseb. var. lanceolata Kuntze, Rev. Gen. Pl. 3:7. 1898. Tiro: BRASIL. Mattogrosso, Corumba, Jul 1892 (fl), O. Kuntze s.n. (LECTÓTIPO, aquí designado: B 10 0242735! e en sue Capparis malmeana Gilg, Engl. Bot. Jahrb. 25 Beibl. 60:2 98. Capt pruinosa f. mal (Gilg) Hassl pert. Spec Nov. Regni Veg. 12:254. 1913. Tipo: PARAGUAY: a. Risso pr Rio Apao. In sil ] “20 Sen 1893 p G. Malme 1000 (LecroTiro, aquí designado: B 10 0242733! [F foto neg. 5807 en NY!, WIS!]; isóripo: 5: S). Fn] 11; ión original de C ; ; Griseh ] t i 5 que esta especi taba basada en E CH LE E ? L r 66 t tani i Texas 2(1) una colección de Lorentz & Hieronymus proveniente de Dragones, donde floreció en Agosto. En el herbario GOET, el Eon O Lorentz & G. Hieronymus 601, es la única colección identificada como “Capparis speciosa Gr. n. sp., proveniente de Dragones y ee con flores en M Capparis adds var a pror plicado del mismo tipo de Capparis speciosa (Lorentz & G. I | ba en el herbario de mias tualmente en NY. Este duplicado fue identificado por antes en letra gianni como cane speciosa Gris. O DOMES e EE ” Esta leyenda concuerda con "folia oblonga,” el carácter propuesto para esta variedad en original por lo que no hay duda de que la var. Reno de | dide ME ocu en el mismo po de Capparis speciosa. En el herbario GOET hay dos c como Ca das como tipos, Lorentz & Hieronymus 227, con fecha Nov 1873 y Lorentz & Hierony 1182, con da 7 Dic 1873. En la publicación original de Capparis pruinosa Grisebach no mencionó colección tipo alguna, solamente “ubi floret Novembri.” Lorentz & Hieronymus 227 está escrita a mano “Capparis pruinosa Gr. n sp,” y su fecha de colección concuerda con la fecha referida en la publicación original, por tanto esta última colección es aquí designada lectótipo. En K, la colección K-220553 tiene dos etiquetas que leen “Capparis pruinosa Gr. Salta” y “Argentine Comm. Dr. Grisebach, Dec. 1878.” Aunque esa es una colección histórica, sin embargo carece de mayor información y no se puede determinar si es un duplicado del lectótipo. Nombres locales —ARGENTINA: Meloncillo (Venturi 9807), naranjo del monte (Gómez 1953), naranjillo (Grisebach 1879, public. orig.), payaguá-naranja, sacha limón (Ruiz et al. 10540), sacha naranja, sacha sandía (Gómez 1953). BOLIVIA: Alcaparro (Nee & Coimbra 35565), coca de chivo (Nee 51204). PARAGUAY: Bola verde (Spichiger & P. Loizeau 1519); i'tsá'jituk [pl: i'tsájituki], 'tso'xuá'nuk [pl: tso'xuá'nkil (T. Gragson 153, Manxuj); Pachyajua naranja (Spichiger & Loizeau 1504). Usos.—Los frutos de Anisocapparis speciosa al madurar caen al suelo y la pulpa en el interior se vuelve muy suave, de color naranja brillante. Éstos se parten a lo largo de las suturas al presionarlos con los dedos, la pulpa es consumida directamente y la piel es desechada. La consistencia de los frutos es como la de los mangos (Mangifera indica L.), y tienen un sabor agridulce, entre mango y naranja (I. Gragson 153). Anisocapparis speciosa es principalmente utilizada para forraje. El ganado come las hojas y frutos de esta especie durante todo el año, aunque está reportado que las vacas se atoran al intentar tragar el fruto (Vargas et al. 2000). Fenología.—Floración nocturna (Vogel 483). Interacciones biológicas. —Las flores de Anisocapparis speciosa son visitadas por por abejas melíferas y grandes avispas negras (Vargas et al. 2000, Nee 51204). Sus frutos son alimento de mamíferos, como los venados Mazama | y M. goauzoubira (Stallings 1984; Rivero et al. 2005). Así como en otras especies de Capparaceae de frutos grandes en América (Iltis et al. 1996), se presume que los frutos de Anisocapparis speciosa en el pasado sirvieron de alimento para una megafauna extinta en la actualidad. E tati ARGENTINA. Prov. Corrientes: Dept. Capital, by the road side ca. 1 Em S of Paso Lessoa, 22 Ago 1951 EM T.M. a 1147 (NY). Prov. Salta: Dept. General Güemes, 5 km NW de F E ta Juana Azurduy (ruta Prov. 52), 25°08'S 61°52'W, 13 Dic 1999 (fl), R. Fortunato, A. D'Agostini, A. Martínez & H. pus 6391 (MO, NY); Pasando G , en el km 1440 de la ruta, 34 m, 20 Nov 1974 (1D, V. Ruiz, R. Roncalia & À. macau 10540 (NY). nis Guachipas: Alemanía, 1300 m, 4 Nov 1929 (£D, S. Venturi 9807 (MO, NY). Dept. Orá Ruta Nac. 81, beside rail e, 20 Nov 1978 (fl, fr), S. R ize 3543 (MO, NY). Prov. Santiago del Estero: Dept. Choya, 12 Se W de Villa la Punta en dirección a Frías, 28°23'S 64°46'W, 257 m, 30 Nov 1995 (ID), R. Fortunato & pon 5153 (NY, US). BOLIVIA. Dept. Chuquisaca: Prov. Luis Calvo, Estancia Acapulco, 300 m, 20°20'S 06"15'W, 5 Ene 1995 (fl), L. Sánchez 55 (USZ, ex Tropicos). Dept. Cochabamba: Puente Arce, 1800 m, 18°35'S 65°10'W, 11 Apr 1963, D. Ugent & M. Cárdenas 4884 S SEN Prov. Narciso Campero, geleed di km Hacia sp 1770 m, 17 Oct Ed (tl, SS Kä E Soe ion (NY, EI WIS). Dept. Sant Andrés Ibáñez, at quebrada, 1 at NN on dirt road to Paurito, 17°50'S 62°57'30"W, 345 m, 5 Ene 1996 (fr), M. Nee 46704 (MO, NY); Prov. E a 5 km hacia Santa Cruz, 1950 m, 28 Sep 1981 (fl), S.G. Beck 7070 (NY, WIS); 11 km (by air) SW of Comarapa on road to Chilón, 17°59'S 64°35'W, 1600 m, 7 Ago 1987 (fl), M. Nee & G. Coimbra 35565 (MO, NY); 5.5 km N of center of Saipina on dirt road to Chilón, valley of Río Ge SE ER E 1405 m, 8 Dic 2005 (fD, M. Nee, D. Villaroel & O. Colque 53664 (NY). 0.5 km SE of turnoff (to Pulquina), | from San ro to o and Mairana, 18°03'3"S 64°25'30"W, 1575 m, 29 Dic 1995 (fl), + M. Nee 46608 (NY). St h der Kal , unweit San Isidro, le Ostkordillere Boliviens, Blüten nocturn, EE bis nh da Filuriemes weiss, starr, — 13 Ene 1965 dus 5: nu 483 (A, F, WIS, Se Prov. rs ein Pozo del Tigre, 135 km E del empresa AB 7°34'S 61°57'W, ca. D Cornejo et al., New South Ameri f Capparaceae 67 250 m, 28 Dic 1994 (fn), S. Ortiz & G. Castro 106 (NY, USZ). Prov. Cordillera, 1 km N of YPFB Planta de Gas at the Río Grande, 18°10'S 62°54'W, 350 m, 3 Dic 1989 (fl, Gen M. Nee 37899 (MO, NY, WIS); 1 a E Se Camiri- M oc road, on dirt road toward Eiti, 19°54.39'S 63°31.59'W, 950 m, 8 Nov 2000 (fl, f j.), M. Nee 51302 (MO, NY); al Aba 7 km NNE of Ipati, 19°38'S 63°33'W, 980 m, 1 2000 (fD M. Nee 51342 (MO, NY);2.5kmEofC idad Salinas f ff oft! iri-Cuevo hwy, 20°13'S 63°26'W, 880 m, 6 Nov 2000 KEN M. Nee 51204 (MO, NY); Alto Parapetí , bosque natural, en depr de lad Om, 8 Ene1982 (fl, fr), R. de Michel 156 (WIS). Prov. Fl S de Mataral d to Vallegrande, 18°09'S 64°13'W, 1400 m, 15 Dic 1990 (fr), M. Nee 40285 (NY, USZ). Y km (by air), 10.2 km (1 y road) NNW of Mataral on road to San Juan del Potrero, 18° 02'45"S 64°14'25"W, 1475 m, 30 Ene 1994 (ir), M. Nee & I. Vargas 44794 (MO, NY, WIS). PARAGUAY. Dept. Boquerón: Mariscal Estigarribia, 22°01'49"S 60°36'52"W, 250 m, 15 Dic 1998 (fl, y fr), E. Zardini & L. Guerrero 49966 (MO, NY); Estancia Copel, 22?12'S 61°54'W, 150 m, 19 Sep 1985 (ID), R. Spichiger & P. Loizeau 1519 (MO, NY). Dept. Presidente Hayes: Estancia Pozo Favorito, 59°53'W 23°7'S, 150 m, 19 Sep 1985 (fl), R. Spichiger & P. Loizeau 1504 (MO, NY); 8 km before 25 Leguas, Trans Chaco hwy SE of Filadelfia, 22°45'S 59°45'W, ca. 200 m, 2 Oct 1985 OI, A. Gentry, L. Pérez & D. Brunner 52003 (MO, NY); Chaco, vic. Manxuj, 21°48'S 61°41'W, 15 Jul 1993, T. Gragson 153 (MO) 2. Monilicarpa X. Cornejo & H.H. Iltis, gen. nov. (Figs. 3, 5, 6). Teus: Capparis tenuisiliqua Jaca. Frutex vel arbores, pubescentia simple vel glabra, folia simple, spiralata, subopposita vel 3-subverticillata, decidua vel sempervirens, inflorescentia racemosa, terminalia, floris cum calyx ed l-seriatis, pes EE erectis, pons) in nectariis cupularibus vel disciformibus insertiis, pollen striato-reticulato, f torulosibus , dehiscentibus, cum semina elipsoidea, 1-linearia disposita, embryo convoluto. Subarbustos, arbustos o arbolillos, mono a multicaules, con pubescencia de pelos simples o glabros. Hojas simples, espiraladas y con peciolos de similar longitud o subopuestas y 3-subverticiladas y subsésiles, pulvínulo presente o ausente. Inflorescencias en racimos terminales, acrópetos; frecuentemente con brác- teas lineares, hasta 13 mm, localizadas en- y cerca de- la base del pedúnculo de la inflorescencia. Cáliz gamosépalo, 1-seriado, sépalos 4, isomórficos, triangulares, en la antesis erectos y fusionados hacia la base, provisto al interior de un pequeño hipantio que contiene un nectario floral cupular o disciforme de bordes libres o fusionado con el hipantio, con o sin 4 apéndices alternipétalos. Corola de aestivación siniestrorsa- torsiva, pétalos 4, fusionados en el borde superior del nectario. Estambres exsertos, ca. 15 a 30, glabros, insertos en dos alternos sobre un corto y ap expandido andróforo, anteras basifijas, granos de polen tricolpados, prolados, con una ornamentación estriada-reticulada. Frutos cápsulas lineares-torulosas a moniliformes, con ápice delgado-atenuado, dehiscentes o Eee MEME esencialmente sin $1] pulpa, con 1 hasta 2232 [según Ruiz-Zapata 2002: 150]) semill psoid puest una hilera, testa presente, embrión verde o EH crema, amO ía Monilicarpa se teriza por f brácteas lineares, hasta de 13 mm, localizadas en y cerca de la base del pedúnculo de las estacas flores con cálices gamosépalos que contienen un pequeño hipantio, nectarios florales cupulares o disciformes, en cuyos bordes están fusionados los pétalos, filamentos de los estambres insertos en dos verticilos alternos, granos de polen poseen una ornamentación estriada-reticulada (Figs. 5, 6, única entre los seis tipos de polen observados entre = EE de hojas simples en el Neotrópico), y frutos lineares-torulosos a moniliformes, de ápices dos, que contienen pequeñas semillas elipsoides dispuestas en una hilera, separadas entr sí por las constricciones de las paredes del fruto. El nectario floral cupular o disciforme de Monilicarpa está localizado alrededor de un androginóforo central y fusionado al interior de- y recubriendo a- un pequeño e inconspicuo hipantio. Debido a la pres- encia de este pequeño panio; el cáliz de Capparis viridiflora Kunth (Humboldt et al. 1821: 92), uno de los sinónimos aquí citados de liqua, originalmente fue descrito como: “turbinato-urceolatus.” El nectario floral de Monilicarpa en sus bordes y entre las bases de los pétalos posee cuatro apéndices erec- tos, dentados o triangulares (Monilicarpa tenuisiliqua) o rudimentarios en forma de inconspicuos lóbulos o ausentes (M. brasiliana), como se aprecia en la lámina de Capparis brasiliana en Flora Brasiliensis (Eichler en Martius 1865: 279, tab. 62, hg 1:3, nótese did superior a ia del cáliz. En contraste, Cappa- ridastrum y Cynophalla, | géneros restantes de las Capparaceae yI p América, tienen flores con cuatro nectarios, apéndices o escamas nectarias y mox insertos sobre un receptáculo usualmente plano. Adicionalmente, Monilicarpa se diferencia de Capparidastrum por tener los filamentos de los estambres insertos en dos verticilos (vs. filamentos de los estambres insertos en varios verticilos), y los 2 cm 0.5 mm L I d.i dien adis f Texas 2(1) 2cm Fic. 3. Monilicarpa X. Cornejo & H.H. Iltis: Monili il (Jacq.) X. Cornejo & H.H litis. A. Diag ] ali | pétalo frontal ido, Fa 1 ža l e D 7 L IL i a A L IL” ye D " | L | tr on Jr “esa NY) R r T LI F T V J i n A J E A D [| E a HII FE pe A L'I ir Sc oda NY £ rm: A A In ES ranh d L] ` r d po] 7 WIS) aa spa i spe £m 3 fa dl y Y Cornelia R, H H tie n mnm " J a "e FI H I E A y Fil Fr J: L'I in tn FADD NY E ms J Lr FA > 3857, WIS) ru [« up put used Corneio et al Nev , WIS). B. escala 1 um (A. de 10 um (Vogel 483 X Cornejo & HH Iltis: A escala Ei Michel 156, WIS). 10 pm. B. escala 1 um (Gentry et al. 10992, ) Y Cornejo & HH tic À escala Fic. 5. Mi WIS). 70 t tani i Texas 2( frutos lineares a ilif de ápices delgado-atenuado, esencialmente sin pulpa, con semillas elipsoides dispuestas en una hilera (vs. lentos lar pepos, oblongos, cilíndricos, ovados u esféricos, con ápices + redondeados, pocas veces agudos, pulpa! abundante hasta escasa y semillas cocleadas, dispuestas en dos ó más hileras). Por sus frutos capsulares lineares-torulosos Monilicarpa se parece a Cynophalla, pero a más de los caracteres genéricos previamente mencionados, Monilicarpa fácilmente se diferencia de este último género por poseer flores con cálices 1-seriados (vs. cálices 2- seriados, decusados-imbricados), y vegetativamente por la ausencia de las glándulas nectarias axilares extraflorales (vs. presentes en Cynophalla) y la diferente filotaxia. Algunos géneros distantemente relacionados a Monilicarpa como Atamisquea Miers, Capparicordis Iltis & Cornejo y Colicodendron Martius, también presentan tazas nectarias o nectarios cupulares, pero éstos fácilmente se diferencian de Monilicarpa, principalmente por presentar indumento de escamas lepídotas o tricomas estrellados, diferente tipo de polen y distintos tipos de frutos capsulares o amfisarcos, globosos hasta oblongos. Frutos lineares-torulosos también están presentes en algunas especies de Quadrella DC.) J. Presl, un género ampliamente distribuido en las Indias Occidentales y América Central hasta el Sur de Beie hacia is norte y norte ee Reno y Venezuela hacia el sur (Iltis & Cornejo 2008). Pero Monilicarpa a de glabra o presentar tricomas He: s tricomas adt Has ] sb heus: BEN DEEG , por lap nrecencia de florece conn cá c la base (vs. flores dialisépalas), hipantio presente Es ausente), nectario cupular o distilomme en cuyos bordes o pared externa se fusionan los pétalos (vs. presencia de cuatro escamas nectarias y pétalos insertos en un receptáculo plano). Etimología.—El nombre Monilicarpa, alude a los delgados frutos moniliformes a linear-torulosos que presentan las especies de este nuevo género. Monilicarpa y Capparidastrum son inermes (Cornejo & Iltis 2005b) y no presentan tallos generalmente espinosos, como ha sido reportado (Inocencio et al. 2006: 122, 125, ver Capparis sect. Capparidastrum y Capparis brasiliana). CLAVE DE LAS ESPECIES DE MONILICARPA le Plantas eodd is EE hojas a espiraladas a lo largo del tallo; nectario ida cupular fusionado verde nilicarpa tenuisiliqua . Plantas sempervirentes, hojas subsésiles, subopuests y/o 3-subverticiladas; nectario floral a e de bordes libres; semillas con embrión crem Monilicarpa brasiliana 2.1. Monilicarpa tenuisiliqua (Jacq. A e Cornejo a FE H. e = nov. (Figs. 3A-C, 5). Basonmo: Capparis tenuisiliqua Jacq., Enum. Pl. Carib. 24. 17 i jue, Tellur. 109. 1838. Uterveria tenuisiliqua Jacq.) Bertol., Nov. Hort. Bonon. 2:8. 1839. Pad ai tenuisiliquum (Jacq.) Sech Genera Flowering Pl 2:310. 1967. Tiro: COLOMBIA: “Carthagenae frequens" (Lecróripo, aquí designado: Jacquin tab. 105 [en Select. Stirp. Amer. 2, 1763]). Capparis obovatifolia Kunth, Nov. Gen Sp. 5:92. 1821. Tiro: VENEZUELA: Sucre, “crescit prope Cumaná. Fructificat Septembri,” A. Humboldt & A. Bonpland 93 (Lecróripo, aquí designado: P!) Capparis viridiflora Kunth, Nov. Gen. Sp. 5:92. 1821. Tiro: VENEZUELA: Sucre, “crescit prope C 4, in umbrosis. Fl Septembri," A. ens Bonpland 569 (Lecróriro, aquí designado: P!) C Turcz., Bull. Soc. Nat. Mosc. 27:327. 1854. Tipo: VENEZUELA: Carabobo, Puerto Cabello, Jun 1841 (fl), H. Funck & L. Schlim 522 (LECTOTIPO, designado: : BM!, LD!, P! iie garciae Dugand, Caldasia 1:45. ba Tio: COLOMBIA: A Mun. Tocaima, Heda. Chucundá, matorrales, 16 Jul 34 (fr), H. ee 3109 (goróriro: COL-28357, foto de COL en WIS!, isóripo: NY!) lr... | € Ja L L a [| . IA” A ( mE solum [Macbr.] X farnaln & H H lle: ) d PS | FA b | HI [| 11 A r1 FILLE od ES "E [| Y L secado en el proceso de herborización. Cornejo et al New South A Í FC 71 Los lectótipos de Capparis tenuisiliqua, C. obovatifolia, C. viridiflora y Colicodendron obovatum son aquí des- ignados debido a que ninguna colección fue referida bajo estas especies en las respectivas publicaciones originales. En el herbario US, H. García-Barriga 3109 (15933661) no constituye un isótipo de Capparis Garciae, ésta procede de una localidad distinta, Purificación, y fue colectada en diferente fecha, 7 Jul 1934. Monilicarpa tenuisiliqua es una especie de hojas deciduas, su follaje se regenera durante la época de floración (Smith 2100). Nombres locales. —COLOMBIA. Maretiro, Nudo, Palo de Agua (Dugand 1941: 44). VENEZUELA. Guari- aro (Aristeguieta 7014), taparito (Galué 167) Usos.—Para curar enfermedades, también para hacer cruces en Semana Santa (L. Aristeguieta 7014). Especímenes representativos. TRINIDAD & TOBAGO: Patos Island, coastal bank, 13 Mar 1920 (fI), N. Britton, T. Hazen & W. Medelson 533 (NY); ibidem, 5 Sep 1931 (fl), R. Williams 12509 (NY). Chacachare, hillside, 3 Apr 1921 (fl), N. Briton, W. Freeman & Sir E Watts 2683 (NY). COLOMBIA. Bolívar: Vic. Cartagena, 1920 (fl), Bro. Heriberto 362 (NY). Magdalena: Santa Marta, 3 mi. N of Bonda, 250 ft., 28 Apr 1898-1899 (ID, H.H. Smith 2100 (NY[2]), 12 Jul 1898-1899 (fr), H.H. Smith 843 (NY); S of Santa Marta, arid coastal belt, 5 Apr 1927 (fl), E. Killip & A. Smith 21082 (NY). Don Jaca, 20 Jan 1930 (st), S. Record 69, 80 (MAD, NY). Parq. Nac. Nat. Tayrona, Ensenada de Neguanje, 9 Sep 1976 (fr), G. Lozano & R. Schnetter 2777 (NY), 21 Sep 1976 (fr), G. Lozano & R. Schnetter 2926 (NY); dry slopes over- looking Caribbean, 11°20'S 74°02'W, 50 m, 23 Ago 1986 (fr), A. Gentry & H. Cuadros 55485 (NY). Supra Honda, Río Seco, Dic 1852 (fr), I. Holton s.n. (NY). VENEZUELA. Anzoátegui: Fundo Lagunita de Flores, ca. caserío Curatiquiche, Barcelona, Mar 1969 (fl), L. Aristeguieta 7014 (NY); Alrededores de Píritu, 18 Ago 1965 (fr), G. Agostini 530 (NY). Atlántico: Entre Leña y Candelaria, 11 Ene 1941 (st), A. Dugand & R. Jaramillo 2787 (COL, NY). Barranquilla, May 1927 (fl), Bro. Elias 167 (NY). Carabobo: near Morón, rd. from Puerto Cabello to San Felipe, near sea level, 25 Apr 1920 (fD), H. Pittier 8809 (NY). Around El Palito, near Puerto Cabello, 0-30 m, 2 Jul 1913 (1), H. Pittier 6427 (NY); ibidem, 1921(fr), H. Pittier 9077 (NY); Puerto Cabello, 28 Sep 1874 (ID, O. Kuntze 1747 (NY[2]); ibidem, 24 Jun 1917 (fl, fr), H. Curran & M. Haman 1168 (NY); Distr. Colina, carr. Matarura- Guaibacoa, 19 Ago 1983 (fr), T. Ruiz & T. Ruiz 4124 (MY, NY), Distr. Federal. Tacagua, May 1953 (f). L. Aristeguieta 1714 (NY). Fila de Machado, al O de Cabo Blanco, 10-100 m, 5 Ene 1964 (st), G. Agostini, M. Farinas & E. Castellanos 123 (NY); around Caracas, near El Portachuelo, 800-1000 m, 4 May 1922 (fl), H. Pittier 10308 (NY). Puerto La Cruz, e 1961 Non E O 4642 (NY). Mun. Vargas, poete Catia La Mar, Escuela nav ix: 10936'N 0740 5 m, 22 Mar 1988 (1D), N Í ^). Falcón: Dist. Silva INal de Chichiriviche, 10º54'N 68"16— 1 W, 1-10 m, 30 Ago 1974 un d mall € B a 110864 (NY); Distr. Colina, carr. Mea See 120 m, 20 Dic1981 (fl, n T. Ruíz & E. Rondón 3641 (MY, NY); Distr. Federación, Caserío los Botalones, ía Churuguara, T. Ruiz 6 F. Rondón 3823 (MY, NY). Guárico: Altagracia de Orituco- -Chaguaramas, Mar 1966 (tD, L. a 6057 (NY[2]). Lara: ds Terepaima, 25 Abr 1968 (fl), G. Ferrari, L. Cárdenas & G. Bunting 313 (MY, v. near Md Comercial de Leones, 30 Apr 1974 (1D, 540 m, A. Gentry, G. Morillo & B. de Morillo 10992 (NY, WIS); Distr. Palavecino, | repaima y Cabudare, 500—900 m, 5, 10 Ago 1970 (fr), J. Steyermark, F. Delascio, G. K. & E. Dunsterville 103633- A(NY). Mérida: DR V. near Estanques, 470 m, 9 Ene 1965 (fr), F. Breteler 4559 (NY). Nueva Esparta: Island of E El Valle, 13 e 1901 id O. Miller & J. Johnston 9 (NY); ibidem, 150 m, 21 Jul/8 Ago 1903 (fl, fr), J. Johnston 6 (NY). Sucre: de Cumaná, I iz y Cumaná, Jul 1964 (fr), L. Aristeguieta 5372 (NY). Tachira: S of la Mulata, near Venezuelan-Colombian border, 72°27'30"W 7°50'30"N, 300 m, 13 Nov 1979 (fr), J. Steyermarh, R. Liesner & A. González 120231 (NY). Zulia: Distr. Urdaneta, hcda. El Zanjón, km 10 entre Porterito y Los Claros, 5 Oct 1982 (st), N. Galué 167 (NY); Distr. Miranda, vía que conduce entre las carr. Lara-Zulia y Coro-Maracaibo, que desvía sobre la Lara-Zulia en km 8 SE del Puente sobre el lago, 4 Mar 1979 (fl), G. Bunting 7183 (NY); Distr. Betijoque, carr. Agua Viva-Valera, en km 4 al S de Motatán, lado derecho de la vía, ca. 400 m, 31 May 1981 (fr), G. Bunting 9938 (NY) 2.2. Monilicarpa brasiliana (Banks ex DC.), X. Comas x E H. Ilis, Po nov. v. (Figs. 3D-E, 6). Basónmo: Capparis brasiliana Banks ex DC., Prodr. 1:249. 1824. C h., Gen. Fl. Pl. 2:310. 1967. Tipo: BRASIL: Brasilia prope Rio de Janeiro, 1768, J. Banks & D. Solander s.n. (LECTOTIPO, aquí do BM!; isóriro: MO-16117209). So brasiliana var. longipes A. Gray, in Wilkes, C., U.S. Explor. Exped. 15:68. 1854. Tiro: BRASIL: Guanabara, Rio de Janeiro, 1 C. Wilkes et al. s.n. (Lecróripo, aquí designado: US- 634131). Pn —: Turcz, Bull. Soc. Nat. Mosc. 36:552. 1863. Tipo: BRASIL: Brasilia, Sellow 700 (B?). Nombres locales.—BRASIL. Feijáo-de-boi (Fuks & Costa e Silva 2000: 6). E tivos. BRASIL. Bahia: | $ 'Bahia-Vale dos Rios, Paraguacu/Jacuipe, ca. 39º05'W 12°32'S, 40-120 m, Sep 1980 (m. Pedro do Cavalo et al. 738 — lambe, Rod. BA-415 M M MM Rn 20,4 km 15?16'1" 40°25'17"W, 3 Nov 2000 (fD, J. Jardim, S. de Santana & F. Juchum 3159 (NY Morro Grande, a 3 km W da saida da cidade na rodovia para Jerónimo Monteiro (BR-482), 9 Dic 1994 (ID, J. Pirani, M. Magenta & A, Cd 3519 (NY, WIS). Rio de Janeiro: Rio de Janeiro, Guanabara, Recreio dos Bandeirantes, Ago 1961 (fr), A. Duarte 6244 (MO, NY); Morro da Gamboa, Cabo Frio, 7 Oct 1968 (fD, D. Sucre 3794 (NY); Entre Barra de S. Joao e Tamoios, 28 Sep 1964 (fI), E. Santos, B. Flaster (1103) & C. Pereira 2066 (WIS [3]. Fc 6 | v H Es GPA (Banks ex DC.) X. Cornejo & H.H. litis: A. escala 10 um. B. escala 1 um (Santos et al. 2066, WIS). ETERNA, TU 10 um. B. escala 1 um (Mori € Kallunki 5379, WIS). Cornejo et al New South A j 73 CLAVE DE LOS GÉNEROS DE CAPPARACEAE DE HOJAS SIMPLES GLABROS O CON PUBESCENCIA SIMPLE EN AMÉRICA ] NI + + fl | | | | | - " | A H J y I | t fl | | ; li tos hori | | | culo, escama nectarias pocas veces rudimentarias o más bien a ausentes; hojas E E Cynophalia selo con peciolos de similar longitud, pulvínulo | ); con em clorofílico, verde Gees (DC) J. Pres] Nactarinc evtrafínralac a + rari lee el) florales carnosos disp! lctme sobre el receptáculo y usualmente redondeados o escamas nectarias erectas o nectarios florales cupulares o disciformes; hojas subo s y de SE Jong itlae a lo ago! de las ramas (a Mente no DoD en s especímenes ae Acció! ys Del + I af F ameno blanco o cr rem amarillo overde (en Monilicarpa t liqua) J i fl i LAPIS a qus. I | Call) - MAICI | 1 l M Se TI los ab eem. mu ui: qul e EE a cae + | ] HII | J l PEN AAA A A m | UE dos ee (en vivo), c d let A il hasta algo asimétricos. 3. Brácteas d e- y/o cerca de- la base del pedúnculo de las inflorescencias, si a Void ms usualmen te < r mente redondeadas, usualmente insertas sobre un receptáculo plano y/o a veces oros d la Dase del inis d pétalos insertos en un receptáculo E pane HE ER los apre L| ew mns fo LI a antesis, ia vie A E anu OLOTO | I Seas toada. fre , te oblong | g con apices + mios pocas veces agudos, con pulpa 3 ( n vivo, que reduce o desintegra al secar) y con semillas cocleadas o reniformes dispuestas en dos ó más hileras cotiledones simétricos Capparidastrum Hutch. Rrarctoaac da ien rarcana Am! Ay’ | ] | : E] ` E n ^ i TO itd» uc O UEETCa uc | I 3 eares, hasta 1 tarios floral | liscifi dentro d ño hi filamentos ER con tación estriada-reticulada; frut ilif lineares-torulosos, ECH ápices delgado- 1.4 T ` ‘ hn e. | E las constricciones de las paredes del futo, cotiledones algo asimétricos (jacknife) Monilicarpa X. Cornejo & HH Iltis J ] ex A recs cd ] ] | £l os os sépalos Elos de menos de la mitad ae mano que los dos sépalos internos; nectarios forales constituidos por cuatro escamas episépalas fuertemente dimórficas (dos menores y dos mayores); gra- nos de E con amame ción finamente reticulada; semillas EE EE dad anisocótilo e d T | | ¡INEA | ROT muy SECH O ausente DC ee X. Cornejo & HH Iltis AGRADECIMIENTOS El autor desea expresar su agradecimiento al Herbario GUAY de la Facultad de Ciencias Naturales de la Universidad de Guayaquil, Ecuador, donde en 1997 se inició el estudio de esta familia para Flora de Ecuador (en prep.), y al Departamento de Botánica de la Universidad de Wisconsin, Madison, donde fue Honor- ary Fellow (Oct 2004—Mar 2006), estudiando las colecciones de Capparaceae en el Herbario WIS, para el tratamiento taxonómico de esta familia de Flora Mesoamericana (manuscrito sometido). Los herbarios que enviaron sus colecciones en calidad de préstamo y/o obsequio son: B, BM, BRIT, CAS, DS, DUKE, F, GH, GUAY, K, MÁ, MEXU, MICH, MO, NY, OXF, P, QCA, QCNE, RSA, SP, TEX, U, UC, US, WASH, XAL. Los herbarios visitados son QCNE, MO, SEL y US. Jochen Heinrich de GOET y los curadores de K concedieron al primer autor las M que ues a los tipos ES E Mark A. Wetter (WIS) diligentemente colaboró con el manejo d | Michael Nee (NY) gentilmente proveyó de una 74 | al A Li | L ] BA lexas 2( fotografía en vivo de Anisocapparis speciosa, publicada en la figura 2 de este artículo. Un revisor anónimo realizó constructivos comentarios del manuscrito. REFERENCIAS Cornejo, X. & H.H. Ins. 2005a. Studies in the Capparaceae XXIII. Capparis coimbrana, a new species from Bolivia. Brittonia 57:155-161. Cornejo, X. & H.H. Wm, 2005b. Studies in the Capparaceae XXVI. Capparis bonifaziana, a new species and Western ecuadorian sister to the mostly amazonian C. macrophylla. Novon 15:393-404 Cornejo, X. & HH. luis. 2006. New combinations in Capparaceae sensu stricto for Flora of Ecuador. Harvard Pap. Bot. 11:17-18. Cornejo, X. & H.H. luris. 2008a. Two new genera of Capparaceae: Sarcotoxi and Mesocapparis stat. nov., and the reinstatement of Neocalyptrocalyx. Harvard Pap. Bot. 13:103-116. Cornejo, X. & H.H. Ins 2008b. New combinations in South American Capparaceae. Harvard Pap. Bot. 13: 117-120. Cornejo, X. & H.H. Iris. 2008c. The reinstatement of Capparidastrum Hutch. Harvard Pap. Bot., manuscrito sometido. DUGAND, A. 1941. El género Capparis en Colombia. Caldasia 2:29-54. Eicher, A.W. 1865. Capparideae. In: Martius, C.EP. von, ed. Flora Brasiliensis 13(1). München. Pp. 237-292. ERDTMAN, G. 1960. The acetolysis method. A revised description. Svensk. Bot. Tidskr. 54:561-564. FRANCESCHINI, M. & S. Tressens. 2004. Morphology of fruits, seeds and embryos of Argentinian Capparis L. (Cappar- aceae). Bot. J. Linnean Soc. 145:209-218 Fuks, R. & M.B. Costa E Stan, 2000. Capparis L. (Brassicaceae Burnett) do estado do Rio de Janeiro. Albertoa 1:1-12. Gomez, S. 1953. Capparidáceas Argentinas. Lilloa 13:279-341. GnisEBACH, A. 1879. Symbolae ad floram argentinam. Zweite Peara nung EE E Planzen, Göttingen. Hatt, J.C., KJ. Systma & H.H. Iris. 2002. Phylogeny of Cay plast sequence data. Amer. J. Bot. 89:1826-1842 HUTCHINSON, J. 1967. The genera of flowering plants (Angiospermae) 2:303-317. Clarendon Press, Oxford. Iris, H.H., L. J. Cumana, R.E. DELGADO & G. AvManp. 1996. Studies in the Capparaceae XVIII A new giant-fruited Cap- paris (C. muco) from eastern Venezuela. Novon 6:375-384. me, H.H. & X. Cornejo. 2005. Studies in the Capparaceae XXII. Capparis sclerophylla, a novelty from arid coastal Peru and Ecuador. Novon 15:429-437, Iris, H.H. & X. Cornejo. 2007. Studies in the Capparaceae XXX. Capparicordis, a new genus from the neotropics. Brittonia 59:245—254. Ims, H.H. & X. Cornejo. 2008. Studies in the Capparaceae XXVII. Synopsis of Quadrella, a Mesoamerican and West Indian genus. Novon (en prensa). Inocencio, C., D. Rivera, C. Ogon, E ALCARAZ & J.A. BARENA. 2006. A systematic revision of Capparis section Capparis (Capparaceae). Ann. Missouri Bot, Gard. 93:122-149. KUNTH, C.S. 1821. Capparideae. In: Humboldt, FW.H.A. von, Bonpland AJ. A. & Kunth, C.S. Nova genera et species plantarum [quarto ed.], 5. Paris. Pp. 82-98. Rivero, K, D. Rumiz & A. Taper. 2005. Diff tial habitat use byt tric brocket deer speci cana and M. gouazoubira) in a seasonal Chiquitano forest of Bolivia Mammalia 69:169-183. Ruiz-ZAPATA, T. 2002. Capparaceae del Parque Nacional Henry Pittier, Venezuela. Ernstia 12:137-172. STALLINGS, J.R. 1984. Notes on feeding habits of Mazama gouazoubira in the Chaco Boreal of Paraguay. Biotropica 16:155-157. VARGAS, |., A. LAWRENCE & ME Otazu. 2000. A es y ar! de Santa Cruz, Bolivia. Guía de campo. o. Editorial FAN, Gees Cruz de E Sierra, Bolivia. TRA V "ds. Ll I I] E E Ir A REVISION OF COLICODENDRON (CAPPARACEAE) Xavier Cornejo Hugh H Itis The New York Botanical Garden Department of Botany 200th St. and Kazimiroff Ave. University of Wisconsin, 430 Lincoln Drive Bronx, New York 10458-5126, U.S.A. Madison, Wisconsin 537 A, jo@nybg.org, joguayagmail SE ABSTRACT The South American g rod: J 1 (C pp ) ; 1:3 1 3 J J T1 EO 1.1 g Togi 1 ip Brazil, and C. scabridum a Seem. from western Ecuador and Perú, pus two new Tp herewith ae 1) Colicodendron bahianum X. Cornejo & H.H. Iltis, restricted f Bahia, eastern Brazil; and, 2) Colicoden- dron valerabellum H.H. Iltis, T. Ruiz & G.S. Bunting, a local endemic f he dry shrubland 1f aio in the State of Trujillo, Venezuela. The new species are illustrated, a map and a key to pecies of Colicodend I ith provi RESUMEN Se valida s. ] jd 1 1 Ta t., gé ear : sq Amóárina dal Suir € "» 1; COS y se discuten sus relaciones con géneros afines. Colicodendron está | tro especies: Las previamente establecidas Colicodendron yco Mart., distribuida al noreste de B Ly C. E anth) Me , localizada en los b v xeroffticos al te de Ecuad de Perú; más d 1) Colicademdon bahianum X. Core & H.H. Iltis, sólo conocida de los bosques humedos del e a Bahía, al este de € Y, 2) Colicodendron valerabellum H.H. Iltis, T. Ruiz € G.S. Bunting, una endémica local e Valera, en el Estado Trujillo, Venezuela. Se ilustran las nuevas especies y se provee de una clave y inita para las ane de PME Key Wonps: Colicodendron, Capparaceae, Brazil, Ecuador, Perú, Venezuela, endemics During our studies of the neotropical species of a s.l. (Capparaceae), carried out to produce a generic realignment of this extremely t became clear that Colicodendron Mart. is a well defined South American genus of stellate dhrubs mid trees, a by having a l-seriate valvate calyx with a dentate or lobed nectary-dish or nectary-cup, and thick-walled, indehiscent amphisarcous fruits. Colicodendron was established by Martius (1839: 25), for two Brazilian species: Colicodendron yco Mart., tl ic type, and C. longifolium Mart. (the latter recently placed in Neocalyptrocalyx, as N. longifolium [Mart] X. mae & H.H. Iltis [2008a]). Subsequently, Seeman (1852: 78) correctly transferred to Colicodendron: Capparis scabrida Kunth, but also the unrelated Capparis avicennifolia Kunth, and Capparis pulcherrima Jacq. (the latter as Colicodendron pulchellum Seem., nom. reject.), as well as the quite unrelated glabrous or simple- haired Capparis subbiloba Kunth (=Cynophalla flexuosa [L.] J. Presl). Two years later, Turczaninow (1854: 321—328) erratically described three species in Colicodendron, all of them synonyms of unrelated species of Capparis s.l., a contribution that obfuscated any clear concept of that genus. Colicodendron was treated as a subgenus of Capparis L. s.l. (Eichler 1865: 272), and later as a section of Capparis (Pax & Hoffmann 1936: 181). In both mentioned infrageneric taxa, the members of Colicodendron were mixed up in a heterogeneous assemblage with the Mexican Capparis angustifolia Kunth (2 Quadrella angustifolia [Kunth] H.H. Iltis & X. Cornejo), and the South American C. crotonoides Kunth and C. tweediana Eichler (both members of Capparicordis H.H. Iltis & X. Cornejo, a recently described stellate neotropical genus, Iltis & Cornejo 2007). Most recently Hutchinson (1967: 309), in his attempted generic realignment of Capparis s.l., made seven additional, but all inappropriate combinations to produce a poorly defined — Colicodendron. In this work, a revision of Colicodendron is presented. The genus is validated and amended, its mor- phological limits are defined, and the relationships with the morphologically closely related genera are discussed. J. Bot. Res. Inst. Texas 2(1): 75 — 93. 2008 76 t tani itute of Texas 2( Colicodendron Mart., Flora 22, Bleibl. 1:25. 1839, emend. X. Cornejo & H.H. Iltis. Tee: Colicodendron yco Mart. Destrugesia Gaudich., Bot. Voy. Bonite, t. 57. 1844-1846. Terr: Destrugesia scabrida Gaudich. Evergreen shrubs or trees, covered by stellate to echinate tric! d hout. Leaves simple, short-petiolate to sessile, opposite to spirally or castes Inflorescences terminal, "T and/or axilar, a panicle, raceme or racemose spike, rarely a solitary axillary flower (in Colicodendron yco); floral bracts filiform or subulate, deciduous. Flowers with calyx 1-seriate, valvate, + cup-shaped, the 4 sepals free, + equal, distinct and entirely enclosing the corolla in bud, their margins and tips touching each other from early bud until near anthesis, at the base fused forming a hypantium, coated by a 4-dentate nectary dish or nectary cup or a 3-lobed nectary cup within. Petals 4, imbricate or torsivus in bud, sessile. Stamens 14 to 60, the filaments geniculate or spirally arranged in bud, at anthesis exserted, inserted on an expanded flat androgynophore, anthers basifixed. Fruits amphisarcous or pseudoamphisarcous (in C. scabridum), woody, thick-walled and multilocular, bearing white- or yellow-embryoned seeds, ca. 1-2 cm, cotyledons ca. 1-1.5 mm wide, convolute, testa crustaceous. Due to the stellate pubescence, 1-seriate valvate calyx with the lobes covering the corolla from early bud until near anthesis, and the presence of a nectary cup or nectary dish, Colicodendron is related to the recently described neotropical genus Capparicordis H.H. Iltis & X. Cornejo. However, Colicodendron differs from the latter by having elliptic, lanceolate, oblong to oblanceolate, pinnately nerved leaf blades (vs. cor- date to reniform or suborbicular and basally subpalmatly-nerved leaf blades); a 4-dentate nectary dish or 3-lobed nectary cup (vs. a nectary cup with four filiform to triangular-filiform appendages); higher number of stamens, 14 to 60 (vs. 4 to 8); and larger indehiscent thick-walled amphisarcous fruits, with an inflexible woody pericarp bearing larger seeds, ca. 1-2 cm (vs. smaller dehiscent fruits, with a soft and flexible, thinly coriaceous pericarp, which splits and falls off at maturity, bearing smaller seeds, ca. 5-7 mm, dispersed by birds, Cornejo pers. obs., 2002). Colicodendron is also related to the South American genus Calanthea (DC.) Miers, as was correctly stated by Martius in the original publication (p. 27, cf. with Capparis pulcherrima Jacq., the type of Calanthea [DC.] Miers). However, Calanthea mainly differs from Colicodendron by the very distinctive calyx with open aestivation, in which the not valvate linear-ligulate sepals are widely spaced from each other, exposing a valvate corolla from the very young bud; and by the seeds with barely folded, very thick cotyledons, 4-7 mm wide (Cornejo & Iltis 2008b). Colicodendron could be also related to the stellate to peltate, Central American-West Indian genus Qua- drella (DC) J. Presl. However, Colicodendron differs from the latter by having flowers with a hypanthium coated by a nectary dish or nectary cup (vs. flowers without hypanthium, but with a flat ptacle and four scales inserted on it in Quadrella); and indehiscent, ellipsoid to globose thick-walled woody amphisarcous fruits with larger seeds (vs. capsular, dehiscent, linear or oblong and few times to ovoid, with seeds distinctively smaller, usually arranged in a single row in Quadrella). Colicodendron was previously known by two widely disjunct endemic species: Colicodendron yco trom eastern Brazil, and C. scabridum (Kunth) Seem., restricted to the dry forests of western Ecuador and adjacent Peru. In this work, two additional South American endemic species, both with whorled subsessile leaves and beautiful inflorescences are added to this genus. 1. Colicodendron bahianum X. Cornejo & H.H. Iltis, sp. nov. (Figs. 1, 2, 7). Tree: BRAZIL. Bana: Antiga rodovia e liga a Estacion Ecologia do Pau — Brasil a Sta. Cruz, 7 km ao NE da estacao, ca. 12 km ao NW de Porto Seguro, 167235 39%8M, Regiao de Mata Higrofila Sul Baiano, ca. 80-100 m, 14 Aug 1979 UM), S.A. Mori, L.A. Mattos Silva & S.A. Euponino 12710 (HOLOTYPE: CEPEC,; isotypes: NY, US, WIS) Colicodendrum species novum cum pubescentia stellata dia ina l-seriati tariu p laril t fructus a reU Epio a E | J Aes "al xecccc RR as E : 1 Tae: 1 SM. f. — faji t petiolatis), infl tii iis (nec paniculatiis), staminis 30 ad 42 (nec 14 ad 20), EE 1.6-2 cm longiis, glabtits (tiet. gy DEE? 3.5—5 cm longiis et stellas pubescentiis). hing shrul lender treelets, 1-6 m x 4 cm dbh; terminal | hes densely stellate-echinate; Spar sely | stipules absent. Leaves subverticillate, grouped in whorls at the end of the terminal branches, sessile or Cornejo and litis, Revision of Colicodendron Colicodendrum bahianum X. Cornejo & H.H. Iltis Det.: Xavier Cornejo (NY), 2007 The ew York Bitanical Garden PLANTS GF BAHIA, BRAZIL dense forest, not common. H ti Cabicha: Est... Ecoógica Pau Grant 14 km MW ql gid i > eveiche rain forest, att. 23 S8, 39.15 Y Sparsely branching serub p ; hign; fls. clear yellow ous, visic ad Zei hermit naming bird. No. 25010 Y July 1984 Cottecies Dy SEI kL. Webster Oelerrsinead b Distnibutert E the Botany Department Ha der. E B. os sois Mais — ty Kabara Em 1 Fl: d I L Le WE Cornein& HH lltic A T H IL L LP | L | A1 AA a ya a | R A £l | A AL d (A, B. G. Webster 25010, DAV). 78 t tani i Texas 2(1) subsessile, with blades narrowly oblanceolate to narrowly elliptic, 13—40 x 3-7(-10) cm, stiff, coriaceous (alive), chartaceous (dry), attenuate and revolute at the base, acute and apiculate to sometimes shortly rostrate at the apex, shiny green (alive) with an impressed to often sulcate midvein, nerved and glabrous above, paler green (alive) and with a strongly prominent glabrous or echinate-stellate midvein beneath, margin inconspicuosly thickened, with 15 to 25 major upward-arching lateral nerves on each side of the midvein. Inflorescences terminal and subterminal racemes, sometimes cauliflorous in older wood, densely stellate-echinate; axis short, 1-5 cm, stout, bearing to 25 flowers at the same time. Floral bracts linear, 3—7 mm, deciduous, densely echinate-stellate. Pedicels 1.8—4 cm, articulate to the axis. Sepals oblong-elliptic or ellipsoid to lanceolate, 12-22 x 6-8 mm, abaxially keeled, mainly toward the base, + acute at apex, yellow (alive), rusty brown (when dry), stellate-echinate abaxially; hypanthium 4—6 mm deep, coated by a subtet- ragonal 4-dentate nectary dish within, nectary appendages broadly deltoid to semiorbicular, 1.5 x 2.5-4 mm, glandular, stellate. Petals torsivae in bud, apparently inserted at the SE e the nectary cup, oblong to obovate-oblong, 20-28 x 7-12 mm, widely obtuse to rounded at apex, longi lly nerved, sulfur yellow and erect at anthesis (Mori, pers. com.), dull rusty-brown (when dry), stellate on both sides. Stamens 30 to 42, filaments 2-3 cm, geniculate in bud, glabrous, inserted on a white-stellate-tomentose expanded torus- like androgynophore, anthers ca. 2 mm, white (alive). Gynophore 1.6-2 cm, glabrous; ovary oblong, 5-6 x 2-3 mm, longitudinally sulcate, glabrous; stigma sessile, capitate. Infructescences with gynophores ca. 2-2.5 x 0.3-0.5 cm, pedicels ca. 3-3.5 cm. Amphisarcous oblong-ellipsoid, 7-12 x 2-4 cm, subpendulous; fruit wall 3—7 mm thick, mature seeds 1.3-1.7 mm in diam. Colicodendrum bahianum is somewhat related to the equally yellow-flowered C. yco, is allopatric in the dry Caatinga to the north (Fig. 7). However, the more mesophytic C. bahianum can be easily distinguished by dense whorls of subsessile leaves, arranged toward the end of the terminal branches (Fig. 1A, vs. sub- opposite and pedicellate leaves, more spaced out along the branches). The inflorescences of Colicodendrum bahianum are short dense racemes (vs. larger and longer, paniculate inflorescences, or rarely solitary axillary flowers in C. yco), bearing flowers with erect (vs. reflexed) sepals at anthesis, higher number of stamens (30 to 42 vs. 14 to 20 in C. yco), smaller anthers (ca. 2 mm vs. 4 mm), shorter glabrous gynophores (1.6-2 cm vs. 3.5—5 cm, pubescent) and glabrous (vs. DUE ovaries. Due to the whorled subsessile leaves, Coli to the herein also described Colicodendron valerabellum H.H. Iltis, T. Ruiz & C 3x Sura (for differences see comment under the latter species). Phenology.—Collected in flower in February, July, August, November and January, and fruit in August. The sulfur yellow flowers are visited by hermit hummingbirds (Webster 25010). Cytology.—Unknown. Distribution —Eastern Brazil, restricted to the southern extension of the moist or wet evergreen Atlantic rain forests (ca. 2000 mm) into southeastern Bahia State (Fig. 7). Conservation concern.—Habitat is under intense human pressure from both agriculture and wood har- vesting, especially for so decorative species Colicodendrum bahianum (Scott Mori, per. comm., Sep 1979; “It was one of the most beautiful species that I’ve ever seen”). Images of this potential ornamental species are available at the New York Botanical Garden Web site (NYBG, 2003). PARATYPES. BRAZIL. Bahia: Mun. Santa Cruz Cabrália, mata costeira, 15 Jul 1966, R.P. Belem & R.S. Pinheiro 2580 (F, UB, WIS). Est. Ecológica Pau Brazil, 14 km NW of Porto Seguro, evergreen rainforest, 39°15'W 16°23'S, ca. 80-100 m, 20 Jul 1984, G.L. Webster 25010 (CEPEC, DAV, WIS). Porto Seguro, Parq. Naca. Monte Pascoal, 15 Jan 1973 (f1), T.S. Santos 2692 (CEPEC, NY, WIS); ibidem, trail to peak, upper slopes and top of Monte Pascoal, 40°34'29"W 15°15'53"S, 250—536 m, 14 Nov 1996, W. Thomas et al. 11256 (NY). Estrada a Santa Cruz, Itaju do Colônia, 15 Jan 1971 (fl), T.S. Santos 1329 (CEPEC, WIS). Guaratinga, | lant de Cacau, 10 Aug 1966, R.P. Belem & R.S. Pinheiro 2749 (UB, WIS). nr n Fazenda Independencia, Rod. Itaimbe, Potiraguá, ca. 15 km a partir da BR 101, 15?42'12"S 39°34'31"W, 6 Feb 2004, P. Fiaschi et al. 2283 (CEPEC, NY [NY photo in WIS]). Pastaria, Itaimbe — Potiraguá, 10 Nov 1967, R.S. Pinheiro 414 & T.S. Santos 77 (CEPEC, UB, WIS). Poir mb o km. 10, 29 Jan 1972, a Pinheiro 1801 [1802] (CEPEC, WIS); km 10 da rod. bé; 24 Jul 1973, R.S. Pin! (CEPEC, WIS). E E bela, + km 15, 28 Nov 1970, E. de Mello Filho 2988 & M. Emmerich 3526 (CEPEC [CEPEC photo and [im at WIS). “Mun. Itabello, Bahia" [there is no such place, perhaps a mistake I 1 È 11 more closely related Cornejo and Iltis, Revision of Colicodendron 79 for Itabun C } the CEPEC herbarij is | ted], Cascalheira, mata pluvial, 13 Aug 1995 (fl, fr), C. Hafschibach M. Hatschbach & J.T. Motta 63068 8 (US [US photo at WISI). 2. Colicodend bridum (Kunth) S ot. Voy. Herald 78. 1852. (Fig. WO Keier doc: i scabrida Kunth, Nov Gen. Sp. 5:95. 1821. EECH Rafin.. Sylva Tellur. 112. 1838. Dest ] er e Been geg De 57. s Ge scabridum Dueb Hutch, Gen FI PI. 2:309 I9 hom. illeg. Tyee: i, without data, Hun i (LECTOTYPE, d P [microfiche at NY!]; soLecroTYPE: B-W 10055-01 0! [B-W photo 9478 at WISI. , nomen , Voyage Capparis angulata Ruiz & Pav. ex DC., Prodr. 1:253. 1824. Colicodend | (Ruiz & Pav.) Hutch., Gen. Fl. Pl. 2:309. 1967. Capparis canina Tafalla, nomen. Tree: E Guayas, Bie 1799 (fb, “Zapote de perro," Ruiz s.n. (=Tafalla) (Lecroryre, designated photo 29269 at WIS! 10 0242738! [B photo 342 at WISI], G! [G fragm. at WIS!]). car gaudichaudiana Eichl. in Mart, FL. Bras. 13:273. 1865. Coli (Eichl.) Hutch., Gen. Fl. Pl. 2: 309. 1967. U: Piura, Paita, Voyage de la Bonite, Jul 1836 (fl, fr), M. (audithaud n. eege designated Boe B 10 0242739; IsoLEC- TOTYPES: B 10 0242737! [“misit 1841"]; F 8765471, F 8941681, F [fragm.] 6091301, G [G photo 8472 at WISI], PI). Nomenclatural note. —The herein selected lectotype of Cap; brida housed at P, has a label with a correct origin of the specimen, *in America equatoriale," and the number *19? added. The B isolectotype of Cap- paris scabrida consist of a branch with a terminal inflorescence, with the two leaves, one inflorescence, and floral pieces detached and remounted. That isolectotype, originally only identified as Capparis, was named Capparis speciosa (a Willdenow's nomen nudum, non Capparis speciosa Griseb. [1879], = Anisocapparis speciosa [Griseb.] X. Cornejo & H.H. Iltis), as reads the label which is at the upper left corner of the photo 9478, now in a separate sheet (B-W 10055 -00 O). That label has the following handwritten information: "Polyandria Monoginia Capparis speciosa racemi(?) terminali, foliis ovato-lanceolatis obtusis mucronatis supra nitidis glabris subtus tomentosis foliola elliptica. Habitat in America meridionali." The discussed isolectotype later was correctly identified by Eichler himself, in a handwritten label of determination, which reads: “Capparis (Colicodedron) scabrida H.B.K. Multi-branched shrubs to trees, to 10 m and 25 cm dbh, erect to low and widely spreading; the stem when injured produces a gum (Little 6725); terminal branches complanate or + tetragonal, densely stellate throughout. Leaves spirally-alternate, few times opposite-decussate, petioles 8-30 mm, lacking pulvini; blade coriaceous (rigidly coriaceous when dry), lanceolate to oblong, (6-)8-23 x 2-7 cm, acute to rounded, sometimes emarginate, apiculate at apex, cuneate to rounded, sometimes retuse at base, olive green (alive), but drying to a bright sulfur-yellow in herbarium material, thinly stellate-tomentose when young, soon glabrescent and smooth or scabrous above, pale grayish or “ashen” greenish-yellow and densely pale stellate-tomentose with a prominent midvein beneath; (7312-23 lateral nerves on each side of the midvein. Inflorescences simple, densely corymbose racemes, solitary and terminal or lateral in the axils of leaves, or branched, terminal racemes compounded into complex, stout, (sub)erect, corymbose panicles, to 30 cm, densely stellate, each raceme with only 1-4 open flowers at its end at one time. Floral bracts filiform, 5-12 mm, soon deciduous. Pedicels 2.5—4 cm, densely brown stellate and aparey as MR dark brown 1 , in perfect fl gynophore to greenish-brown, densely stellate. Fl off-centered within the calyx cup, arching out n upward. Sepal lobes broadly ovate to ancla 10-15 x 3-8 mm, at anthesis cucullate-ascending, acute at the apex, densely brown stellate on both sides, eventually caducous, broadly inserted on the rim of a wide bowl-shaped calyx cup, 9-13 x 6-8 mm, with a 3-lobed nectary cup coating the hypanthium within. Petals narrowly to broadly ovate, oblong or elliptic, 15-23 x 10-15 mm, reflexed and outrolled at the tip at anthesis, widely cuneate to truncate at the sessile base, obtuse to rounded at the apex, white to cream or yellowish, sometimes greenish, stellate on both sides. Stamens 24 to 35, borne in a ring on top of the short 2-3 mm androgynophore, filaments (3.525—7(-8) cm, stellate and dendroid, adnate to each other for 0.5-2 mm at the base; anthers 3-4 mm, white (alive). Gynophore 5-8 cm, creamish- Wille, Ra to purple (alive), sparsely stellate; ovary ovoid to ellipsoid, 4-6 x 3-5 mm, densely stellat te, sessile. Infructescences with gynophores 7-10 x 0.4-0.6 cm, pedicels 2.5—3.5(-4) cm. Pseudo T oblongoid, ovoid, or ellipsoid, slightly asymmetrical, 8-15 x 5-9 80 J | tanical Insti Texas 2(1) cm, pendulous, + umbonate at the apex, densely stellate, when mature exhibiting 8 longitudinal + sulcate (dried) lines (usually 4 major, indicating the valves); fruit wall fibrous, 6-10 mm thick, pulp bright orange at maturity, insipid; seeds 15 to 60, subspherical-reniform, often strongly beveled, ca. 1-2 x 1-1.5 cm, sur- rounded by a densely orange testa infiltrated by many hairs from the crustaceous reddish-brown testa; embryo yellow. The amphisarcum is a simple, indehiscent fruit, provided with a thick, woody to subwoody fruit-wall, which retains the shape upon removal of sarcocarp, and never splits in valves after maturity (Spjut 1994: 23. 37). In Colicodendron scabridum, the fruits at maturity fall on the ground to eventually decompose and release the seeds, leaving usually four thickly-subwoody, elliptical, abaxially convex, unattached, persistent valves. That fruit type of Colicodendron scabridum, which doesn't fit in any of the fruit types known (Spjut 1994), is herein proposed as Pseudoamphisarcum (pseudo + amphisarcum) X. Cornejo (mod. nov, which means a false amphisarcum. Local names.—ECUADOR: Sapote (Spanish, Little 6725), sapote de campo (Spanish, Van den Eynden et al. 1999: 42), sapote de perro (Spanish, Cornejo & Bonifaz 7583), sapote gomoso (Spanish, Steyermark 54842); Zapotillo (Spanish, Brandbyge 42789); Zapote de perro (Spanish, Acosta-Solis 8521, Mille 1940, Madsen 64103). PERU: Sapote (Spanish, Díaz & Baldeón 2374), sapote de perro (Spanish, Woytkowski 5672). Zapote (Spanish, Vargas 42), Zapotillo (Spanish, Simpson & Schunke 567). The bright orange color of the mature fruit pulp of Colicodendron scabridum strongly resembles the color of the pulp of a locally (in Ecuador) well known edible fruit called “Sapote” (Quararibea cordata |Bonpl.] Vischer, Bombacaceae). This is the origin for the several similar and derived local names of Colicodendron scabridum. “Sapote de perro,” one of the local names, refers to Pseudalopex sechurae Thomas. This is an endemic canid locally known as “Perro de monte" (Spanish) or *Sechura fox” (English), that has a distributional pat- tern similar to that of Colicodendron scabridum in the dry coastal areas of Ecuador and Peru. When the food is really scarce, the Pseudalopex sechurae facultatively eats the fruits of Colicodendron scabridum (Bruning 1985; Cornejo 2005). Zapotillo (Spanish), a town located in a dry forest at the Province of Loja in Southwestern Ecuador, owes its name to the local name of the locally well known Colicodendron scabridum. Phenology.—Flowering mostly between February to October, fruiting mostly September to December. A single tree 4-6 m high, produces 20 to 60 flowers at anthesis per night. Within the nectary cup, three droplets of nectar start to be produced at 19:00 h., each droplet from below of each nectary lobe. By 19:30 h, the nectary cup is full of nectar. But by the next morning at 6:00 h, the flowers don't have nectar anymore, perhaps due to the intense bat activity during the night. The nectar is secreted during one night per flower only. The flowers live one night only and don't produce any scent (Cornejo field obs. in Bahía de Caráquez, Ecuador, July 2004). Cytology.—n = 8 [!Pazy, icu: Manabi: Iltis Ena e ee "E Iltis unpublished ms). Distribution.—Colicodend nent of dry to very dry or xerophytic areas of western Ecuador and Perú, where is a dune former. This species ranges from sea level in Ecuador from the Province of Manabí, to ca. 2500 m in the mountains of the Province Loja, and in Perú south to Dept. Ancash (Fig. 7). Ecological interactions and field observations.—The flowers of Colicodendron scabridum are visited by the common domestic honey bee, Apis mellifera L. Dozens of buzzing bees can be heard from under a tree when the nocturnal flowers are just about to open. The honey bee's activity peaks in and around the flowers of Colicodendron scabridum is between 18:30 to 19:30 h. The domestic bees re-start re-visiting the flowers the next morning, as well between 6:00 to 8:00 h. Nocturnal, presumably pollen predator bees such as Halictidae (Megalopta?), also visit the flowers at 19:00 h. While both species of bees touch the anthers and the ovaries, it seems however that the true pollinators are bats. From 20:00 to 5:50 h, many small bats frequent the flowers of Colicodendron scabridum, which are exposed on long peduncles high above the foliage. The bats beat their wings hovering in the air while licking the nectar from each flower during less than one second, ne Cornejo and litis, Revision of Colicodendron 81 as hummingbirds are apt to do. The bats do not hold themselves from the flowers as happens to the African Bignoniaceae, Kigelia africana [Lam.] Benth. (seen in Bahía de Caráquez, Ecuador, the same locality where Colicodendron scabridum was studied by the senior author). When leaving the flower, the bat shakes the inflo- rescence, presumably helping pollination. It is interesting to note that only the individuals of Colicodendron scabridum growing in total darkness are visited by bats. These conspicuously have a higher fruit productiv- ity per tree than the remaining individuals located around the nocturnal lights along the roadsides in the same locality. Additionally, the flowers of Colicodendron scabridum are also visited by Amazilia amazilia, a hummingbird who visits the old (previous night's) flowers between 7:00 to 8:00 h, and the new flowers just about to open at ca. 18:00 h. Finally, the flowers of Colicodendron scabridum are ly visited by wasps, coleoptera and several species of ants (Cornejo field obs. in Bahía de Caráquez, Ecuador, July 2004). Uses.—The fruits have been reported as edible for humans and as medicinal against colds and coughs. They are commonly eaten by cattle and donkeys (C. Cerón et al. 22424, Van den Eynden et al. 1999: 42, Cornejo field obs.), while the wood is used for fuel and to make handicrafts (Cornejo & Bonifaz 7583, Van den Eynden et al. 1999: 42). Finally, a "glue" for paper can be obtained from the cut, gum-oozing ends of the stems (Cerón 11876). Conservation biology.—A characteristic common species in the dry landscape, it does not appear to be in danger of extinction. — Specimens studied. ECUADOR. na a de EE 6 Jun 1955, E. Asplund 16569 (S); 10 m, Apr 1944, M. Acosta-Solis 8521 (F); km 8 carr. Bahía-Tosagua, Uni seco tropical, intervenido, 80°32'W 00°45'S, 30 m, 31 Oct 1997, L d IN 1 X. Cornejo & C. Bonifaz 5852 (AAU, GB, GUAY, WIS [ ] DNA]; 20 a Chone, i ical, 80°25'W 00?38'S, 100 m, 4 Jun 1989, C. Cerón et al. 6174 (MO, QAP, QCNE, WIS); Rene a la Isla Corazón, 80°22'W 0094055, ca. 190 m, 18 Feb 1994, X. Cornejo & C. Bonifaz 1739 (GUAY). 1 km NW of San Vicente, ca. 10 m, 9-12 Jul 1977, H.H. Itis & M. Iltis E-228 Ke IS); Cantón Sucre, 21-22 km S of Leonidas Plaza, 80°25'W 0?46'S, ca. 100 m, 16 Sep 1993, G. Webster et al. 30666 (DAV, QCNE); À Playa, deciduous thorn scrub vegetation, ca. 10 m, 9 Jul 1977, H.H. Iltis & M. fltis E-182 (WIS). 10 km SE of Bahía de Caráquez, at km 11-14 along road to Chone near the highest point, ca. 150 m, 8 Jul 1977, H.H. Itis & M. Itis E-243 B, SC Ms dei n El Recreo, ca. Ka m, di Eggers 15528 E PR). Entre La Salina y Chone, 26 Jul 1945, M. Acosta-Solís 10638 (F); town, common on dry tal hills, 7 Aug 1980, B. Hansen et al. 7980 (AAU, RSA, SEL, USF, WIS); EE Ee road, ca. 1 km S of La Pila, grazed an: forest, ca. 200 m, 4 May 1985, G. Harling & L. Andersson 24821 (GB, QCA, WIS); Jipijapa, 80°35'W 1°21'S, 30 m, 25 Mar 1981, B. Sparre 20020 (GH, S); 10 km N e Ree 180 m, 29 Dec 1961, C. Dodson & Thien 1786 (MO, WIS); 11 km W of Puebloviejo, 22 km E of Manta, ext ly drv scrub wi , ca. 100 m, 28 Oct 1974, A. Gentry et al. 12203 (MO, WIS); 14 km SW of Manta on new road to San Lorenzo, 80°47! W 1*01'S, 300 m, 2 Jun 1997, D. Neill & M. Asanza 10706 (MO, Se aba between Puerto s ge ege 80º48'W 1°32'S, 70 m, 29 Sep 1984, J. Brandbyge 42789 (AAU, MO, NY, QCA); | Machalilla, Nort , monte espinoso Tropical, 80°48'W 1°31'S, 70 m, 3 Apr 1994, X. Cornejo & C. Boni- faz 2326 (GUAY, WIS); comuna Agua Blanca, 80942" W 1º35'5, 30 m, 7 Jan 1994, X. Cornejo & C. Bonifaz 1147 (GUAY, QCNE): ibidem, 350 m, 21 May 1995, X. Cornejo & C. Bonifaz 3900 (GUAY); 80°44'W 1°31'S, 125 m, 2 Aug 1990, C. Cerón et al. 11671 O E e 25 um ici e Dee a pura de Las Sillas, C. Cerón et al. 22424 (QAP); Joa hacia Mero Seco, d cint 80^41"W 1°25'S, ca. 420 m, 6 Sep 1991, C. Josse 638 (AAU, QCA); camino a la Playa de Los Frailes, ca. 25 m, 29 Jul 1992, M. (Env et al. 966 (NY, QCA, QCNE Din mee Los dui ca. 2 m, 23 jan 1991, A. Nude 6 S Josse 72703 (MO, QCNE); Río Blanco—Vuelta Larga, 80°47'W 01°29'S, 150 m, Sep 19 Parque Nacional Machalilla, 100 m, 25 Nov 1990, P. Mena & M. Garcia 87 (NY, SE WIS); Estero Seco, bosque muy seco tropical, 0°37'W 01°39'5, 150 m, 29 Nov 1993, X. Cornejo & C. Bonifaz 924 (GUAY); without locality, 80°32'W 0°04'S 200 m, 9 Jul 1978, G. Webster 2262 (MO). Guayas: Between Daule and Lomas, 9 Oct 1952, F. Fagerlind & G. Wibom 501 (LD, MO, ER Gei Guayaquil,” Mar 1865, R. Spruce 4403 (K); Ad Bahía et prope Guayaquil, Apr 1942, L. Mille 1940 (US); Guayaquil—Playas, Feb 1965, G. Sánchez 1 (GUAY, WIS); Casas viejas, 22 km W of Guayaquil on road to Salinas, ca. 60 m, 25 Sep 1981, C. Dodson & P. Dodson 11508 (SEL); El Azúcar, bosque muy seco tropical, 10 m, 19 Jul 2002, X. Cornejo & C. Bonifaz 7583 (AAU, GUAY, WIS); between Progreso and Playas, ca. 30 m, 27 Oct 1958, G. Harling 3096 (S); 10 km W of Progreso, ca. 20 m, 29 Oct 1958, G. nies 3100 (5); along road between Progreso and Baños San Vicente, 16 Apr 1941, H. K. Svenson 11501 (BKL); al ta Elena, 50 m, 13 Mar 1982, C. Dodson & P. Dodson 12943 (QCNE, SEL. WIS); Cerca a Playa Rosada ca. 1 m, Jul 1993, X. Cornejo & C. Bonifaz 275 (GUAY, WIS); Chanduy, “in litore Maris Pacifi,” R. Spruce 6403 (BM, CGE, Kj); E of m 20 Sep 1952, F. Fagerlind & G. Wibom 220 (S); Muey, May 1978, F. Valver- de 23 (GUAY, MO, SEL); Island of Puná, 24 Aug-8 Sep 1836 [voyage of H.M.S. Sulphur], G. W. Barclay 422 (BM, NY, WIS); Aug 1847, B. C. Seeman 915 (K); Kee Puná,” Oct 1892, H. eg 14/35 (GH, K, LE, M, US); igen 1852, muna Ch Isla Puná, path from Puná Vieja to C E º08'W 02°49'S, ca. 50 m, 4 Nov 1987, J. Madsen 64103 Yanzün, 80°03'W 02?44'S, 50 m, 2 Ost 1987, 1 Madsen 63987 SES QCA, OCNE). El Oro: Papa San coa Hcda. Montecarlo, 22 Mar 1967, C. Pineda 89 (GUAY, MO, S); entre Arenillas y Puerto a Dec 1978, L. Albert de Escobar 913 (QCA, WIS); camino entre Puerto Pitahaya y Arenillas, 15 Apr-15 May 1979, L. All 1248 (NY, QCA, WIS); at Arenillas, 6 km S, 25 Jun 1943, E. Little 6725 82 t tani i Texas 2( (NY, US); 2 km S de Chacras cerca de la entrada a Balsalito en la carretera de la Reserva Militar de Arenillas, suelo Paleustalf, rojo con profundidad variable, 80°14'W, 3°33'S, 200 m, 16 Feb 1997, H. Vargas et al. 1158 (WIS); ca. 1 km N of Huaquillas, 1 May 1980, G. Harling & L. Andersson 18813 (GB, WIS); Huaquillas, dry thorn scrub forest, 80°13'W 3°27'S, 5 m, 7 Apr 1980, L. Holm-Nielsen 22842 (AAU). Loja: Cantón Saraguro, caserío Chayasapa, 79°12'W 4°17'S, 2500 m, 5 Sep 1990, C. Cerón 11876 (MO, QAP, QCNE, WIS); 10 km N El Empalme on Macará- Celica road, 800 m, 12 Feb 1991, Kessler 2463 (AAU); Rd. Macará-Loja, 16 km S of Catamayo, 79°23"W 4º05'S, 1400 m, 13 Feb 1987, J. Bohlin et al. 1331 (GB, QCA); between Loja and Portovelo, 3-6 Oct 1918, J. Rose et al. 23327 (NY, US); El Saucillo, Las Casas Viejas, 80°11'20"W 4°15'40"S, 360 m, 15 Feb 1996, Van den Eynden et al. 615 (LOJA, QCNE); above Catamayo to- wards Loja, very dry slope, ca. 1650 m, 9 Oct 1955, E. Asplund 18077 (B, K, LD, NY, 5); between El Tambo and La Toma, 1000-2200 m, 3 Sep 1923, A. Hitchcock 21330 (GH, NY, US); carr. La Toma-El Tambo, te espinoso premontano, 79°25'W 4°02'S, 1500 m, 26 Dec 1991 (fr), D. Rubio et al. 2287 (MO, QCNE, WIS); between Gonzanamá y La Toma, 1400 m, 21-2 3 Jul 1959, G. Harling 6065 (5); Catamayo, 1 km SE de La Toma, 79°22'W 4°00'S, 1500 m, 6 Sep 1994, E. Little et al. 252 (NY, QCNE, WIS); Cerca a Catamayo, 79°21'W 4°00'S, 1238 m, 22 Mar 1994, X. Cornejo & C. Bonifaz 2148 (AAU, GUAY, QCNE, WIS); Rd. Catamayo-La Toma, ca. 1500 m, 24 Apr 1980, G. Harling & L. Andersson 18600 (AAU, GB); dry rocky desert hills above La Toma, ca. 1675 m, 24 Oct 1943, J. Steyermark 54842 (GH, NY); Catamayo valley, Lehmann 4939 (F, K, LE); along the Río Guayaba, 4800 ft, 8 Oct 1944, W. H. Camp E-651 (NY, WIS[2]); Río Guayabas, W of Catamayo (La Toma), 1300 m, 10 Feb 1945, F. Fosberg & M. Giler 22933, 22934 (WIS); E side of Catamayo valley, of Catamayo, 1690 m, 10 Feb 1945, F. Fosberg & M. Giler 22927 (WIS), near la Toma, 79°04'30"W 4°00'00"S, 1300 m, 20 Oct 1965, D. Knight 439 (LE, WIS); ibidem 14 Feb 1965, D. Knight 609 (WIS); La Toma, 1400 m, R. Espinoza 537 (F, NY); SW Catamayo, valley along Río Catamayo, 79°21'W 3°59'S, 1355 m, 10 Jul 1989, L. Dorr & I. Valdespino 6584 (NY, QCA, QCNE, WIS); La Toma-El Tambo, km 5-11 s e roads), vic. Río Catamayo, dry rocks with cacti, 79°21'W 04°03'S, 1300-1500 m, 15 Oct 2000, J. Madsen & J. Gálvez 7441 NY); Rd. San Pedro de La Bendita-Catamayo, km 6.5 (4 km from Catamayo), 79°23'W 3°57'S, ca. 1325 m, 16 Jul 1996, G. P. Lewis et al. 2451 (K, LOJA, QCNE); Cerro Villonaco, 1800 m, 13 Feb 1985, G. Harling 6» L. Andersson 22023 (GB, NY, QCA, WIS). PERU. Tumbes: Prov. Tumbes, Dtto. Pampas del Hospital, Huapalas, 100m, 26 Jun 1964, J. Vargas 42 (NY); Dtto. San Juan de la Virgen, along Quebrada Las Peñas, betw. San Juan de la Virgen and Matapalo, 6 Jan 1968, D. Simpson & J. Schunke 567 (F, NY). Piura: Prov. Piura, Piura, 1928, O. Haught F-69 (F); Prov. Paita, Paita, D'Urville (Lesson?) s.n., 1825 (B [2]; 25 Jun 1914, Mr. & Mrs. J.N. Rose 18513 (NY); ibidem, 25 Jul 1901, R. Williams 2927, 2928 (NY); Prov. Sullana, Sullana, 260 m, 16 Apr 1987, C. Díaz & S. Baldeón 2374 (MO). Lam- bayeque: Lambayeque, 1150 m, 8 Jan 1964, P. Hutchinson & J. Wright 3518 (MO); Lambayeque, km 698 betw. Lima and Sullana, chief dune former in desert, sea level, 5 Oct 1959, B. Maguire & C. Maguire 44389 (F, MO, NY). Cajamarca: Prov. Celendín, Balsas-Celendín rd., 1-5 km from Balzas, 78°04W 6°50'S, 910-1160 m, desert slopes, Rio Marañon Valley, 23 Feb 1984, D. Smith 6145 (MO, NY); Río Marañón Valley, Celendin-Balsas rd. 25 km and onward from Celendín, 78°05'W 06515, 1000-2500 m, 16 Jul 1983, D. Smith & I. Sánchez 4330 (MO, NY); Prov. San Ignacio, San Ignacio, 600—700 m, 78°46'W 5°12'S, 16 Apr 1996, J. Campos y P. Díaz 2615 (MO). La Libertad: Prov. Trujillo, Trujillo, 15 Apr 1988, C. Díaz & S. Baldeón 2784 (MO), Cerro Campana, 800 m, 25 Oct 1983, A. Sagástegui 10965 (MO); ibidem, 100 m, 14 Jul 1973, A. López & E. Araujo 7978 (MO, NY); Hee A 215 m, 10 ad 1949, N. Angulo 1250 (NY); base del Cerro Campana, 60 m, 23 Oct 1983, A. Sagástegui s.n. (NY); i , 150-200 m, 24 Sep 1957, P. Hutchinson 1358 (NY); Prov. Pacasmayo, Pacasmayo, 7924 d 7335, 120 m, 15 Jul 1985, D. Smith- 4221 (MO); just S of San Pedro de Lloc, 100 m, 31 Dec 1963, P. ÉD rd SC Y paid 3352 (F, G, GH, HEID, K, LE, MO, MICH, M, NY, P, 5, US, USM); ca. 96 km N of Trujillo on P . 60 m, 16 Oct 1984, M. Dillon & M. Whalen 4014 (F, MO); 84 km N of Trujillo, 3 Mar 1973, A. Richardson 2029 NY. Ancash: 78°06'W 10º11'S, 11 Oct 1958, K. Rahn 318 (MO, ex Tropicos). 3. Colicodendron valerabellum H.H. IItis, T pus Y G. Si Er ER mou a (Figs. m dl VENEZUELA: Edo. Trujillo, Distr. bar "Arbusto de 3 j y caediza al tacto. Ped ] dari tario cl inclinad pecto al ej li Bot ect E WW ás] T "X 1 E 1 3 de 64 col t | at, , gineceo gris fiado en las fl li inadas. La infl i I igas y 'pegones' EE "Se discat Iy "n en un área de 3 m? [3 km??]," Carretera vieja Valera-Motatán, saliendo de Valera por el barrio La Floresta. Selva decidua muy intervenida, 550 m, 70°36'W 9?20'N, 16 Mar 1991 (f1, y fr), T. Ruiz-Zapata & C. Benítez de Rojas 4671 (HOLOTYPE: MY; ISOTYPES: K, MO, US, VEN, WIS; photocopies together with individual leaves E IRBR, MER, NY, PORT, 5, He. Colicodendrum species novum cum pubescentia stellata, calyx valvatis 1-seriatis cum nectarius cupularibus et fructus pachycarpus, oro 1 1 paa un Jefta Laat 1 1 1; : : E 1 lar f PNE SUNL [p d ? [3 oppositiis), infl ii iis ( iculatiis), staminis (35—)50—64(-70) (nec 14 ad 20), gynophoriis ca. 8-9 cm longiis, (nec oo 11; s RSS f. e 1 1-1 gynophoriis 3.5-5 cm longiis), g ds: 1 1 1 BEP E +1 [iCal LINE Evergreen palmoid treelets [or in damaged plants with one or t base, some of these decumbent and sending forth 1 to 3, erect, leaf- and sometimes flower-bearing lateral- horizontal shoots], to 5 m long and 3—5 cm dbh often crowded (colonal?); young branch(es) dark-brown echinate or echinate-stellate; stipules absent. Leaves subsessile to short-petiolate, to 1.5 cm, grouped in crowded whorls at the end of the terminal | hes exposing mostly leafless internodes, nodes of leaf inser- tions ca. 3-6 cm, swollen, blades oblanceolate to narrowly elliptic, (16-)20-50 x 4—9 cm, stiffly coriaceous Cornejo and Iltis, Revision of Colicodendron £ og RT | J 1 d hahi Y Corneio & H H Iltis A. | ituri í it fruits. C. Infructescence. (A. Hatschbach. errs 63068, US. B- C. de Mello Filho 2988, CEPEC). and hard but somewhat flexible (alive), chartaceous (dry), attenuate but rounded-retuse to subcordate at the base, acuminate widely obtuse at the apex, margin i land revolute, lustrous green (alive) with a impressed to sulcate midvein, prominently and e rugose De, bullate), with the blade surface between the deeply impressed midrib and main nerves prominently convex and smooth, ap- P ANA A. A em wd A ON H We x RA AE gs Pase Dr wa) kr tiet N E 2 E ES si, Fara eech: es” E BA E > Ê ER TE B P Ed Y ‘ar eS) ue EG A A KSC OR xl * Md Lats Se SR XA E 7: E BAD 7 AN, A EPR V á A KR Viu |. "2 fi Way T e A Oe, yj Fic. 3. Colicodend, ferabellum H H. Iltis, T. Ruiz €: G.S. Bunting A. Habit. B. A leaf blade. C. À A A [a | L all = ELI Jal L a, a a. HI |, Ahnva these, inflorescence bearing (sub)sessil flowerbuds. E-F. Immature fruit. (A. After photo of Bunting & Trujillo 13120. B-D. Iltis et al. 30547, WIS. E-F. Ruiz & Villafañe 4764, MY). A—E, drawings by Marian Firmani; F, by T. Ruiz Z. al H L E | bel D Cornejo and Iltis, Revision of Colicodendron B Fic. 4. Colicodend, ferabellum H.H. Iltis, T. Ruiz & G.S Bunting A. Scd ti ti f flower. nl, bk a d s I d F c M | L . i E al F1 I LEN | E A Eet E D b re FI I LI a FI I L a? IBI E Mala flawar ee LI [UM d r Li LJ d d d d = | A F Al all | L || E ia F1 Le a 1 J || a FI a F1 L L ZA a | (A-D. litis et al. 30547). A, C by Manara; B, D by Iltis. rently glabrous (glal t) above; paler g (alive) and with a st y prominent midvein and densely stellate enla 14 to 26 pairs ge lateral nerves. Inflorescences — racemose spike, stiffly erect and densely EE umes od eee rough to the touch, arising from the center of a leaf whorl, axis eventually elongatin cm x 6-10 mm tick, stout, bearing 20 to 100 or more subsessile, rusty- brown (alive) — ECH but anly 2 to 5 flowers opening each night in successive acropetal circles. Floral bracts subulate, ca. 5-7 mm, deciduous, densely brown-echinate. e 2-3 x 3-4. : mm. Sepals ovate to lanceolate, 15-25 x 8-12 mm, + acute at apex, densely yellow-brown (alive) echinat ially; hypanthium ca. 2-3 mm depth, coated by a 4-dentate nectary dish within, nectary appendages deltoid, ca. 3-4 x 3-4 mm, emarginated to bifid at the apex. Petals apparently inserted at the edge of the nectary cup, at anthesis obovate to obovate-oblong, 25—32 x 10-15 mm, widely obtuse to truncate at apex, longitudinally nerved, bright yellowish or cream (alive), densely pale-brown echinate-stellate abaxially. Stamens (35—)50—64 (—70), filaments (46-7 cm, stellate at the base, inserted on a expanded torus-like androgynophore, át anthesis projecting forward in a narrow phalanx, anthers 3.8-5 mm. Gynophore 8-9 cm, densely stellate: ovary 6—7 x 4-5 mm, subglobose, 8-ribbed, densely stellate: stigma sessile, truncate, discoid. Infructescences a stout gynophore, ca. 10 mm wide. Amphisarcous subglobose, ca. 9 x 7 cm (immature), subpendulous, apex umbonate with an apically stigmatic depression at the umbo, densely rusty-brown echinate, yellow when aml Hum H.H. Iltis, T. Ruiz & G.S. Bunting. A. Terminal two leaf whorls of unbranched, 4 m tall treelet, showing conduplicate 1 | Fi L ITL" | p P = Da! E des R E A E J L A llaafl f t ht. Carmen F. Benit ] y d DU a ed d El li f r A IL E ag L a m Fal L a HE IE E L | n m [| gt | EI L ki E I L] LI - re J rr Li la L [| E E [| | a L E EL | EE J ra FS leaf bases. E , g | ln E Fig’ a | m2 Fr 1 | HE A T inflorescence. (A—D, pl Cornejo and Iltis, Revision of Colicodendron 87 Fic. 6. Colicodend, lerabellum H.H litis, T. Ruiz & G.S Bunting F Rigid, | | y ; y lil H Twoi i fruit k f riai l, stout, erect peduncle (F-G, hot by litis of itis ef al 30547, fi H, photo by G. Bunting). ripe (fide a local informant, Iltis et al. 30547); fruit wall thick, pulp whitish or cream, seeds numerous (ca. 307), reniform, 20—25 x 13-14 mm (immature), seed coat red-purple (fide Bunting). Colicodendron valerabellum differs widely from C. yco by the characters written in the Latin diagnosis. Due to the leaves arrangement in crowded whorls at the end of the terminal branches (Fig. 3A, 5A) and unbranched inflorescences (Fig. 3C), Colicodendron valerabellum seems more closely related to the Brazilian C. bahianum (Fig. 1A). However, Colicodendron valerabellum differs from the latter by having spicate inflo- rescences with a larger central axis, 10—30 cm (Fig. 3C, vs. racemose, with shorter central axis, 1—5 cm, Fig. LA), bearing subsessil (vs. 1.8-4 cm, pedicellate) flowers, longer gynophore, 8-9 cm (vs. 1.6-2 cm), subglobose (vs. oblong) ovaries; subglobose fruits (Fig. 6H, vs. oblong-ellipsoid, Fig. 2) and by its localized distribution, in northwestern Venezuela (vs. eastern Brazil, Fig. 7). Phenology.— Colicodendron valerabellum apparently begins to flower in November, not long after the end of the rainy season, and continues to bloom into April or May (or *does it flower twice a year, once in May and again in November?”), with older hermaphrodite flowers beginning fruit enlargement with the com- mencement of the rainy season at the beginning of May, forming young fruits by July, “when new growth was just starting," and completing their growth by the following January or February. The terminal inflorescence of Colicodendron valerabellum is apparently an extension of the apical mer- I .£ «| Das H Im LI PR £ 88 J Texas 2( istem that continues to grow from the center of the upper-most leaf whorl. It is, TIENE er, a structure most lly sessile or subsessile unusual in many respects, not the least of which is the fact that its fl on the peduncle axis, with the pedicels very short and thick, so that this structure, rather than a raceme, could better be called a racemose spike. The remaining species of Colicodendron and even of the neotropical Capparaceae s.s. have flowers borne on well developed pedicels. In this comparatively specialized species, this loss of pedicel is linked to a basically primitive, racemose inflorescence, while its three relatives, C. bahianum, C. yco and C. scabridum, have pedicellate flowers in racemes or panicles, presumably the derived condition. Correlated with this loss of pedicel in C. valerabellum may be the great length, thickness and stiffness of the peduncle, truly spike-like in the non-botanical meaning of the word, ultimately becoming a full centimeter thick and to 40 cm long, which eventually decompose. Around the stubby remains of the old inflorescence in the center of the leaf whorl, several axillary buds, densely covered with subulate bracts, may be initiated. These in turn may produce 1 to 3 replacement axes, essentially by continuing the unbranched growth upward and the production of another flowering spike, or else lead to the branching of the stem into 2 or 3 axillary branches. Although variable in particulars among individuals, the paran terminal Bp bear flowers in the following sequence: the lowest floral buds soon abort, leaving only pedice Isca duncle (these buds are usually naked, but in some inflorescences subtended by up to 20 or so small leaves rapidly decreasing in size upward and spaced over a distance of some 10 cm). Moving upward, the next 3 to 6 flower buds, well spaced and the first to burst into bloom, develop into fully functioning hermaphrodites, in which the many stamens are exceeded for about 1 cm by a gynophore bearing its well-developed ovary. Finally, towering above the circle of hermaphrodite flowers, and densely aggregated at first into a narrowly conical inflorescence, are some 20 to 100 staminate flower buds, with only 2-3(-5) of these blooming at one time, beginning at base and continuing upward. The whole inflorescence thus has a long-extended flowering period lasting perhaps a month, and in the process gradually elongating the peduncle to its maximum length. Once the staminate flowers finish blooming, they soon wilt and fall off, leaving only the symmetrically placed, oval pedicel scars on the peduncle. If the hermaphrodite flowers at the base of the spike become fertilized and commence fruit develop- ment, the peduncle above them eventually dries up and tually decomposes. But if these hermaphrodite flowers are not fertilized, not only do they fall off, but also the peduncle then often retains its stiff, thick structure to eventually produce a new set of hermaphrodite flowers near its tip some 30 cm or more up from its base. This double insurance that seeds will eventually be produced may be the key to understanding the function of so massive a structure. Clues to the enigmatic reproductive morphology of Colicodendron valerabellum must lie in its pollina- tion agents. Of the eight main attributes of the bat pollination syndrome (Chiropterophily) cited by various authors (Pijl 1961: 51; Faegri € Pijl 1966: 114; Vogel 1958: 492), the following six would seem to apply to Colicodendron valerabellum: 1) Nocturnal anthesis, the flowers lasting only one night. 2) Flowers and inflo- rescences strong enough to carry the weight of a bat (subsessile, often many flowers on the strong pedicels, thick calyx, powerful peduncle, i.e., racemose spike axis). 3) Honey abundant, mucilaginous, accessible (to sphingid moths as well). 4) Flower (corolla) yellowish or creamy. 5) Pollen abundant. 6) Inflorescence exposed high above the foliage. Biological interactions. According to a local informant, the fruits of Colicodendron valerabellum are eaten by deer (Iltis et al. 30547, WIS). Why the big fruits of Colicodendron valerabellum should be borne high above the leaves is a curiosity, when in its close relatives, as in most Capparaceae, heavy fruits hang down. Or were the fruits/seeds of Colicodendron valerabellum dispersed internally by the now extinct megafauna as the giant Gomphotheres (Janzen & Martin 1982), as suggested for Neocalyptrocalyx muco (H.H. Iltis, Cumaná, Delgado & Aymard) X. Cornejo & H.H. Iltis, another giant-fruited Capparaceae of northern Venezuela and Colombia! (Iltis et al. 1996: 379, see Capparis muco). Cornejo and Iltis, Revision of Colicodendron 89 Cytology.—Unknown. Distribution. — Colicodendron valerabellum is known only from one limited metapopulation of several, scattered minute populations, with perhaps a total of 20 to 40 plants, growing in extremely disturbed, open tropical deciduous forest remnants and thickets on top of 10—20 m high, steep and badly eroding road cuts and local trash dumps scattered for 3 to 7 km north of Valera (654 m) along and immediately above the old highway to Motatán (ca. 500 m), in northwest Venezuela (Fig. 7). Growing in seasonally dry, semi-deciduous to deciduous open forest or woodlands of medium-sized trees, the *bosques higrófilos y mesófilos de las sierras en la región Andina" (Hueck 1960), or in degraded secondary thorn thickets (matorral). Conservation biology.— Colicodendron valerabellum appears to be close to extinction, but with all known populations restricted to the edge of the old yet easily accessible Valera-Motatán highway, and the rest of the landscape barely explored, the possibility exists that the extensive woodlands covering the hills above the road may wel! hold more extensive stands. In fact, this is also suggested by the presence of a common name, "fruto de venado” (fruit of the deer), since rare species are generally not honored with common names. The few plants known to exist are certainly in trouble reproductively: except for the inflorescence with two young fruits photographed by Bunting in 1983, the two somewhat older fruits collected by Ruiz-Zapata in 1997, and the five hermaphrodite flowers with developing ovaries at the tip of an inflorescence collected by Ruiz-Zapata in 1998, no other fruits, young or old, have ever been located despite assiduous searching. Perhaps, like many a taxon at the end of its evolutionary career, C. valerabellum is not reproducing well. Negative human influences such as firewood gathering, land clearing, and grazing by cattle as well as the unknown but negative impact of pesticide spraying can also not be discounted. What is clear, in any case, is that the limited fruit production in all the ined populations (by H.H. Iltis) may be due to one or more of the following: 1) The often densely crowded stems may be clones, and, if self-incompatible, cannot form fruit unless cross-pollinated; 2) land clearing may have reduced genetic diversity to the point of no return; 3) the population is inbred, reducing seed production; 4) populations of the pollinating agents (sphingid moths, bats, or hummingbirds) have been much reduced by pesticide application to the local maize fields; 5) the species is adapted to an undisturbed summergreen open wood- land periodically cleared by fire, and now these relictual stands have to compete with weeds in a liana-rich, secondary community induced by human disturbance; 6) A remarkable number of Capparoids in the region, have extra-large, more or less woody, indehiscent fruits that, as one of us suggested (Iltis et al. 1996) may have had Gomphotheres, now extinct, as their dispersal agent (Janzen & Martin 1982). ParaTYPES. VENEZUELA: Edo. eee Dtto. Mw carr. ien Keen Motatán, E-facing slopes of Río Motatán Valley, foothills of the Cordillera de Mérida of the Andes, on to 20 m ab the old highway from Valera to Motatán, ca. 4 km N of Valera (1.4 km N of entrance to old Gen sen. new bur [ca 36'W 9°21'N], 600 m, 10 Jan 1991 (y fl), H.H. Iltis, T. Ruiz & C. Benítez 30547 (WIS); 7 km N. of Valera Q km. S of Mod H.H. T et al. 30548 we MY, US, bd carr. SER en us matorral deciduo, 600 m, 31 Mar 1996 (fD), T. Ruiz & Villafañe 4740 (1 de Valera, para conucos [maize fields] y también el uso como basurero, 435 m [?], 10 Jan 1997 (f1., fr), T Ruiz & Villafañe 4764 (MY): carr. vieja, en matorral deciduo, 590 m, 31 Jan 1998 (old e T. SS VLRO 6 Rada 4801 (MY, WIS), 1 Moy 1998 (FD, T. Ruiz, Hernández & Noguera 4802 (MY), carr. vieja, ca. 3 km N de \ , en ladera a la izquier da de la vía, en zona de bosque oe n alterado, ca. 450 m, 6 Jun 1983 (y fr), G. Bunting & Trujillo 13120 (WIS), carr. vista Agua Viva-Valera 3.5 km debajo e la vía, 500 m, 28 Nov 1982 (FD, A. Stoddard s.n. (MY, NY [2], WIS); carr. Vieja, zona perturbada, 9 May 1980 Sch? E Simeon & A. Stoddard 14 (US, VEN). 4, Colicodendron yco oe Ra ar plns l: 225. ee (Fig. 7). Capparis yco (Mart.) Eichler in NE s Bras. . 1865. Tyree: BRAZII Rio de Contas etc. Provinciae Bah usque in P. Min. Ger., Sep-Oct, C. Martius s.n. (LECTOYPE, dienen fee M [M photo at WIS!]; isoLecrorvres: BM, K, M). C M ( Griseb., 1879), cited in Eichler in Mart., Fl. Bras. 272. 1865. Brazil, Bahia, pU E Jacobina, 1839 (fl), J. a 2564 NY 387633! (tom C. Meisner add NOTE: J. Blanchet 2564 B 10 02427271, it has been determined by error (by H.H. Iltis 1 yI pparis ) Many branched shrubs to trees, 2 to 6 m tall; terminal branches complanate, angulate, often + subtetrago- nal, densely stellate. Leaves opposite, petioles 0.5-2 cm, lacking pulvini; blades coriaceous, lanceolate to aia Texas 2(1) 80° 70° 60° 50° 40° b] : d : A si E M 1 d À AS sg . Colicodendron 10* 1 TA ji WS: C. valerabellum e X6 2 MS ne 1 0º 4 : $ A | Ys gen ul f Y 3 A 7 eet ra ; H . s À i m S , M SÉ À E EA 7 E c A c À nme no RE E df J AMAZONIAN FOREST = ^ “a | CAATINGAS o e pU cem TS d p E) Ss L . A . 3 nj é ` d Li C. scabridum H i ! 10º ^ y X Pus Em Ab © X F Ü i D a ! à : i A 1 S ^N GEME Ng -f Ge SÉ P co $t» a O Dor es | € ^" Ps 20° L } | i 1000 km De 7 ra? | AS E eng F d AA A TL | J L g [do fal E I Je j LU) deciduous, xeric, tal lland C , Prov. Manabí, Ecuador, th through tl d y H b of tal Ecuad tl t y | " £n E r , Prov A h; ac la al [| y, i FI A Jė In! y | j ,E | Jal D n AA e , Prov. Amazonas, Perú. 2) Colicodend lerabellum H.H. iltis, T. Ruiz & G.S. Bunting: A highly local endemic, restricted to a stretch of road ca. 3-6 km north of Valera | g ld hial to Motatán, Est Tujillo, Venezuela, i tod but : lecid t i evergreen, tropical, I | yll ] y f t id (and I thicket , at elevation of ca. 600 m. 3) Colicod ] y M ti | Brazili f Bahi | |, barely, Ceara), in the Caatinga, and d ht-frequented land, where tl getation i phyti , summer-deciduous, and drought- adapted. 4) Colicod 7 (. Cornejo & H H Iltis, r tricted t ist and wet fi ts at SEof Bahia State, at Eastern Brazil. (Distribution of C. oblong or oblong-elliptic, 9-25 x 3-8.5 cm, usually retuse to cordate at the base, acute to widely obtuse and apiculate, few times rounded to truncate or notched at the apex, margin entire to sinuate, dark green glossy (alive) and glabrescent above, whitish, whitish-grey, pale green or pale tan (alive), with prominent nerves and densely stellate beneath; (8210—16 pairs of lateral nerves. Inflorescences axillary and terminal paniculate, rarely solitary axillary flower (Pinheiro 1453), densely stellate. Floral bracts linear-subulate, 5—10 mm. Pedicels 1-2 cm, articulated. Sepals 1-1.8 x 4—7 mm, the lobes acute to obtuse at apex, reflexed at anthesis, cream or dull-yellow, densely stellate, deciduous; nectary a 4-dentate cup, the teeth 3—5 x 2-4 mm triangular, acute. Petals oblanceolate to oblong, 15-30 x 5-7 mm, at anthesis all together erect resembling a tubular corolla, reflexed at the upper third, yellow to orange-yellow, stellate adaxially, densely stellate abaxially. Stamens 14 to 20, filaments 3.5—4 cm, stellate at the base, inserted on a stellate androgynophore anthers ca. 3-4 mm, yellow (alive). Gynophore 3—5 cm; ovary ovate, 3-5 mm, both densely stellate; stigma Cornejo and litis, Revision of Colicodendron 91 capitate-disciform. Infructescences with gynophores 3-5 cm, pedicels 1-4 cm; amphisarcous ellipsoid, 6-10 x 4-6 cm, subpendulous, apex umbonate with an apically stigmatic depression at the umbo, densely stellate, bearing 10 to 20 seeds, testa light brown, crustaceous, glabrous. Icones.—Colicodendron yco is illustrated in detail in Flora Brasiliensis (Eichler in Mart. 1865, tab. 60, see Capparis yco). Local names.—Icó (Portuguese, R. Santos & A. Castellanos 24350), icó branco (Portuguese, Laurénio et al. 262), icó de cavalo, icó preto (Portuguese, Costa e Silva 1995: 75). Biological interactions. —The base of the flower is visited by small ants (Harley 27133). Cytology.—n = 8 [!Pazy, BRAZIL. Banta: Mori et al. 11218, WIS, Iltis unpublished ms). Distribution and ecology.—Widespread in the northeastern Brazilian states of Bahia and Pernambuco (and, barely, Ceara), in the Caatinga, a semi-arid (300—800 mm/year rainfall), hot (23°-28° C yearly average) hytic, summer-deciduous, and drought-adapted = and drought-frequented land, where the vegetation i phy (Costa e Silva 1995: 74). lied. BRAZIL. P ] Mun. Belém de Sáo F isco, ilha do Meio, 18 Jul 1967, E. idi "i cs Mun. Cabro- , chapada ão bó, entre Cabrobó e Terra ae 15 E 1971, D Andrade-Lima et al. 767 (IPA); Mun. Inajá, R ánica em decomposicáo 15 Sep 1995, M. Tschá et al. 236 (NY, PEUFRX 16 Sep 1993, arenítico, solo OT? M. Tschd et al. 247 (NY, PEUFR); ibidem, 9 Dec 1995, A. Laurénio et al. 262 (NY, PEUFR); Mun. Petrolina, Fazenda Santa Maria, proximo ep 1963, a divisa com Santa Maria da Boa Vista, 3 Sep 1968, G. Carvalho 54 (HST). Bahia: Mun. Vitoria da Conquista e Jequié, 26 S R. Santos & A. Castellanos 24350 (NY); Mun. Jequié, Estrada que liga Jequié a Lafayete Coutinho, ca. 11-17 km a W de Jequié, caatinga herbáceo-arbustiva, 19 Nov 1978, S. Mori et al. 11207, 11218 duis. Jequié, 28 Sep s K. d 6 H. PU Mee /9-262 ad. De do Castanháo, 14.51 BR 1 Jequié para Milagres, 20 i 1965, A. D rascaria Corujáo), then 7.3 km W of BR 116, 40?11. 35 W 13º 57'S, SE MEL 575 m, , 19 oe 2001, W. Mona & S. Sant Ána 12527 (NY); 40°11.46'W 13°56.52'S, 600 m, high el pecies, 23 Oct 2001, W. Thomas et al. 12567 (MO, NY); Mun. Juazeiro, N end of Pee eee at E 11 kms i Barrinha rm 52 km N of Senhor do Bonfim) Ss n 1993, W. dien et al. 9632 wD Serra | d Povoado at Fazenda Pasto Bom, 42°12'W 10?00'S, Q da Jacobina, 1839, J. Blanchet 2564 (B, NY); : Mun: Rui Barbosa, 8.5 km from Rui B Alagoas to Itaberaba, e tall, partly deciduous seasonal pd 40°23'W 12°15'S, 29 Jan 1993 J Kallunki & J. Pirani 392 NY): Mun. ado, 41°50'W 13?38'S, 650 m, Caatinga, 12 Dec 1988, R. Harley et Rio de Contas, 7 ! de na Estrada para Li u al. 27133 (MO, NY); Serra do Rio de Gare. by the en of Rio M just N of Livramento d ca. Aldd wW SE ca. 460 m, 20 Jan 1974, R. Harley et al. 15339 (MO, NY); Mun , ca. 30 km na E Grosso, caatinga, 14 Mar 1990, A. de Carvalho & J. Saunders 2765 (CEPEC, NY; Lagoa da Eugenia S end near a ca. 10940 3 39?43"W. 300 m, 21 Feb 1974, R. Harley 16284 (NY), Quemaidas, 9-11 Jun 1915, J. Rose & P. Russel 19843, 19871 (NY); 5 km al S de Ichu, camino a Tanquinho, ca. 39°09'W 11°48'S, ca. 300 m, a 1997, Arbo M.M. et al. ere (CTES, NY); 30 km N of Serrinha, ca. 39°00'W 11°30'S, 2 Nov 1972, T. Botter & Da Fonséca 2704 (NY[2]); B legre km. 27, caatinga, 8 Jul 1971, R. Pinheiro 1453 (CEPEC, NY); Salgada, 15 May 1918, H. Curram 277 (GH, T KEY TO THE SPECIES OP COLICODENDRON the | long the branches, SE petiolate. e cup, adrogynophores 1. Many branched tree 2. Leaves opposite; rowers eee, sepals reflexed at anthesis, nectary a ntered, gyno} -5 cm, densely stellate; petals 5-7 mm wide; E Bra E p» A spirally, rarely subopposite on the same branch; flowers brown to greenish-brown, sepals erect anthesis, nectary a 3-lobed cup, anon ye pnorss Ol entered gynophores 5-8 cm, sparsely stellate; p 10-15 mm wide; W Ecuado Peru C. scabridum dt | whorled at the end of the terminal branches, h.l | E nl | + pa +) C. yco 1. | to sparsely sessile to > cm petiolate 3. Inflorescence a raceme with the flowers distinctively pedicellate, pedicels 1.8-4 cm, relatively laxly ar- ranged on a short axis, 1-5 cm; gynophore 1.6-2 cm, ovary oblong; fruit oblong-ellipsoid; eastern Brazi C. bahianum 3. Inflorescence a racemose spike with the Towels SE pedicels g 3 mm, n, gensely arranged along the axis, 10-40 cm; gynophore 8-9 cm, ov Venezuela, near Valera C. valerabellum Rejected names Colicodendron anceps Shuttl., nomen in Chapman, Fl. South. U.S. 32. 1860. - Quadrella jamaicensis (Jacq.) J. Presl. 92 Colicodendron angustifolium (Kunth) Hutch., Gen. Fl. Pl. 2:309. 1967. = Quadrella angustifolia (Kunth) H.H. Iltis & X. Cornejo. Colicodendron avicenniafolium (Kunth) Seem., Bot. Voy. Herald 78. 1852. - Beautempsia avicennifolia (Kunth) Gaudich. ipod crotonoides (Kunth) Hutch., Gen. Fl. Pl. 2:309. 1967. - Capparicordis crotonoides (Kunth) H.H. Iltis & X. D Bees lepidotum Turcz., Bull. Soc. Nat. Mosc. 27:327. 1854. = Quadrella odoratissima Jacq.) Hutch. Colicodendron longifolium Mart., Flora 22:26. 1839. = Neocalyptrocalyx neum (Matt, ye X. Cornejo & H.H. Iltis Colicodendron obovatum Turcz., Bull. Soc. Nat. Mosc. 27:327. 1854. (Jacq.) X. Cornejo & H.H. Iltis Colicodendron pulchellum Seem., Bot. Voy. Herald 78. 1852, nomen - Calanthea pülcumma Cac) M Colicodendron salicifolium (Griseb. ) Hutch., Gen. Fl. Pl. 2:309. 1967. = Sarcotoxicum salicifolium e X. Cornejo & H.H. Iltis. Colicodendron subbilobum (Kunth) Seem., Bot. Voy. Herald 78. 1852. = Cynophalla flexuosa (L) J. Presl. Colicodendron tweedianum (Eichl) Hutch., Gen. Fl. Pl. 2:309. 1967. = Capparicordis tweediana (Eichl.) H.H. Iltis & X. Cornejo Uncertain name Colicodendron obliquifolium Turcz., Bull. Soc. Nat. Mosc. 27:328. 1854. We have not seen Linden 1370, the cited type specimen of Colicodendron obliquifolium, collected from “Nova Grenada, prov. Pamplona at ripas Rio Zulia” (Colombia). However, Colicodendron obliquifolium, originally described as “glabrum...” evidently does not belong to Colicodendron. ACKNOWLEDGMENTS Thanks are due to the following herbaria for sending their collections as loan or gift for this study: AAU, B, BKL, BM, CEPEC, CGE, CTES, DAV, F, GB, GH, GUAY, HEID, K, LD, LOJA, M, MO, NY, PEUFR, PR, QCA, QCNE, RSA, SEL, US, USF, USM, WIS. The first author had the pleasure to visit the herbaria: BKL, GUAY, LOJA, MO, QAP, QCA, QCNE, SEL and US. Ludwig Martins, Ulrike Starck and Hans-Joachim Esser pro- vided valuable help with the types housed at B and M, respectively. R. Vogt (B) confirmed the Willdenow's handwritten on the ase. e the pip n of Capparis scabrida (B-W 10055-00 0). The second author gratefully acl | | fthe University of Wisconsin-Madison Graduate School, the O.N. and E.K. Allen UW pieibacam Fund and the Consejo de Desarrollo Científico y Humanístico de la Universidad Central de Venezuela for financial help and sponsorship of the 1991 *Capparaceae Expedition" to Venezuela. Carmen Benítez de Rojas made valuable comments of Colicodendron valerabellum. The drawings and photo are mostly the talented artwork of Kandis Elliot, Marian Firmani (Fig. 3A—E), Bruno Manara (Fig. 4A, C), T. Ruiz Z. (Fig. 3F), and George Bunting (Fig. 6H, photo). REFERENCES Brunina, DE 1985. All around the Sapote bush. Animal Kingdom, New York Zoological Society. Pp. 33-35. Corneso, X. 2005. La vegetación del Estuario del Río Chone. einenowlas 6: 87- 94, Cornejo, X and HH. om, 2008a. Two new genera of Capparaceae: 5 nd M pparis stat. nov., and the reinstatement of Neocalyptrocalyx. Harvard Pap. Bot. 13:103-116. Cornejo, X and HH. litis. 2008b. New combinations in South American Capparaceae. Harvard Pap. Bot. 13:117-120. Costa e SiLvA, M.B. 1995. Estudos taxonómicos sobre o género Capparis L. (Capparaceae Juss.) em Pernambuco- Brasil. Dissertação apresentada ao Programa de Pós-Graduação em Botánica da Universidade Federal Rural de Pernambuco. Recife, Brasil. 141 pp. (Thesis, unpubl.). EicHLER, A.W. 1865. Capparideae. In: Martius C.FP von, ed. Flora brasiliensis 13, tab. 51-65. Pp. 237-292. FaEGRI, K and L. van DER PuL. 1966. The principles of pollination ecology. Pergamon Press, Toronto. Hueck, K. 1960. Mapa de la vegetación de la República. de Venezuela 1: 2,000,000. Instituto Forestal Latinoameri- cano de Investigación y Capacitación. Mérida, Venezuela. HUTCHINSON, J. 1967. The genera of flowering plants (Angiospermae). Clarendon Press, Oxford. Pp. 2:303-317. Cornejo and Iltis, Revision of Colicodendron 93 Ins, H.H., LJ. Cumaná, RE DeLGADO, and G.C. AvmarD. 1996. Studies in the Capparidaceae XVIII: A new giant-fruited Capparis (C. muco) from eastern Venezuela. Novon 6:375-384, ILTIS, H.H. and X. Conejo. 2007. Studies in the Capparaceae XXX. Capparicordis, a genus from the Neotropics. Brittonia 59:246-254. JANZEN, D.H. and PS. Martin. 1982. Neotropical anachronisms: The fruits the Gomphotheres ate. Science 215:145-164. Pax, F. and K. HOFFMANN. 1936. Capparidaceae. In: Engler, A. & Prantl, K, eds. Die Natürlichen Pflanzenfamilien, ed. 2, 17b. Leipzig. Pp. 146-223. Pui, L. van DER. 1961. Ecological aspects of flower evolution Il: Zoophilous flower classes. Evolution 15:44-59. SEEMAN, B. 1852. The Botany of the Voyage of H.M.S. Herald under the command of Captain Henry Kellett, R.N., C.B., during the years 1845-51. London. SPJUT, RW. 1994. A systematic treatment of PRU types. Mem. New York Bot. Gard. 70:1-182. THE New York BOTANICAL GARDEN. 2003 | Plant Science Center. The CV. Starr Virtual Herbarium, New York; http:// nybg.org/vl Spes ER nce aed b March 2008, 12:00 SIUS amo N. 1854. A | | | priman partem herbarii iani, nunc universitatis caesareae charkowiensis. Bull. Soc, Imp. Nat. Moscou 2 271 -372. VAN DEN EvYNDEN, V., E. Cueva, and O. Cabrera. 1999. Plantas silvestres comestibles del Sur del Ecuador-wild edible plants of southern Ecuador, Quito. Voce, S. 1958. Fledermausblumen aus Südamerika. Osterreich. Bot. Zeit. 104:491-530. 94 J lof the Botanical R h Institute of Texas 2(1) BOOK REVIEW WALTER S. JUDD, CHRISTOPHER S. CAMPBELL, ELIZABETH A. KELLOGG, PETER E STEVENS, AND MICHAEI J. DONOGHUE. 2007. Plant Systematics: A Phylogenetic Approach (ed. 3). (ISBN 978-0-87893-407-2, hbk.). Sinauer Associates, Inc., 23 Plumtree Road, Sunderland, Massachusetts 01375, U.S.A. (Orders: www.sinauer. com, fax 1-413-549-1118). $94.95, 620 pp., ca. 325 illustrations, 25 color plates, 8 1/2" x 11". Chapters: 1. The Science of Plant Systematics 2. oe and M ae of in on BINOS Historical Background 4. Taxonomic Evidence: Structural and Biochemical Characters 5. Molecular Systematics 6. The Evolution of Plant Diversity 7. An Overview of Green Plant Phylogeny 8. Lycophytes, Ferns and Their Allies, and Extant Gymnosperms 9. Phylogenetic Relationships of Angiosperms Appendix I. Botanical Nomenclature Appendix II. Specimen Preparation and Identification For the third time, we look at plant classificati ith tl d d scientists Judd je UM Kellogg, zem ens, Dee oe ion E J $ Lai 1 1 4 A + £s 11 a >) J YF. ado VaL È e tools of science are all evolving. ER thie II it hold ything Jarai about a families, floral formulas, ned Se E A : Ea T " ] el + E E aea 1 1 ] g p I j : , figures, phy E g acter states dal di | DNA ] bl ic phyl ] iti In addon ] ided al itl E o i I + oO L i F d Eu [e] pictures, tal ] f diagnostic feat Í bfamilies, a nice glossary, and a DVD. f. 4 +1 Te [ 1 TE ola z 1 + T] D ] 1 1 5 7 L f+} +1 [ A s Tas GL 1 O 1 D id a H +1 Lo 1 + T eo 1 ee = 1 1 1 1 p ] = M 1 - g y phylog I , and how we have historically handled the task of plant classification. Tl 1 1 1 t ] 1 t 4l las kg, ve a Al cuim tend in (E , g p groug p why a group Į The authors 1 ] 1 1 1 F.L 1 1 7 ] 1 ] 1 1 1 rr I 4 [e] id LA 1 c 1 Os 11 TYNI A f and anatomy as eatures. Since mc printing ja neo cw oe of P Mean some new EE of grouping have evolved. For the non-flowering plants, (1) f ph; l formerly within Ophioglossales, now constitutes its own order, M iales; (3) Polypodiales i bcategory of the Leptosporangiate Fe Ferns” o of Peng the equivalent, meaning tl famili f Pol lial lefined in tł À editi bum to Osmundaceae, formerly fi EUMD 2 within Der is e dales). Regarding classificati ithin tl gios] rhe “Magnoliid compre is identified I M liales, Laurales, Canellales, and Pi les f recommend this rexibook lora any classes that are botany-related, for any research projects Gen are Gët E and any naturalist groups that are botany-related. Plants are wonderful: let people know more about them.— Virginie H. Raquet, BRIT Research Associate, 13380 Plan-de-Cuques, France, vraquet@brit.org J. Bot. Res. Inst. Texas 2(1): 94. 2008 REVISION OF GONOLOBUS S.S. (APOCYNACEAE: ASCLEPIADOIDEAE) IN THE WEST INDIES Alexander Krings Herbarium, Department of Plant Biology orth Carolina State University Raleigh, North Carolina, U.S.A. 27695-7612 ABSTRACT Á revision of Gage s.s. (Apocynaceae: Asclepiadoid Gonolobinae) in the West Indies is provided based on recent evidence of zl 1 et, 1 1 ] 3n. T -anal Jata T : : jul. 3 ET : ] D pport ixi hyl 5 [s eo I 5 1:11 E 5:3 ] f£. 11 4 3 Tl ti tat f 1 island SE Keys, a I p E E ally assessed using IUCN and NatureServe criteria. Gonolobus stapelioides is neotypified. Two new combinations are made in Matelea: M. dictyopetala and M. pubescens. RESUMEN col a 1.1 KA e Ai ee 1 TENE on 1: A 1 Tad: nr 3 1 1 1 1 T I + L que soportan la maarab lha dal mea ka cada EA | ] 1 Lata ai E ds e PE J: S e 1 iz " 1 L I o Ue rM M PI NS I "s sola isla). S rtan cl , descripciones, de las d S lúa el do d I ] | los criterios ge IUCN y de NatureServe. Se neotipifica Gonolol pelioides. Se hacen d binaci Matelea: M. dic- tyopetala y M. pubescens. INTRODUCTION f lass Dep 1] fA A 7 E WE | ^ About fifty sp subtribe Got inae (Apocynaceae: Ásclepiadoideae) occur ronthe islands comprised by the Greater and Lesser Antilles, the Bahamas E and E and Aruba and the Netherland An- tilles. Evidence from the chloroplast (Rapini et al. 2003; Liede-Schumann et al. 2005; Rapini et al. 2006; Krings et al. 2008) and nuclear genomes (Krings et al. 2008) supports the monophyly of the subtribe. Genera referred to the subtribe with M centanon o in dees GE ees Phera DC., Gonolobus Michx., Matelea Aubl. (incl. Ibatia Decne., Jacaima Rendle, P ] EECH Decne.), Macroscepis Kunth, and Metalepis Griseb. (Fontella & Schwarz 1981a: Rinze 1995; Died: 1997; Morillo 1997; Rapini et al. 2003; Liede-Schumann et al. 2005; Rapini et al. 2006). Subtribal position has been most controversial kers have also placed within a broad Cynanchum L. (Woodson 1941; Spellman for Metalepis, which some 1975; Sundell 1981). Recent evidence places it basal to a well-supported Metastelmatinae-Oxypetalinae- Gonolobinae clade (Liede & Kunze 2002; Liede-Schumann et al. 2005). The last regional treatment of Gonolobinae is now over 100 years old (Schlechter 1899) and a number of new species have recently been described by various workers, including Britton, Krings, Liogier, Proctor, Spellman, Urban, and Woodson (see Krings 2005b, 2006, 2007). Most members of the subtribe have never been monographed (Rapini et al. 2003) and the circumscription of most genera remains poorly defined. The Gonolobus-Matelea complex in particular is in desperate need of taxonomic revision. However, the size of the complex (ca. 200-350 sppJ requires an approach focusing on smaller, more tractable subgroups. Considering the support for a monophyletic Gonolol s. based on recent evidence from chloroplast (trnL-F, rps16) and nuclear (LEAFY) data (Krings et al. 2008), recent t ic changes in West Indian taxa (Krings 2005a—d), and that several species have been published from the area since the last comprehensive treatment over a hundred years ago (Schlechter 1899; Krings 2006, 2007), a revision of the genus in the West Indies seemed appropriate. This revision treats all ten West Indian Gonolobinae species referable to Gonolobus s.s. (i.e., characterized by the presence of laminar, dorsal anther appendages, winged follicles, and a synapomorphic guanine; Krings et al. 2008). In addition, two new combinations are made in Matelea: M. dictyopetala and M. pubescens (see Doubtful and excluded names). | Rat Rac Inet Tayac 2(1): oc —138. 2008 METHODS This treatment is based on critical study of ca. 250 specimens of the fifty some known species in West Indian Gonolobinae, obtained in part through: (1) loan requests from ninety herbaria—of which sixty-hve responded with either loans, digital images, or negative search results (see Acknowledgements), (2) visits to BM, BSC, DUKE, HAC, HAJB, IJ, K, UCWI, UPRRP, US, and P, and (3) forty-eight days of field work by the author in the Bahamas (Long Island), Cuba, Dominica, Jamaica, Puerto Rico, St. Lucia, and St. Vincent. The treatment is also informed by analyses of sequences of portions of the chloroplast (trnL-F, rps16) and nuclear genomes (LEAFY) of selected accessions (see Krings et al. 2008). The species concept employed is an amalgam of the Phylogenetic Species Concept (PSC) of Nixon and Mee (1990) and the Evolutionary Species Concept (ESC) as Hoc by Wiley (1978), namely that: a species is a lineage of ancestral-descendent populations, which tained their identity from other seh] lineages, and sieh are diagnosable by a unique combination of character states in Ge individuals. Although the PSC has been critiqued as a non-historical concept—and tl tially failing to identify instances of phenotypic homoplasy (Baum & Donoghue P een has not been accepted universally (Luckow 1995). The concept has been useful both as t efinition and operational delimitation criterion. However, as the application of the PSC, and its ONE EE EE EEN ge (PAA; Davis & Nixon 1992), requires study of populations, its application is limite uniquely from a population, or worse, from a single collection a Bis e also In the absence of ODDS multi- porn accessions ini extant and historical species, the po EE E here is to rely on obse I with the general phy! due to a variety of biological BE + e L L underlying tl hologi ide al integrity of species (McDade 1995). As a result, character states of each lone papules a I tative were assum ed fixed a together used to form a population profile for PAA. Specimens representing each distinct aggregated population profile were considered to belong to distinct species, even if represented by only single collections, if they exhibited unique, qualitative morpho- logical character states, unknown from other population profiles. As with any hypothesis, concepts of these pee are open to modification and reinterpretation in light of SCH new collections. tion status oft d | lly evaluated by applying the criteria articulated in version 3.1 of the International Union for on of Nature and Natural Resources (IUCN) Red List Categories and Criteria (IUCN 2001) and version 6.1 of the NatureServe Conservation Status Ranks (NatureServe 2006). RESULTS Ten species of Gonolobus are recognized from the West Indies, here defined to include the Bahamas, the Greater Antilles, and the Lesser Antilles (excl. Aruba, Bonaire, Curacao, Trinidad, and Tobago; Fig. 1). All ten species are endemic to the region. Descriptions of the species are provided following a discussion of morphology and keys to the species. Corona morphological terminology follows Liede and Kunze (1993) and Kunze (1995): Ca = faucal annulus (corolline corolla); Cd = dorsal anther appendage; Ci = interstaminal gynostegial corona; C(is) = fused staminal and interstaminal gynostegial corona; Cs = staminal gynostegial corona. Species are arranged alphabetically. IUCN criteria justifying an assigned conservation category are listed following each category. Following Franz et al. (accepted), taxon concept mapping is provided to facili- tate databasing. The operators «, =, and > are used to indicate whether a given taxon concept is respectively narrower than, equal to, or broader than a previously published concept. The symbol + is used to indicate the misapplication of a name to a concept. Herbarium abbreviations follow Index Herbariorum (Holmgren & Holmgren 1998-present). Book abbreviations follow TL-2 (Stafleu & Cowan 1976-1988) and journal abbreviations B-P-H (Lawrence et al. 1968) and B-P-H/S (Bridson & Smith 1991). Author abbreviations follow Brummitt and Powell (1992). Krings, R ici "Ta A WARS a G. aer LATA ES G. stellatus mA G. martinicensis G. youroumaynensis Fic. 1. Distributi TAXONOMIC HISTORY The genus Gonolobus was erected by Michaux in 1803 based on a taxon from t] theastern United States— Gonolobus suberosus (L.) R.Br. Reveal and Barrie (1992) reviewed the complicated nomenclatural history of the type species and Krings and Xiang (2004, 2005) its t y. About 318 names have been published in Gonolobus. Estimates of species numbers in the genus vary from 100 to 150 (Rosatti 1989; Mabberley 1997: Stevens 2001). The degree of variation is largely the result of still poorly known tropical taxa and differ- ences regarding generic limits. Woodson (1941) considered that the genus Gonolobus should contain plants characterized by only none: oo EE EE dorsal anther Ap pensas. M" smooth, angled or A 1 ] 1 by glandular ] trichomes, ni winged follicles, anthers lacking laminar dorsal appendages, wi muricate follicles. Unconvinced that laminar dorsal anther appendages should serve as a generic character and citing examples of smooth [but not angled or winged] fruits in Matelea, Shinners (1950) argued against Woodson's generic concept and later included twelve of Woodson's (1941) Gonolobus combinations in Matelea (Shinners 1950, 1964). Drapalik (1969) essentially followed Shinners by maintaining the type of Gonolobus Oe. G. suberosus, syn. G. gonocarpos (Walter) L.M. Perry) in Matelea, although noting its morphological distinctness vis-a-vis other subtribal members in the southeastern United States. Taking a broader geographical persp f logy, Rosatti (1989) argued for renewed circumscription of Gonolobus based on the presence y. laminar deal anther append- ages. Most recently the concept of Gonolobus as characterized by short, capitate-glandular, short acicular, and long acicular trichomes, laminar dorsal anther appendages (typically), and smooth, winged follicles has been used by Stevens (2001). However, glandular hairs, although thought characteristic of Matelea by Woodson (1941), also appear without circumscriptional value in the Gonolobus-Matelea question, being pres- ent in both the type of Gonolobus and numerous species lacking dorsal anther appendages (Rosatti 1989). Nonetheless, the monophyly of Gonolobus, whether narrowly or broadly circumscribed, was supported in a recent analysis of chloroplast and nuclear data (Krings et al. 2008). Although parsimony is equivocal on 98 | loft tanical Insti Texas 2( whether laminar dorsal anther appendages evolved once in the most recent common ancestor of Gonolobus s.l., or once in Gonolobus s.s. and once in the Fimbristemma Turcz. group within Gonolobus s.l. (Fig. 2), the character remains restricted to the Gonolobus s.l. clade within Gonolobinae. Interestingly, winged follicles appear to have arisen twice independently and can be considered sy napomorphic for both Gonolobus s.l. (Krings et al. 2008), as well as for a mostly West Indian ocellate-petaled complex that likely includes Matelea acuminata (Griseb.) Woodson, M. bayatensis (Urb.) Woodson, M. correllii Spellman, M. costata (Urb.) Morillo, M. nipensis (Urb) Woodson, M. oblongata (Griseb.) Woodson, M. pusilliflora L.O. Williams, M. rhamnifolia (Griseb.) Krings, and M. tigrina (Griseb.) Woodson. None of these latter taxa exhibit laminar dorsal anther appendages. This putative complex did not evolve within the Gonolobus s. L clade and may deserve indepen- dent generic recognition. Members of the complex tend to share morphological characters apparently rare in Gonolobus s.l., such as reticulate, ocellate corollas. A no E Gonolobus s.l. can be characterized by the pium e two synapomorphic indels in LEAFY, as well as winged follicles (Krings et al. 2008). A p! bus s.s. can be characterized by a synapomorphic transversion in ml. F (thymine to guanine) not sated by any of 154 other Asclepiadoideae species sampled across Secamoneae, Ceropegieae, Marsdenieae, and Asclepiadeae. The former circumscrip- tion (i.e., Gonolobus s.1.) essentially agrees with that of Woodson (1941), Rosatti (1989), and Stevens (2001), and stands in contrast to that of Shinners (1950) and Drapalik (1969). To remain monophyletic however, Gonolobus s.l. must include the Matelea denticulata (Vahl) Fontella & E.A. Schwarz complex. Considering the morphological and molecular distinctions between the three lineages in the Gonolobus s.l. clade (Fig. 2), recognition of three separate genera currently seems preferable to me. MORPHOLOGY A discussion of the morphology of species of Gonolobus s.s. (hereafter simply Gonolobus, unless otherwise specified) is provided below. The discussion concentrates on West Indian Gonolobus species, but when pos- sible provides a broader context of variation within West Indian Gonolobinae. Growth form.—Species of West Indian Gonolobus are twining vines, as are all currently known members of West Indian Gonolobinae (Fig. 3A). Older stems may become somewhat woody, but no collection or live plant has been seen by the author that approached 1 cm or greater in diameter. Individuals may climb as high as a few meters in gaps and openings. Latex.—Latex is known for the following species: Gonolobus iyanolensis Krings, G. jamaicensis Rendle, G. stapelioides Desv. ex Ham., G. stellatus Griseb., G. stephanotrichus Griseb., G. waitukubuliensis Krings, and G. youroumaynensis Krings. It has been primarily described as either milky or white, observations confirmed by the author in the field for G. iyanolensis, G. EE G. E MAN G. Beer G. tar + waitukubuliensis, and G. youroumaynensis. Latex was described by collectors a Puerto Rican collection of a young sterile vine referred to G. las (Acevedo- Rodrigues e» nos 7785, US!). Published studies of latex chemistry in Gonolobinae are unknown to the author, although such inquiry would likely yield interesting results based on studies of other members of Asclepiadoideae (see Uses). Latex chemistry has been applied to solving taxonomic problems in Cynanchinae (Liede et al. 1993; Liede & Kunze 2002). Pubescence.—Most | f subtribe Gonolobinae, exhibit at least two distinct t frequently more. West Indian species of Gonolobus exhibit Du types: (1) multi-cellular, sharp, eglandular, (2) uni-cellular, sharp, eglandular, (3) uni-cellular, glandular-capitate, and (4) papillate (Fig. 3B). Papillae appear to be restricted to the adaxial surface of the corolla lobes—typically borne only on the right side of the lobes. The remaining trichome types may be found anywhere else on the stem, leaves, inflorescences, or flowers, usually in combination. Multi-cellular, um aes Home found on stems are primarily AAI er D types, but IS EP EO OC KE retrorse to retrorse-spreading internodally, but may | opposite petioles. In contrast to some other members of West adimi Gonolebinas É g., M. coniecit Bees, M. linearipetala Alain, M. phainops Krings, M. rhynchocephala Krings, M. torulosa Krings), stem trichomes in West Indian Gonolobus are not borne in two distinct lines. Krings, R iet £r al vu TT r A | " Cd laminar Cd absent Fimbristemma group (e.g. G. calycosus, G. fimbriatiflorus, G. stenosepalus) Rest of Gonolobus s.s "M." denticulata Gonolobinae De > M al af | Er no ik SI A SA , sal I I L: | J L L (fi Kring t al. 2008) E Esa "La. d £ Fars. IE NOS) r A ITT A al 24750 IM anv Ad A ye Is fh rj 2 Dodson 15450, NY) Ca f: | | f H Ci int + H | " mm] t e P J I ai d ge; = onalohirs M. = Matelea dd dy fie MAA s >= ww > Leaves.—In general, species of Gonolobus exhibit simple, opposite, membranous, cordate leaves, al- though a few taxa, such as the Jamaican G. stellatus, G. jamaicensis, and G. stapelioides, exhibit truncate to cuneate leaf bases (Fig. 3A & C). The leaves of G. jamaicensis and G. stapelioides tend to be coriaceous. Leaf apices vary from acute to acuminate. Leaf size and shape may be influenced by position on the stem (e .g., basal vs. terminal) and habitat (e.g., ge vs. Pes interior; sun vs. shade) (Krings, pers. obs.). Laminar colleters.—1n West In G } ies, as in all West Indian Gonolobinae, two or more colleters are borne on the adaxial midrib near the eegen of the leaf base and the petiole apex (Fig. 3D). Colleters vary from deltoid to lanceoloid. Stipular colleters.—In West Indian Gonolobus species, a single colleter is borne on each side of the petiole at its extreme base (Fig. 3E). Colleters vary from deltoid to lanceoloid. Inter-petiolar (borne along a ridge or line between two opposing petioles at a node) colleters are rare and infra-petiolar (borne within the width of a petiole at its base) are apparently absent. Stipular colleters in other West Indian Gonolobinae vary in position, but frequently appear to be associated with the stem rather than the petiole. Inflorescences.—Inflorescences of most West Indian Gonolobinae species, including all Gonolobus species in the area, are extra-axillary and racemose (umbelliform inflorescences appear to be restricted to Haitian Matelea crispiflora (Urb.) Jiménez). Inflorescences can be borne at nodes with immature leaves 100 LI de lla ET ae M1! Fic 3 I r Lak. A T Ak E ir rduccin RT 4 f Miralo charn E A / F He I fa al La tenk 4 nt Toc! walled AN Pu A il Ee roctratual C Variatinn a X / A r ree V3 (gf f i f (cordate: G. i 1 j te: G sfellatus). D. | H iG ES lleters (G. d D d F ` d d e n ep 4 G "m D a7 | zs A H P: Fr ot L BS SR BEES | Pal Ilag if ot. | — vor t Ki f 1 ki f |) d + d ` F f E Mae Infe f£ WER: ig! t* ctanelinidoc) Ki—iil. Variati f | | y g (i G stapelioides; X T F T 7 ti + F ii G dussii iii: G. iamaicensis). L. Liaulat inal tof Genial iG martinicensis). M. V. ti ] F J i kl kel J À i re " e os puis Te A £ A en LAT I £I la dE san tg asi ebtaeh DC Je dieta ll J bat (left: G. rostratus; right: G. stellatus; see also Ki-ii, L). N g (G. ste) ) y j lensis) and entire (6. stephanotrichus). C f | f ll caudicle; Ci g t: Cd P" rato Pe 4 + Da nli A O = porum. we = d D UI | RA AL TA DA Se) Krings, n EN Ze | p | H al war & | ye 101 as short as 13 cm long. The inflorescence is indeterminate and a series of flowers are produced on lateral pedicels from a single axis, the oldest at the base and the youngest at the apex (Fig. 3F). The axis is frequently significantly contracted and may not measure more than 1 cm in some species. In West Indian Gonolobus, bracts are frequently borne at pedicel bases, although they tend to be small and caducous in most species. For additional a review of inflorescence structure in other members of Asclepiadoideae, see Liede and We- berling (1995). Based on specimens of West Indian Gonolobus seen in the course of the present study, one to three flowers may be open at a time. No phenological studies in West Indian Gonolobus have been undertaken and the life span of individual flowers remains unknown. Based on prior studies in Gonolobinae (see Lipow & Wyatt 1998; Krings 1999) and informal observation, it is unlikely that individual flowers last much longer than a week. In Gonolobus suberosus of the southeastern United States, individual flowers were open (2-) 4.61 (-10) days (N = 175; s.d. = 1.53) and inflorescences had open flowers for (2 8.96 (-23) days (N = 57; s.d. = 4.63) (Lipow & Wyatt 1998). Aestivation. Sane EEN of West nd Mid. species are imbricate and dextrorse in bud (Fig. 3G). Imbricat t also i of West Indian Gonolobinae, but not all other genera exhibit dextrorse buds. Calyces.— Calyces of Gonolobus species consist of five, green sepals (Fig. 3H). Apices are g lly nar- rowly obtuse to rounded. Adaxial surfaces are uniformly glabrous. Abaxially surfaces vary from pubescent to glabrous. Frequently, the distal pedicel pubescence (when present) creeps onto the calyx base. Calycine colleters.—West Indian Gonolobus species, as most members of West Indian Gonolobi bear 1-2 colleters in each sepal sinus us 3D. The colleters are DT to cds Corollas.— Corollas inr Indian Gonolobinae, including all G I in the area, are 5-lobed (Fig. 3)). Corollas are late to campanulate at the base with lobes spreading to reflexed. The corolla lobes of SR DCH are EM An exception is Jamaican G. stapelioides, which exhibits wavy and recurved EE EES Corolla lobes of Gonolobus in the area nd overlap at the base and frequently bear a minute gl g in the sinus. Adaxial corolla coloration of West Indian Gonolobus species varies from anio green (most taxa) to burgundy tinged (G. stephanotrichus) to maroon (G. jamaicensis). Pubescence of the adaxial surfaces, when present, is limited to the faucal annulus and surrounding corolla lobe bases, and the right side of the corolla lobes. Corolline coronas: Faucal annuli.—Faucal un sensu Liede and Kunze (1993), non sensu Woodson and Moore 1938 (see Kunze 1990, 1995) —are nular thickenin gs of the tubular por- tion of the corolla (Fig. 3Ki-iii, L). Endress and Bruyns (2000) considered the Ca and Cc (corolline corona) of Liede and Kunze (1993) homologous, although Fishbein (2001) and Kunze (2005) explicitly rejected this hypothesis. Kunze (2005) noted that early ontogenetic EES of even taxa such as G. lasiostemma (Hemsl.) Woodson, which exhibits a Ca of five large fleshy lobes int inally (similar to Jamaican G. stapelioides), show a closed annular meristem at the initial stage. Faucal annuli (Ca) are distinctive in West Indian SCH species and generally much more developed in members of this genus than in other West Indian Kunze (1995) found faucal annuli present in continental Gonolobus spp., as well as Matelea lanata (Zucc.) Woodson, and Matelea dictyantha Woodson, and absent in Matelea tinensis (T. Mey.) Pon- tiroli and Fischeria spp. (all Gonolobinae). He also noted that outside Gonolobinae, al annuli were found mainly in some Stapelieae. The faucal annulus occurs immediately to the outside of the gynostegial corona Oe, closer to the corolla lobes than the center of the flower). Annuli may be "interrupted" and reduced to a ridge opposite each corolla lobe sinus (e.g., G. martinicensis Decne.; Fig. 3L) or “uninterrupted” and well- developed (e.g., G. jamaicensis; Fig. 3Kiii). They are frequently pubescent, but not always, in contrast to the always glabrous, cns APR coronas. Gynostegial ost a oronas are quite varied in West Indian Gonolobinae (Fig. 3Ki-iii, L), bare can bedescubed tly in terms of staminal (Cs) and interstaminal (Ci) segments. The coronas have been interpreted as a “fusion” Games the Cs and Ci (Liede & Kunze 1993), although it remains un- 102 j tl tanical Insti Texas 2( clear whether the combined structure in fact represents a true fusion of disparate elements (see Endress & Bruyns 2000). In Gonolobus, the Cs region typically is raised and ridge-like, often meeting the lower portion of the laminar dorsal anther appendages (Cd). There are no instances of free, ligulate Cs segments as seen in other West Indian Gonolobinae taxa, such as M. ovatifolia (Griseb.) Woodson or M. pentactina Krings, or three-dimensionally complex Cs segments as seen in M. tamnifolia (Griseb.) Woodson and M. oblongata. Ci segments are single and cup-like in most West Indian Gonolobus species, and double (i.e., ligulate) only in G. iyanolensis and G. martinicensis (Fig. 3L). The interstaminal ligules of the double coronas of the latter two species are not positionally homologous with the staminal ligules seen in numerous species of Matelea, such as West Indian M. ovatifolia and M. pentactina, or continental M. pubiflora (Decne.) Woodson. However, some Matelea species, such as M. maritima (Jacq.) Woodson, also exhibit interstaminal ligules. In the Ascle- piadoideae in general and West Indian Gonolobus in particular, the Ci region appears to serve as a holding cup for nectar secreted from a primary nectary on the flanks of the filaments insi de the guide rail and in the stigmatic champer (Christ & Schnepf 1985, 1988). Among West Indian Gonolobus species, defined second- ary nectaries are evident—at least as far as can be determined from herbarium specimens—only in a few species (e.g., G. dussii Krings). Kunze (1995, 1999) also found defined secondary nectaries in continental G. chloranthus Schltdl. and G. fraternus Schltdl., as well as the gonolobinoid Matelea reticulata (Engelm. ex A. Gray) Woodson and M. argentinensis. Secondary nectaries are epithelial, consisting of enlarged epidermal cells enriched with cytoplasm (Kunze 1995). Epithelial nectaries on the filament flanks appear to be wide- spread in Asclepiadoideae (Christ & Schnepf 1985; Kunze 1991; Kunze 1995). The gynostegial stipe is generally edentate, although G. jamaicensis, G. stephanotrichus, and G. yourou- maynensis have distinct teeth or “notches”. The stipal teeth or “notches” of all three species are distinct in shape and occur in idi locations. Their homology is unclear. Laminar dorsal dages.—Laminar dorsal amtner EE (Cd sensu Kunze 1995; Fig. 3Ki-ii, L-M) are restricted to Gondlabus s.l. or s.s., although it remains u whether they evolved once in the most recent common ancestor of the former, or once in the ancestor of the latter and in the ancestor of the Fimbristemma group within Gonolobus s.l. (Krings et al. 2008). Parsimony is equivocal on the question (Krings et al. 2008). The only other genus exhibiting sr anther iM within Gonolobinae (and Asclepiadoideae, fide Kunze 1995) is Fischeria, but the apr ther than laminar (Murphy 1986; Vethacke 1994; Kunze 1995). Laminar dorsal MA vary in shape from truncate to rounded to strongly, divergently bi-lobed. They tend to be spreading, but are conspicuously (and spectacularly) erect in mature flowers of G. stapelioides (Fig. 3Ki). Style-heads.—Style-heads vary conspicuously in West Indian Gonolobinae, but tend to be essentially planar (varying from somewhat convex to somewhat concave) in Gonolobus species in the area (Fig. 3Kii, L). West Indian eons Lundi PED lack terminal style-head appendages, in contrast to other West Indian ies such as itima, M. corynephora, M. rhynchocephala, and M. torulosa (see Krings 2006). Pollinaria.—Pollinaria in Asclepiadoideae consist of a secreted corpusculum and two connected pollinia (pollen sacs; Kunze 1994). The two pollen sacs are the result of a reduction in anther locules from four in the more primitive Secamonoideae to two in the more derived Asclepiadoideae (Kunze 1996; Rapini et al. 2003: Liede-Schumann et al. 2005; Rapini et al. 2006). The pollinia are borne essentially horizontally in West Indian Gonolobus species—as they are in most members of the Gonolobinae—and should be considered broad, rather than long (Kunze 1995; Endress & Bruyns 2000; Fig. 3N). Only a portion of each pollinium is fertile and hyaline, sterile portions are easily visible. The corpusculum in the Gonolobinae (based on studies of Matelea reticulata and M. argentinensis) consists of solid side walls and a thick, three-layered floor (Kunze 1994). Kunze (1994) suggested that the formation of the side-walls originates in two primarily separated strands of secretion which eventually merge into one compact wall. Caudicles are present (Fig. 3N), but Kunze (1994) noted it was difficult to determine whether they are extensions of the outer flanks or new ad- ditions. He suggested the latter, based on the lack of staining with either hematoxylin or safranin, although Krings, Revisi f Gonolobus in the West Indi 103 noting that caudicle initiation in the two Matelea species occurs very early in ontogeny and directly from the corpusculum, rather than apart from it. Follicles.—Follicles in West Indian Gonolo! ies (as in most continental G 2001), when known, are 5-winged (Fig. 30), MM -— murications may be seen d the wings of some taxa (e.g., G. stellatus). Some continental species of Gonolobus also exhibit 4-winged (G. albomarginatus (Pittier) Woodson) or apparently 3-winged (G. incerianus W.D. Stevens & Montiel) follicles (Stevens 2001). Winged follicles are also exhibited by other members of West Indian Gonolobinae, such as M linearipetala, and the putative members of an ocellate complex M. acuminata, M. bayatensis, M. correllii, M. nipensis, M. oblongata, M. rhamnifolia, and M. tigrina. Seeds.—Seeds of West Indian Gonolobus species, as of most West Indian Gonolobinae, are essentially planar and pyriform in outline (distinctly plano-convex in Matelea maritima and M. rubra (H. Karst.) Spell- man & Morillo). Distal margins vary from entire to dentate or crenate (Fig. 3P). Comas are translucent. Tal fide Stevens HABITAT In general, West Indian Gonolobus species occur in small gaps or openings in mid-elevation, moist or wet forests on limestone derived soils. They are absent from high-elevation dwarf forests, coastal thickets, man- groves, marshes, or dry forests. Occurrences are infrequent to rare and likely limited by available habitat. Small gaps in minimally disturbed forests appear vital to seedling germination as vines are rarely encoun- tered elsewhere. Changes in gap dynamics or forest condition, thus could have profound impacts on the maintenance of Gonolobus populations. With the exception of G. youroumaynensis, the species have not been found along roadside edges, where other vines (particularly Ipomoea L. spp. and Dioscorea L. spp.) are gener- ally superior competitors and quickly form thick, tangled thickets. Gonolobus youroumaynensis was found along a roadside patch of vegetation in St. Vincent, although it remains unclear whether the population is competitive there or merely persistent following a colonization event. A population of Gonolobus iyanolensis persists on top of Gros Piton in St. Lucia among boulders. POLLINATION As there have been no pollination biology studies on West Indian Gonolobus species, their pollinators remain unknown. In fact, pollinators appear to be documented for only two species of Gonolobus range wide: G. fraternus (Mexico and Central America) and G. suberosus (United States). Kunze (1999) studied the pollination ecology of G. chloranthus and G. fraternus, but was able to observe putative pollinators only on the latter. Pollinarium uptake was confi ] for two diptera species (Calliphori- dae, Tachinidae), a bee (Apidae), and a wasp (Vespidae) (Kunze 1999). Other observed visitors included Apis mellifera Linnaeus (Apidae) and unidentified butterflies (Lepidoptera) (Kunze 1999). However, pollinarium removal by these could not be verified. Corpuscula were removed by the wasp and bee by their legs, and by the bee and the flies by their probosces. Foraging behavior differed among the observed species, with the wasp and bee prefering upper regions of the studied vine, and the flies, the middle and lower. E (1969) dis P Ke (all Chloropidae) on flowers of G. suberosus, but it remains unclear whether they s due to their size (1. om mm) and lack of pollinaria on ee 1 Lipow and Wyatt (1998) also Se G. suberosus, but despite daily and nightly searches were pollinators. Gonolobus suberosus appears to DESCH considerable variation in the numbers of flowers per individual plant, the number of fl , and the number of infl individual plant (Lipow Sr Wyatt 1998). Only a mall fraction of P monitored by Lipow and Wyatt (1998)—less than 596—were pollinated and per flower removal rate of pollen was only 0.31. This rate was less than previous reports in other asclepiads (Table 1). Ollerton and Liede (1997) noted that subtribe Gonolobinae appears distinct in being predominantly dipteran pollinated, although this claim was based on only five Matelea species in the southeastern United MM UT t tanical i Texas 2( Tase 1. Per flower rates of pollen removal and pollinium deposition within Asclepiadoideae (arranged by per flower rate of pollen removal). Species Per flower rate of Per flower rate Source pollen removal of pollinium eposition Asclepias exaltata L. 0.23-1.38 0.39-0.66 Queller (1985); Wyatt & Shannon (1986); Broyles & Wyatt (1990, 1995) Gonolobus suberosus (L.) R.Br. 0.31 0.043 Lipow & Wyatt (1998) Matelea reticulata (Engelm. ex A. Gray) Woodson 0.54-0.94 -- Liede (1994); Krings (1999) Asclepias curassavica L. 0.62-1.7 0.13 Wyatt (1980); Willson & Melampy (1983); Wolfe (1987) Asclepias syriaca L. 0.62-1 7 -- Willson € Rathcke (1974) Asclepias quadrifolia Jacq. 0.81-241 -- Chaplin & Walker (1982) Asclepias tuberosa L. 1.64 0.71 Wyatt (1978) Funastrum pannosum (Decne.) Schitr. 243-248 0.57-1.52 Kunze & Liede (1991) Funastrum clausum (Jacq.) Schltr. 2.64 2.16 Kunze & Liede (1991) Asclepias solanoana Woodson 2.91-43 1.23-1.81 Lynch (1977) 1.4 E 1] s } fel pe AS | States (Drapalik 1969). The claim remains mostly true today, although g VE species of Fischeria and Gonolobus each has been added (Skutch 1988; Kunze 1999). Two additional studies examined species also studied by Drapalik (1969) and found additional dipteran floral visitors (Liede 1994; Krings 1999). Kunze (1999) found flowers of G. fraternus polyphilic and noted that they were not specialized for a single kind of pollinator. The ASCLEPOL database (http://www.uni-bayreuth.de/departments/planta2/ research. wgl/ pollina/as_pol_d.html) currently also lists pollinators for G. argentinensis T. Mey. (1 sp. of Calliphoridae, Diptera), but this species has most recently been recognized in Matelea. CH INSECTS HOSTED There are no published records of insects hosted by West Indian Gonolobus species known to the author. However, a few records of primarily dipterans and lepidopterans have been published for continental spe- cies in the genus. Castrejon-Ayala and Camino-Lavin (1991) reported Gonolobus sorodius, nom. nud., as host for papaya fruit fly (Toxotrypana curvicauda Gerstaecker)—the principal insect pest of commercial papaya (Carica papaya L.). Norrbom (unpubl.) suggested this plant may be Gonolobus barbatus Kunth. Baker et al. (1944) reported another asclep host plant of papaya fruit fly—a wild species known as talayote or talayotillo in northeastern Mexico (Santa Engracia and Cañon de Rosario). Castrejón-Ayala (1987) suggested the plant may be a species of Gonolobus and Landolt (1994) suggested perhaps G. erianthus DC. Gonolobus salvinii Hemsl.—commonly called champeron in Guatemala—may yet be another host for the papaya fruit fly (Norrbom & Muñiz, unpubl.). Capinera (2005) reported “Gonolobus sp." among hosts for the saltmarsh caterpillar (Estigmene acrea (Drury))—a species ranging from Canada to Central America. An unidentified species of Gonolobus was reported as host for Erinnyis obscura obscura (Fabricius) —a sphingid found in Amazonas, Brazil (Silva Motta & Xavier-Filho 2005). There are a number of reports of Gonolobus laevis Michx. serving as host plant for various lepidopterans, including Danaus gilippus Cramer and D. plexippus (Linnaeus), however G. laevis belongs to the Cynanchinae, not Gonolobinae, and should be recognized as Cynanchum laeve (Michx.) Pers. (Liede & Tauber 2002). SEED DISPERSAL The follicles of known West Indian Gonolobus species are smooth and uniformly 5-winged, dehiscing along a solitary suture between two wings to release the comose seeds. Seeds are very light (Table 2) and can travel on slight breezes. Unfortunately, mature follicles have been do cumented by collections for only three Krings, R ESI Er ILL H al 3AF PE | je 105 of the ten species of West Indian Gonolobus (Table 2). Immature follicles are known for G. stapelioides (see Additional specimens examined). The paucity of fruiting collections is echoed in West Indian Gonolobinae as a whole. Follicles have been collected for less than half of the estimated fifty species in the West Indies (Krings, unpubl.). The low rate of fruit collections may be due to naturally low levels of fruit set in the subtribe (Lipow & Wyatt 1998), as well as collector habit, flowers being necessary for most species level determinations. Lipow and Wyatt (1998) suggested that low fruit set in Gonolobinae may be attributable to pollen limitation. Less than 5 % of flowers monitored were pollinated and only a single fruit matured from 352 flowers on 13 plants (Lipow & Wyatt 1998). CHROMOSOMES There are no published reports of chromosome numbers in West Indian Gonolobus or Gonolobinae. Previ- ously published reports for continental Gonolobinae, including continental Gonolobus, show the same basic chromosome number x = 11 that predominates the subfamilies Asclepiadoideae, Periplocoideae, and Se- camonoideae (Albers & Meve 2001). Only 6% of 672 species surveyed in the three subfamilies by Albers and Meve (2001) were polyploid and deviations (reductions or increases in chromosome numbers) were found only in the Asclepiadoideae. No deviations within the Gonolobinae from 2n = 22 were found by Albers and Meve (2001), although only five species in four genera were included from the subtribe in their survey. USES As there are no published accounts of human uses of West Indian Gonolobus species known to the author, a survey of uses in other areas may be of interest to the reader. The Ecuadorian endemic and IUCN Vulnerable (VU) listed Gonol onus zip mane monto has been reported to be used by the Saraguro native people as a medicinal plant (sp ifi ), under the name sacha ango (Pitman 2003). Unspecified medicinal uses are also reported for boum yucatanensis (Woodson) W.D. Stevens, a taxon described in Trichostelma Baill. (Durán García et al. 1997). Standley and Williams (1969) noted that the tender, young fruits of Gonolo- bus species are commonly used as vegetables in Guatemala, and may be available in local markets under the name cuchamper, although this name has also been applied to the fruits of related genera. Specific uses are noted by Standley and Williams (1969) for G. salvinii (tender follicles eaten raw when almost mature, and cooked and eaten at almost all stages of growth) and G. stenanthus (Standley) Woodson (young follicles boiled or otherwise cooked with sugar to make sweetmeats or dulces). Stevens (2005) noted that "most, if not all, species of Gonolobus have fruits that are eaten when young, either raw or cooked." Numerous studies have examined potential medicinal properties of latex in Asclepiadinae (Asclepias L.: Liggieri et al. 2004; Calotropis R.Br.: Rasik et al. 1999; Kumar et al. 2001; Dubey & Jagganadham 2003; Shivkar & Kumar 2003; Ahmed et al. 2004; Alencar et al. 2004; Kumar & Shivkar 2004a & b; Shivkar & Kumar 2004; Al-Mezaine et al. 2005; Arya & Kumar 2004, 2005; Iqbal et al. 2005; Pahdy & Kumar 2005; Rajesh et al. 2005; Roy et al. 2005; Sehgal & Kumar ae ovate et 2005; ae et al. 2006; Kumar et al. 2006; Ramos et al. 2006a & b; Sehgal et al. 2006), Metast | (Fi n E. Fourn.: Morcelle et al. 2004; Philibertia Kunth: Sequeiros et al. 2005), and Oxypetalinae T Brot.: Priolo et al. 2000; Gaig et al. 2005; Morrenia Lindl.: Vairo Cavalli et al. 2003), ER no ass studies are known to the author in Gonolobinae. CONSERVA TION Human disturbance disproportionately affects infrequent plant species more than common taxa as further reductions of already low population si y not be sustainable (Lawton 1993; Casagrandi & Gatto 2000; Hendrix & Khyl 2000; Benitez Walvis & Martinez Ramos Sch West ud Gonolobus species, as many other species of Asclepiadoideae (Kunze & Liede 1991), appear to be quite i t, if not rare, and are in need of serious survey (particularly in Cuba and the Lesser Alles to Ste a conservation status However, some provisional remarks are possible, based on study of collections and field observations as- sociated with this study. [| [| £ sl Dos H In La as P ET ajai a F 106 J e D RAJ + | UH SCT Fab a E E:xol " r * L L ] is LE E | Te H 2 | EC (e lightest to € seed. S each species. Species Average seed weight Length! Width1 [N = number of seeds sampled] G. Ee (Acevedo- 0.0035 g [N=10] (7.1) 7.4 (7.8) [N=10; s.d.=0.2066] (3.4) 4.23 (4.7) [N=10; s.d.=0.4083] 11445, UPRRP) rigue. G. a (Bellingham 0.0044 g [N=81] (4.6) 5.0 (5.6) [N=10; s.d.=0.2914] (2.5) 2.93 (3.5) [N=10; s.d.=0.2627] Mr G. iyanolensis (Graveson s.n, 0.0116 g [N=26] (8.8) 9.92 (10.5) [N=10; (4.9) 5.38 (5.9) [N=10; s.d.=0.3084] NCSC) s.d.=0.4848] J I J + J WE BE ae I 4 'Format for length and width ts = (Min) Avg (Max) [number of | f furt! tud , but preliminary The habitat requirements of West Indian Gonolol in need of evidence suggests that most eS are Ke limited to Small gaps or openings in mid- dyi, moist or wet forests. They appear to be p long forest edges. The combined infrequency of occurrence of individuals, poor competitive SCH long edges, ely low pollination and fruit set rates (see Pollination and Seed Dispersal SE) ane meee d SE SE or openings in specific forest community types, suggests that Gonolol erable to loss of habitat. This is of particular concern consid- ering the high rate of island endemis — the species. Eight of the ten West Indian Gonolobus species are endemic to single islands (Fig. 1). Gonolobus dussii occurs on two islands and G. stephanotrichus occurs on three islands. Seven of the ten West Indian Gonolobus species are known from five or fewer collections or localities (Table 3). None of the remaining three species appears to be known from more than ten localities. Provisional estimates of conservation status according to the criteria outlined in Version 3.1 of the IUCN Red List Categories and Criteria (IUCN 2001) and Version 6.1 of NatureServe Conservation Status Ranks (NatureServe 2006) are provided in Table 3. Provisional estimates are based on historical collections, the number of known localities, whether taxa are known from protected areas, and inferred habitat or popula- tion changes. These estimates need to be followed by targeted survey to better establish the conservation status of the species. TAXONOMIC TREATMENT Gonolobus Michx., Fl. Bor.-Amer. 1:119. 1803. Tere: G. macrophyllus Michx., nom. illeg. (Vincetoxicum gonocarpos Walter; Gonolobus gonocarpos (Walter) Perry (LecrorwrE, designated by Perry 1938) = Gonolobus suberosus (L.) R.Br. Rhizomatous, perennial vines. Leaves opposite, blades simple, membranous or coriaceous, apices obtuse, acute, or acuminate, bases mostly cordate, but cuneate to rounded in some taxa; colleters present on the adaxial surface at the base of the midvein. Stipular colleters 2, one borne at the base of the petiole on each side. Inflorescences extra-axillary, racemiform. Floral aestivation imbricate, dextrorse. Sepals 5; colleters 1—2 per sinus. Corolla lobes 5, mostly uniformly colored with various shades or green to reddish or ma- roon, sometimes multi-colored and basally maroon or dark purplish and apically green (e.g., G. suberosus), not ocellate, lobes overlapping at the base, glandular emergences present in the sinuses. Faucal annulus (Ca) present, sometimes interrupted and geg ede in ius staminal—or Séier interstaminal (e.g., G. stapelioides) —position. Gynostegial corona inal (Cs) and interstaminal parts (Ci); Cs not foliolate or free, not ligulate; Ci ligulate or not. Laminar dorsal m appendages (Cd) present, reflexed, spreading, or erect. Style-head flat to slightly concave or convex, lacking a defined protuberance or elongate terminal appendage. Pollinaria with pollinia borne horizontally or essentially so. Follicles winged. Seeds essentially plane, distally entire or dentate, comose. 100—150 species (10 spp. in the West Indies as here defined). Perry (1938) chose Gonolobus macrophyllus Michx.—the first of three species listed by Michaux (loc. cit.)—as the lectotype for the genus Gonolobus. However, Michaux cited Vincetoxicum gonocarpos Walter as a Krings, Dn LA ZC e Hz) H D IT & | I: 107 TABLE 3 Number of collecti | localities, distribution, and isional conset tion stat PASSING ONDE spe- J Klat C Cies liti ,then Fue? of collections í IUCN ranks. IUCN conservation M are global. Critically endangered taxa are unknown from protected areas. Species No. of No. of Distribution Provisional conservation localities collections status G. absalonensis 1 1 Martinique Critically endangered (CR); GH NH G. martinicensis 1-3 3 Martinique Critically endangered (CR); GH NH G. youroumaynensis 1-2 3 St. Vincent Critically endangered (CR); G1 N1 dussii 5 3 Guadeloupe Critically endangered (CR); GH NH and Martinique (both islands) G, ivanolensis 2 5 St. Lucia Endangered (EN); G1 N1 G. stellatus 4 4 Jamaica Vulnerable (VU); G3 N3 G. jamaic 4 5 Jamaica Vulnerable (VU); G3 N3 G. See eee 4-8 17 Dominica eli (VU); G3 N3 G. stapelioides 5-7 13 Jamaica east concern (LC); G4 N4 G. stephanotrichus 7-10 19 Greater Antilles pe concern (LC); G4 N4 (all islands of occurrence) GH: Possibly Extinct (species)—Missing; bud from only historical occurrences but still some hope of Maia G1: Critically Imperiled—At very high risk of extinction due to extreme rarity (often 5 or fewer populations), very steep declin or other factors. G3: Vulnerable—At moderate risk of extinction due to a restricted tange, relatively few GE (often 80 or fewer), t MIO Sa praag declines, or other factors. G4 : Appar ntly Secure but not for long-term concern other factors. NH: Possibly Extirpated (Historical) —Species o! | historically in the nation or state/province, and «herbs is some possibility that it may be rediscovered. Its Bus may no have been verified in the past 20-40 years. N1: Critically Imperiled—Critically imperiled in the nation or state/province ie cause of extreme rarity (often J or fewer OCCURANCE or because of some actor ) Suena i ni steep one Making it N2- Vulnera la especiall ly | PO AC a to a restricted EN relatively few populations (often 80 or fewer), recent and widespread declines, or rother factors making it vulnerable to SEH N4: Apparently Secure—Uncommon but not rare; some cause for long-term concern due to declines or other facto J : +l Tati :11 Pes + synonym for his G. macrophyllus, Perry (1938) proposed the combina- tion G. gonocarpos as the correct name for Reen sG. — M Recent study found the entity variously known as G. gonocarpos or G. macrophyllus cons pecific with G. suberosus (Krings & Xiang 2004, 2005). The latter name has priority following Reveal and Barrie (1992). Some Robert Brown names have sometimes been incorrectly dated. In 1810, a preprint of his "On the Asclepiadeae, a Natural Order of Plants separated from the Apocinae of Jussieu" was made available. This same work later appeared in the first volume of the 1811 Memoirs of the Wernerian Natural History Society (Stearn 1960; Stafleu & Cowan 1976; Mabberley 1985). Although sometimes incorrectly cited from the first reading of the paper in 1809, communication at public meetings does not constitute effective publication (Forster 1991; Greuter et al. 2000; McNeill et al. 2006). Names should also not be ascribed to the 1811 appearance of the Memoirs of the Wernerian Natural History Society but rather to the preprint released in 1810. Thus the combination Gonolobus suberosus (L.) R. Br. should be cited as above, rather than in Mem. Wern. Nat. Hist. Soc. 1:35 (1810). As Brown's 1810 combination in Asclepiadeae is validly published (by definite association of the final epithet with the genus name), it takes priority over the Schultes combination in Systema Vegetabilium 6:59 (1820). KEY TO THE SPECIES OF GONOLOBUS IN THE WEST INDIES . Corolla lobes with lateral margins revolute; faucal annulus (corolline corona or Ca) interrupted, strongly EE DS two mounds opposite each corolla lobe, appealing il Oppose each corolla sinus, : pp g ( spent), white, to 4.2 m long, i li tly and sharply bilobed 6. E stapelioides I TU + dorsal at ti IK 108 Journal of the Botanical R h Institute of Texas 2(1) £l | | . hi4 1. Corolla lobes essentially plan or (a) co or i nterupted wen i mos developed into a shallow Mage gehs each corolla colored, < 2 mm long, bi LI al "EMI | GJ ] LII | apices ounce truncate, ied a acini or rounded-bilobed (divergently and sharply bilobed only in G. stellat. tus s). 2. F lus of lla ( lli Ca) an Unine upra Ine Cons pie uoo Tae d c M tall), pubescent along the entire rim or oi each corolla : gynostegial corona Sa fused into an erect ring, ore from ms by the faucal annulus. 3. Gynostegial stipe edent 3. -o stipe dentate. Adaxial corolla SC purple-red to marroon; gynostegial stipe teeth appearing as notches, borne on the lower portion of the column, just above the upwardly rising segment of each Cs; Jamaica . G. jamaicensis 4, due corolla Opes various shades of green; E I ud ee not appearing as BEE ES o on ofthe colum | y 9. G. waitukubuliensis of each Cs; Cuba, Hispaniola, Puerto 8. G. stephanotrichus 2. Frauen mate of corolla (paroline corona or rca peace? to an ER nage ll y RAN tall; I EX FÉ | ] E iL £. | j al corona t bas ally fused into an erect ring that i view by 5. Interstaminal gynostegial corona double (i.e, a narrower upper ligulate corona lobe occurring on top of the broader lower corona). Corolla ro SC lla tul | | | tl izontal length of low | | tegial corona (Ci) dE base of supe to o lobe: apex l; 1- 1 A mm, Vi row upper uae corona lobe ‘ofthe Gi abruptly g cP G. iyanolensis the stipe base, dorsally ridged 6. corolla slender, base Br corolla Tube Fonds See EE longer than broad; horizontal , narrow upper ligulate corona lobe ofthe Ci ending essentially at the stipe base, dorsally plane or sometimes slightly mp raised into a shallow bum 5. G. martinicensis 5, deck gynostegial corona single. ae stipe with a single tooth DROW each anther. along th g; teeth d stipe borne on lower portion of the column, justa bove the upwardly risi Cs 8. G. stephanotrichus 8. Faucal annulus pubescent only in the staminal position; ert of gynostegial stipe borne on upper O of the column, just below the anther 10. G. youroumaynensis 7. P dd stipe edentate. pub dorsal anther appendages (Cd) truncate or rounded 2. G. dussii “Gs a (Ci) smooth, lacking raised | ps; laminar dorsal anther appendages (Cd) emarginate. 10. Leaf bases uniformly cuneate; laminar dorsal anther appendages (Cd) slightly emarginate to e d ana EE bilobe 7. G. stellatus eaf | dorsal anther ap lag led bilobed 1. G. absalonensis REGIONAL KEYS TO THE SPECIES OF GONOLOBUS GREATER ANTILLES 1. Corolla lobes witl gi lute; faucal lline corona or Ca) interrupted, strongly developed into t ds opposite each corolla lobe, Deer pposite eac a sinus, pul t; lami lorsal t| | t ( li t), white, pi li g tly ES | ly bilobed, dch Y | to 4.2 mm long; Jamaica 1. Corolla lobes essentially plane, sometime reflexed, but not lute; f | lus ( lline corona or er + J SÉ m A +! A ro | IER I lH a Hi l^ I lal | lorsal ant lages (Cd) d ling or sj ling, vari y colored, apices rounded, truncate, d-bilobed (divergentl | sharply bilobed only in G. stellatus), « 2 mm long. ilg emma: or rounde | (corolli Ca): En ‘uninterrupted ring, EIERE iid Os 4 er 9mm ally tall), pubescent along the entire rim or only OPPOSE E e each Ker fused into an erect ring, view by Krings, Revision of Gonolobus in the West Indi 109 3. Adaxial corolla lobes purple-red to marroon; gynostegial stipe teeth appearing as notches, borne on the lower portion of the column, just above the upwaraly rising segment of each Cs; Jamaica 4. G. jamaicensis d 3. ^ 2. H ll l ] E ` " j . ic _instea truncate, borne on the lower portion of the column, just above the upwardly rising ce of each a, Hispaniola, Puerto Rico G. stephanotrichus Cs 2. STE gece of TA (corolline corona or Ca) reduced to an interrupted Ge distinct St apposite each corolla lobe sinus, tufted pubescent to glabrate or, if uninterrupted, then very shallow tall; gynostegial corona not basally fused into an erect ring that is obscured SC view by de faucal an- nulus 4. Leafl if ly te; gy tegial stit lentate; lami | | anti D lages (Cd) slightly emarginate to divergently and sharply bilobed; Jamaica. 7. G. stellatus 4. Leaf bases cordate; gynostegial stipe with a single tooth borne on lower portion of the column, just above the upwardly rising segment of each Cs; laminar dorsal anther appendages (Cd) truncate or rounded; Cuba, Hispaniola, Puerto Rico 8. G. stephanotrichus LESSER ANTILLES 1. Faucal annulus of corolla oles corona or cal an di ring, conspicuously raised (to 0.5 mm tall), Cc ddl XI only PI LAIR bel Po A densely folded avicallvibe basally fused into a ring, ol | from view by the faucal lus; gy | E 1 id mm tall a emerginate; EE 9. G. waitukubuliensis corolla lobe sinus, tufted pubescent to abae or, if uninterrupted, then very y shallow, PM 0. 08 r mm elt and glabrous, except to 0.16 mm tall and tufted pubescent opposite each corolla lobe sinus; gynostegial corona neither Ee densely folded nor basally Gees by a faucal annulus; gynostegial stipe < 1 mm tall; Cd rounded or truncate (emarginate in G. absalon 2. mestarina Ee corona double (e, a narrower upper ligulate corona lobe occurring on top of i SPIONAT coronas "ho: E e ). 3. E bus | lat | las | broader than | i | length of lower interstaminal SE corona (ci from base of stipe to Seen apex t 1- d 4 mm, Tr row upper ligulate corona lobe of the Ci abruptly g the stipe base, dorsally ridged; St. Lucia Ro iyanolensis 3. Corolla slender, base of corolla fuge elongate Campanula EE EH Dn broad; horizontal length of int 7 mm, narrow upper ligulate corona lobe ofthe Ci ending essentially atthe stipe base, dorsally plane or sometimes s raised into a shallow bump; Martinique 5. G. martinicensis 2 Hide gynostegial corona single. 4. Gynostegial stipe with a single tooth ca. 0.2 mm long below each anther; St. Vincent 10. G. youroumaynensis 4. Gynostegial stipe edentate. 5. le oe Ge a ui mm ML E i Dime ll; interstaminal gynostegial corona (Ci) with two distinc | ant! ppendag (Cd) truncate or rounded; ECG and Martinique 2. G. dussii 5: coroll lones men aa: mm ong: Cs < 0.3 mm tall; interstaminal gynostegial corona (Ci) smooth, | dorsal anther appendages (Cd) rounded bilobed; Martinique 1. G. absalonensis 1. Gonolobus absalonensis Krings, Syst. Bot. 32:181. 2007. (Fig. 4). Tyee: MARTINIQUE: Absalon, Pres la cascade, May 1910 (fl), Herb. d'Alleizette s.n. [4801?] (HoLotyee: L!). Herbaceous perennial vines. Latex unknown, presumably white. Stems glal to very sparsely pubescent, both short, capitate-glandular, and longer, sharp, eglandular trichomes present, the latter retrorse internod- ally and antrorse-appressed along an indistinct, horizontal ridge between two opposite petioles and/or just above the node, to 0.5 mm long; nodes sparsely pubescent, piane held EE ues Leaf blades ovate to oblong-ovate, (3.4-)6.6-8.6 cm x (1.3-)3.4—4.5 cm, api tly acuminate narrowly obtuse, to 0.7 cm SCH bases deeply or shallowly cordate, margins entire, adaxial surface sparsely strigose, a rere } T NEA, MR 1 A +] abaxial y StT igose with shar B; eglandular t along y or AK Fic. 4. G | ic A | 1 infil R Open flower. C. E f tl Il (C ), gy t gi | , and style-head nr r 4 ral EE J A 1 Liri TER A 5 NI t- fetia L J A Y! E I ol. linarium. Based on Herb. d'Alleizette s.n. [48077] (L). Ca = faucal lus of corolla; Ci = interstaminal gynostegial g ; Cd = laminar J | Al J F ud S n E as 71 A H |] a LEM | A ge; Co = corpusculum ; Cs gy g g veins, colleters 2, 0.5-0.9 mm long; petioles 3.6—5.7 cm long, glabrous or very sparsely pubescent on all sides, capitate trichomes to 0.1 mm long, sharp, eglandular trichomes ca. 0.3 mm long, antrorsely-appressed or -ascending; stipular colleters 2, ca. 0.3 mm long, one borne on each side of the petiole base. Inflorescences racemiform, peduncles 4.1—5.4 cm long, glabrate to very sparsely pubescent, capitate, as well as sharp, eglandular trichomes present, the latter antrorsely-appressed or —ascending, found primarily at the apex, ca. 0.3 mm long; pedicels 1.5-1.7 cm long, more pubescent at the apex than at the base, capitate-glandular trichomes ca. 0.05 mm long, longer, sharp, eglandular trichomes antrorsely-appressed or —ascending, ca. 0.13 mm long, bracts linear-lanceolate, ca. 0.9 mm x 0.21 mm long, caducous, adaxial surface glabrous, abaxial surface coarsely pubescent, trichomes sharp, eglandular, ca. 0.26 mm long, antrorse. Calyx lobes 5, n a a E e Ial "oa xl 3AF TO J; 111 Krinas g5, lanceolate, 1.95 mm x 1 mm, apices narrowly obtuse or acute, margins glabrous, abaxial surface pubescent mostly at the base, sharp, eglandular trichomes, antrorsely-ascending or -appressed, to 0.25 mm long; col- aa 1 pes sinus. Corolla lobes 5, lanceolate to ovate, 4.8—6.7 mm x 2.5-2.8 mm, MÀ es at Es ES a g frequently present in the sinus, adaxial glabrous, abaxial surface (cora hme corona or in a shallow, uninterrupted ring, ca. 2.5 mm diam, ca. 0. 08 m mm tall, EN SS where raised opposit h corolla lobe sinus, then to ca. 0.16 mm tall and short hispi d; g5 gynostegial corona fused al (Cs) and interstaminal parts (Ci), single, 5-lobed, erect- or prostrate- MET 0.4—0.6 mm tall; anther guiderails without appendages; laminar dorsal anther appendages (Cd) 0.48-0.7 mm x 0.5-0.8 mm, emarginate; style-head ca. 3.2 mm diam, stipe 0.3-0.5 mm long, not toothed. Pollinaria: corpuscula ca. 0.18 mm long, pollinia borne horizontally, ovate, ca. 0.9 mm x 0.3 mm. Follicles unknown. Phenology.—Flowering in EN Distribution. — Gonolobus | is is endemic to Martinique (Fig. 1). Its habitat is poorly known, but likely primarily middle elevation rainforest based on the requirements of most congenerics in the Lesser Antilles. The type was apparently collected near a waterfall. Provisional conservation status —IUCN: Critically endangered (CR) —Bla,biii. NatureServe: GH NH. Taxon concept mapping.— « G. martinicensis sensu Duss (1897); « G. martinicensis sensu Schlechter (1899); < G. martinicensis sensu E (1988); < G. martinicensis sensu Fournet cd d Notes.—Gonolobus al llest-flowered Lesser Antillean (corolla lobes reaching only to 6.7 mm) and one of tl Gonolol ies known from Martinique. It can be disti "P from other Gonolobus species on Martinique by the e following a 1. Corolla lobes to 6.7 uu long, both surfaces glal ostegial corona (Ci) single, smooth, lacking dorsal anther appenaa (Cd) emarginate 1. G. absalonensis . Corolla lobes > 9. 5r mm n long, e either | ial (Ci) net or if single, then with defined ae bumps medially: laminar dorsal anther “appendages (Cd) rounded or runcate 2 Interstamina eee corona mg 2. G. dussii de ninal gy e (i.e. a narrower upper ligulate corona lobe occurring on top of the broader lower corona) 5. G. martinicensis 2. Gonolobus dussii Krings, Syst. Bot. 32:183. 2007. (Fig. 5). Tere: GUADELOUPE: Chemin de la Soufrière, alt. 500 m, s.d. (fl), Quentin 732 (HOLOTYPE: Pl; Isotype: GH!). Herbaceous perennial vines. Latex unknown, presumably white. Stems glabrate to very Sparen PRESSE both short, capitate- ae and longer, sharp, eglandular tricl I t, the and antrorse-appresse g an indistinct, horizontal ridge | two opposing petioles and/or just above the node, to 0. 5 mm long; nodes pubescent or glabrous, alatid field apparently absent. Leaf blades ovate to oblong-ovate, (3.3-)5.7-10.7 cm x (0.9-)2.5-6.1 cm, apices gradually acuminate with the acumen narrowly obtuse, to 2 cm ee bases deeply or spud d cordate, margins entire, adaxial surface Ge or S 1 glabrate or mo Mis aer + LE als m —— A i PA e win snar P, eglandular t stri gose, major and minor veins, colleters 2-3, 0.4-0. 6h mm be petioles (1.3-)2.1-5.5 cm long, glabrous or noe pubescent on all sides, capitate trichomes to 0.09 mm long, sharp, eglandular trichomes 0.3-0.4 mm long, mostly antrorsely appressed or ascending (some spreading and a very few retrorse); stipular colleters 2, ca. 0.3 mm long, one borne on each side of the petiole base. Inflorescences racemiform, peduncles 0.7-2.3 cm long, glabrate to very sparsely pubescent, capitate, as well as sharp, eglandular trichomes present, the latter antrorsely-appressed or -ascending, found primarily at the apex, ca. 0.3 mm long; pedicels 1.8—2.2 cm long, more pubescent at the apex than at the base, capitate-glandular trichomes ca. 0.08 mm long, longer sharp, eglandular trichomes ucc or -ascending, ca. 0.3 mm long, bracts linear-lanceolate, ca. 1.5 mm x 0.17 mm long, caducous, ad glabrous, abaxial surface ly pubescent, trichomes sharp, eglandular, ca. 0.2 mm long, antrorse. Calyx lobes 5, beso to elongate triangular (widest at or just above the base), 3.9-5.8 mm x 1-1.5 mm, apices obtuse to narrowly rounded, margins glabrous, abaxial surface coarsely pubescent at the base and variously toward the apex, trichomes antrorsely-appressed or 112 A ~~ en LM EE == TRA PS PRE ARO um ET = eme ~ Pn is. - Fic. 5, Gonolobus dussii. A. | | infi B. Fl howing | pul f b Do lobes. C. Faucal | Beie corolla (Ca), g , and style-head. D. Gynostegial f f inal (Cs) le-head remove | ppend 1). F Pollinarium. A- ` based on Duss 3775 (NY). C and EF based on Duss 4565 (NY). » based on add 7 32 (P). Ca l = staminal gynostegial corona segment: Krings, R isi fG lobus in the West Indi 113 -ascending, to 0.5 mm long; colleters 1 per sinus. Corolla lobes 5, du lanceolate to elongate triangular, (10 Kevan: mm x 3.23.7 mm, ERE | pping at the base, a gland lling frequent ly present in the sinus, adaxi face glabroi face with coarse, haen, eglandular homes antrorsely-appressed or -ascending, De at the base and variously toward the apex, ca. 0.36 mm long; faucal annulus (corolline corona or Ca) interrupted, a distinctly raised ridge opposite each corolla lobe sinus, short-hispid or glabrate; gynostegial corona of fused staminal (Cs) and interstaminal parts (Ci), single, erect-undulating, two distinctly raised and rounded mounds borne in the interstaminal position; anther guiderails without appendages, laminar dorsal anther appendages (Cd) ca. 0.8 mm wide, rounded or emarginate; stvle-head 2.3-2.4 mm diam, stipe ca. 0.45 mm long, not toothed. Pollinaria: corpuscula 0.18—0.3 mm long, pollinia borne horizontally, ovate, ca. 0.5-1 mm x 0.25-0.35 mm. Follicles unknown. Phenology.—Flowering in April, June, and October. Distribution.—Gonolobus dussii occurs in Guadeloupe and Martinique (Fig. 1). Its habitat is not well known, but it appears to have been collected from riversides and at mid-elevations (480—500 m). Provisional conservation status. IUCN: Critically endangered (CR)— Bla,biii. NatureServe: Guadeloupe (GH NH); Martinique (GH NH). Taxon concept mapping. — < G. martinicensis sensu Duss (1897); < G. martinicensis sensu Schlechter (1899); « G. martinicensis sensu Howard (1988); « G. martinicensis sensu Fournet (2002). Notes.—Gonolobus dussii can be distinguished from G. youroumaynensis of St. Vincent by the short pedicel trichomes, the edentate stipe, and the adaxially glabrous corolla lobes. The gynostegial coronas of G. absalonensis, G. dussii, and G. youroumaynensis are most similar to each other among Lesser Antillean Gonolobus species, but G. dussii uniquely exhibits two distinctly raised and rounded mounds in the inter- staminal position (Fig. 5). The mounds appear very close and near ridge-like in Martinique specimens, as well as Duss 3775 of Guadeloupe. Additional collections are needed to determine the extent of variation in the coronas and to evaluate whether additional taxa are represented. Additional i ined Bord de! Noire, 29 Jun 1895 (£D, Duss 3775 (MO-fragment, NY); Pointe-Noire, 20 Oct 1895 (£D, Duss 3714 (NY-2 sheets). MARTINIQUE: Environs a St. Pierre, 1884 (fD, Duss “1862, 4565" (NY); Pointe-Fine, prés de la riviere Calabre, Apr 1900 (fl), Duss 4565 (NY). 3. Gonolol lensis Krings, Syst. Bot. 32:185. 2007. (Fig. 6). Tre: ST. LUCIA: Gros Piton, 9 Aug 1996 [*9/8/76"] (£D, CEU 107 (HOLOTYPE: GH. Herbaceous oes vines. Latex white. ge S BAbrous or glabrate, potki short, capitate-glandular, and lon- 1 D ger, sharp, eglandular t nternodally, and antrorse-appressed along an indistinct, horizontal EU bet two opposing dicm to 1.2 mm long; nodes pubescent or glabrous in age, gland field sometimes present. Leaf blades ovate to Be to 8 cm x 5.4 cm, apices obtuse or short-acuminate with the acumen obtuse, bases cordate, margins entire, adaxial surface glabrous abaxial surface glabrous or with sharp, eglandular trichomes — SEN the major veins, colleters 2— 4, 0.37-0.65 mm long; petioles to HE 2 cm long, glabrous or glabrate, capitate-glandular trichomes very sparse if present, sharp, eglandular tricl apex if present, to 0.9 mm long; stipular colleters 2, ca. 0.3 mm long, one borne on each side of the petiole base. Inflorescences racemiform, peduncles 2.1-2.8 cm long, glabrous or glabrate, sharp, eglandular hairs antrorsely-appressed or KEE most Ge Dear apex, ca. 0.7 mm long; pedicels 2.8-3.1 cm long, glabrate, capitate-glandular tri 1 throughout or only near the apex, ca. 0.09 mm long, longer, sharp, eglandular hairs antrorsely- appressed or -ascend- ing, occurring "d near the apex, 0.3-0.9 mm long, oa linear-lanceolate, ca. 2.8 mm x 0.5 mm long, caducous, adaxial surface glabrous, abaxial surface ly pubescent, trichomes sharp, eglandular, ca. 0.4 mm long, antrorse. ae lobes 5, lanceolate (widest distin ly above the base, ca. 1.5-2 mm above base), 6.1-6.6 mm x 1.6—1.8 mm, apices obtuse, margins glabrous, abaxial surface glabrous or glabrate with the few trichomes present, coarse, antrorsely-ascending or -appressed, to 0.4 mm long: colleters 1 per sinus. Corolla lobes 5, lanceolate, 10.2-17.3 mm x 2.7-4.8 mm, slightly lobed at the base, a glandular swelling frequently present in the sinus, adaxial surface pubescent along the right half, trichomes flat, ribbon-like, a rr P . IR dida e £ 114 | l of tl hl Texas 2(1) AK De A "m" P.L H a D A [| (EBEN. R Onan flawar C FI L D J . | L EA Ia Em le nr Faneal annulus ofthe corolla (Ca) t | G | Ef mal (Cd and i timinal ffi TE SE ES hazd cc 1 AA J + » L D ` ei r , and style-head E t UM SEY eU ^ g g pper ligulat lobe of the Ci. F. To f gynostegial (style-head removed). G. Cross-section of corolla tube howing position of coroll 1 gynostegial | ts. H. Pollinarium. I. Follicle. J. Seed. A-B based on Johnny 57 et al. (UPRRP). Cand E based on Graveson 107 (GH). D and F-H based on Krings 1373 et al. (NCSC). |-J based on G. (NCSC). Ca = faucal lus of corolla; Ci — interstaminal gr Vw MM NA À f Sens be É, k Jg Pr d vd Krings, D E E e LL D ail TT £d pe 115 0.18-0.26 mm long, abaxial surface glal f ] lus ( lline corona or Ca) interrupted, a distinctly raised ridge, opposite each corolla lobe sinus, short hispid; nose corona of fused staminal (Cs) and interstaminal parts (Ci), double, 5-lobed, prostrate, narrow upper ligulate corona lobe of the Ci appressed to the lower, abruptly ending in a scooped out depression before reaching the stipe base (horizontal length of lower interstaminal gynostegial corona (Ci) from base of stipe to lobe apex 1.1-1.4 mm), dorsally ridged; anther guiderails apparently without appendages, laminar dorsal anther appendages (Cd) 0.5-0.6 mm wide, rounded or truncate; style-head ca. 2.5 mm diam, stipe ca. 0.23 mm long, not toothed. Pollinaria: corpuscula 0.16—0.18 mm long, pollinia borne horizontally, ovate, ca. 0.5-0.7 mm x 0.25-0.35 mm. Fol- licles ovoid, 10-11.6 cm x 3.3-4 cm, 5-winged, appearing glabrous, but with minute papillae or capitate trichomes evident at high magnification; seeds pyriform, compressed, not plano-convex, 8.8—10.5 mm x 4.2-5.9 mm, glabrous, margins dentate, coma to 3.7 cm long. Phenology.—Flowering in March, July, August, and September. Fruiting in January. Distribution.—Gonolobus iyanolensis is apparently endemic to St. Lucia (Fig. 1), occurring in seasonal deciduous woods from 180—400 m in elevation. An established population occurs on Gros Piton. Roger Graveson (pers. comm.) noted having seen vines away from the Pitons, but that they have not established themselves well. Woodland boundaries and recent forest openings appear to be the prefered habitat. Provisional conservation status. — IUCN: Endangered (EN)—Bla,biii. NatureServe: G1 NI. Taxon concept mapping. — < G. martinicensis sensu Schlechter (1899); < G. martinicensis sensu Howard (1988 Notes.—Roger Graveson (pers. comm.) noted that the collection date is i t on the label of his col- lection Graveson 107 (GH) and should instead be coe ree Gonolobus iyanolensis is i liately recognizable among Lesser Antillean Gonolol inal gynostegial corona (Ci) The ony other species that shares this corona pe in the a area is G. martinicensis. (Fig. 8). In contrast to the robust-flowered G. iyanolensis, G. martinicensis exhibits distinctly more slender flowers with elongate-campanulate bases (ca. 2 mm long) that are longer than wide (as wide as long or wider than long in G. iyanolensis) and a narrower gynostegial corona (see key). The distinct dorsal ridge of the narrow upper ligulate corona lobe of the Ci in G. iyanolensis has also not been seen in G. martinicensis, which is either plane in the homologous position or exhibits a a sightly raised bump. Addit ined. ST. LUCIA: C] in, edge of clearing, Esa 1987 (fr), Slane 1056 (GH); Soufrière, Gros Piton summit, facing Saunier: 5 Jul 2005 (D, Johnny 57 with S l and Sealys (UPRRP friére, Gros Piton summit, facing Soufriére, 13 Sep 2005 OD. Graveson 1248 with Smith (UPRRP); Gros Piton, 11 Mar 2006 d. Kings 1373 with Graveson and Smith (NCSC). 4. Gonolobus jamaicensis Rendle, J. Bot. 74:345. 1936. (Fig. 7). Tire: JAMAICA: near Vinegar Hill, climbing up trees to height of 20 ft, 4200 ft elev., 5 Jun 1896 (fl), Harris 6368 (HOLOTYPE: BM!). Herbaceous perennial vines. Latex white. St pul t, both short, capitate-glandular, and longer, sharp, eglandular trichomes present, the latter spreading or retrorse internodally, and antrorse-appressed along an MOLDE horizontal ridge between two opposing petioles and/or just above the node, to 1.8 mm long; ] t, gland field apparently absent. Leaf blades elliptic to ovate, (2—)3.9-9.7 cm x (1.3-)1.8-6.6 cm, TR acuminate, acumen narrowly obtuse, to 1.1 cm long, bases cuneate, rounded, truncate, or cor- date (frequently all base types present on a single individual vine), margins entire, adaxial surface glabrate, abaxial surface sparsely pubescent to glabrate, sharp, eglandular trichomes relatively dense along the major and minor veins, colleters 2, ca. 0.45 mm long; petioles 0.9-3.5 cm long, pubescent on all sides, capitate trichomes to 0.13 mm long, sharp, eglandular trichomes ca. 1.25 mm long, mostly spreading or retrorse, less dense than the capitate trichomes; stipular colleters 2, ca. 0.29 mm long, one borne on each side of the petiole base. Inflorescences racemiform, peduncles to 4.3 cm long, pubescent, capitate, as well as sharp, eglandular trichomes present, the latter spreading, retrorse, or antrorsely-ascending or —appressed, ca. 0.5 mm long; pedicels 1.2-1.4 cm long, evenly pubescent throughout, capitate-glandular trichomes ca. 0.08 mm long, longer sharp, eglandular trichomes antrorsely-appressed or -ascending, ca. 0.4 mm long, bracts linear-lanceolate, ca. 1.7 mm x 0.5 mm long, caducous, adaxial surface glabrous, abaxial surface coarsely 116 | l of ti tanical Insti Texas 2( AK Fic. 7. Gonolobus jamaicensis. AE | I infl R Open flower C Sepal D Calycine colleter. E. F | tl Il , gynos- tegial corona, and style-head. F. Detail of faucal | b tegial stipe. G. Laminat dorsal anter f appendage: i Corpusculum : and subtending anther wings: l. Pollinarium. wë a on n Kelly et al 10207 (Tc CD). A | on Proct I ta "1 H 1 i a " Y Hi Ye WWE a y Bei kÄ kl kl Po = pollinium. pubescent, trichomes sharp, eglandular, ca. 0.3 mm long, antrorse. Calyx lobes 5, lanceolate, 4.3-7.45 mm x 1.6-2.2 mm, apices obtuse, margins glabrous or sparsely ciliolate, abaxial surface pubescent at the base and glal toward the apex, trichomes antrorsely-appressed, largest to 0.5 mm long; colleters 1 per sinus. Corolla lobes 5, lanceolate, 10—13 mm x 3.8—5 mm, slightly overlapping at the base, a glandular swelling frequently present in the sinus, adaxial surface pubescent along the right, abaxial surface glabrous; faucal Krings, n HESS Es Lat H AL LAJ a. | ys 117 annulus (corolline corona or Ca) a continuous ring, bilobed opposite each anther, ca. 0.93 mm tall, short- pubescent and papillate throughout; gynostegial corona of fused staminal (Cs) and interstaminal parts (Ci), single, erect-undulating, interstaminal secondary nectaries apparently absent or at least not formed into conspicuous bumps or mounds; anther guiderails without appendages, laminar dorsal anther appendages (Cd), ca. 2.3 x 1.7 mm, spreading, apices truncate; style-head ca. 3.3 mm diam, stipe ca. 1 mm long, den- tate, teeth appearing as notches, borne on the lower portion of the column, just above the upwardly rising segment of each Cs. Pollinaria: corpuscula ca. 0.18 mm long, pollinia borne horizontally, ovate, ca. 1.2 mm x 0.3 mm. Follicles unknown. Phenology.—Flowering in March and June. Distribution.—Gonolobus jamaicensis is endemic to Jamaica. It is known only from montane rainforests from 1025-1200 m elevation. Provisional conservation status —IUCN: Vulnerable (VU)—Bla,c. NatureServe: G3 N3. Taxon concept mapping.— = G. rostratus sensu Schlechter (1899); = G. jamaicensis sensu Adams (1972). Notes.—Gonolobus jamaicensis was proposed by Rendle (loc. cit.) as a nomen novum for the Jamaican endemic t Schlechter (1899) treated as G. rostratus (Vahl) R.Br. Not only did Robert Brown never make the combination attributed to him (see Mem. Wernerian Nat. Hist. Soc. 1:35. 1810), but the true Gonolobus rostratus (Vahl) Schult. is only known from Trinidad (and perhaps South America). It is based on the type of Cynanchum rostratum Vahl. Because Rendle corrected a misapplication, his name—G. jamaicensis—is a new species name, not a nomen novum as he mistakenly stated. A nomen novum is an avowed substitute (replacement name) for a validly published but illegitimate name, the type of which would be the same as that of the name which it replaced. Addit ined. JAMAICA: 1886 (fl), Hart 968 (MO D single l 12 flowers]; US); John Crow Mountains, eastern Jamaica, SE a Millbank, 17 Mar 1992 (fl), Kelly, Iremonger and REA team 10207 (TCD). Portland P. i N of Hard the waterfall, 22 Jun 1952 (fl), Proctor 6828 (A, IJ); 0.5 mi N of Hardwar Gap, uphill from the trail above the “Waterfall,” 3 Ms 2006 (st), Krings 1393 with Suiter and Proctor (IJ, NCSC) 5. Gonolobus martinicensis Decne. in DC., Prodr. 8:595. 1844. (Fig. 8). Tw: MARTINIQUE: s.d. (fl), Pleé s.n. (LecroTwrE, designated by Krings 2007: Pb. "Gonolobus scandens (Aubl.) Urb.," Repert. Spec. Nov. Regni Veg. 16:151. 1919, nom. illeg. (see Nicolson 1991). Herbaceous perennial vines. Latex unknown, presumably white. Stems villous to glabrate, both short, capitate-glandular, and longer, sharp, eglandular trichomes present, the latter retrorse or spreading inter- nodally, and antrorse-appressed along an indistinct, horizontal ridge between two opposing petioles, to 1.9 mm long; nodes pubescent or glabrous in age, gland fields sometimes present. Leaf blades ovate to oblong- ovate, to 6.4 cm x " 9 cm, SR obtuse, FR to E acuminate with pa acumen to 1.4 cm long, bases date, margins entire , adaxial ui al LIO SC , abaxial t along the veins, trichomes D Ge (and longer sharp, eslandolar hor EECH done the midvein), colleters 2, 0.7 mm long; petioles to 4 cm long, villous, capitate-glandular trichomes sparse but throughout, sharp, eg- landular trichomes ubiquitous, mostly antrorse-ascending and spreading, to 1 mm long; stipular colleters 2, ca. 0.24 mm long, one borne on EES Se e the SE base. Inflorescences racemiform, peduncles 1.5-2.1 cm long, villous with sharp, egland ly- or retrorsely -appressed or —ascending/-descending and spreading, ca. 0.44 mm long, capitate trichomes throughout; pedicels to 1.9 cm long, less pubescent than peduncles, capitate-glandular trichomes throughout, ca. 0.09 mm long, longer, sharp, egl antrorsely- appressed or -ascending, throughout, though most dense at the apex, ca. 0.47 mm long, bracts linear-lanceolate, ca. 1.6 mm x 0.17 mm long, caducous, adaxial surface glabrous, abaxial surface coarsely pubescent, trichomes sharp, eglandular, ca. 0.2 mm long, antrorse. Calyx lobes 5, oblong lanceolate, 4.5-5 mm x 0.8 mm, apices obtuse or acute, margins sparsely ciliolate, abaxial surface glabrate with the few trichomes present, coarse, antrorsely-ascending or -appressed, to 0.3 mm long; colleters 1 per sinus. Corolla lobes 5, lanceolate, 9.7-11 ine mm x id 5 mm, ES lobed at the base, a glandular swelling frequently present in the sinus, adaxial ight half, trichomes flat, ribbon-like, ca. 0.2 mm long, abaxial L o 118 Corolla lobe PE ; » n ri L e aa s..l r I I ifn\ tanial Fic H F IL A I J R H C 1 EP, bi 2l and T P g dnmiml ictula-haad ramavad) EG Pe | EE 1 n4 Sa ol ff- A 4 ban + style- -head. D gy g ) y g fonda hand vam nado el lalat Inhanfthafi El : J J (Cd G Pollinarinm ROS on Herb Mart af Suppl Nr. 63 (MO). Band ES based on Leber sn (M). c4 D and G based on L.C. Rich. Herb EE hntillanum ( ) wt a A ee? VPE 119 Krinas -F F surface glabrate with a few, sparse, sharp, eglandular trichomes near the base, middle, and margin along the tip; faucal annulus (corolline corona or Ca) interrupted, a distinctly raised ridge, opposite each corolla lobe sinus, short hispid; gynostegial corona of fused staminal (Cs) and interstaminal parts (Ci), double, 5-lobed, prostrate, narrow upper ligulate corona lobe of the Ci appressed to the lower, ending essentially at the stipe base (horizontal length of interstaminal gynostegial corona (Ci) from base of stipe to lobe apex < 0.7 mm), dorsally plane or sometimes raised into a shallow bump; anther guiderails apparently without appendages, laminar dorsal anther appendages (Cd) 0.2 mm wide, rounded or truncate; style-head 1.8-2.3 mm diam, stipe ca. 0.28 mm long, not toothed. Pollinaria: corpuscula 0.1-0.15 mm long, pollinia borne horizontally, ovate, 0.45—0.6 mm x 0.24-0.26 mm. Follicles unknown. Phenology.—The months of flowering are unknown. Distribution. —Gonolobus martinicensis is apparently endemic to Martinique (Fig. 1). Its habitat is un- known, but likely primarily middle elevation rainforest based on the requirements of most congenerics in the Lesser Antilles. Provisional conservation status.—IUCN: Critically endangered (CR)—Bla biii. NatureServe: GH NH. laxon concept mapping.— < G. martinicensis sensu Duss (1897); < G. martinicensis sensu Schlechter (1899); < G. E sensu EE (1988); G. martinicensis sensu Fournet (2002). bl AUR Lesser nU Wee species Notes. inicensis is immediately gni 41 byi its double (or ligulate) ) interst l gynosteg egial corona (Ci). The only type in the area is C. iyanolensis. (Fig. 6). In contrast to the robust-flowered E iyanolensis, G. martinicensis exhibits distinctly more slender flowers with elongate-campanulate bases (ca. 2 mm long) that are longer than wide (as wide as long or wider than long in G. iyanolensis) and a narrower gynostegial corona (see key). The distinct dorsal ridge of the narrow upper ligulate corona lobe of the Ci in G. iyanolensis has also not been seen in G. martinicensis, which is either plane in the homologous position or exhibits a slightly raised "iJ A dditi ined. MARTINIQUE: L.C. Rich. Herb. Guyanensi-Antillanum, s.d. (fl) (P-2 specimens [G. caribaeus Rich., nom. shed Tr s.d. (fl), Lieber s.n. (M). 6. Gonolobus stapelioides Desv. ex Ham., Prodr. Pl. Ind. Occid. 32. 1825. (Fig. 9). Tex JAMAICA. PORTLAND PARISH: N of Hardwar Gap, uphill from the trail above the "Waterfall," ca. 3900 ft, montane rainforest, vines growing over trailside shru s and in small forest openings, population J1, 3 Mar 2006 (fl), Krings 1395 with Suiter and Proctor (NEOTYPE, here designated: P!; ISONEOTYPES: BM!, Y!, NCSC!, NYD. Fischeria cincta Griseb., Fl. Brit. WI. 421. 1862 Gonolobus cinctus (Griseb.) Benth. & Hook. f. ex B.D. Jacks., Index Kewensis 1:1054. 1895. TYPE: JAMAICA: Higson s.n. (HOLOTYPE: K!) Herbaceous perennial vines. Latex white. Stems pubescent, both short, capitate-glandular, and longer, sharp, eglandular trichomes present, the latter retrorse internodally, and antrorse-appressed along an indistinct, horizontal ridge between two opposing petioles and/or just above the node, to 1.3 mm long; nodes pubescent, gland field apparently absent. Leaf blades elliptic to ovate, 3-9 cm x 1.2-4.5 cm, apices acute or gradually acuminate with the acumen narrowly obtuse, to 0.75 cm long, bases cuneate, EE or truncate (fre- quently all base types present on a single individual vine), margins entire, adaxial surface ly strigose, abaxial surface sparsely pubescent to glabrate, sharp, eglandular trichomes relatively dense Alene the major and minor veins, colleters 2(3?), ca. 0.5 mm long; petioles 1-2.3 cm long, pubescent on all sides, capitate trichomes to 0.06 mm long, sharp, eglandular trichomes ca. 1.1 mm long, mostly spreading or somewhat retrorse, less dense than the capitate trichomes; stipular colleters 2, ca. 0.3 mm long, one borne on each side of the petiole base. Inflorescences racemiform, peduncles to 2.5 cm long, pubescent, capitate, as well as sharp, eglandular trichomes present, the latter spreading or retrorse, ca. 0.6 mm long; pedicels to 2.5 cm one puse a capitate-glandular trichomes ca. 0.05 mm long, longer sharp, eglandular appressed or -ascending, ca. 0.5 mm long, bracts Sieste 2.83.7 Ap pl SML AI NA ALE, mm x Q.45- 0. 5mm one uns adaxial surface glabrous, abaxial surface coa sely pubescent, trichomes sharp, eglandular, 0.36-0.4 mm long, antrorse. Cale lobes 5, linear-lanceolate, 7-8.5 mm x 2-2.3 mm, 120 | tl tanical Texas 2( A Corolla lobe Fic. 9. Gonolot tavelioides. A. | i infl Bi. O fl (adaxial view). Bii. O fl (abaxial view). C Sepal D Calycine col- U = f T A Li latar E r E || Eal il A nl and ctula-haadl rP Ff a? cri ^ Marat r a q rf Ioa a 1 did ia VW KI b "FAT J + í Ia hanai rannan la J E 1 D | Pollinarium. J. Follicle. K. Seed. J-K based on Bellingham 1487 (BM), rest based on Krings 1394 (NCSC). Aw = anther wings; Ca — faucal annulus VI PMS Y Wd WAN WI C corona segment; Po = pollinium; Sh = style-head. Krings, n RS Ze lak H PR | (TT A Jj: 121 apices acute to narrowly obtuse, margins glabrous or very sparsely ciliolate, abaxial surface weakly and finely pubescent at the base and glabrous toward the apex, trichomes antrorsely-appressed, to 0.9 mm long; colleters 1 per sinus. Corolla lobes 5, oblong-ovate, 12-15 mm x 4—6.5 mm, slightly overlapping at the base, a glandular swelling frequently present in the sinus, margins and apices strongly recurved, adaxial surface papillate-pubescent along the right, abaxial surface essentially glabrous; faucal annulus (corolline corona or Ca) interrupted, a pronounced twin-peaked thickening opposite each corolla lobe, apparently reduced opposite each corolla lobe sinus, Los See corona Wi fused staminal (Cs) and interstaminal parts (Ci), single, erect-undulating, int tly absent or at least not formed into conspicuous bumps or mounds; anther guiderails silio apenas ses laminar dorsal anther append- ages (Cd) white, ca. 4—4.2 x 1.8—2 mm, erect to spreading, apices bilobed; style-head 3.6-4.8 mm diam, stipe ca. 1.6 mm long, not toothed, though with white ribs present at the base of each anther. Pollinaria: corpuscula ca. 0.2 mm long, pollinia borne horizontally, ovate, ca. 1.5 mm x 0.41 mm. Follicles fusiform, 5-winged, 1.7 x 7.2 cm. Seeds pyriform, compressed, not plano-convex, 4.6—5.7 x 2.53.5 mm, glabrous, margins dentate, coma to 2.4 cm long. Phenology. XE deeg EEN June. Fruiting in June. Distribution. onolobus star is endemic to Jamaica. It is known only from montane rainforests from 1200-1660 m elevation. Provisional conservation status.—IUCN: Least concern (LC). NatureServe: G4 N4. laxon concept mapping.— = G. stapelioides sensu Schlechter (1899); = G. stapelioides sensu Adams (1972 Notes.—An appropriately labeled red type folder is present at P, but was found empty during visits in 2004 and 2006. In the absence of the specimen, G. stapelioides is neotypified here. Schlechter (1899) noted that Hamilton (loc. cit.) likely mixed up the localities when describing G. stapelioides and G. virescens Ham. The locality of the former was given as ER ana that of the EEN as Jamaica. However, neither entity is known by any other speci from t] land indicated in t tol In addition, Hamilton's description of ovate and acute corolla pubes would be an inaccurate decctiption of the Jamaican entity to which Schlechter (1899) and subsequent workers have applied the name G. stapelioides. However, Hamil- ton’s corolla lobe description is a very nice match of the type of G. virescens. Thus, it appears that while the localities were mixed up, Hamiiton’s protologues clearly allow association of the name G. stapelioides to the Jamaican entity and G. virescens to the Tobagoan entity as Schlechter (1899) proposed. Gonolobus stapelioides is immediately recognizable by the white Cd segments that are fully vertical at maturity (Fig. 9E, F Additional specimens examined. JAMAICA. Portland Parish: Vinegar Hill, 21 Feb 1894 RE ps 5561 (MO [a single leaf and a single flower]; Vinegar Hill, “6.2.95” [1895] (fl), Harris 5561 (BM); Hardwar Gap, 25 Feb 1964 Adams 12705 (M, MO); Immediately Ridge of the Blue Mountains, between John Crow Peak and Morce's Gap, 15 Se 1989 (fI), Bellingham 1156 (BM); 0.5 mi N of Hardwar Gap, near "The Waterfall,” 14 Jan 1980 (fl), Proctor 38534 (IJ); 0.5 mi N of Hardwar Gap, uphill from the trail above the “Waterfall,” 3 Mar 2006 (D, Krings 1394 with Suiter and Proctor (IJ, NCSC); 0.5 mi N of Hardwar Gap, uphill from the trail above the “Waterfall” 3 Mar 2006 (fl), Krings 1396 with Suiter and Proctor (IJ, NCSC). St. Andrew Parish: Grand Ridge of the Blue Mountains, E of the summit of John Crow Peak, 30 Apr 1990 (fl), Bellingham 1211 (BM); Grand Ridge of the Blue Mountains, immediately E of John Crow Peak, 11 Jun 1991 (fl-sheet 1; fr-sheet 2), Bellingham 1487 (BM-2 sheets). St. Thomas Parish: Monkey Hill, S spur of Mossmans Peak, 16 Jun 1952 (fl), Proctor 6803 (BM, IJ). 7. Mignon stellatus ed nd. Brit. Ke Men 1862. (Fig. 10). Tire: JAMAICA: Manchester, Knockpatrick, Nov [??] (fD), Purdie s.n. (HOLOTYP Herbaceous perennial vines. Latex white. St | t to glabrate in age, both short, capitate-glandular, and longer, sharp, eglandular trichomes Presa. the latter retrorse- appressed or -spreading internodally, and antrorse-appressed along an indistinct, horizontal ridge between two opposing petioles and/or just above the node, to 1.25 mm long; nodes pubescent to glabrate, gland field apparently absent. Leaf blades elliptic to oblong-elliptic, 2.2-11.5 cm x 0.8—4.2 cm, apices acute or gradually acuminate with the acumen narrowly obtuse, to 1.4 cm long, bases cuneate, margins entire, adaxial surface sparsely strigose, abaxial 122 AK De 10 6 Hatuc A leaves R Open flower. C Sepal D.C lleter E. F | | ftl || , 9y t gi | „and style- head fm ET | + -p | H (fren (t | ) Hi- -ii. inar dorsal anti ] 1. Pollinarium. J. Follicle A C- D and J based on Proctor 29378 (BM). Band i ii based on Pures. n. (K). E r JEL J D 1117032 ( 1) anther wings; gy g ; P A " P» I Pa 4 A I 4 | + B | See fy up Sh chy le basal corpusculum; LS d d d e PO = poimium, > tyie-nead. Krings, n aa if IL H al TT ai A 123 surface glabrate, trichomes sharp, eglandular, very sparsely scattered along the major and minor veins, colleters 2, 0.2-0.4 mm long; petioles 0.8—5.6 cm long, sparsely pubescent on all sides to glabrate, capitate- glandular trichomes, ca. 0.1 mm long, sharp, eglandular trichomes, ca. 0.6 mm long, antrorsely-appressed or -ascending; stipular colleters 2, ca. 0.25 mm long one borne on each side of the petiole base (rarely on the stem). Inflorescences racemiform, peduncles 0.7—3 cm long, sparsely pubescent to glabrate, capitate, as well as sharp, eglandular trichomes present, the latter antrorsely-appressed or -ascending, distributed throughout, ca. 0.8 mm long; pedicels 1.2-2.3 cm long, eer evenly Eessen SE apex to base, capitate-glandular trichomes ca 0.7 mm long, longer, sharp, eglandular tricl ascending, ca. 0.26 mm long, bracts linear to linear-oblong, ca. 1.35 mm x 0.24 mm, caducous adaxial meer: pubescence glabrous, abaxial surface coarsely pubescent, trichomes sharp, eglandular, ca. 0.22 mm long. Calyx lobes 5, lanceolate, 7-11 mm x 1.7-2.7 mm, apices obtuse, margins glabrous, abaxial surface very sparsely coarsely pubescent at the base, glabrous toward the apex, trichomes antrorsely appressed or -ascending, to 0.6 mm long; colleters 1-2 per sinus. Corolla lobes 5, linear-lanceolate, 15-18 mm x 4-5.3 mm, slightly lobed at the base, a glandular swelling frequently present in the sinus, adaxial surface densely strigillose, not papillate- pubescent, on the right half, abaxial surface glabrous; faucal annulus (corolline corona or Ca) interrupted, a raised bump or ridge opposite each corolla lobe sinus, pubescent; gynostegial corona of fused staminal (Cs) and interstaminal (Ci) parts, prostrate-undulating, single, secondary nectaries in int | position absent; anther guiderails without appendages, laminar dorsal anther appendages (Cd) 1.7-2.1 mm wide, slightly emarginate to divergently and sharply bilobed; style-head ca. 4.3 mm diam, stipe ca. 1.7 mm long, edentate. Pollinaria: corpuscula ca. 0.25 mm long, pollinia borne sono any ovate, ca. 1.7 mm x 0.48 mm. Follicles (immature) fusiform, 5-winged, but with small the wings, 7.1-7.3 cm x 1.6-1.7 cm. Seeds unknown. Phenology.—Flowering in March and July. Fruiting in July. Distribution.—Endemic to Jamaica, G. stellatus is known only from montane rainforests from about 390-460 m in elevation. Provisional conservation status. —IUCN: Vulnerable (VU)—Bla,c. NatureServe: G3 N3. Taxon SC mapping.— = G. stellatus sensu Schlechter (1899); = G. stellatus sensu Adams (1972). Additi ined. JAMAICA. Portland Parish: Hog Hill study , ca. 4 mi S of Sherwood Forest, N edge of John Crow Mtns., 1976-78 (st), Kelly JCM 1490 (IJ); Gorge f tl River, 25 Jul 1967 (fl & fr), Proctor 29378 (BM, Ij). St. Ann Parish: County of Middlesex, 1 Mt. ft. Diablo, woods, 9 Mar 1936 (fL, une 11032 (GH). 8. Gonolobus stephanotrichus EUR poe Pi. Cub. 177. 1866. (Fig. 11). Tr: CUBA: 1860-1864 (fl & fr), Wright 2969 (Lecrorwe, designated by Kring GOET!-fl; svvrvegs: BREM!, MO, G!, GH!, HACI, K!, NY!, P!, UC!, US). Herbaceous perennial vines. Latex white (rarely watery). Stems pubescent, both short, capitate-glandular, and longer, sharp, gedoe trichomes present, the latter EE spreading, or retrorse internodally, and antrorse-appresse E indistinc E horizontal ridge bet WO opposing petioles and/or just above the node, to 1.08 mm long; nodes pul t, gland field apparently absent. Leaf blades lanceolate, ovate, or oblong-ovate, 2-14.4 cm x 0.7-7.7 cm, apices gradually or eege acuminate with the acumen narrowly obtuse, to 2 cm long, bases deeply cordate, margins entire, adaxial surface sparsely but evenly strigillose to glabrate, abaxial surface sparsely but evenly strigillose, sharp, eglandular trichomes scattered along the major and minor veins, to ca. 0.58 mm long, colleters 3-4, ca. 0.4 mm long; petioles 0.9—7 cm long, evenly pubescent on all sides, some trichomes capitate, to 0.1 mm long, but mostly sharp, eglandular, antrorsely ascending or spreading, to 0.67 mm long; stipular colleters 2, ca. 0. 39 mm long, one borne on each side of the petiole base (rarely on the stem). Inflorescences racemiform, peduncles 0.18—0.4 cm long, sparsely pubescent to glabrate, capitate trichomes apparently absent, sharp, eglandular tric] appressed or -ascending, distributed throughout, ca. 0.4 mm long; pedicels 0.25—0.89 cm in done relatively evenly pubescent from apex to base, capitate-glandular trichomes very sparse, ca 0.07 mm long, longer, sharp, eglandular trichomes antrorsely-ascending, ca. 0.43 mm long, bracts linear-lanceolate, ca. 4.9 mm x 0.48 mm, caducous, adaxial surface glabrous, abaxial surface coarsely pubescent, trichomes sharp, eglandular, , antrorsely- 124 H Fic. 11. Gi AK A fl Bi Or fl (adaxial vi ). Bii Of fl ial view). C Sepal D Calycine colleter. E. F f tl H gy g , and style-head F. Detail f gy l gi | fused staminal (Cs) and interstaminal (Ci) segments (top view) il of gy g | | basal tooth. H. Pollinarium. 1. Follicle. J. Seed. A-D based on Wright 2969 (NY). E-H based on Acevedo-Rodríguez 10596 (US). |-J based do-Rodríguez 11445 (US). Aw = anther wings; Ca = faucal annulus of corolla; Ci = interstaminal D LEN | FF ad | 5 J | al J "al [| fm A Sh = style-head. UI peo. wae Ei Krings, n S £r lal H Al IT 2 jo 125 0.22-0.47 mm long, antrorse. Calyx lobes 5, lanceolate, 2.7-6.7 mm x 1.6-2 mm, apices obtuse, margins glabrous, abaxial surface sparsely but coarsely pubescent, trichomes antrorsely appressed or -ascending, to 0.4 mm long; colleters 1 per sinus. oe eee 5, elongate triangular or lanceolate, 8.7-14 mm x 2.4-3.7 arfaro gla Brame | 1] g frequent ly present in the sinus, adaxial si mm, slightly lobed at the base, a g abaxial surface pubescent wie coarse, sharp, Eolmdulsr. antrorsely-appressed or -ascending boss ca. 0.7 mm long; faucal annulus (corolline corona or Ca) annular, pubescent; gynostegial corona of fused staminal (Cs) and interstaminal (Ci) parts, prostrate-undulating, single, secondary nectaries in interstaminal position absent or at least not developed into distinct bumps or mounds; anther guiderails without append- ages, laminar dorsal anther appendages (Cd) ca. 0.6 mm wide, truncate; style-head ca. 2.6 mm diam, stipe ca. 0.73 mm long, teeth not appearing as notches, instead truncate, borne individually on the lower portion of the column, just above the upwardly rising segment of each Cs. Pollinaria: corpuscula ca. 0.2 mm long, pollinia borne horizontally, narrowly ovate, ca. 0.84 mm x 0.3 mm. Follicles ovoid, 5-winged, 9.3-9.5 mm x 2.6-2.9 mm. Seeds pyriform, compressed, not plano-convex, 7—7.8 mm x 3.4-4.9 mm, glabrous, margins entire, not dentate, coma to 2.7 cm long. Phenology.—Flowering in January, February, May, June, and October. Fruiting in April, November, and September. Distribution.—Gonolobus stephanotrichus occurs in Cuba, Hispaniola, and Puerto Rico (Fig. 1). It has primarily been collected in mid-elevation (340—1100 m), moist to wet forests. Provisional conservation status.—IUCN: Least concern (LC). NatureServe: Cuba (G4 N4); Dominican Republic (G4 N4); Puerto Rico (G4 N4). laxon concept mapping.— = G. stephanotrichus sensu Schlechter (1899); = G. stephanotrichus sensu Liogier (1957, 1994, 1995); = G. stephanotrichus sensu Acevedo-Rodríguez (2005) Notes.—Gonolobus stephanotrichus was recently “re-discovered” from Cuba, from where it had been known only by collections sometime between 1860 and 1864 (Krings et al. 2005). Additional illustrations of G. pus from Puerto Rico can be found in Acevedo-Rodríguez (2005). Readers should note however, that the g ium pictured in fig. 36 D should be rotated so that the Cd are opposite the corolla Os lobe sinuses and not — the corolla lobes themselves as pictured. Additional specimens examined. CUBA: Cuba Orientali, 1856-1857 (fl, fr-imm), Wright 407 (GH; blue MR n Orientali, 1861 [1860 on white label, but with a one marked through the zero] (fl, fr-imm), Wright 164 (S). Santiago de C Maestra, Parque Nacional Turquino, within 3 km of Campamento Joaquín on trail from the Alto de Naranjo, 3 June 2005 (fl-imm), ead Areces, and Lazcano s.n. (HAJB, NCSC); Sierra Maestra, Parque Nacional Turquino, 2-3 km downhill from Campamento Joaquín in direction of the Alto de Naranjo, 2 June 2005 (fl-imm), sai Areces, and Lazcano s.n. (NCSC). DOMINICAN REPUBLIC. Sen Cordillera Septentrional, Mora, Colonia de Jamao, 21 May 9 (fl), Ekman H12568 (K). Puerto Plata: estribo sur de Isabel de Torres, 16 Oct 1976 (ID, Liogier and a 25627 (JBSD). San a Cordillera Central, Loma La Humeadora, ladera este, en el nacimiento del Arroyo Derrumdadero, 20 Apr 1994 (fr), Jiménez, Mejia, ¢ and Mee 1304 a dd us munda n Barrio Guilarte, Guilarte Forest Reserve, ipis trail, 21 Nov 1997 (fr), R (U re , 18 Jan 1996 (st), Acevedo-Rodríguez and Axelrod 7785 (US); Río Abajo State Forest, along ae trail, 12 Jan 1999 (fl), Acevedo-Rodríguez 10596 (UPRRP, US); Rio Abajo State Forest, 2 Feb 1999 (fl), Acevedo- Rodriguez 10814 (UPRRP, US); Rio Abajo State Forest, along Igartua trail, 20 Apr 2006 (st), Krings 1390 (NCSC); Río Abajo State Forest, 20 Apr 2006 (st), Krings 1391 (NCSC); Río Abajo State Forest, 20 Apr 2006 (st), Krings 1392 (NCSC). Orocovis: Toro d One Recreation area, 11 E 000 GE Acevedo-Rodríguez 11445 (UPRRP, US); Toro Negro State Forest, near uphill del Bolo, 21 Apr 2006 (st), i ains 1387 A Toro SE State Forest, Sendero Piscina, sa 21 e 2006 60, Krings 1388 (NCSC); Toro N d on opposite from Rt ping , 21 Apr 2006 (st), Kings 1389 (NCSO. 9, o no E Syst. Bot. 32:187. 2007. (Fig. 12). Tr: DOMINICA: In sylvis ad Laudat, May 1882 (fl), Eggers 7 Gl; sores: FRI, Gl, GH!, JE!, MI, P!, WI, Z!). Herbaceous perennial vines. Latex white. n lides or le both short, capitate-glandular, and - | J]: ginternodally, and antrorse- longer, sharp, eglandular t appressed along an indistinct, horizontal idge between a opposing sde alí just above the node, to 0.64 mm long; nodes pubescent or glabrous in age, gland e sometimes presen Leaf Pads ovate to oblong-ovate, (2.1-)6.7-10.4 cm x (0.933.2-5.7 cm , apices g 1] y OL abr uptly acuminate w ith the acumen 126 Fic. 12. Gonolot Ae dl L nr Ab R Flower. € . Raised e as Al fai nc , d d Led a D ` D Sab £ - s fada Land eamnuadt Erf i t removed) E I gy (sty ) g] | Zen H bal L E k Ae 0 FAA R and D- elements. G ge (Cd). H Pollinarium. A and C 99 (M) H based rJ [| [| ER P a x m " | 2. PF | J | al A F al F ET wf E rta min gynostegial corona segment. wee së E eee Krings, n La. Ze | H FW | "TAI a | H 127 acute, to is BE cm | long, bases deeply or shallowly cordate (rarely rounded, but then other leaves present with argins entire , adaxial surface glabrate or sparsely strigose, abaxial surface glabrate or more densely toen with sharp, ebe trichomes scattered along the major and minor veins, colleters 2, (0.4—)0.7-0.9 mm long; petioles (0.9-)2.2-5.4 cm long, glabrate or sparsely pubescent primarily along the adaxial side and near the apex on the abaxial side, trichomes mostly capitate, to 0.09 mm long, with some Sues OM s oe i nomie: mixed in, 0.2-0.4 mm Dons stipular colleters 2, ca. 0.3 mm long, one EE 4.1-5.2 cm long, glabrous or very mE pubescent with bodscssitsteandetspo exlandulsr] t, the latter antrorsely-appressed or —ascending, most E E near apex, ca. 0.3 mm nisus re 14-3.4 cm long, pubescence pronounced at apex and essentially absent elsewhere or sparsely to densely, relatively evenly pubescent from apex to base, capitate-glandular trichomes ca. 0.08 mm long, longer, sharp, eglandular hairs antrorsely- appressed or -ascending, ca. 0.3 mm long, bracts linear-lanceolate, ca. 0.8 mm x 0.2 mm, caducous, adaxial surface glabrous, abaxial surface ly pubescent, tricl harp, eglandular, ca. 0.17 mm long, antrorse. Calyx lobes 5, sublanceolate to elongate triangular (widest at or just above the base), 2.2-5.3 mm x 0.8-1.4 mm, apices obtuse to narrowly rounded, margins glabrous, abaxial surface glabrous or pubescent primarily at the base and variously toward the apex, trichomes, if present, antrorsely-ascending or -appressed, to 0.27 mm long; colleters 1 per sinus. Corolla lobes 5, narrowly lanceolate to elongate triangular, 4.2-11.2 mm x 1.83.4 (3.7) mm, slightly lobed at the base, a glandular swelling frequently present in the sinus, adaxial surface glabrous, abaxial surface glabrous; faucal annulus (corolline corona or Ca) a distinctly raised ring, ca. 0.5 mm high, short-hispid along the entire rim or only opposite each corolla lobe sinus; gynostegial co- rona of fused staminal (Cs) and interstaminal (Ci) parts, single, fused at the base into an erect ring, 5-lobed, lobes appearing strongly folded; anther guiderails apparently without appendages, laminar dorsal anther appendages (Cd) 0.6—0.9 mm wide, emarginate or truncate; style-head 2.8-3.14 mm diam, stipe 1-1.5 mm long, not toothed. Pollinaria: corpuscula 0.14—2 mm long, pollinia borne horizontally, ovate, ca. 0.6—0.85 mm x 0.3—0.38 mm. Follicles unknown. Phenology.—Flowering in March, May, June, and July. Distribution.—Gonolobus waitukubuliensis is endemic to Dominica (Fig. 1). It has been found primarily in middle elevation rainforests (ca. 609-1067 m; 2000-3500 ft). Like most West Indian Gonolobus taxa, G. waitukubuliensis appears to require small gaps in mature forests. It does not appear to be able to compete in large gaps or roadside edges in which other vines quickly become dominant. It is rather infrequent to rare, especially in mature forests with few gaps. Provisional conservation status. —IUCN: Vulnerable (VU)—Bla,c. NatureServe: G3 N3. laxon concept mapping.— « G. martinicensis sensu Schlechter (1899); « G. martinicensis sensu Howard (1988); « G. martinicensis sensu Nicolson (1991) Notes.—Gonolobus waitukubuliensis is immediately gnizable among Lesser Antillean Gonolobus species by the combination of a distinctly raised, completely circular faucal annulus (ca. 0.5 mm tall), an erect gynostegial corona that appears quite folded apically, and a relatively tall gynostegial stipe (1-1.5 mm). It shares with G. absalonensis, G. iyanolensis, and sometimes G. martinicensis, the absence of hairs from the abaxial corolla lobe surface. The trichomes of the pedicel are fine and not as long (ca. 0.3 mm) as in G. iyanolensis (0.3-0.9 mm Additional specimens examined. DOMINICA: Mar 1882 a Eggers s.n. ii Ge 1882 SS A 1100 (K); May 1882 1), Eggers 1700 (MO); s.d. OD. Imray 263 (K); St. John/St. Peter, Hiki May 6 (st), a 1375, 1376, 1377 (NCSC); St. John/St. Peter, Syndicate hiking trail, 3 May 2006 We Kring me ne St. John/St Peter, Hiki 2006 (st), Krings 1380 (NCSC); St. John/St. Peter, Hi ] , 4 May 2006 (st), Krings — P Si EI Peter, im trail to d AOS 4 id ~ b Krings 13 1382, 1383, 1384 (NCSC); St. John/St. Peter y 2006 (st), Krings 1385 (NCSO): St. ohne Peter, Edd to ome eos 4 May 2 2006 6 (0, Krings 1386 (NCSC); St. ms = Haut Jean, 24 Jun 1965 (fl), Webster 13505 (BM, U 10. Gonolobus y Krings, Syst. Bot. 32:191. 2007. (Fig. 13). Tec: ST. VINCENT: Orange Hill Estate, roadbank in — and Hs pea gege, uphill from fork in road to the aes Water and Sewage Authority water intake PD 1274 Al Y A | 1. site, Kiss- -me, 14 Mar 2006 cf, Krings 13/4 with Spt inger 1 : , NY US! 128 Sepal G : : De 12 " IL j H A 1 ee + | D r g L Cc r. | | £ al rr ZEFA J*'.4 A H |) a, EF! Ta BS €— gel corolla (Ca) , and style-head. D Gy g (Ci) ( yl head ) a E E se | E n a mana’ ET £ | 4..L L H LP £ H J 4 sn 1 " G.D f ftl tegial st (Cd). 1. Pollinarium. A-B and D based on Howard 19584 (GH, NY, US). C AE de Se INFO Fa £ p re Dad " 1 [| "n sal he f^ PC PAN | and E 9 pringer (NCSC) > gy g gment; Cd = laminar dorsal al J Fr sas Pa I + Krings, R isi fG lobus in the West Indi 129 Herbaceous perennial vines. Latex white. Stems glabrate to pubescent, both short, capitate-glandular, and longer, sharp, eglandular trichomes present, the latter throughout, spreading, or retrorse internodally, and antrorse-appressed along an nq. horizontal ridge between two opposing petioles and/or just above the node, to 1.2 mm long; nod t, gland field apy tly absent. Leaf blades ovate to oblong-ovate, (3.0-)5.4-9.6 cm x (1.2-22.4-6. 3 cm, apices gradually or Da acuminate d the acumen pou] obtuse, to 1.5 cm long, bases deeply or shallowly cordate, margin r sparsely strigose, abaxial surface glabrate or more densely strigose with sharp, ea E scattered along the major and minor veins, colleters 2-4, 0.35-1 mm long; petioles (1.22)2.7—5.3 cm long, spreading pubes- cent on all sides, some trichomes capitate, to 0.09 mm long, but We SC E 1-1.2 mm one stipular colleters 2, ca. 0.26 mm long, one born peduncles 0.7-3.1 cm long, capitate, as well as sharp, nie wi homes present, the latter mostly ad ing, but also some antrorsely-appressed or -ascending, distributed throughout, 0.7-0.9 mm long; pedicels 1.3-1.8 cm long, relatively evenly pubescent from apex to base, capitate-glandular trichomes ca 0.08 mm long, longer, sharp, eglandular trichomes mostly spreading, but some antrorsely-appressed or -ascending, ca. 0.9 mm long, bracts linear-lanceolate, 1.1-1.7 mm x 0.2-0.25 mm, caducous, adaxial surface glabrous, abaxial surface coarsely pubescent, trichomes sharp, eglandular, ca. 0.53 mm long, antrorse. Calyx lobes 5, sublanceolate to elongate triangular (widest at or just above the base), 3.8-5.8 mm x 1-1.3 mm, apices obtuse to narrowly rounded, margins ciliate or glabrate, abaxial surface densely and coarsely pubescent at the base and variously toward the apex, trichomes spreading and antrorsely appressed or -ascending, to 0.8 mm long; colleters 1 per sinus. Corolla lobes 5, narrowly lanceolate to elongate triangular, (7.5-)8.7-10.1 mm x 1.9-3.1 mm, slightly lobed at the base, a glandular swelling frequently present in the sinus, adaxial surface pubescent on the right half, trichomes capitate, ca. 0.1 mm long, abaxial surface pubescent with coarse, sharp, eglandular, antrorsely-appressed or -ascending trichomes, particularly at the base and vari- ously Miis ES BEE 0.36- B 5mm PNE faucal annulus (corolline corona or Ca) interrupted, a raised bump corolla lobe sinus, pubescent or glabrous; gynostegial corona of fused MEME ES and interstaminal aL parts, prostrate-undulating, single, a small ne borne near the ae guiderails VV ithout appendages, laminar d ] 5 A (Cd) ca. 0.5 mm wide, — or truncate; style-head 2.83.95 mm diam, stipe 0.48-0.7 mm p bearing a single tooth ca. 0.2 mm long below each anther. Pollinaria: corpuscula 0.2-0.23 mm long, pollinia borne horizontally, ovate, 0.6-0.9 mm x 0.32—0.45 mm. Follicles unknown. Phenology.—Flowering in February and March. Distribution.—Gonolobus youroumaynensis is apparently endemic to St. Vincent (Fig. 1). It occurs on the windward side of the island at the foot ER a Soufriére in the Orange Hill Estate area. Vines were found take amidst banana and pigeon pea plantations. Vines were not found growing along the road to the water i along the trail to the crater of La Soufrière. However, only a half day was spent searching this trail and they could have been overlooked. Provisional conservation status. — IUCN: Critically Endangered (CR)—Bla,biii. NatureServe: G1 NI. Taxon concept mapping.— < G. martinicensis sensu Schlechter (1899); < G. martinicensis sensu Howard (1988) Notes.—Gonolobus youroumaynensis is readily recognizable among Lesser Antillean Gonolobus species by the combination of a single, prostrate-undulating gynostegial corona exhibiting a small bump towards the base of the Ci, a toothed stipe, and long, spreading trichomes (to 0.9 mm) found on stems, peduncles, and pedicels. Additional specimens examined. ST. VINCENT: Mar 1890 (fl), Smith and Smith 382 (BM, E, GH, K, NY); Orange Hill Estate, 10 Feb 1980 (fl), Howard 19584 (BM, GH, NY, US). DOUBTFUL AND EXCLUDED NAMES The following list includes names that are excluded from Gonolobus s.s. based on their morphology (e.g., 130 J | of the Botanical R h Institute of Texas 1(2) lack of laminar dorsal anther appendages), and names of uncertain status. Two new combinations are made below: Matelea dictyopetala and M. pubescens. Gonolobus bakeri Schltr., Symb. antill. 7(3):341. 1912. Tee: CUBA: Cult. in Santiago de las Vegas, May 1907, Baker 7286 OLOTYPE: location unknown Gonolobus bakeri is apparently only known from the type collection, which has not been located despite query of nearly one hundred institutions known to house West Indian collections. If the specimen was at B, it may have been destroyed in WW II. The position of G. bakeri appears uncertain. Schlechter does not describe dorsal anther appendages in the protologue. Combined with his note that the species is "mit G. variifolius Schltr. [now Matelea variifolia (Schltr.) Woodson] am náchsten verwandt," there is reason to believe that G. bakeri may better belong in Matelea. Matelea variifolia clearly lacks laminar dorsal anther appendages and also exhibits suborbicular corolla lobes. It is known only from Puerto Rico (Schlechter 1899; Acevedo- Rodríguez 2003, 2005). As the single specimen representing G. bakeri was cultivated in Cuba, it may be that the species is merely a pubescent form of M. variifolia. Among West Indian Gonolobus s.s. (if not the entire genus), G. bakeri would be the only species with suborbicular corolla lobes. Gonolobus bayatensis Urb., Symb. antill. 9(3):420. 1925. Tre: CUBA: Oriente, Bayate, 9 Oct 1914, Engström 3056 (LECTOTYPE, designated by Krings 2008: St. isoLecrorvrE: NY!) = Matelea bayatensis (Urb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. Gonolobus bicolor (Britton ër PWilson) Urb., Symb. antill. 9(3):421. 1925. Tree: CUBA: Oriente, Maestra ridge, 1300 m, ul 1922, León 10787 (HoLoTyPE: NY!) = Matelea bicolor (Britton & P. Wilson) Woodson, Ann. Missouri Bot. Gard. 28:256. 1941. Gonolobus oe Urb. & Ekman, Ark. Bot. 20A(5):41. 1926. Tree: Hart: Massif de la Selle, Morne Cabaio, c. 1900 924 (fl), Ekman H1625 an S!, designated by Krings 2008; isoLecrorYPE: IJ! — fragment [single leaf]) = Matelea Ge Ge & Ekman) Krings, comb. n Gonolobus domingensis Alain, Moscosoa 1(3):46. 1978. Tree: DOMINICAN REPUBLIC: Estril del Isabel de Torres, Puerto Plata, alt. 750 m, 16-17 Aug 1975 (ID, Liogier & Liogier 23780 (LECTOTYPE: JBSD!, designated by Krings 20054) i ing (Alain) Krings, Sida 21:2081. 2005. Gonolobus floccosus Bertol., Opusc. Sci. 4:225. 1823. Tree: GUADELOUPE: (HoLotyre: BOLO!) = Matelea maritima (Jacq.) Woodson, Ann. Missouri Bot. Gard. 28:222, 1941. Gonolobus a Schltr. in ee , Symb. antill. 1(2):287. 1899. Tee: CUBA: Wright s.n. (HoLorveE: location unknown) ain, Mem. Soc. Cub. Hist. Nat. "Felipe Poey" 22:120. 1955. Matelea BEE is apparently known only from the type collection, which has not been located despite query of nearly one hundred institutions known to house West Indian collections. Ifit was at B, it may have been destroyed in WW II. Thus, the species can be placed solely based on the protologue and past treat- ments. Although Krings et al. (2005) maintained the taxon in Gonolobus, the combination in Matelea may be more correct based on the glabrous, reticulate corolla lobes and very narrow, linear-lanceolate leaves (0.5-0.7 cm wide)—a leaf form quite atypical for Gonolobus. The description of the corolla lobes as being 9 cm long, is likely a print-setting error to be corrected to 9 mm. Grisebach (1866) did not mention laminar dorsal anther appendages in his protologue for G. tigrinus var. angustifolia and, upon reconsideration, it is unclear to me whether Schlechter (1899) was refering to them or not [italics mine]: *Diese Art ist trotz ihrer habituellen Aehnlichkeit mit G. tigrinus Griseb. von diesem vollstánding verschieden. Die Blátter sind schmaler, die Blúthen grösser, die äussere Corona ganz verschieden und die Schuppen der inneren Corona viel deutlicher vom Antherenrúcken abgehoben, Gei von oben gesehen, in der Mitte nicht ausgerandet, sondern eher verdickt.” 1 am hesistant to pres tinct, as another Cuban Gonolobinae species— G. stephanotrichus—was recently oral in the country m having been known from the island only from the type, collected between 1860 and 1864 (Krings et al. 2005). Geen haitiensis PT. Li, J. South China Agric. Univ. 14:58. 1993. Tre: HAITI: Massif du Nord, St. Louis du Nord, Baron, 950 m, 20 Aug 1925, Ekman H4693 (HoLOTYPE: Sl; ISOTYPES: Bl, US!). = Matelea crispiflora (Urb.) Jiménez, Rhodora Ge 1960, syn. nov. Krings, Revision of Gonolobus in the West Indi 131 Gonolobus haitiensis P.T. Li is a nomen novum for Gonolobus stipitatus Alain (Phytologia 64:345. 1988). It is here reduced to synonymy beneath Matelea crispiflora, as its type (Ekman H4693) clearly belongs with Poicil- lopsis crispiflora Urb. However, see also note for Gonolobus membranaceus Schltr. Gonolobus maritimus (Jacq.) R.Br., Mem. Wern. Nat. Hist. Soc. 1:24. 1810. Te: Herbar. Du Jacquin, 2 Insulae Caribaea, s.d., De Ponthieu Gecrotyre, designated by Krings & Saville 2007: BM!) = Matelea maritima (Jacq.) Woodson, Ann. Missouri Bot. Gard. 28:222. 1941 Gonolobus membranaceus Schltr. in Urb., Symb. antill. 1(2):285. 1899. Tre: HAITI: Poiteau s.n. (HoLorvrE: location unk- nown) = Matelea crispiflora (Urb.) Jiménez, Rhodora 62:238. 1960 The location of the type remains unknown. Ekman H4693 (IJ!) is labelled Gonolobus membranaceus. The specimen consists of mostly stem and 3-4 leaf fragments held in a fragment pocket. It appears to match Poicillopsis crispiflora Urb. Gonolobus membranaceus is likely very closely related to Matelea haitiensis as, based on the protologues, the two appear to differ only in the shape and apices of the leaf blades and calyx lobes. Additional collections and discovery of the type may prove them synonymous. eee GC Urb. , Symb. antill. 9(3): 421. 1925. Tree: CUBA: Oriente, Sierra de Nipe, Loma Mensura, c. 725 m, 11 9 (tr), Ekman 9710 (LecrotTYPE, designated by Krings 2008: S!; isoLECTOTYPE: NY!). = SE nipensis (Urb.) Woodson, "e p: Bot. Gard. 28:226. 1941. Gonolobus ottonis C. Koch & Bouche, Ind. Sem. Hort. Berol. 13. 1855. Te: unknown. Sometimes cited as Gonolobus ottonis Walp. (Ann. Bot. Syst. 5:502. 1859). However, Walpers (1859) clearly cited G. ottonis C. Koch & Bouche and should not be considered author of a homonym. Walpers (1859) repeated Koch and Bouche's citation: “Ex insula Cuba reportavit Otto.” However, Schlechter (1899) thought that the specimen collected by Otto (apparently formerly at B) was from Caracas, Venezuela, not from Cuba. The location of the type is unknown. Gonolobus oxyanthus Turcz., Bull. Soc. Imp. Naturalistes Moscou 25(2):318. 1852. Tre: VENEZUELA: 1845-1852, Funk 2 (G, n. v; B, n. v) fide TROPICOS. Fide Fontella and Schwarz (1981), a synonym of Gonolobus rostratus (Vahl) R. Br. ex Schult. (Systema Veg- etabilium 6:61. 1820). TROPICOS (23 Aug 2006) cited Funk 2 (Venezuela) as the type, but a copy of the protologue could not be located. Specimens were not found on either of two visits to P. d pauciflorus Spreng., Syst veg: E puo um Tyr: HISPANIOLA: St. t, Domingue, Bertero s.n. (LECTOTYPE, designated by 2006: P! lolle P , fiche # 15431]). = Matelea pauciflora (Spreng.) P M ue Missouri Bot. Gard. 28:226. 1941. Gonolobus pubescens Griseb., Fl. Brit. WI. 420. 1862. Ter: JAMAICA. Macfadyen s.n. (Hotoryee: GOET!; : K!) = Matelea pubescens (Griseb.) Krings, comb. nov. eii linis oa iseb., Fl. Brit. WI. 420. 1862. Tre: JAMAICA, Alexander s.n. ( GOET!) = Matelea rhamnifoli eb.) Krings, Sida 21: 1515. 2005. Gonolobus sintenisii Schltr., Symb. antill. 1(2):288. 1899. Tee: PUERTO RICO: Sierra de Luquillo, Jiménez Mts., 9 Jul 1885 (ID, Sintenis 1354 (Lecrorvre, designated by Krings 2008: HBG!; ISOLECTOTYPES: BM!, G!, GH!, K!, US!). = Matelea sintenisii (Schltr.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941 Gonolobus stipitatus Alain, Phytologia 64:345. 1988, nom. illeg.; non Gonolobus stipitatus Morillo, 1987. Tye: HAITI: Massif du Nord, St. Louis du Nord, Morne Baron, 950 m, 20 Aug 1925, Ekman H4693 (HoLoTYrE: S!; isotypes: B!, US!) = Matelea crispiflora (Urb.) Jiménez, Rhodora 62:238. 1960. See discussion under Gonolobus haitiensis. Gonolobus tigrinus Griseb., Mem. Amer. Acad. Arts 8:520. 1863. Tree: CUBA: Holguín, 19 Mar, Wright 1667 (notormr: GOET, n.V.; SYNTYPES: MO [image online!], G!, GH!, K!). = Matelea tigrina (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941 132 Joumal of the Botanical R h Institute of Texas 1(2) ERU d Griseb. var. SEET ipid ee pl. Cub. 177. 1866. Tre: CUBA: M du (HoLoTYPE: location c. Cub. Hist; Nat. “Felipe Poey” 22:120. 1 o See Gonolobus grisebachianus Gonolobus variifolius Schltr., Symb. antill. 10: i 1899. Terr: oa. RICO. Sierra de Luquillo, Jiménez mts., Jun 18 Sintenis 1653 (Lecrorvre: NY!, designated by K , GH!, MO!, S!) = Matelea variifolia (Schitr.) Zeen Ann. Missouri Bot. Gard. 28:226. 1941. Gonolobus virescens Desv. ex Ham., Prodr. pl. Ind. occid. 32. 1825. Tee: JAMAICA” Desvaux s.n. (HOLOTYPE: P!). Rendle (1936) considered G. virescens Desv. a species incerta. Schlechter (1899) noted that there was likely a label mix-up between G. stapelioides and G. virescens, and that the latter was likely collected from Tobago and not Jamaica. The morphology of the flowers of Desvaux s.n. (P!) places it in Marsdenieae Benth., rather than Gonolobinae. ACKNOWLEDGMENTS I thank the following institutions and their staff for facilitating access and collecting permits: Jardín Bo- tánico Nacional (Havana), Centro de Inspección y Control Ambiental (CICA), Delegaciones del Ministerio de Ciencia, Tecnología y Medio Ambiente (CITMA) de Santiago de Cuba y Granma, Regiones Militares de Santiago de Cuba y Granma, and Delegación del Ministerio del Interior de Santiago de Cuba, departments of Forestry of Dominica, St. Lucia, and St. Vincent, National Envi t and Planning Agency of Jamaica, University of Puerto Rico, Río Piedras Herbarium. I thank the curators and staff of the following herbaria for searching, or providing access to or loans of their collections: B, BG, BH, BKL, BM, BOLO, BR, BREM, BSC, BUF, C, CGE, COLO, CR, DUKE, E, F, FI, FLAS, FR, FTG, G, GH, GOET, H, HAC, HAJB, HBG, IA, IJ, ISC, JBSD, JE, K, L, LD, LE, LINN (Linnean and Smithean), M, MICH, MIN, MO, MSC, NCU, NEU, NSW, NY, O, OXF, P, PH, RSA, S, U, UBT, UC, UCWI, UPRRP, UPS, US, USF, TUR, WILLI, WU, Z. I am also grateful to the following for assistance in the field in (1) Cuba: Fabiola Areces, Julio Lazcano, Mino Leyeba, Anel Matos Viñals, (2) Jamaica: Dale Suiter, George Proctor, Davian Campbell, (3) St. Lucia: Roger Graveson, Melvin Smith, and (4) St. Vincent: Carlton Thomas, Fitzroy Springer. For help with logistics I thank Frank Axelrod, Jenny Cruse-Sanders, Saara DeWalt, Miguel García-Bermüdez, Amos Glasgow, Eric Hypolite, Ka- lan Ickes, Arlington James, Lenoire (Karen) John, Brian Johnson, Ruth Knight, Nancy Osler, Hainson Paul, Cornelius Richards, Ricardo Valentin, and Wendy Worley. 1 thank linguist Lise Winer (McGill University) for providing English *eye dialect" phonetic renditions for Amerindian names for Dominica, St. Lucia, and St. Vincent. 1 thank Nico Franz (UPRM), Bob Peet (NCU), and Alan Weakley (NCU) for sharing an accepted, but unpublished manuscript on taxon concepts. Pedro Acevedo (US), David Goyder (K), and Bruce Hansen (USF) provided thoughtful reviews of the manuscript. This research was sponsored in part by grants from the American Society of Plant Taxonomists and the Field Museum of Natural History. REFERENCES AceveDO-RODRIGUEZ, P. 2005. Vines and climbing plants of Puerto Rico and the Virgin Islands. Contr. U.S. Natl. Herb. 51:1-483. 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Biotropica 19:86-89. Woobson, RE, Ja. 1941. The North American ee Ann. abu BOE Gard, ao 193-244, Woobson, R.E. JR. and J.A. Moore. 1938. Vascular anatomy a | | gy poc Bull. Torrey Bot. Club 65:135-166. Wyatt, R. 1978. Pollination and fruit set in Asclepias: A reappraisal. Amer. J. Bot. Ge Gen 851. Wyatt, R. 1980. The impact of nectar-robbing ants on the pollination system of ls savica. Bull. Torrey Bot. Club 107:24-28. Wyatt, R. and TR. SHANNON. 1986. Nectar production and pollination of Asclepias exaltata. Syst. Bot. 11:326-334. INDEX OF NAMES AND TYPES IN WEST INDIAN GONOLOBINAE (APOCYNACE- AE: ASCLEPIADOIDEAE), INCLUDING FOURTEEN NEW LECTOTYPIFICATIONS, ONE NEOTYPIFICATION, A NEW NAME, AND A NEW COMBINATION Alexander Krings Herbarium, Department of Plant Biology North Carolina State University Raleigh, North Carolina 27695-7612, U.S.A. Alexander_Krings@ncsu.edu ABSTRACT Jh] : ra : 14.4 F. A A A ex AE fA Aecleniadoid ) in the West Indies. The following fourteen taxa are e lepia Gonolobus bayatensis Urb., G. TM NUM Schltr, G. alar Schltr., G. dictyopetalus Urb. Henan e ee es G. ds die Ms G. sintenisii Schltr., G. ida Mai: G. Pc AN Nie G. ET Schltr., Ibatia mollis i I nd P S i5 neotypified A new Griseb., P G ,an name -— a new ios in Mateiea Pu are respectively proposed for Jacaima parvifoli PN and J. costata (Urb.) Rendle var. goodfriendii Proctor. RESUMEN Se en EES y su localización de taxa e n ids b ee Ee en las Indias de Se G atus Schltr., G. dictyopetal & Ekman G manih , C apsk Ue , G. sintenisii Schltr G tigrinus PM _G AEN ,G — ied ee mollis on Poicilla costata Urb., Poicill y ir nueva en Malelta Aubl. — Jecam jarol Proctor y à costata (Urb.) Rendle var. goodfriendii E respectivamente. INTRODUCTION Subtribe Gonolobinae (A Asclepiadoideae) j| bout fifty in the West Indies, here defined to include the cies and Lesser Antilles: the Bahamas, ee Kë Tobago, Aruba and the Neth- erland Antilles, and the Cayman Islands. Evid plast (Rapini et al. 2003; Liede-Schumann et al. 2005; Rapini et al. 2006; Krings et al. 2008) and nuclear genomes (Krings et al. 2008) supports the monophyly of Gonolobinae. Generic circumscriptions remain poorly defined and it is likely that the broad concept of Matelea Aubl. sensu Woodson (1941) is not monophyletic. Genera referred to the subtribe with representation in the region include Fischeria DC., Gonolobus Michx., Matelea (incl. Ibatia Decne., Jacaima Rendle, Poicilla Griseb., Poicillopsis Schltr., Ptycanthera Decne.), Macroscepis Kunth, and Metalepis Griseb. (Fontella & Schwarz 1981a; Kunze 1995; Liede 1997; Morillo 1997; Rapini et al. 2003; Liede-Schumann et al. 2005; Rapini et al. 2006). Subtribal position has been most controversial for Metalepis, which some work- ers have also placed within in a broad concept of Cynanchum L. (Woodson 1941; Spellman 1975; Sundell 1981). Recent evidence places it basal to a well-supported Metastelmatinae-Oxypetalinae-Gonolobinae clade (Liede & Kunze 2002; Liede-Schumann et al. 2005). The last regional treatment of Gonolobinae is now over 100 years old (Schlechter 1899) and a number of new species have been described by various workers, including Britton, Krings, Liogier, Proctor, Spellman, Urban, and Woodson (see ne 2005b, 2006, 2007, as well as the present study). Recent effort as part ofa l tudy of Gonolobinae and treatment for the Flora de la República de Cuba (Krings 2005a—d, 2006, 2007 Runes et al 2003, iun & Fantz 2006; Krings & Saville 2007), has revealed ES a number of lectoty] Publishing and compiling these desi tely allows for greater reene of some issues ium is possible in a flora format. The son ofa Pa index to all names in West Indian Gonolobinae seems additionally useful considering the scattered nature of the literature and historical errors in citation. | Rat Rec Inst Tavac 2(1): 130 163. 2008 140 | tl tanical h Insti Texas 2( Fourteen lectotypes and one neotype are designated here. A new name and a new combination in Matelea are proposed for Jacaima parvifolia Proctor and J. costata (Urb.) Rendle var. goodfriendii Proctor. METHODS Names in West Indian Gonolobinae were compiled from searches of the TROPICOS database (http://www. tropicos.org) for species with West Indian types, as well as from review of pertinent literature (i.e., Grisebach 1862, 1866; Duss 1897; Schlechter 1899; Millspaugh 1902; Boldingh 1909, 1914; Woodson 1941; Moscoso 1943; Cheesman 1947; Alain 1957; Gooding et al. 1965; D'Arcy 1967; Adams 1972; Little et al. 1976; Correll & Correll 1982; Proctor 1984; Howard Ee We 1991; Liogier 1994, 1995; Fournet 2002; Acevedo- Rodríguez 2005). Protologues were compiled from held at the North Carolina State University Libraries or obtained through inter-library loans. Subsequently, queries were made to ninety institutions known to house West Indian collections—of which sixty-five responded with either loans, digital images, or negative search results (see Acknowledgments). Visits were also undertaken to BM, BSC, DUKE, HAC, HAJB, IJ, K, UCWI, UPRRP, US, and P. Nearly half of the species names in West Indian Gonolobinae can be attributed to three German work- ers: August Heinrich Rudolf Grisebach (1814—1879), Friedrich Richard Rudolf Schlechter (1872-1925), and Ignatz Urban (1848-1931). The former worked from the University of Góttingen (GOET) and the latter two from the Botanical Garden and Museum Berlin-Dahlem (B). The general philosophy followed herein, is that for taxa described by Grisebach, applicable specimens judged original material at GOET are considered holotypes—although, in a strict sense, Grisebach did not designate them as such, although his use of them is evident by his notes and annotations. However, typi- fication of Grisebach names based on collections by Charles Wright (1811-1885) can be problematic, as collections cited under a single collection number frequently represent gatherings from different localities and different times (Howard 1986, 19892). The philosophy adopted herein is that Grisebach names based on Wright collections should be lectotypified when (1) it can be shown that specimens distributed under a single number represent gatherings from different localities at different times, and/or (2) no specimen of a Wright number remains extant at GOET. When it cannot be shown that gatherings came from different localities and different times, duplicates of a Wright number designated lectotype are considered isolecto- types (see Krings & Fantz 2006). When it can be shown that gatherings came from different localities and different times, duplicates of a designated lectotype are treated as syntypes. Urban described West Indian taxa largely based on specimens received by him from Erik Ekman (Cuba and Hispaniola), William Harris (Jamaica), and Walter Elias Broadway (Trinidad, Tobago, and Grenada). For these taxa, holotypes are presumed to have remained at B until their destruction in a fire in World War IL. Thus, lectotypes are chosen for Urban names from duplicates of numbers cited in his protologues. RESULIS Overview.—Names of novelties in West Indian Gonolobinae can be attributed to seventeen workers (Table 1). The majority of taxa were described during four phases of intensive interest in the region that occurred at ca. 20-30 yr intervals over the past 140 yrs. The first phase was dominated by Grisebach and based on the collections of Wullschlagel (Antigua), Charles Wright (Cuba), Imray (Dominica), and Alexander, Higson, Macfadyen, and Purdie (all Jamaica). Important works resulting from activity during this phase include the Flora of the British West Indian Islands (Grisebach 1862) and the Catalogus Plantarum Cubensium (Grisebach 1866). The second phase was dominated by the activity of Schlechter—based primarily on collections by Broadway (Grenada; Trinidad & Tobago), Eggers (Tobago), Poiteau (Haiti), Sintenis (Puerto Rico), and Wright (Cuba)—and Urban—based primarily on the collections of Broadway (Tobago), Buch (Haiti), Ekman (Cuba & Hispaniola), and Harris ww d e most important relevant work resulting from this later period of activity is Urban's multi-volume S; e Antillanae (1898—1928). The third phase was dominated by Liogier (a.k.a. Hermano Alain), ND SES more West Indian SE SES novelties based on his own collec- tions than any other worker before or since Additional species described by Liogier are based on collections Krings, tal A) H fe Lat? s [| E la 141 TABLE 1. Auti | | f novelties at the speci k in West Indian Gonolobinae, based on types from the region. Author(s) No. of spp. names Author(s) No. of spp. names published published Alain s "A.H. Liogier" 8 Proctor, G.R. 1 Bertolon 1 Richard, A 1 Britton, i ^ Seel PG. Wilson 1 Schlechter, ERR 9 Decaisne, J. 3 Spellman, D.L. 1 kman, E. 1 Sprengler, C.PJ. 2 Grisebach, A.H.R 12 Urban, l. 9 Hamilton, W. 1 Vahl, M. 2 Jacquin, von, N. 1 Woodson, Jr., R.E. 3 rings, 10 by Ekman (Hispaniola) and Howard (Dominican Republic). Important works resulting from his efforts or contributions include floras of Cuba, Hispaniola, and Puerto Rico (Alain 1957; Liogier 1994, 1995). The fourth phase of interest is the result of current efforts undertaken by the present author (Krings 2005b, 2006, 2007). New discoveries during this phase are based on collections by Eggers (Dominica), Ekman (Hispaniola), Graveson (St. Lucia), Krings and Springer (St. Vincent), Liogier (Dominican Republic), Marcano and Jiménez (Dominican Republic), and Quentin (Guadeloupe). No new names in West Indian Gonolobinae resulted from R.A. Howard's efforts toward his Flora of the Lesser Antilles (Howard, 1989b). List.—Names are listed index style and include legitimate, illegitimate, and misapplied names. They are arranged strictly alphabetically by basionym or combination. The most recent combination for a respective name follows the basionym (3). Currently accepted names are in bold or specified under “Notes.” Under the name, the locality and/or collector the type is given as it appears in the protologue. Type specimens seen and verified by the author are indicated by an exclamation point (‘!’) following the herbarium of deposit. The very few specimens not seen are cia by ‘n.v. on n Loc. not cited” signifies that the dee of Hel : Holmgren deposit was not indicated in t] Herbariu m t Index & Holmgren 1998- present). Book EE follow TL-2 (Stafleu & Cowan 1976-1988) " una abbreviations B-P-H (Lawrence et al. 1968) and B-P-H/S (Bridson & Smith 1991). Author abbreviations follow Brummitt and Powell (1992). Asclepias maritima Jacq., Enum. Syst. Pl. 17. 1760. = Matelea maritima (Jacq.) Woodson, Ann. Missouri Bot. Gard. 28:222. 1941. Protologue: Neither specimens, nor locality cited. Type: Herbar. Du Jacquin, 2 Insulae Caribaea, De Ponthieu (Lectotype designated by Krings & Saville 2007: BM). Notes: Only a single sheet of Asclepias ala was located that could qualify as belonging to the origi- nal material. Eight fragments, all Asc] itima, are mounted on a single sheet held at BM and barcoded (BM000834432). A handwritten dies on the top left of the back of the sheet reads "Herbar. Du Jacquin." followed by “2 Insulae Caribaeae. De Ponthieu.” Based on the note, the fragments appear to represent two elements, but they cannot be delineated. A dehisced follicle is mounted along with leaves in relatively good condition and a few flowers in poor condition. Fruits and flowers are not mentioned by Jacquin in the pro- tologue, and these elements may represent the collection later combined with the vegetative fragment used for the protologue. Consid that this sheet app t the only remaining collection of Asclepias maritima from the herbarium af Jacquin, it was desieneted lectotype by Krings and Saville (2007). Cynanchum CC Vahl, Eclo g Amer. 2:23. 1798. = Matelea denticulata (Vahl) Fontella & E.A. Schwarz, Bol. M ot. Munic. 46:4. 1981. Protologue: ' "mer in Cana: von Rohr,” loc. not cited. Type: Guiana, von Rohr120 (noLotrrE: C [IDC microfiche photo: Vahl herbarium nr. 17:1II, 1!]) 142 | ti tanical h Instit Texas 2( Notes: Most recent authors have accepted this entity under the name Matelea denticulata (Vahl) Fontella & E.A. Schwarz. However, in a recent phylogenetic analysis based on chloroplast and nuclear data, the taxon emerged in a clade of Gonolobus s.l. (Krings et al. 2008) and should be recognized as Gonolobus denticulatus (Vahl) W.D. Stevens in a broad interpretation of the genus. EE hirsutum Vahl, Eclog. Amer. 2:24. 1798. = Matelea hirsuta (Vahl) Woodson E.E. Cheesman, FI. nidad 2:170. 1947. en “Habitat in insula Trinitatis. von Rohr,” loc. not cited. Type: Trinidad, von Rohr 92 (noLorYpE: C [IDC microfiche photo: Vahl herbarium nr. 17-111, 3!; ISOTYPE: M). Notes: An annotation on von Rohr 92 indicates that it was studied by Ignatz Urban. d maritimum (Jacq.) Jacq., Select. Stirp. Amer. Hist. 83, t. 56. 1763. = Matelea maritima (Jacq.) oodson, Ann. Missouri Bot. Gard. 28:222. 1941 Notes: Kee aos have mistakenly attributed the combination in Cynanchum to Linnaeus” Syst. Nat. (1767). However Linnaeus (1767) clearly referenced Jacquin (1763). In Syst. Nat. (1767), he wrote: *C. caule volubili, fol. ap BH Se tomentosis, pedunc. Aggregatis. Mant. 54.” The Mantissa plantarum (Mant.) pul lix to volume 2 of his Syst. Nat. (1767). In the Mantissa, Linnaeus (1767) provides further SE cc to Jacq. amer. 86, t. 56, and describing corollas of C. maritimum as ‘atropurpurea. Although C. maritimum is described on pages 83 and 84 in Jacquin’s Select. Stirp. Amer. Hist. (1763), rather than on page 86 as cited by Linnaeus (1767), the illustration reference (plate 56) is correct and is not inconsistent with plants known most recently as Matelea maritima in the West Indies. Thus, it is clear that Linnaeus accepted Jacquin's combination and was not himself proposing a new one. Continued citation of Cynanchum maritimum L. is incorrect. See Asclepias maritima Jacq. Cynanchum maritimum (Jacq.) L., Syst. Nat. 2:192. 1767, err. cit. [2C. maritimum (Jacq.) Jacq., see above]. Notes: Linnaeus (1767) never made this combination mistakenly attributed to him by subsequent workers. Cynanchum rostratum Vahl, 2 Bot. 3:45. 1794. = Gonolobus rostratus (Vahl) Schult. in Roemer & Schultes, Syst. Veg. 6:61. 1820. Protologue: “In insula Trinitatis legit Dn. v. Rohr,” loc. not cited. Type: Trinidad, (von Rohr?) Hb. Liebmann (HOLOTYPE: C [IDC microfiche photo: Vahl herbarium nr. 17:111, 51]; isoryre: BM!). Notes: Although Stafleu and Cowan (1986) ven that wane S SE Geen are ipis on the Forsskál material at C; G is not listed in the i IDC fiche of the Forsskál herbarium. However, there is a specimen in the Vahl befharium corresponding to Cynanchum rostratum. The specimen label identifies it as having been part of the Hb. Liebmann at some point, however the hand, identifying the collection as Cynanchum rostratum, belongs to Vahl. Von Rohr is not indicated on the label. Cheesman (1947) mistakenly considered *Gonolobus rostratus R.Br. sens. Griseb Fl. 420" a synonym of *Matelea viridiflora (G.EW. Meyer) Woods.” However, the latter clearly corresponds to Matelea denticulata (Vahl) Fontella & E.A. Schwarz based on Cheesman’s description (e.g., “callyx] lobes lanceolate,” "corolla [...] lobes ovate,” “anthers without dorsal appendages”) and not to Gonolobus rostratus sensu Grisebach (1862) (e.g., “calyx [...] segments ovate,” "corolla [...] segments lanceolate-linear”). Grisebach’s (1862) taxon (also from Trinidad) is clearly referable to G. rostratus (Vahl) Schult. The two taxa are difficult to confuse by anyone familiar with them. The combination in Gonolobus is to be attributed to Schultes as Robert Brown never made the combination erroneously attributed to him by some authors such as Grisebach (1862) and Schlechter (1899). Cynanchum stellatum Vell., Fl. Flumin. 121; 3, t. 80. 1829. = Fischeria stellata (Vell.) E. Fourn. in Mart., Fl. 85. Protologue: “Habitat silvis maritimis," loc. not cited. Type: Vell., Fl. Flum. 121; 3, t. 80. 1829. (LecroTYrE, designated as type by Murphy 1986). Krings, AF &l je: r LL: H | £ Ja 143 i Cynanchum viridiflorum G. Mey., Prim. Fl. Esseq. 141. 1818, nom illegit., non Sims, 1817. Protologue: “in fruticetis plantationis Hamburg,” loc. not cited. Type(s): The type could not be located. Notes: A synonym of Cynanchum denticulatum Vahl fide Howard (1989b). Fischeria cincta Griseb., Fl. Brit. W.I. 421. 1862. = Gonolobus cinctus (Griseb.) Benth. & Hook. f. ex B.D. Jacks., Index Kewensis 2:1054. 1895. Protologue: “Jamaica!, Higson,” loc. not cited. Type: Jamaica, Higson s.n. (HOLOTYPE: K!). Notes: A synonym of Gonolobus stapelioides Desv. ex Ham. fide Adams (1972). Fisheria havanensis Decne. in DC., Prodr. 8:601. 1844. Protologue: “in insula Cuba (Ramon de la Sagra, n. 556). (v.s.h. DC)” Type: Cuba, Ramon de la Sagra 555 [556 fide Decne. in DC., Prodr. 8:601.] (HoLorYrE: G-DC [IDC microfiche Candolle Prodromus Herbarium, fiche + 1542!]), photos at MO, n.v., F, n. v). Notes: A synonym of Fischeria scandens DC. fide Murphy (1986). Fischeria multiflora Decne. in DC., Prodr. 8:601. 1844. Protologue: “in Brasilia (cl. Martius) (v.s.h. reg. monac.).” Type: Brazil, Martius s.n. (HOLOTYPE: M, n v.; ISOTYPE: M, n.v.). Notes: Murphy (1986) considered F. multiflora Decne. a synonym of E stellata (Vell. E. Fourn. The latter is the only species of Fischeria she recognized from Trinidad. Fischeria multiflora Griseb., Fl. Brit. W.I. 421. 1862, err.cit. [=F. multiflora Decne.]. Notes: In contrast to Grisebach himself (loc. cit.), Schlechter (1899) believed that Grisebach's F. multiflora did not belong to Decaisne's Brazilian species by the same name, but rather to F. havanensis Decne. Chees- man (1947) accepted this as well. Murphy (1986) considered F. havanensis Decne. a synonym of F. scandens DC., but did not recognize this taxon from Trinidad, the locality from which Grisebach (loc. cit) cited his F. multiflora (based on Purdie and Crueger collections). Murphy (1986) recognized only F. stellata (Vell) E. Fourn.—under which she submerged F. multiflora Decne.—from the island. Fischeria scandens DC., Cat. Pl. Horti Monsp. 112. 1813. Protologue: “Hab. imiliter in America meridionali. Habuimus ex horto Madritensi, t 1, > e h. monsp. inedit. Type: America meridionali, hort. monsp. ined., t. 67 (HoLoTyPE: MPU, n.v.). Notes: Fischeria scandens is the only species of Fischeria recognized by Murphy (1986) from the Greater Antilles. Fischeria stellata (Vell) E. Fourn. in Mart Fl. Bras. 6:301. 1885. See Cynanchum stellatum Vell. Gonolobus absalonensis Krings, Syst. Bot. 32:181. 2007. Protologue: “Martinique. Absalon, Près la cascade, Mai 1910 (fl), Herb. d'Alleizette s.n. [4801?] (HOLOTYPE: LD.” Type: Martinique, Absalon, Près la cascade, May 1910, Herb. d'Alleizette s.n. [4801?] (HoLotyre: L!). Gonolobus bakeri Schltr. in Urban, Symb. Antill. 7(3):341. 1912. Protologue: “Hab. in Cuba cult. in Santiago de las Vegas: C.F. Baker n. 7286, flor. m. Majo 1907” loc. not cited. Type: Location of type is unknown. No additional matching material was located from which a lectotype or neotype could be designated. See Krings 2008) for a discussion of this taxon. Gonolobus bayatensis Urb., Symb. Antill. 9(3):420. 1925. = Matelea bayatensis (Urb.) Woodson, Ann. Mis- souri Bot. Gard. 28:226. 1941. 144 Journa Protologue: “Prov. Oriente prope Bayate, m. Oct. flor.: Arth. Engström in herb. Ekman n. 3056,” loc. not cited. Type: Cuba, Oriente, Bayate, 9 Oct 1914, Engstróm 3056 (LECTOTYPE, here designated: S!; ISOLECTOTYPE: NY). Notes: Two sheets of Engström 3056 are extant —one at S and one at NY. The sheet at Sis superior in condition to the NY sheet, exhibiting much more floral and vegetative material and is therefore designated lectotype. See Krings (20050) for additional notes on this species. Gonolobus bicolor (Britton & P Wilson) Urb., Symb. Antill. 9(3):421. 1925. = Matelea bicolor (Britton & P. Wilson) Woodson, Ann. Missouri Bot. Gard. 28:236. 1941. See Marsdenia bicolor Britton & P. Wilson. Gonolobus broadwayae Schltr. in Urb., Symb. Antill. 7(3):340. 1912. Protologue: “in Trinidad, ad St. Clair Experiment Station inter Bambusas volubilis: W.E. Broadway n. 2743," loc. not cited; ‘ad Santa Cruz: López (Herb. Broadway n. 2419), loc. not cited. Type: Trinidad, López 2419 (Lectoryre, here designated: Z!). Notes: Of the two syntypes cited in the protologue, only López 2419 (Z) appears to be extant. Gonolobus broadwayae Schltr. is a synonym of G. rostratus (Vahl) Schult. Gonolobus ciliatus Schltr. in Urban, Symb. Antill. 12):282. 1899. Protologue: "Tobago, in silvis prope Frenchfield, alt. 400 ped., Oct.: Eggers 5561,” loc. not cited; "Trinidad, prope La Ventilla: Crüger sn. loc. not cited; Venezuela, loc. not cited. Type: Tobago, Eggers 5561 (Lecroryre, here designated: Pl; isoLecToYPE: UCWI)). Notes: Unidentified as a type, Eggers 5561 had been filed at P gus Matelea denticulata (Vahl) Fontella & E.A. Schwarz. The Crüger speci could not be located. Gonolol iatus Schltr. is a synonym of G. denticulatus (Vahl) W.D. Stevens. Gonolobus cinctus (Griseb.) Benth. & Hook. f. ex B.D. Jacks., Index Kewensis 1:1054. 1895. See Fischeria cincta Griseb. Gonolobus cinctus (Griseb.) Benth., Gen. Pl. 2:750. 1876, err.cit. [= Gonolobus stapelioides Desv. ex Ham. Notes: This combination is not valid, as the epithet of the basionym—Fischeria cincta Griseb.—was not definitely associated with the genus Gonolobus. Gonolobus denticulatus (Vahl) W.D. Stevens, Phytologia 64:334. 1988. See Cynanchum denticulatum Vahl. Gonolobus dictyopetalus Urb. & Ekman, Ark. Bot. 20A(5):41. 1926. = Matelea dictyopetala (Urb. & Ekman) Krings, J. Bot. Res. Inst. Texas 2:130. 2008 Protologue: Haiti, “Massif de la Selle in Morne Cabaio ad Jardin Bois [...] n. H. 1625,” loc. not cited. Type: Haiti, Massif de la Selle, Morne Cabaio, ca. 1900 m, 24 Aug 1924, Ekman H 1625 (Lectotype, here designated: S!; ISOLECTOTYPE: IJ! — fragment [single leaf]. Notes: Two sheets of Ekman i nd: are known—one at IJ and ui at S. Whereas the sheet at IJ consists of a stem fragment with a singl | more leaf fragment a flower bud in the fragment pocket), the sheet at S elimine in Urbans hand) consists of numerous leaves and flowers and is in good condi- tion. Ekman specimens were sent to Urban at B for critical study (Howard, 1952; Nordenstam et al., 1994). Nordenstam et al. (1994) noted that Urban was to identify and publish on the work of Ekman and return the principal (full) set of vascular and non-vascular plants to ee (S). Nordenstam et al. (1994) also noted that it is obvious that Urban returned the original Stockholm as many Ekman sheets are annotated as types in Urban’s handwriting. Urban Ger Bee one duplicate of the material upon which any of his and Ekman’s publications were based (Nordenstam et al., 1994). Urban published a number of names Krings, (TT) a | | I Fa 1 L* * F Fr B a 145 di in West Indian G lobi | 1 on Ekman material and it can be assumed that holotypes Were kept at B. However, all of Gonolobinae types at B were destroyed in World War II, necessitating lectotypification. Gonolobus domingensis Alain, Moscosoa 1(3):46. 1978. = Matelea domingensis (Alain) Krings, Sida 21:2081. 2005. Protologue: “Loma Isabel de Torres, Puerto Plata [...] Alain & Perfa Liogier 13780 (SDM, NY); en bosque, id. A. & P. Liogier & N. Melo 23348 (SDM).” Type: Dominican Republic, estribo sur del Isabel de Torres, Puerto Plata, alt. 750 m, 16-17 Aug 1975 (fl), Alain & Perfa Liogier 23780 (LectotYrE designated by Krings 2005d: JBSD)). Notes: See Krings (2005d) for a detailed discussion of lectotypification of this taxon. Gonolobus dussii Krings, Syst. Bot. 32:183. 2007. Protologue: “Guadeloupe. Chemin de la Soufrière, alt. 500 m, Quentin 732 (HOLOTYPE: P!-fl; isorvre: GH!- Type: Guadeloupe, Chemin de la Soufrière, alt. 500 m, Quentin 732 (HoLotyre: P!-fl; isorvee: GHI-fl). Gonolobus ekmanii Urb., Symb. Antill. 9(3):422. 1925. = Matelea ekmanii (Urb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. Protologue: "Prov. Oriente in Sierra Maestra inter Rio Yara et Rio Palmamocha cr. 1300 m. alt., m. Jul. flor., n. 14453,” loc. not cited. Type: Cuba, Ekman 14453 (Lectotype, here designated: S!; isoLecTOTYPE: NY!). Notes: Two sheets of Ekman 14453 are held at S. One sheet holds a nicely, openly pressed flower in its frag- ment pocket. Considering that the condition of both sheets is otherwise comparable, the sheet with the openly pressed flower in the fragment pocket is here designated lectotype. As accession numbers are the same on both sheets, the lectotype can be identified by the label header: "Mus. Botan. Stockholm. Plantae Indiae Occidentalis N:o 14453" [sic]. Gonolobus floccosus Bertol., Opusc. Sci. 4:225. 1823. Protologue: “[...] ex Guadalupa, et Portorico,” Anonymous (loc. not cited). Type: Guadeloupe, Anonymous s.n. (HOLOTYPE: BOLO). Notes: Á synonym of Matelea maritima (Jacq.) Woodson. Gonolobus grenadensis Schltr. in Urban, Symb. Antill. 7(3):339. 1912. Protologue: “in Grenada ad Annandale, W.E. Broadway n. 34777 loc. not cited. Type: Grenada, St. Mark Parish, Wooded hillsides near Victoria, elev. 100—300 ft, 24 Nov 1957, G.R. Proctor 17225 (neorYrE, here designated: UI Notes: Based on Schlechter's (1912) description, Gonolobus grenadensis Schltr. is a synonym of Gonolobus denticulatus (Vahl) W.D. Stevens. Schlechter (1912) considered it a close relative to G. ciliatus Schltr. (also a synonym of G. denticulatus), from which he distinguished it only by larger faves and flowers. Howard (1989b) also treated G. grenadensis as a synonym of G. denticulatus, although confusing it with G. martinicen- sis in key and description (though not in illustration). Unfortunately no original Bate of G. grenadensis could be located. Proctor 17225 (IJ) appears to be the only extant specimen of this taxon from Grenada. It is here designated neotype. Gonolobus grisebachianus Schltr. in Urban, Symb. Antill. 1(2):287. 1899. = Matelea grisebachiana (Schltr.) Alain, Mem. Soc. Cub. Hist. Nat. “Felipe Poey" 22:120. 1955. Protologue: “Cuba, loco speciali haud indicato: Wright s.n.” loc. not cited. Type: Location of type is unknown. No additional matching material was located. Gonolobus haitiensis P.T. Li, J. South China Agric. Univ. 14(1):58. 1993. Type: Haiti, Massif du Nord, St. Louis du Nord, Morne Baron, 950 m, 20 Aug 1925, E.L. Ekman 4693 (Ho- LOTYPE: Sl: ISOTYPES: B!, US). 146 Notes: A nomen nov. for Gonolobus stipitatus Alain, Phytologia 64:345. 1988, non Morillo, 1987. Krings (2008) considered G. haitiensis a synonym of Matelea crispiflora (Urb.) J. Jiménez Alm Gonolobus iyanolensis Krings, Syst. Bot. 32:185. 2007. Protologue: “St. Lucia. Gros Piton, 9 Aug 1996 [*9/8/76"] (fl), Roger Graveson 107 (HOLOTYPE: GH!” Type: St. Lucia, Gros Piton, 9 Aug 1996 [*9/8/76"], Roger Graveson 107 (HoLoTYPE: GH). Notes: Roger Graveson noted that the collection date is incorrect on the label of the type and should be 9 Aug 1996 instead (Graveson, pers. comm .). Gonolobus jamaicensis Rendle, J. Bot. 74:345. 1936. Type: Jamaica, near Vinegar Hill, climbing up trees to height of 20 ft, 4200 ft elev., 5 Jun 1896, Harris 6368 (HOLOTYPE: BM. Notes: Gonolobus jamaicensis was proposed by Rendle as a nomen novum for e CR a taxon Schlechter (1899) attributed to G. rostratus (Vahl) R.Br. Robert Brown never made the combination attribute to his Memoirs of the Wernerian Natural History Society 1:35. 1810. The true Gonolol tratus (Vahl) Schult is known only from Trinidad in our area and is based on the type of Cynanchum rostratum Vahl. Because Rendle corrected a misapplication, his name—G. ee a new species name, not a nomen novum as he incorrectly stated. A nomen novum is an avowed substitut t name) for a validly published but illegitimate name, the type of which would be the same as that m the name which it replaced. Gonolobus maritimus (Jacq.) R.Br., Mem. Wern. Nat. Hist. Soc. 1: 24. 1810. = Matelea maritima (Jacq) Woodson, Ann. Missouri Bot. Gard. 28:222. 1941. See Asclepias maritima Jacq. Gonolobus maritimus (Jacq.) R.Br. ex Schult. in Roemer & Schultes, Syst. Veg. 6:59. 1820, err. cit. [=G. mar- itimus (Jacq.) R.Br.] Notes: Brown (1810) validly and effectively published the combination Gonolobus maritimus (Jacq.) R.Br. The sometimes mistakenly cited combination Gonolobus maritimus (Jacq.) R.Br. ex Schult., Syst. Veg. 6:59. 1820, is an error in citation as Schultes clearly accepted Brown's combination, correctly additionally citing Jacquin. See Asclepias maritima Jacq. Gonolobus martinicensis Decne. in De Candolle, Prodr. 8:595. 1844. Protologue: “in Martinicae sylvis umbrosis, (cl. Plée.) Tourimibi Caribeorum Plum. et Surian 821 (v.s.h. Mus. par) Type: Martinique, Pleé s.n. (Lectotype designated by Krings 2007: PD. Notes: Two syntypes were cited in the protologue, but only Pleé s.n. (P) could be re-located. Plum. et Surian 821 could not be found. Gonolobus membranaceus Schltr. in Urban, Symb. Antill. 12):285. 1899. Protologue: “Haiti, loco speciali haud indicato: Poiteau s.n.,” loc. not cited. Type: The location of the type remains unknown. No additional matching material was found. Notes: Ekman H4693 (IJ!) is labelled Gonolobus membranaceus. The specimen consists of mostly stem and 3-4 leaf fragments held in a fragment pocket. It appears to match Poicillopsis crispiflora Urb. Gonolobus mem- branaceus is likely very closely related to Matelea haitiensis as, based on the protologues, the two appear to differ only in the shape and apices of the leaf blades and calyx lobes. Additional collections and discovery of the type may prove them synonymous. Gonolobus nipensis Urb., Symb. Antill. 9(3):421. 1925. = Matelea nipensis (Urb) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941 Protologue: "Prov. Oriente in Sierra de Nipe ad pedem Loma Mensura in savannis pinetorum umbrosis cr. 725 m. alt m. Jul. flor.: n. 9710,” loc. not cited. | js m" p= H I £ B a 147 Krinas W i" hd] Type: Cuba, Oriente, Sierra de Nipe, Loma Mensura, ca. 725 m, 11 Jul 1919, Ekman 9710 (Lectotype, here designated: S!; ISOLECTOTYPE: NY). Notes: Ekman 9710 (S) is a superior s relative to Ekman 9710 (NY) in that it bears many more leaves, inflorescences, and flowers. It is here designated lectotype for Gonolobus nipensis. Gonolobus pauciflorus Spreng., Syst. Veg. 1:846. 1824. = Matelea pauciflora (Spreng.) Woodson, Ann. Mis- souri Bot. Gard. 28:226. 1941 Protologue: “Hispaniola, Bertero,” loc. not cited. Type: Hispaniola: ‘St. Domingue, Bertero s.n. (Lectotype designated by Krings 2006: P!; isoLecToTYPE: G-DC [IDC microfiche Candolle Prodromus Herbarium, fiche 1543!]). Notes: Bertero collections were among specimens in the fragmented Sprengel collection acquired by B in 1890, but no Bertero specimen corresponding to G. pauciflorus was found extant at that institution. As neither extant duplicate of Bertero s.n. at either G-DC or P corresponding to G. pauciflorus bears the hand of Sprengel and as no other Bertero material corresponding to the taxon has been located at the other institu- tions queried (see Acl led ts), itis assumed that the holotype was destroyed. Consequently, having been able to examine the Se at P, but only the microfiche of the specimen at G-DC, Krings (2006) designated the sheet at P lectotype for G. pauciflorus. Gonolobus pubescens Griseb., Fl. Brit. W.I. 420. 1862. = Matelea pubescens (Griseb.) Krings, J. Bot. Res. Inst. Texas 2:131. 2008 Protologue: “Jamaica, Macf.” loc. not cited. Type: Jamaica, Macfadyen s.n. (HOLOTYPE: GOET!; Isotype: KT). Notes: The GOET specimen consists of an unmounted section of stem with five nodes, but only three de- veloped leaves. A fourth developed leaf and a fragment of a fifth are loose in the pocket. A smaller fragment pocket contains floral parts. Macfadyen s.n. (K) is a superior specimen relative to Macfadyen s.n. (GOET) in bearing many more leaves, as well as an openly pressed flower kept in the fragment pocket. Both the K and GOET specimens are annotated in Grisebach’s hand. Gonolobus rhamnifolius Griseb., Fl. Brit. W.I. 420. 1862. = Matelea rhamnifolia (Griseb.) Krings, Sida 21:1515. 2005. Protologue: “Jamaica!, Al., S. Anns, near Moneague,” loc. not cited. Type: Jamaica, R.C. Alexander s.n. (HOLOTYPE: GOET!). Notes: See Krings (2005a) for a discussion of the morphology of this taxon. The type locality was visited on an expedition to Jamaica by the present author and Dale Suiter in March 2006. However, the area was unfortunately flooded (houses under 10—15 feet of water) and inaccessible. The species was not observed on any of the access roads up to the flooded valley. Dr. George Proctor noted collecting a specimen in a narrow patch of woods between a road and a pasture. The area appears to be in similar condition currently, with many pastures separated from rural roads by narrow patches of woods. Gonolobus rostratus (Vahl) Schult. in Roemer & Schultes, Syst. Veg. 6:61. 1820. Notes: Schultes (1820) cited Vahl's protologue of Cynanchum rostratum (see above) and attributed the com- bination in Gonolobus to Robert Brown. However, as Brown never made the combination in Gonolobus that has been attributed to him, Schultes should be considered the author of the valid combination. See Cynanchum rostratum Vahl. "Gonolobus rostratus" auct. non (Vahl) Schult.: Schlechter in Urb., Symb. Antill. 1:284. 1899. Notes: Schlechter (1899) mistakenly attributed a Jamaican entity to the name Gonolobus rostratus (Vahl) R.Br., for which Rendle later proposed the name G. jamaicensis (see above). Robert Brown never made this combination attributed to him (see Memoirs of the Wernerian Natural History Society 1:35. 1810). See also Cynanchum rostratum Vahl. "Gonolobus scandens (Aubl.) Urb.," Repert. Spec. Nov. Regni Veg. 16:151. 1919; nom. illeg. 148 | tl tanical h Inst Texas 2( ] 4 Ankh] Hietaira A ] + Ta Ta 434 E 4 Notes: Based on an invalid polynomial 2 (Tabl. Nom. Lat.):23. 1775. 260. See Nestor (1991) for a discussion. Gonolobus geet Urb. Ld aped noe Regni Veg, D 151. 1919; nom. illeg. Notes: A sul y | is Decne. (Prodromus Systematis Naturalis Regni Vegetabilis 8:595. 1844) Gonolobus sintenisii Schltr. in Urban, Symb. Antill. 12):288. 1899. = Matelea sintenisii (Schltr.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941 Protologue: “Portorico, Sierra de Luquillo, in regione superiore montis Jimenes in silvis, Jul. 1885: Sintenis n. 1354; in silva primaeva montis Andubo prope Adjuntas, Jun. 1886: Sintenis n. 4643,” loc. not cited. Type: Puerto Rico, Sierra de Luquillo, Jiménez mts., 9 Jul 1885, Sintenis 1354 (Lecrorvre, here designated: HBG!; ISOLECTOTYPES: BM!, G!, GH!, K!, US!). Notes: Among the syntypes, Sintenis 1354 (HBG) is chosen as ce as Hic EE bears flowers in relatively superior condition to the duplicates under this number. The t ly at Gand GH are sterile. The K specimen bears a single flower in bud. Specimens of Sis 4643 eld not Te located. Gonolobus stapelioides Desv. ex Ham., Prodr. Pl. Ind. Occid. 32. 1825. Protologue: “Habitat in fruticetis circa Alfred Hall, prope Scarborough, Tobago, cum Echite quinquangulare etc. ubi florentem inveni mense Octobris (V. et S. vi" loc. not cited. Type: Jamaica, Portland Parish, 0.5 mi N of Hardwar Gap, uphill from the trail above the “Waterfall,” ca. 3900 ft, montane rainforest, vines growing over trailside shrubs and in small forest openings, population J1, 3 Mar 2006, A. Krings 1395 with D. Suiter and G.R. Proctor (neotype: P!, designated by Krings 2008; iso- neotypes: BM!, Ij!, NCSC!, NY). Notes: Schlechter (1899) noted that there was likely a label mix-up | the Jamaican entity G. stapelioides and the Tobagoan G. virescens Ham. It would indeed be unlikely for G. stapelioides to be limited in distribu- tion to Jamaica and Tobago. The taxon is not represented by any other specimens from Tobago. Similarly, G. virescens is not known from Jamaica by another specimen. A neotype is proposed as the holotype could not be found. Gonolobus stellatus Griseb., Fl. Brit. W.I. 420. 1862. Protologue: “Jamaica!, Pd., Manchester, rare, near Knockpatrick,” loc. not cited. Type: Jamaica, Purdie s.n. (HOLOTYPE: GOET!; isotypes: BM!, K!). Notes: Purdie s.n. (K) is the best specimen of the three extant sheets. It | leaves, and an openly pressed flower. The specimen also bears an annotation in Grisebach's hand. The specimen at BM is also in good condition, bearing a nicely preserved flower. However, this specimen does not bear annotations in Grisebach's hand. The specimen at GOET consists entirely of fragments (two leaves and immature flowers in bud; no stem). $ £1 Gonolobus stephanotrichus Griseb., Cat. Pl. Cub. 177. 1866. Protologue: “Cuba or. (Wr. 2969 [...],” loc. not cited. Type: Cuba, 1860-1864, Wright 2969(LectorYrE, designated by Krings & Fantz 2006: GOET!; syntypes: BMI, BREM!, G!, GH!, HACI, K!, MO, NY!, P!, UC!, USD. Notes: The Wright collection of G. stephanotrichus at S bears the number 164 and does not appear to be part of the type collection. See Krings and Fantz (2006) for a detailed discussion of the lectotypification of G. stephanotrichus. Gonolobus stipitatus Alain, Phytologia 64(5):345. 1988, nom. illeg.; non Gonolobus stipitatus Morillo, 1987. = Gonolobus haitiensis P.T.Li, J. South China Agric. Univ. 14(1):58. 1993. Protologue: “Haiti: On hard limestone, Massif du Nord, St. Louis du Nord, on top of Morne Baron, Jul 20, 1925, Ekman 4693 (noLorvpus: S).” Type: Haiti, Massif du Nord, St. Louis du Nord, Morne Baron, 20 Aug 1925, Ekman H 4693 (HoLoTYrE: Si; ISOTYPES: B!, USD. Krinas W wi P “Gonolobus suberosus” auct. non (L.) R.Br.: Spreng., Syst. Veg. 1:846. 1824. Notes: Cited as Gonolobus suberosus Spreng. by ge 11099) aw Mierer oan. SIUE Sprengel (1824) clearly cited Robert Brown as author of th bi G Cynanchum suberosum L. Although the typification of C. suberosum L. i lex (Drapalil RR EN a & ERE n the basionym has most recently been recognized to apply to the single | United States (see Krings & Xiang 2004, 2005). Schlechter (1899) and Woodson (1941) considered Spren- gel's G. suberosus to be a synonym of what is now known as Matelea maritima (Jacq.) Woodson. Sprengel's description is not inconsistent with their decision. Gonolobus tigrinus Griseb., Mem. Amer. Acad. Arts 8:520. 1863. = Matelea tigrina (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. Protologue: “[Wright] (1667); loc. not cited. Type: Cuba, Holguín, 19 Mar, Wright 1667 (Lectotype, here designated: GOET, n.v., fide Howard 19892; SYNTYPES: MO [image online!], G!, GH!, KI Notes: Specimens distributed under Wright 1667 were collected by Wright from at least two different lo- calities at two different times (GH: *in coffee fields, Josephina, Nov 1"; GOET: *Holguin, 19 Mar"). Krings (20050) mistakenly stated that Howard lectotypified G. tigrinus based on Wright 1667 at GOET. However, Howard's *lectotypification" (19892) appeared in a microfiche appendix, not in print, and thus is not ef- fectively published and has no standing in nomenclature. Rankin & Greuter (2000) reported a similar case in Antillean Aristolochia. Gonolobus tigrinus Griseb. var. angustifolius Griseb., Cat. Pl. Cub. 177. 1866. Protologue: "Wr. [Wright]? loc. not cited. Type: The location of the type remains unknown. No additional matching material was found. Notes: This taxon was recognized at the specific rank by Schlechter (1899) as G. grisebachianus Schltr. Gonolobus tobagensis Urb., Repert. Spec. Nov. Regni Veg. 16:37. 1919. Protologue: “in Tobago in districtu The Widow nominato, m. April. flor, W.E. Broadway no. 4467 [loc. not cited].” Type: Tobago, the Widow, 28 Apr 1913, WE Broadway 4467 (Lectotyre, here designated: BM!). Notes: Among the great bulk of collections received by Urban from Erik Ekman (Cuba and Hispaniola) and William Harris Jamaica), are specimens sent to him by W.E. Broadway from Trinidad, Tobago, and Grenada. Holotypes are presumed to have remained at B until their destruction in a fire in World War II. There is no indication on Broadway 4467 (BM) that it was seen by Urban, but as the only apparently extant sheet of the number cited by Urban in the protologue, it is here designated lectotype of G. tobagensis. Gonolobus variifolius Schltr. in Urban, Symb. Antill. 1(2):286. 1899. = Matelea variifolia (Schltr.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. Protologue: “Portorico, Sierra de Luquillo, in regi n. 1653, loc. not cited; ‘montis Cienega prope Adjuntas, Jun. 1886: Sintenis n. 4687,” loc. not cited. Type: Puerto Rico, Sierra de Luquillo, Jiménez mts., Jun 1885, P. Sintenis 1653 (Lectotype, here designated: NY; ISOLECTOTYPES: G!, GH!, MOI, P!, USD; P. Sintenis 4687 (syntyee: K). Notes: Sintenis 1653 (NY) is here designated as lectotype for Gonolobus variifolius Schltr. as, unlike the other specimens, it bears a diversity of leaves, a number of inflorescences, and flowers. 1 D montis Jimenes in fruticetis, Jun. 1885: Sintenis Gonolobus viridiflorus Schult. in Roemer & Schultes, Syst. Veg. 6:61. 1820.; nom. illeg., non G. viridiflorus Nutt. (1818) Protologue: “Meyer,” loc. not cited. Type: unknown. Notes: Schultes (1820) clearly cited Meyer's basionym, Cynanchum viridiflorum. However, C. viridiflorum G. Mey. is illegitimate, being a later homonym of C. viridiflorum Sims (1817). The basionym epithet is not avail- able in Gonolobus due to the earlier G. viridiflorus Nutt. (1818). Most recent authors have accepted this entity [| E al n H Il m LI dee o £T. 150 Jo t Texas 2( under the name Matelea denticulata (Vahl) Fontella & E.A. Schwarz (Boletim do Museu Botanico Municipal 46:4. 1981.), although, as it emerged in a clade of Gonolobus s.1. (Krings et al. 2008) the name G. denticulatus (Vahl) W.D. Stevens is preferable. Gonolobus waitukubuliensis Krings, Syst. Bot. 32:187. 2007. Protologue: “Dominica. In sylvis ad Laudat, Mai 1882 (fl), Eggers 728 (HoLotyPE: HBG! [2 sheets]; ISOTYPES: FR!, G!, GHI, JEI, M!, W!, Z0." Type: Dominica, in sylvis ad Laudat, May 1882, Eggers 728 (noLorvre: HBG! [2 sheets]; isotypes: FRI, Gl, GH!, JE! MI, WI, ZÌ). Gonolobus youroumaynensis Krings, Syst. Bot. 32:191. 2007. Protologue: “Orange Hill Estate, roadbank in banana and pigeon pea plantations, uphill from fork in road to the CW.S.A. water intake site, Kiss-me, 14 Mar 2006 (fl), Krings 1374 with Springer (HOLOTYPE: NC SCT IsOTYPES: GHI, K!, NY!, US!, PD. Type: St. Vincent, Orange Hill Estate, roadbank in banana and pigeon pea plantations, uphill from fork in roSad to the Central Water and Sewage Authority water intake site, Kiss-me, 14 Mar 2006, Krings 1374 with Springer (HOLOTYPE: NCSC!; isotypes: GHI, KI, NY!, US!, P Holostemma candolleanum Spreng., Syst. Veg. 1:851. 1824, nom. illeg. Notes: Sprengler (loc. cit.) cited F. scandens DC. in synonymy, thus making hi illegitimate. Schlechter (1899) placed Sprengler's name in synonymy under Fischeria crispiflora (Sw.) Schltr.—a m homonym of F. which he misapplied to the taxon recognized by Murphy (1986) as F. scandens crispiflora (Sw.) K. Schum. DC Ibatia maritima (Jacq.) Decne. in De Candolle, Prodr. 8:599. 1844. = Matelea maritima (Jacq) Woodson, Ann. Missouri Bot. Gard. 28:222. 1941. See Asclepias maritima Jacq. Ibatia mollis Griseb., Cat. Pl. Cub. 177. 1866. s Matelea mollis (Griseb.) Woods., Ann. Missouri Bot. Gard. 28:223. 1941 Protologue: “Cuba occ. - pr. La Concordia (Wr. 2978)” loc. not cited. Type: Cuba, te 2978 D E EE GH!; syntypes: K!, MO! [image online]). Notes: See di under G ling problems of typification of Grisebach names based on Wright collections. Unfortunately, there isno ant Wright material of number 2978 at GOET. However, a specimen does exist at GH, which can be designated lectotype following Howard (1989a). Two elements are mounted on the sheet—one with flowers and one with fruits. As fruits were not mentioned in the pro- tologue, the flowering element of Wright 2978 (GH) is here designated as lectotype for Ibatia mollis Griseb. Other combinations include: Ptycanthera mollis (Griseb.) Schltr. in Urb., Symb. Antill. 1(2):280. 1899. Ibatia muricata Griseb., Fl. Brit. W.I. 421. 1862. Protologue: “Antigua!, Wullschl., Dominica!, Imr., [Guadeloupe! ; Venezuela!],” loc. not cited. Type: Antigua, Cedar Hall, 1849, Wullschlagel (Lectotype designated by Krings & Saville 2007: M! [2 sheets]) Notes: Although not identified as a type, the material at M was the only Wullschlagel specimen from An- tigua found in the course of a study by Krings and Saville (2007) and was designated lectotype of Ibatia muricata. Grisebach's name is a synonym of Matelea maritima (Jacq.) Woodson, Ann. Missouri Bot. Gard. 28:222. 1941 Jacaima costata (Urb.) Rendle, J. Bot. 74:340. 1936. = Matelea costata (Urb.) Morillo, Anales Jard. Bot. Madrid 43:239. 1987 [“1986”]. See Poicilla costata Urb. Jacaima costata (Urb.) Rendle var. goodfriendii Proctor, J. Arnold Arbor. 63:290. 1982. = Matelea costata (Urb.) Morillo var. goodfriendii (Proctor) Krings, comb. nov. Krings, DA bl ys r "Le Li J E I ty 151 Protologue: “St. Ann: Cedar Valley distr., ca. 1.5 mi NE of Cave Valley Square, ca. 2000 ft, Goodfriend s.n. (IJ 65657), May 17, 1977 (flowers) (HOLOTYPE), Goodfriend s.n. (IJ 66697), Jul 1977 (fruit).” Type: Jamaica, St. Ann: Cedar Valley distr., ca. 1.5 mi NE of Cave Valley Square, wooded limestone hills, ca. 2000 ft, 17 May 1977, Goodfriend s.n. (HoLoTYeE: IJ); Jul 1977, Goodfriend s.n. (PARATYPE: IJI). Jacaima parvifolia Proctor, J. Arnold Arbor. 63:291. 1982. = Matelea proctori Krings, nom. Protologue: "Clarendon: Broom Hall hills, 1.2 mi SW of Cave Valley Square, 1800-2000 E M 37887, Jul 9, 1978 (HoLOTYPE).” Type: Jamaica, Clarendon: Broom Hall hills, 1.2 mi SW of Cave Valley Square, 1800—2000 ft, 9 Jul 1978, Proctor 3/887 (HoLoTYPE: GH!; isoTYPE: IJ!). Notes: Á new name is required as a new combination in Matelea is blocked by M. parvifolia (Torr.) Woodson. As Proctor 37887 (IJ) is labelled isotype, it is assumed that the GH duplicate was intended to serve as the holotype. Dr. Proctor accompanied the current author, Dale SEN and ER Campbell, on an expedition to the type locality in March of 2006. At the time tl ibed the area was called Broom Hall and the valley was mostly in pineapple cultivation (Godo pers. comm.). On a change of ownership, the area is now mostly in coffee and under the name Baron Hall. Although the hills maintain forest cover, much is disturbed and has either been converted to coffee or is in the process of conversion. Specimens of J. parvi- folia were not located. There are forests not owned by Baron Hall which are part of the endemic Peckham complex and may retain populations of J. parvifolia, but further research is needed. Lachnostoma maritimum (Jacq.) G. Nicholson, Ill. Dict. Gard. 2:226. 1884. = Matelea maritima (Jacq.) Woodson, Ann. Missouri Bot. Gard. 28:222. 1941. See Asclepias maritima Jacq. Lachnostoma mollis (Griseb.) M. Gómez, Anales Hist. Nat. 23:276. 1894. = Matelea mollis (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:223. 1941. See Ibatia mollis Griseb. Macroscepis hirsuta (Vahl) Schltr. in Urban, Symb. Antill. 1:265. 1899. = Matelea hirsuta (Vahl) Woodson in E.E. Cheesman, Fl. Trinidad 2:170. 1 See Cynanchum hirsutum Vahl. “Macroscepis obovata” auct. non Kunth: Griseb., Fl. Brit. W.I. 421. 1862. Notes: The most recent combination to which Grisebach misapplied the name Macroscepis obovata is Matelea hirsuta (Vahl) Woodson in E. E. Cheesman, Fl. Trinidad 2:170. 1947. See Cynanchum hirsutum Vahl. Marsdenia bicolor Britton € P. Wilson, Bull. Torrey Bot. Club 50:47. 1923. = Matelea bicolor (Britton & P. Wilson) Woodson, Ann. Missouri Bot. Gard. 28:236. 1941. Protologue: “Sierra Maestra, Oriente (León 10787, type; 10788); loc. not cited. Type: Cuba, Oriente, Maestra ridge, 1300 m, Jul 1922, León 10787 (noLoTYPE: NY!; isorYPE: HAC, n.v.); Cuba, León 10788 (paratypes: NY!, US). Notes: Although TROPICOS currently indicates the holotype to be at GH (as of 12 Apr 2006), the specimen is not listed in the GH online catalogue nor was it sent on loan or found in a targeted search by collection manager Emily Wood. In addition, there is a discrepancy in the label data, particularly the altitude of col- lection, for the paratype Léon 10788, suggesting that the sheets at NY and US are not duplicates of the same collection. The NY sheet indicates only “Maestra ridge, 1300 m,” whereas the US sheet indicates “Woods: Sierra Maestra. Region of Pico Turquino. Alt. near 2000 m.” Collector, collector number, and date are the same on both sheets. Matelea acuminata (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:225. 1941. See Orthosia acuminata Griseb. Matelea alainii Woodson, Contr. Ocas. Mus. Hist. Nat. Colegio “De La Salle” 15:23. 1956. 152 Jo | tanical xas 2( Protologue: “Cuba: Oriente: on coastal limestone rocks, Km. 90 between Imías and Cajobabo, Vía Azul, Jan. 12, 1956,” Brother Alain & C.V. Morton 5029 (HT: MO); “on coastal limestone rocks. Km 50 of Via Azul, Jan 10, 1956,” Brother Alain & C.V. Morton 4959 (PT: MO). Type: Cuba, Alain & Morton 5029 (noLotYPE: MO; isotypes: GH}, HAC!, US); Alain & Morton 4959 (PARATYPES: GH!, MOL, 1). Matelea annulata Woodson ex Alain, Brittonia 20:149. 1968. Protologue: “Hispaniola: Dominican Republic: Montiada Nueva, Barahona Prov., 21-25 Aug 1946, R. & E. Howard 8512 (Type: NY; isotype: GH); Howard 8590 (GH)? Type: Dominican Republic, Montiada Nueva, Barahona Prov., 21-25 Aug 1946, RA. & E.5. Howard 8512 (HOLOTYPE: NY! [2 sheets]; isotypes: GH!, MO). Notes: Alain H. Liogier described Matelea annulata based on suggestions from Robert E. Woodson, Jr. and legitimized the latter's nomen in schedula. Three sheets were cited in the PROTOLOGUE: R. & E. Howard 8512 (HT: NY!: IT: GH) and Howard 8590 (PT: GH). The former two contain matching material. However, the latter collection (the paratype), which incidentally was also collected by both Howards, contains material that clearly does not belong to the holotype and isotype. Although also glabrous, leaves of R. & E. Howard 8590 exhibit: (1) more numerous secondary veins, and (2) secondary veins nearly straight (vs. ascending in R.& E. Howard 8512). The seeds associated with R. & E. Howard 8590 are also: (1) significantly longer (11-11.6 mm vs. 5 mm in R. & E. Howard 8512), (2) of a thin papery texture (vs. hardened in R.& E. Howard 8512), and (3) lack the distally swollen margin seen in Howard 8512 (and other West Indian Matelea species). Although published as a paratype, R. & E. Howard 8590 (GH) must be excluded from Matelea ipia It remains "Jr unclear to what taxon it should be referred instead. Typification of M As a final note, in contrast to the indication by Dosis (1968), t the of Matelea lat t solitary They appear to be borne sequentially on inflorescences subtended by a much reduced peduncle. Matelea bayatensis (Urb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. See Gonolobus bayatensis Urb. Matelea bicolor (Britton & P. Wilson) Woodson, Ann. Missouri Bot. Gard. 28:236. 1941. See Marsdenia bicolor Britton & P. Wilson. Matelea borinquensis Alain, Phytologia 61:360. 1986. Protologue: “Puerto Rico: Cerro Pelucho, San Lorenzo, alt. 400—500 m, Mar 8, 1984, Alain & Perfa Liogier, Luis F. Martorell 35111 (holotypus: UPR).” Type: Puerto Rico, Liogier et al. 35111 (HoLoTYPE: UPR, n.v.). Notes: A synonym of Matelea variifolia (Schltr) Woodson fide Acevedo-Rodríguez (2005). This specimen could not be obtained on loan, despite requests. Matelea constanzana J. Jiménez Alm., Rhodora 62:238. 1960. Protologue: “in Sto. Domingo prope Constanza, inter frutices volubilis 1400 m. alt., H. von Tuerckheim n. 3466, flor. Jul. 1910” loc. not cited. Type: The location of the type remains unknown. No additional matching material was found. Notes: Nomen nov. for Poicillopsis tuerckheimii Schltr. in Urb., Symb. Antill. 7(3):339. 1912. Jiménez pro- posed the new name as Schlechter's epithet was unavailable in Matelea. The name is provisionally treated in Matelea, pending the rediscovery and study of the type. Matelea correllii Spellman, Ann. Missouri Bot. Gard. 65(4):1255. 1979 [*1978"]. Protologue: “Bahama Islands. Long Island: ... low places along Queen's Highway about 4 miles north of Clarence Town center, 18 Nov. 1977, D.S. Correll 49112 (MO, HoLotYPE; F, FIG, GH, NY, US, isorvpes)." Type: Bahamas, Long Island, D.S. Correll 49112 (uoLorvre: MO; IsoTYPES: El, FTG!, GH, NY!, USD. Matelea corynephora Krings, Sida 22:942. 2006. Krings, W A) As Fal mL. a [| £ IA 153 di Protologue: “Haiti. Ile la Tortue, la Vallée, top of Morne Barranca, ca. 300 m, 21 Mar 1928 (fl), E.L. Ekman H9740 (HOLOTYPE: SD? Type: Haiti, Ile la Tortue, la Vallée, top of Morne Barranca, ca. 300 m, 21 Mar 1928, E.L. Ekman H9740 (HOLOTYPE: SP). Matelea costata (Urb.) Morillo, Anales Jard. Bot. Madrid 43:239. 1987 [“1986”]. See Poicilla costata Urb. Matelea costata (Urb.) Morillo var. goodfriendii (Proctor) Krings, J. Bot. Res. Inst. Texas 2:150. 2008. See Jacaima costata (Urb.) Rendle var. goodfriendii Proctor Matelea crispiflora (Urb.) J. Jiménez Alm., Rhodora 62:238. 1960. See Poicillopsis crispiflora Urb. Matelea ekmanii (Urb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. See Gonolobus ekmanii Urb. Matelea denticulata (Vahl) Fontella & E.A. Schwarz, Bol. Mus. Bot. Munic. 46:4. 1981. See Cynanchum denticulatum Vahl. Matelea dictyopetala (Urb. G Ekman) Krings, J. Bot. Res. Inst. Texas 2:130. 2008. See Gonolobus dictyopetalus Urb. & Ekman. Matelea domingensis (Alain) Krings, Sida 21:2081. 2005. See Gonolobus domingensis Alain Matelea grisebachiana (Schltr.) Alain, Mem. Soc. Cub. Hist. Nat. “Felipe Poey” 22:120. 1955. See Gonolobus grisebachianus Schltr. Matelea hastata Alain, Brittonia 20:150. 1968. Protologue: “Haiti: Massif des Matheux, Croix des Bouquets, Morne a Cabrits, 18 Jul 1924, Ekman H. 973 (Type: US; isotype: NY)" Type: Haiti, Eeman H 973 (HoLorYrE: US!; isotypes: K!, NY!) Matelea hirsuta (Vahl) Woodson in E.E. Cheesman, Fl. Trinidad 2:170. 1947. See Cynanchum hirsutum Vahl. Matelea linearipetala Alain, Phytologia 64:346. 1988. Protologue: “Haiti: On oligostene limestone, 200 m alt., Massif des Matheux, Thomazeau, Morne à Cabrits, Oct. 24, 1926, E.L. Ekman 7136 (HoLotypus: S).” Type: Haiti, Morne à Cabrits, Ekman H 7136 (HoLoTYPE: St). Matelea maritima (Jacq.) Woodson, Ann. Missouri Bot. Gard. 28:222. 1941. See Asclepias maritima Jacq. Matelea mollis (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:223. 1941. See Ibatia mollis Griseb. Matelea monticola Alain, Sida 20:1645. 2003. Protologue: “Ciénaga de la Culata, Constanza, alt 1600-1700 m, 15-16 Oct 1968, Alain Liogier 13029 (HOLOTYPE: NY; isotype: US); in woods, Cabezada de la Ciénaga de la Culata, Constanza, alt 1650 m, 16 Oct 1968, Alain Liogier 13069 (NY, US)" Type: Dominican Republic, Ciénaga de la Culata, Constanza, alt 1600-1700 m, 15-16 Oct 1968, Liogier 13029 (HoLoTYPE: NY. Dues UE nov. for Matelea sylvicola Alain, Phytologia 22:168. 1971, nom illeg., non L.O. Williams 1968. e housed at US, despite so cited in the protologue. The paratype is also neither listed in the NY "n lose nor included in a ben from NY. Its location remains unknown. [| E sl n.a D In LI E i.a A £T iai AT? 154 Matelea nipensis (Urb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. See Gonolobus nipensis Urb. Matelea oblongata (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. See Orthosia oblongata Griseb. Matelea ovatifolia (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:225. 1941. See Poicilla ovatifolia Griseb. Matelea pauciflora (Spreng.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. See Gonolobus pauciflorus Spreng. Matelea pentactina Krings, Sida 21:1519. 2005. Protologue: “Haiti. Les Roseaux, Massif de la Hotte western group, rocky ledge, ca. 1300 m, 16 Sept 1928, Ekman H 10685 (HoLoTYPE: S!); Massif de la Selle, gr. Crete-a-Piquants, Port-au-Prince, between Carrefour- Martin and Bois d'Orme, ca. 800 m, limestone, 17 Dec 1926, Ekman H 7402 (paratype: Sn." Type: Haiti, Les Roseaux, Massif de la Hotte western group, rocky ledge, ca. 1300 m, 16 5ept 1928, Ekman H 10685 (mozorvpe: S!; isotypes: B!, US!, EHH n.v); Massif de la Selle, gr. Crete-a-Piquants, Port-au-Prince, between Carrefour-Martin and Bois d'Orme, ca. 800 m, limestone, 17 Dec 1926, Ekman H 7402 (PARATYPE: S!, EHH ny). Matelea phainops Krings, Sida 22:948. 2006. Protologue: “Dominican Republic. Vine, up to 1.5 m high, flowers yellowish green, in thickets, about 2 mi W of Oviedo, alt. about sea level, on limestone, 3 Nov 1989 (fl), A.H. Liogier 16617 (HOLOTYPE: GH!; ISOTYPES: NY! USED.” Type: Dominican Republic, about 2 mi W of Oviedo, alt. about sea level, on limestone, 3 Nov 1989, Liogier 16617 (HOLOTYPE: GH!; 1sorypes: NY!, USF). Matelea proctori Krings, J. Bot. Res. Inst. Texas 2:151. 2008. See Jacaima parvifolia Proctor Matelea pubescens (Griseb.) Krings, J. Bot. Res. Inst. Texas 2:131. 2008. See Gonolobus pubescens Griseb. Matelea rhamnifolia (Griseb.) Krings, Sida 21:1515. 2005. See Gonolobus rhamnifolius Griseb. Matelea rhynchocephala Krings, Sida 22:949. 2006. Protologue: “Dominican Republic. Prov. Santiago, Valle del Cibao, Santiago, Hato del Yaque, in thickets, fl. green, fruiting, 15 Feb 1930 (fl & fr), E.L. Ekman H14296 (HOLOTYPE: S!).” Type: Dominican Republic, Prov. Santiago, Valle del Cibao, Santiago, Hato del Yaque, in thickets, 15 Feb 1930, E.L. Ekman H 14296 (mororyre: SD. Matelea rubra (H. Karst.) Spellman & Morillo, Phytologia 34:152. 1976. See Omphalophthalma rubra H. Karst. Matelea rubra (H. Karst.) Aa & Stoffers, Proc. Kon. Ned. Akad. Wetensch., C 84(3):309. 1981. Notes: Matelea rubra (H. Karst.) Aa & Stoffers is nomenclaturally superfluous as the combination by Spell- man & Morillo (loc. cit.) has priority. See Matelea rubra (H. Karst.) Spellman & Morillo. Matelea sintenisii (Schltr) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. See Gonolobus sintenisii Schltr. Matelea sylvicola Alain, Phytologia 22:168. 1971, nom. illeg., non L.O. Williams 1968. See Matelea monticola Alain. Krings, W al H Fal Un D [| r Ba 155 di Matelea tamnifolia (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:225. 1941. See Poicilla tamnifolia Griseb. Matelea tigrina (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. See Gonolobus tigrinus Griseb. Matelea torulosa Krings, Sida 22:951. 2006. Protologue: “Dominican Republic. Distr. Nacional, Los 3 Ojos de Agua, near Santo Domingo, 31 Oct 1959 (fl), E. Marcano ILL Jiménez] 4096 (HoLotYPE: USD." Type: Dominican Republic, Distr. Nacional, Los 3 Ojos de Agua, near Santo Domingo, 31 Oct 1959, E. Marcano [].]. Jiménez] 4096 (HoLotYrE: US). Matelea variifolia (Schltr.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. See Gonolobus variifolius Schltr. Matelea viridiflora (G. Mey.) Woodson, Ann. Missouri Bot. Gard. 28:325. 1941. See Cynanchum denticulatum Vahl (syn. fide Howard 1989b). Matelea viridivenia Alain, Phytologia 22:169. 1971. Protologue: “Cabo Rojo, Pedernales, 4 Nov 1969, Alain Liogier 16637 (4oLorYPE, NY; isorypes: US, GH, P, IJ); about 5 miles E of Cabo Rojo, 8 Feb 1969, Alain Liogier 13620 fruiting specimen (NY); in thickets, km 92 from Bani to Azua, alt 80 m, 3 Nov 1969, Alain Liogier 16598 (NY, US, GH, P); on limestone, los Guanitos, 7 mi E of Cabo Rojo, Pedernales, 13 Nov 1969, Alain Liogier 16961b (NY, US, GH, P, JJ).” Type: Dominican Republic, Cabo Rojo, Pedernales, 4 Nov 1969, Alain Liogier 16637 (HoLoTyPE: NY!); about 5 miles E of Cabo Rojo, 8 Feb 1969, Alain Liogier 13620 fruiting specimen (paratype: NY); in thickets, km 92 from Bani to Azua, alt 80 m, 3 Nov 1969, Alain Liogier 16598 (paratype: NY!); on limestone, los Guanitos, 7 mi E of Cabo Rojo, Pedernales, 13 Nov 1969, Alain Liogier 16961b (paratype: NY!, USF). Notes: The types or paratypes listed in the protologue for GH, IJ, P, and US have not been found. Omphalophthalma rubra H. Karst., Fl. Columb. 2:119, t. 163. 1866. = Matelea rubra (H. Karst.) Spellman & Morillo, Phytologia 34:152. 1976. Protologue: "Habitat litora maris Caribaei prope St. Martam [Colombia], *loc. not cited. Type: Colombia [New Granada], St. Marta, Herb. Karsten (Hotorvee: LE [Sennikov, pers. comm.; n.v., not sent on loan due to budget constraints]). Orthosia acuminata Griseb., Cat. Pl. Cub. 175. 1866. = Matelea acuminata (Griseb.) Woodson, Ann. Mis- souri Bot. Gard. 28:225. 1941. Protologue: “Cuba or. (Wr. 2966),” loc. not cited. Type: Cuba, Oriente, Wright 2966, 1860-1864 (LecroTYPE designated by Krings ër Fantz 2006: GH!; isoLEc- TOTYPES: BM!, G!, HACI [2 sheets], K!). Notes: Krings and Fantz (2006) noted that sheets of Wright 2966 (BM, G, GH, HAC, and K) bear white labels with the dates 1860-64. The mounted field ticket on the GH sheet reads: “Asclepias — Fl. (except the white stigma) green. Farallones San Andre Oct 27.” As Wright 2966 (GH!) contains fifteen inflorescences and is in very good condition; it was designated as lectotype for Orthosia acuminata Griseb. by Krings and Fantz (2006). Wright 2966 (G!) contains four inflorescences and is in superior condition to the HAC mate- rial. Wright 2966 (BMI, G!, HAC!, K!) should be considered isolectotypes. See Krings and Fantz 2006 for additional discussion regarding their philosophy of typification of Wright specimens. Orthosia oblongata Griseb., Cat. Pl. Cub. 176. 1866. = Matelea oblongata (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. Protologue: “Cuba occ. (Wr. 2967)” loc. not cited. Type: Cuba, Wright 2967, 1860-1864 (Lectotype designated by Krings & Fantz 2006: GH!; ISOLECTOTYPES: BM!, G!, HAC!, K!, MO, SD. Notes: Krings and Fantz (2006) noted that sheets of the original material for O. oblongata—at BM!, G!, GHI, 156 | tl tanical Insti Texas 2( HACI, K!, MO, and Si—Aall bear white labels with the dates 1860-64. 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. Krings and Fantz (2006) designated Wright 2967 (GH) lectotype for Orthosia oblongata Griseb., considering the duplicates at G, HAC, K, MO. and S isolectotypes. Wright 2967 (GH) is in good condition, with inflorescences. See Krings and Fantz (2006) for additional discussion regarding their philosophy of typification of Wright specimens. Poicilla acuminata (Griseb.) Schltr. in Urban, Symb. Antill. 5(3):469. 1908. = Matelea acuminata (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:225. 1941. See Orthosia acuminata Griseb. Poicilla costata Urb., Symb. Antill. 6(1):38. 1909. =Matelea costata (Urb.) Morillo, Anales Jard. Bot. Madrid 43:239. 1987 [*1986"]. Protologue: *Hab. in Jamaica juxta viam ad Wareka in Long Mountain ad latus australe, 200 m. alt., m. Nov. fl., m. Jun. fruct.: Harris n. 9590, 10006,” loc. not cited. Type: Jamaica, road to Wareka, Long Mountain, 5 side, 19 Nov 1907 (fl), W. Harris 10006 (LecTOTYPE, here designated: BMI; rsorecrorvee: UCWI), Jamaica, Long Mountain, S side, 600 feet altitude, 21 Jun 1907 (fr), W. Harris 9590 (syntypes: NY!; UCWID. Notes: Harris 9590 (NY) is annotated in Urban's hand. Poicilla mollis (Griseb.) Schltr. in Urban, Symb. Antill. 5(3):470. 1908. = Matelea mollis (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:223. 1941. See Ibatia mollis Griseb. Poicilla oblongata (Griseb.) Schltr. in Urban, Symb. Antill. 5(3):470. 1908. = Matelea oblongata (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. See Orthosia oblongata Griseb. Poicilla ovatifolia Griseb., Cat. Pl. Cub. 177. 1866. = Matelea ovatifolia (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:225. 1941. Protologue: “Cuba or. (Wr. 2965)" loc. not cited. Type: Cuba, Oriente, Wright 2965 (LECTOTYPE designated by Krings and Fantz 2006: GOET!; synryres: BMI; BREM!, G!, GH!, HAC!, K nv., LE n.v., NY!, PI, S!, UCI). Notes: As Krings and Fantz (2006) noted, no field Ge accompany the GOET specimen or any syntype, suggest that the sheet is comprised A + OLI g field tickets oft UE except the GH specimen of at least two collections iade at different times (Mar, Jun), nl three fragments are mounted. The two fragments mounted on the right contain inflorescences; the fragment mounted on the left contains in- fructescences. 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 xu a prn eem of 1600, ALB Ge zero EE to have been crossed out. It is heavily written on in Gri is herein desi illa ovatifolia Griseb. The studied (and matching) duplicately-numbered material in her eer remain syntypes. Poicilla tamnifolia Griseb., Cat. Pl. Cub. 176. 1866. = Matelea tamnifolia (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:225. 1941 Protologue: “Cuba or., in monte Toro pr. Potosi (Wr)? loc. not cited. Type: Cuba, San José, 5 Oct, Wright s.n. (LECTOTYPE, here designated: GH!); Wright s.n. (syntyre: NY! [frag- ment]). Notes: See di ion under Gonolobus tigrinus regarding problems of typification of Grisebach names based on Wright collections. Unfortunately, deren is no extant Wright RE corresponding to Poicilla tamnifolia at GOET. However, a specimen does exist at GH, which can be designated lectotype following Howard (198923): Cuba, San José, 5 Oct, Wright s.n. Poicillopsis crispiflora Urb., Feddes Repert. 19:7. 1923. 2 Matelea crispiflora (Urb.) Jiménez, Rhodora :238. 1960. Protologue: “Haiti prope Furcy 1500 m alt., m. Sept. flor.: Buch no. 1995,” loc. not cited. Type: Haiti, prope Furcy, im Gebüsch, Blth weiss, 1500 m alt., m. Sept. flor., Buch no. 1995 (LecToTYPE, here designated: UD. Notes: Buch 1995 consists of four flowers in two umbellate inflorescences and a few leaves on stems, though most are in a fragment pocket. The single additional collection of this species that matches the protologue is Ekman H1933 (EHH nv., K!, S!): Haiti, Morne de la Selle, Furcy, 1540m, in Buchs [sic] Botan. Garden, 17 Sept 1924. The sheets at K and S are in exceedingly good condition, bearing numerous leaves and flowers. The sheet at EHH has not been seen, but is listed here based on the catalogue provided by: http://www. umce.ca/cours/martin/herbier ekman/recherche pl. Poicillopsis ovatifolia (Griseb.) Schltr. in Urban, Symb. Antill. 7(3):339. 1912. = Matelea ovatifolia (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:225. 1941. See Poicilla ovatifolia Griseb. Poicillopsis tuerckheimii Schltr. in Urban, Symb. Antill. 7(3):339. 1912. See Matelea constanzana J. Jiménez Alm. Ptycanthera berterii Decne. in de Candolle, Prodr. 8:606. 1844, nom. illeg. Protologue: "S. Domingo (Bertero) [...] (v.s.h. DC.).” Type: Hispaniola, ‘St. Domingue,’ Bertero s.n. (HoLoTYPE: G-DC [IDC microfiche Candolle Prodromus Her- barium, fiche & 15431]; isotype: PP. Notes: The name Ptycanthera berterii Decne. is Min as sthig O Ee is ru same as that of Gonolobus pauciflorus Spreng. (see above). Schlechter (189 rthosia acuminata Griseb. (syn. Matelea acuminata (Griseb.) Woodson). Ptycanthera mollis (Griseb.) Schltr. in Urban, Symb. Antill. 1(2):280. 1899. = Matelea mollis (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:223. 1941. See Ibatia mollis Griseb. Ptycanthera oblongata (Griseb.) Schltr. in Urban, Symb. Antill. 1(2):280. 1899. 2 Matelea oblongata (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. See Orthosia oblongata Griseb. Ptycanthera ovatifolia (Griseb.) Schltr. in Urban, Symb. Antill. 1(2):279. 1899. 2 Matelea ovatifolia (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:225. 1941. See Poicilla ovatifolia Griseb. Vincetoxicum acuminatum (Griseb.) M. Gómez, Anales Acad. Ci. Med. Habana 23:276. 1894. = Matelea acuminata (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:225. 1941. See Orthosia acuminata Griseb. Vincetoxicum oblongatum (Griseb.) M. Gómez, Anales Acad. Ci. Med. Habana 23:276. 1894. = Matelea ob- longata (Griseb.) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. See Orthosia oblongata Griseb. Vincetoxicum sintenisii (Schltr.) Britton in Britton & P. Wilson, Sci. Surv. Porto Rico & Virgin Islands 6:100. 1925. z Matelea sintenisii (Schltr) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. See Gonolobus sintenisii Schltr. Vincetoxicum stephanotrichum (Griseb.) Britton in Britton & P. Wilson, Sci. Surv. Porto Rico & Virgin Islands 6:100. 1925 See Gonolobus stephanotrichus Griseb. E £ ul Dos H Inm L 1 PE £T afa\ i7 158 Jou Vincetoxicum variifolia (Schltr.) Britton in Britton & P. Wilson, Sci. Surv. Porto Rico & Virgin Islands 6:100. 1925. = Matelea variifolia (Schltr) Woodson, Ann. Missouri Bot. Gard. 28:226. 1941. See Gonolobus variifolius Schltr. Names of uncertain application or excluded from Gonolobinae. The following names include those for which neither protologues nor types could be obtained, or for which the specimens do not belong in Gonolobinae. Names of the former category have long been placed in synonymy by authors of treatments of West Indian Gonolobinae (e.g., Schlechter 1899; Alain 1957). Cynanchum. crispiflorum Sw., Prodr. 52. 1788. = Fischeria crispiflora (Sw.) K. Schum., Nat. Pflanzenfam 4(2):230. 1895. Protologue: “India Occidentalis, Jamaica,” loc. not cited. Type: Plum., Pl. amer. 216, f. 1. 1759. Notes: According to Murphy (1986), the name Cynanchum crispiflorum Sw. (and various subsequent combina- tions in Fischeria) has been misapplied by various workers to what she recognized as Fischeria scandens DC. Murphy (1986) excluded the type of C. crispiflorum Sw. from Fischeria and recognized only the following two species in the West Indies: F. scandens DC. (Cuba and Jamaica) and F. stellata (Vell.) Fourn. (Trinidad). A current annotation in TROPICOS suggests that the type of C. crispiflora Sw.—the Plumier figure—may be Prestonia agglutinata (Jacq) Woodson (= Allotoonia agglutinata Jacq.) J.F. Morales & J.K. Williams). I am unfamiliar with this latter taxon, but a comparison of the Plumier figure to that of Allotoonia agglutinata (syn. Echites agglutinata) as illustrated in Acevedo-Rodríguez (2005), shows differences in the corolla lobes (linear-lanceolate in Acevedo-Rodríguez (2005) vs. suborbicular in the Plumier figure). Fischeria crispiflora (Sw.) Schltr. in Urban, Symb. Antill. 1(2):268. 1899, nom. illeg. Notes: A superfluous combination; see F. crispiflora (Sw.) K. Schum. Fischeria crispiflora (Sw.) K. Schum., Nat. Pflanzenfam. 4(2):230. 1895. See Cynanchum crispiflorum Sw. Gonolobus crispiflorus (Sw.) R.Br. ex Schult., Syst. Veg. 6:60. 1820. See Cynanchum crispiflorum Sw. Gonolobus ottonis C. Koch & Bouche, Ind. Sem. Hort. Berol. 13. 1855. Protologue: unknown. Type: unknown. Notes: Sometimes cited as Gonolobus ottonis Walp. (Ann. Bot. Syst. 5:502. 1859.). However, Walpers (1859) clearly cited G. ottonis C. Koch & Bouche and should not be considered author of a homonym. Walpers (1859) repeated Koch and Bouche’s citation: “Ex insula Cuba reportavit Otto. However, Schlechter (1899) thought that the specimen collected by Otto (apparently formerly at B) was from Caracas, Venezuela, not from Cuba. The location of the type is unknown. Gonolobus oxyanthus Turcz., Bull. Soc. Imp. Naturalistes Moscou 25(2):318. 1852. Protologue: "Venezuela, Funck 2," fide TROPICOS (protologue n.v.). Type: Venezuela, 1845-1852, Funk 2 (G, n.v.; P, nv.) fide TROPICOS. Notes: Fide Fontella & Schwarz (1981), a synonym of Gonolobus rostratus (Vahl) R.Br. ex Schult. (Systema Vegetabilium 6:61. 1820). TROPICOS (23 Aug 2006) cited Funk 2 (Venezuela) as the type, but a copy of the protologue could not be located. Specimens were not found on either of two visits to P. Gonolobus virescens Desv. ex Ham., Prodr. Pl. Ind. Occid. 32. 1825. Protologue: “Herb. Prof. Desvaux, Jamaica. (S. v.),” loc. not cited. Type: Jamaica,” Herb. Desvaux (HOLOTYPE: P!). Notes: Rendle (1936) considered G. virescens Desv. a species incerta. Schlechter (1899) noted that there was likely a locality mix-up between G. stapelioides and G. virescens, and that the latter was likely collected from Krings, war Al As fe LL? D | E Ba 159 £V Tobago and not Jamaica (see discussion under G. stapelioides above). The morphology of the flowers of Des- vaux s.n. (P) places it in Marsdenieae Benth., rather than Gonolobinae. Lachnostoma ovatum Turcz., Bull. Soc. Imp. Naturalistes Moscou 25(2):318. 1852. Protologue: unknown. Type: unknown. Notes: Cited as L. ovata and as a synonym for Matelea ovatifolia (Griseb.) Woodson by Alain (1957). Neither protologue, nor type could be located. ACKNOWLEDGMENTS I thank the curators and staff of the following herbaria for searching, or providing access to or loans of their collections: B, BG, BH, BKL, BM, BOLO, BR, BREM, BSC, BUF, C, CGE, COLO, CR, DUKE, E, F, FI, FLAS, FR, FTG, G, GH, GOET, H, HAC, HAJB, HBG, IA, IJ, ISC, JBSD, JE, K, L, LD, LE, LINN (Linnean and Smithean herbaria), M, MICH, MIN, MO, MSC, NCU, NEU, NSW, NY, O, OXF, P, PH, RSA, S, U, UBT, UC, UCWI, UPRRP, UPS, US, USF, TUR, WILLI, WU, Z. I also thank Pedro Acevedo-Rodríguez (US), Paul Fantz (NCSC), and an anonymous reviewer for review of the manuscript. REFERENCES ACEVEDO-RODRÍGUEZ, P. 2005. Vines and climbing plants of Puerto Rico and the Virgin Islands. Contr. U.S. Natl. Herb. ADAMS, C.D. 1972. Flowering plants of Jamaica. University of the West Indies, Mona. ALAIN [HNO]. 1957. Flora de Cuba 4. Dicotiledóneas: Melastomataceae a Plantaginaceae. Ocas. Mus. Hist. Nat. Colegio "De La Salle" 16. BoLDINGH, |. 1909. The flora of St Eustatius, Saba and St Martin. Flora of the Dutch West Indian Islands 1:1-321. BOLDINGH, |. 1914. The flora of Curacao, Aruba and Bonaire. Flora of the Dutch West Indian Islands 2:1-197. BRIDSON, G.D.R. and ER Smit. 1991. B-P-H/S: Botanico-Periodicum-Huntianum/Supplementum. Hunt Institute for Botanical Documentation, Pittsburgh. BRUMMITT, R.K. and C.E. PoweLL. 1992. Authors of Plant Names. Royal Botanical Garden, Kew. [http://www.ipni.org/ index.html]. CHEESMAN, E.E. 1947. Asclepiadaceae. In: EE Cheesman and R.O. Williams, eds. Flora of Trinidad and Tobago. Department of Agriculture, Port-of-Spain. Pp. 162-175. CorreLL, D.S. and H.B. CorreLL. 1982. Flora of the Bahama archipelago: including the Turks and Caicos islands. J. Cramer, Vaduz. D'Arcy, W.G. 1967. Annotated checklist of the dicotyledons of Tortola, Virgin Islands. Rhodora 69:385-450. D'Arcy, W.G. 1970. Jacquin names, some notes on their typification. Taxon 19:554-560 DrapaLik, D.J. 1969. A biosystematic study of the genus Matelea in the southeastern United States. Ph.D. disserta- tion, University of North Carolina, Chapel Hill. Duss, Rév. Père. 1897. Flore phanérogamique des Antilles françaises (Guadeloupe et Martinique). Ann. Inst. Co- lonial Marseille 3:1-656. FONTELLA P, J. and E.A. ScHwarz. 1981a. Estudos em Asclepiadaceae XII. Consid sobre os generos Roulinia Decne. et Rouliniella Vail. Bol. Mus. Mun. Curitiba 45:1-12. FONTELLA P, J. and E.A. Schwarz. 1981b. Estudos em Asclepiadaceae, XIII. Novos sinónimos e novas combinações. Bol. Mus. Mun. Curitiba 46:1—10. FOURNET, J. 2002. Flore illustrée des phanérogames de Guadeloupe et de Martinique. Gondwana editions. GOODING, E.G.B., A.R. Loveless, and G.R. Proctor. 1965. Flora of Barbados. HMSO, London. GRISEBACH, A.H.R. 1862. Flora of the British West Indian Islands. Reeve & Co, London. GRISEBACH, A.H.R. 1866. Catalogus plantarum cubensium. Guilielmum Engelmann, Lipsiae. HOLMGREN, PK. and N.H. HOLMGREN. 1 en EE Ge EES? Index Herbariorum. New York Botanical Garden. http://sciweb.nybg dexHerbariorum.as — - [| £ ul n.a H In LI rr £T 160 | Texas 2(1) Howaro, RA. 1952. The Society of Plant Taxonomist's plaque honoring Erick L. Ekman. Bull. Torrey Bot. Club 79:80-84. Howanb, R.A. 1986. Notes on Quiina (Quiinaceae) and /lex species (Aquifoliaceae) in Cuba. Brittonia 38:1 3-16. HowarD, R.A. 1989a. Charles Wright in Cuba, 1856-1867. Chadwick-Healy, Alexandria. Howaro, R.A. 1989b. Flora of the Lesser Antilles, Leeward and Windward Islands 6. Jamaica Plains, Massachu- setts. KRINGS, A. 2005a. A new combination in Matelea (Apocynaceae - Asclepiadoideae) for an endemic Jamaican vine. Sida 21:1515-1517. KRINGS, A. 2005b. A new species of Matelea (Apocynaceae - Asclepiadoideae) from Hispaniola. Sida 21:1519-1523. | KRINGS, A. 2005c. Notes on the Matelea bayatensis-correllii-tigrina complex (Apocynaceae - Asclepiadoideae - Gonolobinae) in the Greater Antilles and Bahamas. Sida 21:1525-1533. Krings, A. 2005d. Lectotypification and a new combination in Matelea (Apocynaceae - Asclepia loideae) for an endemic Hispaniolan vine. Sida 21:2081-2085. KaiNGs, A. 2006. Four novelties and a lectotypification in Matelea (Apocynaceae: Asclepiadoideae) from Hispan- iola. Sida 22:941-953. Kaings, A. 2007. Novelties in Gonolobus (Apocynaceae: Asclepiadoideae) from the Lesser Antilles. Syst. Bot. 32:180-194. KINGS, A. 2008. Revision of Gonolobus s.s. (Apocynaceae, Asclepiadoideae) in the West Indies. J. Bot. Res. Inst. Texas. 2:95-138. KRINGs, A. and PR. Fantz. 2006. Notes on types in Apocynaceae - Asclepiadoideae in Cuban herbaria and four lectotypifications in West Indian Gonolobinae. Sida 22:533-537. KRINGS, A. and A.C. SaviLLE. 2007. Novelties and lectotypifications in the Ibatia-Matelea complex (Apocynaceae: Asclepiadoideae) from northern South America. Syst. Bot. 32:862-871. KRINGS, A. and Q.-Y. Xiang. 2004. The Gonolobus complex (Apocynaceae - Asclepiadoideae) in the southeastern United States. Sida 21:103-116. KRINGS, A. and Q.-Y. XIANG. 2005. Taxonomy of the Gonolobus complex (Apocynaceae - Asclepiadoideae) in the southeastern US: ISSR evidence and parsimony analysis. Harvard Pap. Bot. 10:147-159. KRINGS, A., E Areces BERAZAÍN, and J.C. Lazcano LARA. 2005. New and rediscovered milkweeds from Cuba: Calotropis gigantea and Gonolobus step! trichus (Apocynaceae - Asclepiadoideae). Willdenowia 35:315-318. KRINGS, A. D. THOMAS, and Q.-Y. Sieg 2008. On the generic circumscription of Gonolobus (Apocynaceae, Asclepia- doideae): Evidence from molecules and morphology. Syst. Bot. 38:403-415. Kunze, H. 1995. Floral morphology of some Gonolobeae (Asclepiadaceae). Bot. Jahrb. Syst. 117:211-238. Laweence, G.H.M., A.F.G. Bucheim, G.S. Daniets, and H. DoLEzaL. 1968. B-P-H: Botanico Periodicum Huntianum. Hunt Botanical Library, Pittsburgh. Liebe, S. 1997. Subtribes and genera of the tribe Asclepiadeae (Apocynaceae - Asclepiadoideae) - a synopsis. Taxon 46:233-247. Lens, S. and H. Kunze. 2002. Cynanchum and the Cynanchinae (Apocynaceae - Asclepiadoideae): A molecular, anatomical and latex triterpenoid study. Org. Divers. Evol. 2:239-269, LiEDE-SCHUMANN, S., A. Rapni, D.J. Govber, and M.W. Chase. 2005. Phylogenetics of the New World subtribes of Asclepiadeae (Apocynaceae-Asclepiadoideae): Metastelmatinae, Oxypetalinae, and Gonolobinae. Syst. Bot. 30:184-195 Liocier, A.H. 1968. Novitates Antillanae IIl. Brittonia 20:148-161. Liocier, A.H. 1994. La flora de la Española 6. San Pedro de Macorís, Dominican Republic. Liocier, AH. 1995. Descriptive flora of Puerto Rico and adjacent islands 4. Río Piedras, Puerto Rico. Lig, EL, Jr, RO. Woopsury, and EH. Wabswonrn. 1976. Flora of Virgin Gorda (British Virgin Islands). United States Dept. Agric. For. Service Res. Pap, ITF-21:1-36. MiLLSPAUGH, C.F. 1902. Flora of the island of St. Croix. Field Mus., Bot. 1:441-546. D À 4 h 161 Val A) di Krinas v eg Mon oO. G. 1997. Revisión preliminar de Metalepis Griseb. (Asclepiadaceae). Pittieria 26:65-99. Moscoso, R.M. 1943. Catalogus florae Domingensis. L. & S. Printing Co., New York. Mupp, H. 1986. A revision of the genus Fischeria (Asclepiadaceae). Syst. Bot. 11:229-241. NicoLsoN, D.H. 1991. Flora of Dominica, Part 2: Dicotyledonae. Smithsonian Contr. Bot. 77:1-274, NORDENSTAM, B., R. LUNDIN, and T.A. Zanoni, 1994, Herbaria of Olof Swartz and Erik L. Ekman at Stockholm (S). FI. Greater Antilles Newsl. Proctor, G.R. 1984. Flora of the Cayman Islands. Kew Bull., Addit. Ser. 11:1-834. RANKIN RODRÍGUEZ, 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 RAPINI, A., M.W. Chase, and T.U.P. Konno. 2006. Phylogenetics of South American Asclepiadoideae (Apocynaceae), Taxon 55:119-124, Rapni, A, M.W. Chase, D.J. Govpes, and J. GrirritHs. 2003. Asclepiadeae classification: Evaluation the phylogenetic relationships of New World Asclepiadoideae (Apocynaceae). Taxon 52:33-50. REVEAL, J.L. and FR. Bannie. 1992. Matelea suberosa (L.) Shinners (Asclepiadaceae)—once again. Bartonia 57:36-38. SCHLECHTER, H 1899. Asclepiadaceae. In: |. Urban, ed. Symbolae Antillanae. Gebrüder Borntraeger, Berlin. Pp. 236-290 SPELLMAN, D. 1975. Asclepiadaceae. In: Flora of Panamá, R. Woodson et al. Ann. Missouri Bot. Gard. 62:103—156. SUNDELL, E. 1981. The New World species of Cynanchum subg. Mellichampia (Asclepiadaceae). Evol. Monogr. 5:1-62. STAFLEU, FA. and R.S. Cowan. 1976, Taxonomic literature: A selective guide to | | with dates, commentaries and types. Vol. 1: A-G. Bohn, Scheltema, and Holkema, Utrecht. STAFLEU, F.A. and R.S. Cowan. 1979. Taxonomic literature: A selective guide to botanical publications and collec- tions with dates, commentaries and types. Vol. 2: H-Le. Bohn, Scheltema, and Holkema, Utrecht and W. Junk, ds E [| Lol: Lt ] collections The Hague.v STAFLEU, FA. and R.S. Cowan. 1981. Taxonomic literature: À selective guide to botanical publications and collec- tions with dates, commentaries and types. Vol. 3: Lh-O. Bohn, Scheltema, and Holkema, Utrecht/Antwerpen and W. Junk, The Hague/Boston. STAFLEU, F.A. and R.S. Cowan. 1983. Taxonomic literature: A selective guide to botanical publications and collec- tions with dates, commentaries and types. Vol. 4: P-Sak. Bohn, Scheltema, and Holkema, Utrecht/Antwerpen and W. Junk, The Hague/Boston. STAFLEU, F.A. and R.S. Cowan. 1985. Taxonomic literature: A selective g | with dates, commentaries and types. Vol. 5: Sal-Ste. Bohn, Scheltema, and Holkema, Utrecht/Antwerpen and W. Junk, The Hague/Boston. STAFLEU, RÁ. and R.S. Cowan. 1986. Taxonomic literature: A selective guide to botanical publicati | collections with dates, commentaries and types. Vol. 6: Sti-Vuy. Bohn, Scheltema, and Holkema, Utrecht/Antwerpen and W. Junk, The Hague/Boston. STAFLEU, F.A. and R.S. Cowan. 1988, Taxonomic literature: A selective guide to botanical publications and collec- tions with dates, commentaries and types. Vol. 7: W-Z. Bohn, Scheltema, and Holkema, Utrecht/Antwerpen and W. Junk, The Hague/Boston. WOODSON, JR, RE 1941. The North American Asclepiadaceae. Ann. Missouri Bot. Gard. 28:193-244. opo eger " : | LI: E ] llections INDEX TO NAMES Asclepias maritima Jaco, Cynanchum rostratum Vahl, Cynanchum crispiflorum Sw., Cynanchum stellatum Vell., Cynanchum denticulatum Vahl, Cynanchum viridiflorum G. Mey., Cynanchum hirsutum Vahl, Fischeria cincta Griseb., Cynanchum maritimum (Jacq.) Jacq., Fischeria crispiflora (Sw.) K. Schum., Cynanchum maritimum (Jacq.) L, Fischeria crispiflora (Sw.) Schltr., 162 Fischeria multiflora Decne., Fisc eria multiflora - Fischeria scanden Ge stellata Za ) E. Fourn., Fisheria havanensis Decne., nolobus absalonensis Krings, Gonolobus bakeri Schltr., Gonolobus bayatensis Urb., Gonolobus broadwayae Schltr., Gonolobus ciliatus Sch Geer cinctus EE BEE ct PA a DIAlilenn) Drs nt À Jacaima costata (Urb.) Rendle var. goodfriendii Proctor, Jacaima parvifolia Proctor, Lachnostoma maritimum (Jacq.) G. Nicholson, Lachnostoma mollis (Griseb.) M. Gómez, Lachnostoma ovatum Turcz., Macroscepis hirsuta (Vahl) Schltr., "Macroscepis obovata" auct. non Kunth, atelea acuminata (Griseb.) Woodson, Matelea alainii Woodson, Matelea annulata Woodson ex Alain, Matelea Dayatensis de rb. ) Wee & Hook. f. ex B.D. Jacks., Gonolobus crispiflorus (Sw) R.Br. ex Schult., nolobu s denticulatus (Vahl) W.D. Stevens, Gonolobus dictyopetalus Urb. & Ekman, Gonolobus dom mingen sis Alain E Gonolobus grenadensis Schltr., Gonolobus grisebachianus Schltr., Gonolobus haitiensis P.T. Li, Gonolobus iyanolensis Krings, Gonolobus jamaicensis Rendle, E an (Jacq.) R.Br., (Jacq.) R.Br. ex Schult., Gonolobus martinicensis Decne; Gonolobus membranaceus Schltr., Gonolobus nipensis Urb., Gonolobus ottonis C. Koch & Bouche, Gonolobus oxyanthus Turcz., Gonolobus pauciflorus Spreng. 4 "Gonolobus rostratus" auct. non (Vahl) Schult., Gonolobus rostratus (Vahl) Schult "Gonolobus scandens (Aubl.) Urb., Gonolobus scandens Urb., Gonolobus sintenisii Schltr., Gonolob as Desv. ex Ham., Gonolobus stellatus Griseb., Gonolobu vindi a Griseb., Gonolobus stipitatus Alain, “Gonolobus suberosus" auct. non (L.) R.Br., Gonolobus tigrinus Griseb., Gonolobus tigrinus Griseb. var. LÊ I: Gonolobus tobagensis Urb., Gonolobus variifolius Schltr., Gonolobus virescens Desv. ex Ham., Gonolobus viridiflorus Schult., E waitukubuliensis Krings, Gonolobus youroumaynensis Krings, viam) ma candolleanum Spreng., Ibatia maritima (Jacq.) Decne., Ibatia muricata Gris Jacaima costata due j Rendle, Griseb., Matelea borinquensis Alain, Matelea constanzana J. Jiménez Alm., Matelea costata (Urb.) Morillo var. goodfriendii (Proctor) Krin ings, mallee erispinora (Ur, ) - Senes Alm., t t E.A. Schwarz, AA pv» l dl fl "m o El “1 Matelea domingensis (Alain) Krings, Matelea ekmanii (Urb.) Woo Matelea grisebachiana Scu Am Matelea hastata Alain Matelea hirsuta (Vahl) Woodson, vu linearipetala Alain, Matelea maritima (Jacq.) Woodson, Matelea mollis (Griseb. Woodson, = Ud fo S 3 = e Matelea ovatifolia (Griseb.) Woodson, Matelea pauciflora (Spreng. ) Woodson, į Krin Matelea a (Griseb.) Krings, Matelea rhamnifolia (Griseb.) GER Matelea rhynchocephala Krin Matelea rubra (H. Karst) ds & Stoffers, Matelea sintenisii chi ) Woodson, Matelea sylvicola Ala Matelea tamnifolia iio Matelea tigrina (Griseb.) Woodson, Matelea torulosa Krings, atelea variifolia (Schltr.) Woodson, Matelea viridiflora c. qu oodson, at MP Omphalophthalma s A Karst., d dee Geen Orthosia oblongata Griseb., iid acuminat Jet ) Schltr., Poicilla c Poicilla DEE e? ) Schitr., 3AF a] E F zs z | Krinas x- or, Poicilla oblongata o } Schltr., Poicilla ovatifolia G Poicilla tamnifolia i, Poicillopsis crispiflora Urb Poicillopsis ovatifolia (Griseb.) Schltr., Poicillopsis tuerckheimii Schltr., Ptycanthera berterii Decne., Ptycanthera mollis (Griseb.) Schltr., Ptycanthera oblongata (Griseb.) Schltr., Ptycanthera ovatifolia (Griseb.) Schltr., Vincetoxicum acuminatum (Griseb.) M. Gómez, Vincetoxicum oblongatum (Griseb.) M. Gómez, Vincetoxicum sintenisii (Schltr.) Britton, Vincetoxicum variifolia (Schltr.) Britton, 163 164 as "ES BOOK REVIEWS ELISEO “CHEO” Torres AND Timotuy L. Sawyer, JR. 2005. Curandero: A Life in Mexican Folk Healing. (ISBN 0-8263-3640-x, pbk.). University of New Mexico Press, 1312 Basehart Rd. SE, Albuquerque, New Mexico 87106-4363, U.S.A. (Orders: www.unmpress.com, 1-800-249-7737). $14.95, 170 pp., b/w illustrations, 5 1/2" x 8”. I first became Ge in Ge EE iw E to SH mother's stories about our foremothers' skillful use of medicinal plants. Th f SE 1 C f Zn qe E 8 L LA of South Texas aa AE Mexico. b» ee stories in Dr. T 's bool te with tl it f Southwestern Mexvican-A eee Chapter 10 “How I Was = and Beare by iud is an account o the 1 herbal wisdom of Dr. Torres's mother t! that were handed out freely to sick friends and mil as did my own ta fica: my mother I was already familiar with one of the legendary healers profiled in this book, Don Pedrito Jaramillo, a folk saint whose tombstone near Falfurrias, Texas, reads “The SCHEDE of xad ra E 1 fN » | ES All Lu Sis € ch ] le 1 : got 1 1 P H 1. de this book does much to promote al dnd. and understanding of a beloved foll dicine svstem that exists in various forms b ao See E See States. € i porates el ts of Euro[ d Arabi licine | ht f 1 14 1 1 ] "r1 1; ] H 1 1 d I A CE Writ H I! A the curanderos, allow the reader to und 1 this fi f foll licine f ithin tl lture. Dr. Torres is an expert who teaches summer courses on curanderismo at pue uin ie Nee Mexico. ie Mir would Hi useful ae for in one eer in herbal and folk medicine, as well [t medicine that incorporates spirituality and the mind with leslie, —Marissa N. Oppel, MS, Comditiaign Science Learning Laboratories, Biology Department, El Centro College, Dallas, Texas 75202, U.S.A L. KATHERINE KIRKMAN, CLAUD L. BROWN, AND DONALD J. Leororp. 2007. Native Trees of the Southeast: An Identification Guide. (ISBN 13: 978-0-88192-828-0, pbk.). Tin dud Inc., 133 S.W. Second Avenue, Suite 450, Portland, Oregon 97204-3527, U.S.A. (Orders: www.t com i com, 1-800-827-5622). $34.95, 370 pp., numerous drawings, maps, M sa 6.25" x 9. 25". Contents.—Preface; Acknowledgments: Introduction; deci of Lp VAL MM Scientific and common names; How to use the keys; Identification features of trees; Tree diversi forests; keys to trees ; Winter keys to flowering trees; [Descriptions of 44 plant families]; Some common ede and Alba trees in ilte Southeast; Glossary; Conversion tables; Bibliography; Index This book, Native Trees of the Southeast: An Identification Guide, is divided int tions by family. The authors include a detailed introduc- t hat p t d ith their specific definiti fat ] inf tion on boul to use keys and features to identify trees. The authors f line drawings helps to clearly explain the diversity of tree anatomy, ou concerning = es and branches. A NE A 1 VM 1 "m A -Ta d 1 ] i e] nf. at * Tear ra p d I in ges book to pe ety i d a good source for any amateur or P outdoorsman. Soe authors used oe com- Thev p ji kt measurements for identification to species The ee of a winter and | a great addition. The authors used easy to understand terms in each dn i uis the E ne ofatree. 1 ipi: n images Men os to be is E deeg I ue 1 P any vide: or tn. ten McNew, MS polea, Project a Botanical R h Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S.A J. Bot. Res. Inst. Texas 2(1): 164, 2008 MOLECULAR PHYLOGENETIC ANALYSIS OF THE AMERICAN STIPEAE (POACEAE) RESOLVES JARAVA SENSU LATO POLYPHYLETIC: EVIDENCE FOR A NEW GENUS, PAPPOSTIPA Konstantyn Romaschenko Paul M. Peterson and Robert J. Soreng M.G. Kholodny Department of Bot National Museum of SH History Smithsonian d Washington, DC 20013-7012, U.S.A, peterson@si.edu; QE. Oksana Futorna Institu Botany National Academy of Sciences of Ukraine 01601 Kiev, UKRAINE kromaschenkoggmail.com Nuria Garcia-Jacas Laboratory of Molecular Analysis G. Kholodny Institute of Botany Botanic Institute of Barcelona (CSIC-ICUB) TR Academy of Sciences of Ukraine Pg igdia, 01601 Kiev, E oksana_drofa@yahoo.com 08038 Barcelona, SPAIN ngarciajacas@ibb.csic.es Alfonso Susanna Botanic Institute of Georg (CSIC-ICUB) Pg. de ia, S. n. 08038 Gees SPAIN asusanna@ibb.csic.es ABSTRACT 4 molecular phyl lysis based lastid (trnK-matK, matK, trnH-psbA, and trnL-trnF regions) and nuclear ITS DNA sequences of 68 species representing 1 " n suggests an ee new E pothesis id I y history within the tribe HM Md Anatherostipa, Ptil lently f | e e o LI + subg. Pappostipa OU NU s.l. p letic) that i group to Austrostipa pl bclade referred tl “Major American Clade” (MAC). Within the MAC, New World. Achnatl is the sister group of a grade or clade of Jarava and a Nassella-Amelichloa clade. Jarava subg. P. ic and loselesalstad EA led species of J supporting gnition of th new eg Pappostipa (Speg) Romaschenko, P.M. Ron & M with 23 species; Stipa speci ] 1 as the lectotype. Within tions, and we make ne binati to 1 fer | | Pappostipa rr E o , y D LI D P 1 n £L D 1 ajaj: 1 T H a 2 e LE L d e EE E CD JI-\ D 1 1 D + 4 TD 13 D 1 ] D fc ^ D 1 ] T ^1 á AR a ta LK A DN oe Ax Lu (Phil) Romaschenko, P PE tip D ser E 11 (P ] )R 1 1 P PI I E 1 (P 13D 1 1 42 Ts fo ^ T3 1 1 T» at 1.3 a A El fn VAT 1 7 TA zx a “a Eu LA" ia LL LX a na E E LL Lu "EI (GA Dm D honl (< R henl ] i hirsutissitia (E. A. Roig) Romascheniko, Pappostipa frigida (Phil) daa apposta frigida var. ic parvispíenla (Parodi) Romaschenko, Pappostipa humilis (Cav.) Romaschenko, Pappostipa humilis var. decrescens (Kuntze) Romaschenko, Pappostipa humilis var. ruiziana rio: Romaschenko, es a (E. s NE Car posTpa min i. minuta (F.A. appostipa 11 A Dannactina (Kuntze) R Roig) R hen! Bannon sect. P following new combinations: Pappostipa speciosa (Trin. & Rupr.) Bonnes dese. Pappostipa speciosa var. lie (F.A. . Roig) Romaschenko, Pappostipa speciosa var. parva (F.A. Roig) H E a E pl hen] Pappostipa chry sophy lla f modica (Ey A Roig) Romaschenko, Pappostipa speciosa var. atuelensis (F.A Romaschenko, Pappostipa ia var, a ias m oie Lip ee var. nous bid A. Roig) Sa aschenko, Papp f. horrida (F.A. R PE Pappostipa speciosa var. are (Torres) Romaschenko, Pappostipa inata (Phil 1) Romaschenko, P sagiuata f. rigida (F.A. Roig) Romaschenko, Pappostipa vaginata f. inmersa (F.A. Roig) Romaschenlio, Pippostipa TEE SS laevis (F.A. Roig) ipa vaginata f. contracta (F.A. o raed Pappostipa Reg var. dea i A. Roig) Rom- ics d) Papposti aschenko, Pappostipa vaginata var. dilatata (F.A g PE , Pappostipa hieronymusii (Pilg.) Romaschenko, Papp pa patagonica (Speg.) R henl S Papnastipu maeviae (F.A. Roig) Romaschenko, [ Rat Rac Inct Tava« 2(1): TEL. 192. 2008 166 J i £ ul n.a . Im LJ dh dis ote £T. 2(1) m E (F.A. Roig) R 1 1 11 tii (F.A. Roig) R 1 1 (F. "i Roig) ao Pappostipa ruizlealii (E A Roig) Romain: Pappostipa p (EA. Roig) Romes tiko Pappostipa major (Speg.) Romaschenko, Pappostipa vatroensis (F.A. Roig) Romaschenko, and Pappostipa barrancaensis (F.A. Roig) Romaschenko. RESUMEN e D iu Un análisis filogenético basado en los d lecul ] 1 i lastidiales (trnK-matK, matK, trnH-psbA y trnL- Mina i T. secuencias nucleares del ITS de 68 especies representando 11 géneros, a i e de los EDU nidi y ini ind con eo como un dias adyacente) se presenta como un linaje hace Jarava s.l. polifilético) que (lo que a su vez, un ee hermano para Austrostipa pe un subclado al pe nos como "dado a americano al uw Dentro del MAC el maneta del N ta como un grup plej p de Jarava s.s. y Nassella con el ind i E incloido. E hecho de que Jarava subg. Pappostit filetico y no se relaci el resto del Jarava s.l. apoya epar ) Romaschenko, P.M. Pet & S g g 23 Se di speciosa seleccionado como el ege Dentro delP tipa las d i f idas | do sobre los datos moleculares y mor fológicos. Para la iden del abi subsp. Pappostipa un gé prop onen | j combrmaciones taxonómicas e apropiadas: Pappostipa secc. ` g 8 Romaschenko, Pappostipa (Parodi) Bee ko, P ti (Speg.) Ro masdherko Pappostipa ibarii (Phil) Romaschenko, Pappostipa ibarii f. nallescems (Parodi) beer Pappostipa ibarii var. anomala aes Romaschenko, T3 a ameghinoi Pappostipa ameghinoi (Speg.) Romaschenko, Pappostipa ameghinoi var. digona (Parodi) Romaschenko, var. precordillerana (F.A. Roig) Romaschenko, Pappostipa chubutensis (Speg) Romaschenko, Papposti inä a var. hirsutissima (F.A. GR oeren Pappostipa ios n Romaschenko, Pappostipa ipa var. ee (Parodi) k Romaschen 0, Pappostip ] umilis var. , Pappostipa humilis var. ruiziana mien ia Pappostipa chrysophylla (E, Desv) Romaschenko, Pappostipa chrysophylla f. anhwlla minuta (F.A. Roig) Romaschenko, Pappostipa eon Var. crispula (Kuntze) Romaschenko, and Pappostipa dn sophylla £ É. nda € A. Roig) Romaschenko Dentrode e Pappostipa s secc. ciosa var. RE EE nid Roig) Remaschenko, Pappostipa speciosa var. _atuelensis GA Roig) Romaschenho, Pappostipa ciosa var. parva (F.A. R pec var. ciliata (F.A. Roig) Romaschenko, Pappostipa: eo " horrida (F.A. Roig) dla, Pappostipa speciosa f. Es dita (F.A. Roig) Romaschenko, Pappostipa speciosa var. media (Torres) Romaschenko, Pappostipa vag il) Romaschenko, Pappostipa vaginata f. Heins (EA, na Romaschenko, EE cda f. inmersa iiid "d e l Pappostipa vaginata f laevis (F A R i Pappostipa 3 cont ta (F AR ) | , Pa ppostipa aos Var. a argyroidea (FAR 1 ] , Pappostipa a vaginata dilatata js Roig) R ] | P (F A oig g) [eli H RK A Lu "rh e a za as D fc ^ Ta 1 1 Ta as Romaschenko, ga Lu A o Id T E E KE La ke E [e] Ea A 1 1 RE {Bik DAL.) DB 1 1 I LI quetii (F.A Paisai D 1 ] me EE ain erie E ?" Roig) Romaschenko, Tamos caiz leali E. A. Roig) Romaschenko, Pappostipa EE EE (E op J omaschenko, Pappostip t Y Pappostipa ER (F.A. Roig) Bomascherko.- The tribe Stipeae Dumort. are enparan, cool-season (C; 3) E that ae] occupy somewhat moist to predoptinan dry open grass} teppe pt Antarctica. Worldwide the Stipeae consist of a minimum of 21 Bened and 524—604 species, denen: on how Ásian taxa are recognized (Soreng & Davis 2005). In the New World there are 279 species of Stipeae (Soreng et al. 2008); of these, 222 (79.696) are found in South America. Thirteen genera are indigenous, and nine are endemic to the New World. The Stipeae s.s. (excluding elements of Phaenospermateae Renvoize & Clayton, and subtribes Ampelodesminae Connert and Duthieinae Pilg. ex Potztal) are characterized by having single- flowered spikelets with terminally-awned lemmas where the awn is the result of the fusion between the central and two lateral vascular traces, simple starch grains, florets with two or three linear lodicules, and small-sized chromosomes with a base number of x = 10-12. The Stipeae are placed in subfamily Pooideae (GPWG 2001). Within the Pooideae they are an 'early diverging lineage that arose after the separations of the Brachyelytreae Ohwi, and Lygeeae J. Presl plus Nardeae W.DJ. Koch (Davis & Soreng 2007). Historically, delimitation of genera within the American Stipeae was based on a broad concept of the genus Stipa L. Hitchcock (1935, 1951) accepted three genera in North America: Oryzopsis Michx., Piptocha- etium J. Presl, and Stipa. In South American Stipeae, Spegazzini (1901) recognized only Aciachne Benth., n 1 ' sal ni yl geny CA e Sti J , Pappostipa 167 i Oryzopsis, and Stipa. In the second part of the 20th century, Stipeae taxonomy was characterized by the consecutive resurrection of previously described genera or emendation of taxa considered by Spegazzini (1901, 1925) as subgenera of Stipa. Nassella (Trin.) E. Desv. was restored to generic status by Parodi (1947) and significantly expanded, and thus became the most species rich genus among American Stipeae (Bark- worth 1990; Barkworth & Torres 2001). In considering the distribution of morphological features among stipoid grasses and lemma epidermal features (Thomasson 1978, 1980, 1981, 1982), Barkworth and Everett (1987) suggested that Stipa includes only Eurasian species: Hence the name Stipa should not be used for any American species. In subsequent major rearrangements of the American Stipeae, five species were transferred to Hesperostipa (M.K. Elias) Barkworth. Thirty-seven other American species formerly placed in Stipa or Oryzopsis were shown to share some morphological features with the Eurasian representatives of Achnatherum P. Beauv.; they were transferred to the latter genus (Barkworth 1993). Stipa subg. Anatherostipa Hack. ex Kuntze (Kuntze 1898; Spegazzini 1901, 1925), including the Obtusae group of Parodi (1946), was recently established at the generic rank by Peñailillo (1996), currently with 11 species (Soreng et al. 2003). Nassella now includes 115 and Piptochaetium 35 species, respectively (Soreng et al. 2003). The genus Jarava Ruiz et Pav. was resurrected by Rojas (1997), in the sense of Stipa subg. Jarava (Ruiz et Pav.) Trin. & Rupr. (Caro & Sánchez 1973), and significantly expanded by Peñailillo (2002, 2003) to 55 species by inclusion of Stipa subg. Pappostipa Speg. and S. subg. Ptilostipa. Speg. The expansion of Jarava effectively completed the removal of all remaining accepted eae Oe New wens pede im the Reus Stipa, as anticipated by Barkworth and Everett (1987). However, Sti that such a broadly circumscribed Jarava was probably Hopi lert, and Arriaga p Barkworth (2006) began to revise it by transferring five Jarava s.l. species to a new genus, Amelichloa Arriaga & Barkworth. Parodi (1960) recognized two main morphological leaf types in Spegazzini’s Stipa subg. Pappostipa, represented by Stipa speciosa and S. humilis (Arriaga 1983). The first type (leaf anatomical “type I” of Arriaga 1983) encompassed species with stiff, thick, and usually pungent leaves. The second type (“type II” of Ar- riaga 1983) covere d species with comparatively thin, slender, and non-pungent leaves. Later, Parodi (1960), followed by Roig (1964) and Nicora and Rúgolo de Agrasar (1978), noted that species with the former type had brownish-red basal sheaths and species with the latter type had pale basal sheaths. There have been several attempts to infer phylogenetic relationships among stipoid genera based on morphological evidence (Tzvelev 1977; Barkworth & Everett 1987; Thomasson 1978, 1985) and preliminary molecular analyses (Jacobs et al. 2000, 2007; Cialdella et al. 2007). The origins of Hesperostipa, Jarava s.s., Piptochaetium, and Stipa s.s. have been reported as monophyletic (Jacobs et al. 2000, 2007; Cialdella et al. 2007), but the overall phylogeny of the American Stipeae remains poorly bin E and EOD) Our preliminary study of the Stipeae phylogeny based on ITS sequence data ko et al. 2007) revealed a deep separation between a Euro-Mediterranean clade, EE Sia s.s., and a clade including genera and species of East Asian, Australian, abs nM v pn In addition, Jarava (sensu Peñailillo 2003) was resolved as polyphyletic, with a clade y placed in Stipa subg. Pappostippa, clearly independently derived from the core species of aram S.S. The main objectives of this paper are to present a phylogenetic hypothesis for the American Stipeae and to evaluate the monophyly of Jarava s.l. based on an analyses of plastid (matK, EE trnH-psbA and trnL-trnF) and nuclear ribosomal ITS DNA sequences. We also discuss phological and anatomical characters that support our phylogenetic M RAM METHODS Ta i The majo ft | led in this study were collected by PMP and RJS, and il j Al ity Wi k are posed at t the Smithsonian institution, United States National Herbarium (US). Eighty-one samples representing 66 species and three infraspecific taxa, with duplicates for nine taxa, were included in our analysis (Table 1). ne Aa igata! of the 11 Stipeae genera reported as indigenous for the Americas (Soreng et al. 2003), p gregate genus, Amelichloa. 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Reeder, Ortachne Nees ex Steud.), the North American genus Oryzopsis s.s., and North American species placed in Piptatherum (all other species are Old World), are not included in the present analysis because our preliminary data indicate that they have no effect on the disposition of taxa of concern here. One Asian species of Ptilagrostis Griseb. and five Australian species of Austrostipa : id L. Rone & J. Everett were included to show evidence of the EE eg Subsequent papers will include a much | Sher RE? and Nardus stricta were chosen as outgroups based on the paculie of eo else of phylogenetic relationships in Pooideae (Hilu et al. 1999; Soreng & Davis 2000; GPWG 2001; Davis & Soreng 2007; Soreng et al. 2007). These outgroups were used to root trees in the ITS analyses of the Stipeae (Jacobs et al. 2000; Romaschenko et al. 2007). DNA extraction, lifi ing.—Leaf tissue was homogenized using Qiagen TissueLyser, and DNA was isolated using a BioSprint 96 DNA Plant Kit (Qiagen, Valencia, California, USA, 2005). For some specimens, the CTAB method (Doyle & Doyle 1987) was used. PCR amplifications were performed in MJ Research or PE 9700 thermal cyclers. Genomic DNA was combined with 1x reaction buffer (200 mM Tris-HCl, 500 mM NH) (Bioline Biolase Taunton, Madison, USA) without Mg++, 2 mM MgCl,, 200 mM dNTP's, 1.5pl of Taq polymerase (Bioline Biolase Taunton, Madison, USA), 40 pmol/ul each of forward and reverse primers. The entire nuclear ribosomal ITS regio ified using primers ITS1, ITS2, ITS3, ITS4, ITS5 (White et al. 1990) and ITS5A (Stanford et al. 2000) with the following thermal cycler settings: initial denaturation step of 2 min at 94°C, followed 35 cycles at 94°C for 30 s, 50-55ºC for 30 s, 72°C for 1 min 30 s, and a final extension of 10 min at 72?C. Four chloroplast DNA regions were sequenced: trnK-matK, matK, trnH°V°-psbA, and irnL5'-trnF. The 5'-end of trnK-matK and 5'- matK were amplified separately. The 5'trnK-matK readily amplified using the forward primer trnK3914F (Johnson & Soltis 1995) and newly designed reverse primer trnK6605R (5'-CGAATGATGATACATAGTGC-3) positioned close to the end of the 5'-trnK-matK intron. The reverse complement (i.e. forward version) of the latter primer (trnK660SF; 5'-GCACTATGTATCATC ATICG-3) and the reverse complement of a slightly modified matK1412F primer (Johnson & Soltis 1995; i.e., matK14125R 3- CTGATACATAAGAGTTRTAT-3) were used to amplify hos poe: of Me E end of matK. The DCH ze lified with primers trnHSUS (Tate & Simpso d psbA (Sang etal. 1997). The trnL- trnF Econ (which included a portion of 3'-trnL intron, the 3- trnL exon, and the trnL-trnF intergenic spacer) was amplified using primers 5'trnLU^^(f) and trnFS44(c) [Taberlet et al. 1991]. The amplification parameters for all chloroplast regions were: 94°C, 2 min; 35 cycles of 94°C for 45 s, 58-62°C for 45 s, 72°C for 1 min 30 s; 72°C for 10 min. PCR products were cleaned with ExoSAP-IT (USB, Cleveland, Ohio, USA). DNA sequencing was performed with BigDye Terminator Cycle Sequencing v.3.1 (PE Applied Biosystems, Foster City, CA, USA) according to the following parameters: 80°C, 5 min; 25 or 30 cycles of 95°C for 10 s, 50°C for 5 s and 60°C for 4 min. Sequenced products were analyzed on an ABI PRISM 3730 DNA Analyzer 7900HT (ABD. Sequences were aligned manually using BioEdit v.7.0.5.3 (Hall 1999) and then by eye. All sequences were submitted to GenBank (http://ww bi.nlm.nih.gov); voucher information, and GenBank accession numbers are provided in Table 1. Even though some indels, especially in chloroplast regions, seemed to be very specific for phylogenetic groups, they were not re-coded for this analysis and were excluded along with ambiguously aligned regions. The following gene regions were excluded: trnK-matK with four regions of 2-25 bp in length; trnH-psbA with 10 regions of 1-4 (22) bp in length; trnL-trnF with 31 regions from 1-6 (20) bp; and ITS with 15 regions 1-4 (-22) bp. No regions were excluded from matK. All gaps were treated as missing data. Parsimony analysis was performed using PAUP v. 4.0b10 (Swofford 2000) and PAUPRat v.1b (Sikes & Lewis 2001), which implements the Parsimony Ratchet of Nixon (1999). Parsimony searches were carried out for individual E combinen P and combined plastid and nuclear regions. In searching for the optimal tree, the | d and tree bisection-reconnection (TBR) E > PAR L I A ai ; nL yl g y £A H fat J É Pappostipa 173 was chosen as a branch swapping algorithm. Character states specified as unordered and unweighted. PAUPRat searches were set for generating 1001 most parsimonious trees. Because PAUPRat finds the most parsimonious tree in each iteration, a set of 1001 iterations is recommended by program developers as suf- ficient to derive a consensus tree. All of the most parsi in our analyses were of the same length. These data sets were used to yield the oa consensus trees. Bayesian posterior probabilities were ci Bu Mae ds v.3.01 (Huelsenbeck & Ronquist 2001; Ronquist et al. 2005) and the best ilabl tit was selected using MrModeltest 1.1b (Nylander 2002). The symmetrical model (SYM+I+G; Zharkikh 1994) with gamma-distributed rate variation across sites was selected by hierarchical likelihood ratio tests (hLRT) and Akaike information criterion. Bayesian analysis was initiated with random starti 1 was initially run for 1 x 10° generations with sampling frequency of the chains set to every 100% iteration. If this number of generations was not sufficient for the Markov chains to reach stasis, additional generations were invoked. The standard deviation value of the split frequencies below 0.01 indicates that Markov chains reached their stasis (Ronquist et al. 2005). The number of generations needed to complete the analysis varied from 1 x 10º for individual DNA regions to 4 x 10º in analysis of the combined data. Branches with posterior probabilities (PP) >0.95 were considered statistically significant. For additional information about the stability of phylogenetic groups the branch frequency or parsimony frequency (BF) value was used. The BF value indicates how often a branch appeared in the analysis, i.e., when this value was 100% it means the node appeared unchanged in all iterations (i.e., strict consensus). Bootstrap has a long history of being used as a standard measure when estimating phylogeny and we have included bootstrap analysis. Felsenstein (1985) states, “Bootstrapping provides us with a confidence interval within which is contained not the true phylogeny, but the phylogeny that would be estimated on repeated sampling of many characters from the underlying pool of characters.” We agree with Felsenstein that bootstrap values are measures of repeatability rather than measures of accuracy (see Soltis & Soltis 2003). Regardless of interpretation, bootstrap values are affected by the number of characters supporting the clade of interest and the data set as a whole (Soltis & Soltis 2003). For a dataset with no conflict among characters, at least three characters are needed to provide 95% support (Felsenstein 1985). With real data a greater number of characters may be needed to achieve 95% support (Soltis & Soltis 2003). Thus bootstrap tends to be overly conservative, often underestimating a well-supported clade (Sanderson & Wojciechowski 2000). Bootstrap (BS) values were calculated using PAUP v. 4.0b10 (Swofford 2000) from 10 x 106 fast- heuristic bootstrap replicates with random addition sequence. Sequence divergence was calculated based on the uncorrected “p” distance measure included in Neighbor-joining search settings. + Scanning electron microscopy.—Lemma ultra st was studied using dry mature seeds sampled from herbarium of the following species used in the phylogenetic analysis: Jarava chrysophylla, Jarava speciosa, Jarava nicorae, Jarava pseudoichu, Jarava plumosula, and Amelichloa caudata. Wax from the seed surface was removed by soaking the seeds in xylol solution for six to eight hours. After soaking, seeds were mounted on scanning electron microscopy sample tables. Mounted samples were covered with gold in the vacuum spray gun (JII-4X, Japan). The ultra structure of the lemma was studied using scanning electron microscope Jeol (J5M35C, Japan) at varying magnifications. Light microscopy.—Cross-sections of leaf blades were prepared with the use of a rotary microtome. Intact leaf blades of Jarava barrancaensis, Jarava speciosa var. major, and Jarava chrysophylla from herbarium speci- mens were rehydrated for 10-12 min in distilled water at 100 °C. For studies of the sclerenchyma tissue, the cross-sections were stained with safranin. RESULTS Amplification of the trnK intron was significantly improved when the 5'-end of the intron and the maturase K gene were amplified separately. This enabled us to optimize our protocol for routine amplification of all chloroplast regions, even when old herbarium material was used. With this approach, PCR reactions were 174 | of t! tanical Insti Texas 2( up to 6096 more successful than when the traditional intron primers were used (i.e,, trnK3914F-2R). Conse- quently, the trnK! intron had the highest rate of amplification (92.6%) compared to the non-coding regions trnH-psbA (91.496) and trnL-trnF (8996), which are traditionally considered to be the most easily amplified regions of the chloroplast genome (e.g., Kress et al. 2005). The matK gene had a lower rate of amplification (85.296); lack of matK sequence data for a subset of taxa probably affected branch support in the combined chloroplast analysis. For the most of the silica preserved samples or recently collected herbarium specimens the entire ITS region was easily amplified (10096). A few specimens that lack several chloroplast regions were not included in the chloroplast analysis but we did include them in the combined analysis based on ITS and partial plastid sequences. A small subset of samples with two or more missing chloroplast regions was included in the combined analysis (see Table 1 for details). The plastid data set is missing 11.4% and the combined analysis is missing 996, mostly due to unsequenced plastid regions for some samples. There is no missing ITS data. Analyses of DNA sequences.—The general characteristics of the DNA regions used to infer phylogeny in our data set are detailed in Table 2. The greatest number of potentially parsimony informative characters per region was provided by the nuclear DNA ITS region. The topologies of the trees produced by parsimony and Bayesian analyses for each of the four plastid regions sequenced were not inconsistent with that produced by the combined plastid data. Since the sepa- rate pane datasets d comparatively low numbers ei Ge e cds EE and low levels of nc ombined (Fig 1). The phylog tr nH-psbA and trnL-trnF EE were stronger than other gene regions and were eclighily better oa partly due to their higher Ee of EE SE ds 2) Noc derived from the trnH-psbA and trnL-trnF regions were found in this amd se The only region that ud sae substitutions responsible for separation of the Pappostipa clade of Jarava s.l. was the trnL-trnF region (data not shown). The ITS tree was more than two times longer than the plastid tree and had lower CI and RI values (Table 2). There was relatively minor topological incongruence among clades in the trees generated from the combined plastid DNA-ITS data set (Fig. 3) and the ITS data (Fig. 2). The monophyly of the Stipeae is well supported (BF=100, PP=0.64-1.00, BS=96-100) in all three cla- dograms (Figs. 1-3). Several major clades were consistently resolved in all three analyses (Fig. 1-3). A clade including Achnatherum, Amelichloa, Austrostipa, Jarava s.l., and Nassella [AAAJN] was detected in all three analyses (Figs. 1-3). Within the AAAJN clade a primary deep split between the Pappostipa clade and the clade of Austrostipa plus Achnatherum, Jarava (excluding the Pappostipa clade), Nassella and Amelichloa was maintained in all three analyses (Figs. 1-3). Austrostipa (from Australia) was resolved in all three analyses as sister to a clade of Achnatherum, Jarava (excluding the Pappostipa clade), Nassella, and Amelichloa; this relationship received strong support only in the nuclear and combined analyses. The latter clade is denoted in Figs. 1-3 as the Major American Clade or MAC. Within-MAC, Achantherum was always monophyletic and the sister group of the rest of the clade. A clade of Nassella with Amelichloa nested within it was detected in all three analyses (Figs. 1-3); this clade and always excluded EE of pa Table 2 summar how the highest rate of parsimony inf tive characters ingroup-outgroup and ingroup Séquence Gees for nuclear rDNA ITS region. Analysis of the combined plastid-ITS data set (Fig. 3) showed deep splits among the American stipoids, also detected by the separate analyses, and separation of the Pappostipa clade. The Stipeae genera included in this analysis were resolved as monophyl- etic groups with three exceptions. Ptilagrostis resolved in a polytomy in the plastid analysis, but the species were united in the ITS and combined analyses. Jarava s.l. was divided into a Pappostipa clade and a Jarava s.s. (Jarava excluding the Pappostipa clade) clade (Fig. 3), or grade (Figs. 1&2). Jarava s.s. was resolved as a paraphyletic grade to Nassella (encompassing Amelichloa) in the independent plastid and ITS analyses, but the species were united in the combined analysis. Additionally, as detected in the separate plastid and ITS D L J FEN DL, d £A : Cr: ] d Pappostipa 175 i 3 TABLE 2. SI immary of trnK-matK, matK, trnH-psbA, trni -trnF, H D E I a IS LES ADIA BL TB IA A USANA E Sa Al A trnk- matk trnH- trnl- Combined ITS Combined matk psbA trnF plastid plastid & data nuclear data parle = Ech (range) 571-556 555 585-567 612-541 2315-2231 592-584 . 2904-2271 Num 75 69 75 81 Nu imber re caracters 588 555 663 686 2537/7 640 3177 Number ofin characters 24 22 23 33 97 137 222 Ingroup- BEE divergence (%) 3.62 422 52 7.35 - 14.34 Ingroup divergence (%) 0.79 0.78 0.87 0.84 — 4.3] - Tree length 33 3] 37 56 155 380 519 Consistency index (CI) 0.85 077 0.73 0.68 0.72 0.50 0.55 Retention index (RI) 0.98 0.95 0.96 0.93 0.95 0.82 0.87 Homoplasy index (HI) 0.15 0.22 027 0.32 0.27 0.50 0.44 Rescaled consistency index (RC) 0.83 0.74 070 0.63 0.68 0.40 0.48 analyses, Nassella was only resolved as monophyletic if Amelichloa is subsumed within it. Goodness-of-fit statistics i.e., consistency index, retention index, and rescaled consistency index, for the combined plastid DNA-nrDNA set was higher than for nrDNA set alone. Early diverging clades at the base of the Stipeae included the genera Ptilagrostis (only as a polytomy in Fig. D, Anatherostipa, and Piptochaetium. All analyses su pport d these genera as independent lineages at the 1 2 WS Al Al bs d eu olas current level of taxon sampling (Figs. 1-3) ción of the datasets. Analysis of the plastid data set yielded a consensus Rs with these VEHI and Hesperos- tipa in a polytomy with a large clade including Jarava s.l., Austrostipa, Achnatherum, Nassella, and Amelichloa [AAAJN]. Analysis of the ITS data set yielded a stepwise grade leading to Hesperostipa as sister group to AAAJN. The analysis of the combined plastid-nuclear data set resulted in a fully resolved and a moderately well supported resolved basal clade of ur E as sister group to Anatherostipa and Piptochaetium (Fig. 3; BF=71, PP=0.93), and Hesperostipa ter group to AAAJN (Fig. 3; BF=100, PP=1.00, BS=100). Henceforth, for ease of discussion, we Exil the set of Anatherostipa, Piptochaetium and Ptilagrostis the "basal lineages" (BL in EIS 1-3). The position of these the basal lineages with respect to the core of the tree needs further examinati n, particularly including Old World genera, and is not the focus of the present study. Both of the Piptochaetium groups (Cialdella et al. 2007), Piptochaetium sect. Podopogon (Raf.) Parodi (P. ruprechtianum) and Piptochaetium sect. Piptochaetium (P. panicoides) are represented in the analysis. In all topologies, the Piptochaetium clade had high statistical support (BF=100, PP=0.95-1.00, BS=59-96; Figs. 1-3). Representatives of the two extant geographical lineages of Ptilagrostis chosen for analysis were P. mon- gholica from Asia and P. porteri from the Americas. These species formed a consistent and well supported clade (BF=100, PP=1.00, BS=95) on the ITS tree (Fig. 2) and the combined tree (Fig. 3; BF=100, PP-1.00, BS=97). A phylogenetic relationship between these two species was resolved as a polytomy in the consensus tree from the plastid analysis (Fig. 1). There are two clades adjacent to the basal lineages. One comprises the representatives of the genus Hesperostipa in a clade in all three trees (BF=100, PP=0.99-1.00, BS=99-100; Figs. 1-3). A second clade contains AAAJN members showing deep split (polyphyly) in the genus Jarava: Stipa subg. Pappostipa (Pap- postipa clade) of Spegazzini (1901) elements of Jarava are separated from Jarava s.s. by sequentially diverging clades of Austrostipa and Achnatherum (BF=99-100; PP=0.63, «0.50, 0.99; BS=99, «50, 99; respectively in Figs. 1-3). In combined analyses as well lear DNA ITS sequence analysis, the Pappostipa clade was well supported (BF=100, PP=0.95-1. ER BS- Se 99; Figs. 2 & 3). 4 was sister group to MAC. These clades had high statistical support (BF=100; PP=0.68-1.00; BS=90, «50, 80; respectively in Figs. 1-3) and their relationship as sister clades was consistent. In all analyses, the Ametralian 176 | tani it Texas 2( | SE Anatherostipa hans-meyeri ee atherostipa bomanii 1.00% Anatherostipa rigidiseta p jos e p i mongholi 99 pms Hesperostipa neomexicana 0.99 Lr lesperostipa spartea Hesperostipa comata 98 > Jarava barrancaensis = | 0.64 MEC — —— ! arava speciosa var. major ^ easi Ed e : 1 'a humilis trava humilis * arava humilis rava ibar. arava chrysophylla a sophylla a speciosa var speciosa arava A Pappostipa ; ip ustrostipa pycnostachya ustrostipa scabra pn falcata dustrostipa tenuifolia Achna. m aridum chnatherum occident therum occidentale subsp. TEDS GENS media rava polycla Gei rava leptostachya ' o seudoich S arava Cru 2 3 arava scabrifo Aa arava castellanosii 3 rava pha 1 p assella clarazii fl assella dasycarpa SE prachychactoides sella n sell inconspicua assella D n. | asse pua ussella zacharia Fic. Sé Canna " = D ` L [| [| E is £ Li [| A MAA — mat, Unit psbh, and mtm region Branches in bold are [ tin the strict tree, i.e. branch frequency (BF) equal to 100. , H (BS) | " | | | | | f dt B J i | t i probability (PP) values; DIE L pas BAR han? A H Fa PP | DÉI -Dannnctina rara mE 2-3 | ! SERGE HN In the ITS and combined trees (Figs. 2 & 3) two major clades are resolved (with low support indices) within the Pappostipa clade: "P" that includes Jarava barrancaensis, J. hieronymusii, J. nicorae, J. patagonica, J. speciosa var. major, J. speciosa var. speciosa, and J. vaginata (BF=53, PP=<0.50-0.52); and “C” that includes Jarava atacamensis, J. chrysophylla, J. frigida, J. humilis, and J. ibarii (BF=97-100, PP=<0.50-0.90). Within all three trees (Figs. 1-3), J. speciosa var. speciosa and J. speciosa var. major never share a most-recent common D kh I + al Di, EA H e J E d d f V e Pappostipa 177 f = pecan erectum 92 p= Piptochaetium brachyspermum 0.95 [T Eptochactium panicoides y poeta SEN stipa hans-meyer ata Jarava speciosa var. speciosa ata rava - maj 1.00 69 1.00 See d var. SE? l 0.95 95 T Ja a a tacamensis? arava rigida 0.98 0.99 a E Jee den hyll arava chrysophylla Jarava chrysophylla Jarava c Pappostipa t 3 2 A 1.00 Jar 0.97 Ee Ste Wee falcata 92 Achnatherum occi lhe tale 1.00 Achnatherum GE subsp. pubescens 0.70 Achnatherum parishii 0.88 peius erum rikan rdso onii rum : Jarava ichu MAC 100 1.00 1,87 Jarava ichu ? Jarava plum See? 65 a da Sa £ d $ GRE SEE ck S Ri a Jarava s.s. Masela, Gegen EH nardoides He esia 2 98 Ge la flic Ted assella filicu Js 1.00598 pus Nassella sanluisensis l 0.83 Nassella tenuissima 20.52 Nassella nidulans [| " £4nn1l MN " H H A L J H H al lakta É | MAA Fic. 2. Majority I ITS region. B bold t in the strict tree, i.e., branch f y (BF) equal t 10. Numbers above branches correspond to f T d 3 Clisdsa DF—D ti Jaa f— Dannar dir enga A LE ; 3 DW Dannactinag cart Pannactina GRE HOM d Li FF F ef H +] ] 43 L ]$ 24 pd : +1 7 ancestor. There is a t of J. speciosa var. speciosa in and nuclear trees (Figs. 1 & 2), but our placa data Furtenits provide little information on this discrepancy since there are only one or two apomophies found in the sequences of the trnL-F region. In the three consensus trees (Figs. 1-3) MAC includes: a monophyletic Achnatherum (BF=100; PP=0.70-1.00; BS=84, «50, 76, respectively); a grade of four lineages (Fig. 1), two clades (Fig. 2), or a single clade (Fig. 3) for Jarava; and a single monophyletic clade of Nassella (including Amelichloa) (BF=61-100; PP 178 Brachyelytrum erectum Nardus See ni TI Ptilagrostis por 1.00 Ptilagrostis a SE lica T3 herostipa bomanii Pi 0.99 1.00 Jarava speciosa Y var. speciosa Jarava vaginata? Jarava vaginata Jarav _ Pappostipa 2m rysophylla | Jarava cmt 2 chryso o ee , SU 99 Austrostipa SE TOO bee Austrostipa sca fa Su. falcata ^ Tools Austrostipa pue "7 1.008 Austrostipa tenuifolia 99 Achnath erum occidentale 00 Achnatherum m occidental subsp. pubescens yy die anim x T Achnatherum parishii 0.86 Achnatherum G IDE ncoi 1.00 rava ichu? arava lek tostachya! Jarava E Id Jarava pseudoi Pn e Jarava s.s. Jar Jar rava a polyelada Jarava Nase la depauperata 56 assella SEET Nas sella ru ae dee caes, Nassella Brach ychaetoides L Nassella p Nassella punio vphylla Nassella nardoides Arieh Erloa Are oa Ee 1.00 Amelichloa cla ndestina LAARA | + Fic. 3. Majority DNA trnK-matK, matK, trnH-psbA, and trnl-i prt e S S A pig Fr T f—Dammnetfima cart fh H F Dm Dnnnneftinam sect ct. Pappostipa. Lei dada! a 2 J EEN «0.50, 0.93, 1.00; Figs 2 & 3). Jarava ichu, the type species for the genus, resolved in a clade with J. castel- lanosii, J. leptostachya, J. pseudoichu, and J. scabrifolia in Fig. 1 (BF=100, PP=0.77, BS=87). Three Jarava species not in the plastid analysis, J. hystercina, J. juncoides, and J. plumosa, join the clade with the above set of species in Fig. 2 (BF=100, PP=1.00) and Fig. 3 (BF=100, PP=1.00, BS=58). In the combined analysis Jarava media, J. plumulosa, and J. polyclada are united as a sister clade to the above set (BF=100, PP= 0.58; Fig. 3). The latter three species formed a clade in the ITS analysis (BF=100, PP=0.64) that was sister group to Nassella includ- n | E] P | BL... EA H Es J Pannactina 179 y P f VI LI - d | e ing Amelichloa, but were scattered elements of the grade of Jarava lineages leading to Nassella (including Amelichloa) ds in i e O t Nassella i hyletic (BF=61-100, PP=<0.50-1.00) only if Amelichloa is included. This conclusion is most ge supported in the ITS and combined analyses (BF=100, PP 0.93- I. Q0). In out sensu LDere are two major clades within Nassella. One clade includes N. brachy- chaetoides, N. brachyphylla, N. ES N. dasycarpa, N. depauperata, N. meyeniana, N. pubiflora, N. nardoises, and N. rupestris (with N. inconspicua in Figs. 2 & 3; BF=100, PP= 0.93-0.99, BS=<50-56). The second clade includes Amelichloa species as sister group to the set of N. clarazii, N. filiculmis, N. manicata, N. neesiana, N. nidulans, N. pfisteri, N. éd de N MO N. sellowiana, N. tenuissima, and N. trichotoma (Figs. 2 & 3; BF-100, PP=0.53-0.66, BS=62-10 meli a caudata! as sister group t tl of A caudata?, N. clarazii, N. filiculmis, N. neesiana, N. zer N. trichotoma, and A. clandestina (Fig. T BF=100, PP=<0.50). Amelichloa is supported as monophyletic in the ITS and combined analyses (Figs. 2 € 3; BF=100, PP=100; BS=72-98). Comparative morphology.—We evaluated a series of morphological and anatomical characters including: awn indumentum, form of the callus, lemma epidermis, and anatomy of the leaf blade in transverse section that differ among the DODGE adn of Jarava s.l. Nearly all samples of Jarava s.l. listed in Table 1 were surveyed and re] examples are given in Fig. 4. In addition, features of Amelichloa have also been evaluated. 1 e J. A) 1.1 4 Callus.—In the Pappostipa clade (Fig. 4A-C) the callus i te to acuminate the callus base plane and floret axis is low; callus base is lanceolate; seel See is marginiortit with proximal borders near one another, not fused nor thin, leaving an open foveola (literally, “small depression or scar"; the term used to name the section of conducting bundle at the base of the callus, but not only its open ventral part as it was for species resolved here in the Pappostipa clade by Roig 1964); distal, flat prow of the peripheral ring is not developed or is moderately developed. The foveola is fusiform or lanceolate, usually the same shape as the peripheral ring. In the Jarava ichu group (Fig. 4D) the call hort, truncate, blunt to acute; the angle between the callus base plane and floret axis is higher CS in Pappostipa clade; callus base is obovate or ovate, rarely orbicular or lanceolate; peripheral ring is marginiform to flat, proximal borders not near each other, thinned and not fused, leaving the foveola open; distal, flat prow of the peripheral ring is not developed, inconspicuous or is slightly conspicuous. The foveola is obovate, ovate or orbicular, rarely lanceolate, and usually the same shape as the peripheral ring. In Jarava media, J. plumosula, and J. polyclada, (Fig. 4E) the the callus base plane and floret axis is high; callus base is ovate or acuminate, eege to the M MEM long but obtuse point; peripheral ring is flat or slightly marginiform, proximal borders joined, usually fused and not thinned, closing the foveola; distal, flat prow of the peripheral ring is well developed, characteristically fused at the base. The foveola is oval or orbicular and does not follow the shape of peripheral ring. In Amelichloa caudata and A. cladestina (Fig. 4F) the callus is short, blunt; the angle between the cal- lus pare plane and floret axis is high: callus base is obovate or oblong; peripheral ring is conspicuously , proximal borders not near each other, thinned, leaving the foveola open; distal, flat prow of the pesca ring is not developed or is inconspicuous. The foveola is oval or oblong ellipsoid and usually follows the shape of the peripheral ring. 11 : re 1 ERRA 1. +1 1, 1 Lemma epidermis.—In species of the EE clade (Fig. 4G) the epidermal fundamental cells regularly alternate with silica cells and are all apy tely the same dd or shorter than the silica cells; end walls are straight or slightly uneven, sidewalls slightly si s are oval to oblong rectangular, slightly unequal. In the Jarava s.s. group (Fig. 4H) the — fundamental cells regularly alternate with silica cells and are considerably shorter than the silica cells (designated here as *maize" pattern); end walls are straight, sidewalls thin-walled, lacking silica, straight or slightly ilica bo e t rectangular, often equal. In Amelichloa caudata and A. clandestina (Fig. 41) the dm fundamental cells are usually ir- ba LGG Hra De A AA L [| [| [| Ef A £A D fat A A " In L £n L f NT” Oo A LF searr R e T = F Ff f d L ` H A A i L £n H H hed OA Lia adden TON ( A A n L £ n [Dat DA Lia 131450 fici rr F F E A fa rr A PA n T, A H | L £ I y L (dv? fe on L L ^^ 4777 INT E A ra I L EJ i i rng. Sarena L + kl A fa i ha ^5 mb on L F} -4n72?.£57/117011 FAL A H L £A fe LI TEE d y P OA LI 449590 FIC G [| 3 LE J bal In A Fl E A fa T L al all al [| i n L LH H [| sa IE | FER | H | L al I In nf D d T bk vs. "E A i I I | LE J I I] H [| I la. AL | In J L 4 I, al F T f El re A J A ZE ak HH EI J In A L | alen) j mM + A i: sbb) lr. J D B | £ Lisa + sé E - "I d Li L H od [] J [| [| L JE rn L OA Lila 343343741 [MCV K D Il Ef Lisa m L Ei LI a i zi NM a HJ I IE £n "n re 77727 1101 I n IB | Elli. L st F i a ki ` fa Ki D furrows of Pappostipa chrysophylla. Daum L [| de [| nli [| LA a fat [| D , Phytogeny | , Pappostipa 181 ith silica cells ,and are shorter t telv tl ti the | gt! of the silica cells; regularly alternate end walls are straight or slightly uneven, sidewalls slightly sinuate; ses bodies are oblong, unequal. Leaf blade transection.— Basal leaf blade cross-sections from Jarava barrancaensis, J. speciosa var. major, and J. chrysophylla are shown in Fig. 4J-L, and represent anatomical forms in the Pappostipa clade. We found two distinct types of leaf morphology and anatomy as detected by Arriaga (1983) that correspond with the two subclades P and C in our trees (Figs. 2&3). The P clade has type I leaf blade anatomy (incl. Jarava barrancaensis, J. hieronymusii, J. nicorae, J. patagonica, J. speciosa var. major, J. speciosa var. speciosa, and J. vaginata) with stiff, relatively thick usually pungent leaf apices, adaxial ribs well developed, separated by deep furrows; primary veins (vascular bundles) are numerous (5 or 6), often adaxially and abaxially con- nected by sclerenchyma girders (rarely by Geer EE girders; Fig. 4J); tertiary veins are often abaxially connected by scl | girders; bulliform t at the bottom of all f ; abaxial sclerenchyma is MM ACE dal sitet da is rd (Fig. 4K). The C clade (Figs. 2&3) has type II leaf blade anatomy (incl. Jarava atacamensis, J. frigida, J. humilis, and J. ibarii) with slender, usually non-pungent leaf apices, adaxial ribs not well developed separated by shallow furrows; primary veins not numerous (1-3, rarely to 4), often only abaxially connected by sclerenchyma girders (rarely adaxially connected at the midrib); tertiary veins are AMI not depen by sclerenchyma; bulliform cells are present only close to the midrib; abaxial ; adaxial scleren- E is SE da Or oa ee 1965; giles mia Jaraa lado la (Fig. 4L) differs from li | tertiary veins that are adaxially Fi connected by stletendhymia girders in a compound midrib. TAXONOMY The phyl ic inf tion provided by our DNA analyses coupled with significant morphological differ- ences n the Jarava s.l. clades detected, forces us to recognize the Pappostipa clade as an independently derived monophyletic genus if we are to maintain the monophyletic status of the other genera currently recognized in the AAAJN clade. Here we propose combinations in the new genus Pappostipa for all 14 spe- cies resolved in this clade in our DNA analyses, and Stipa speciosa f. major which is raised to the rank of species in Pappostipa. Eight additional species are transferred to Pappostipa that were formerly included in Stipa subg. Pappostipa by Parodi (1960) and or Jarava s.l. by Peñalillo (2002, 2003), based on their shared morphology with the DNA tested set of species. All currently accepted infraspecific taxa (Peñalillo 2003) in these 23 species are also transferred to Pappostipa. Leaf anatomy types described by Parodi (1960) and Arriaga (1983) correlated with the two Pappostipa subclades (P & C) detected in our DNA analysis. Further review of Pappostipa leaf blade and sheath morphology after our DNA analyses (based on Roig 1964, Nicora Sr Rúgolo de Agrasar 1978, and pers. obs.), continue to support this correlation. Therefore we conclude that Pappostipa can be divided into two morphologically distinct sections of 14 (P. sect. Pappostipa) and nine (P. sect. Chrysovaginatae) species. Pappostipa (Speg.) Romaschenko, PM. Peterson & Soreng, comb. et stat. nov. Basowxw: Stipa subg. Pappostipa Speg., es Mus. Nac. Montevideo 4:111, 45. 1901. Synonym Stipa sect. Papoforeas E. Desv., Fl. Chil. 6:278. 1854. Tyre: Stipa speciosa Trin. € Rupr. (Lecroryre, designated here). Plants perennial, caespitose, sometimes with stolons. Culms 10-70 cm long, erect or geniculate, with two or three nodes, often concealed by sheaths; basal SE E Basal leaves pata UIS cm long, (0.5-)0.7-2.3(2.5) wide, convolute, api t: adaxial sulcate, the furrows 8-13 or only 3-6; abaxial surface alison Seene EE with E E sheaths glabrous or pubescent, open, whitish, pale or stramineous to brownish-red, reddish or purplish; ligules 0.5-1(3) mm long, truncate, membranous, ciliate, or pilose. Flag leaves 4-15 cm long, sometimes deciduous; ligules 2-5 mm long, elongated. Panicles 30-240 mm long, with 5-50 spikelets, often compact, included or exserted in the upper leaf sheath; pedicels 0.5-12 mm long, often hairy. Glumes 15-45 mm long, subequal or slightly unequal, longer than floret, acuminate, hyaline, (1-)3-5-nerved, purplish, green- 182 | tani | Texas 2( ish, rarely plumbeous. Florets fusiform, terete. Calluses 1-4 mm long, often pilose, apex obtuse to truncate; foveolas proximally opened, often the shape of peripheral rings, fusiform or lanceolate; peripheral rings thickened near the margins, flat prow usually not developed. Lemmas 4-16(-17) mm long, chartaceous, P poc with short EE hairs or dorsally pans along the upper % of the lemma, rarely ] the base of the column or dorsally glabrous; pl ickles (hooks) SJ usually present in the: upper — ee up to 1 mm long; apices not fused into a crown. Awns 20-70(-193) mm long, per- sistent, once geniculate; columns 15-33 mm long, sometimes protruding above glumes, plumose, hairs 4-9 mm long; bristles 12-45(-170) mm long, equal to much longer than columns, scabrous, smooth or glabrous, sometimes pungent, pale to chestnut colored or black, straight, rarely sinuous or falcate. Paleas 4-13 mm long, usually subequal to lemmas, occasionally % as long as the lemmas or shorter, linear, linear-cuneate or elongate, hyaline, often dorsally hairy, exceptionally glabrous, two-nerved, rarely nerves extended as short awns, these not conspicuous. Lodicules 3. Stamens 3, anthers yellow or purplish. Pp sect. Pappostipa, sect. nov. Tyre: Pappostipa speciosa (Trin. & Rupr.) Romaschenko. D + Da SRM A SEI? 1 H 4 1 : 1 1.1 PES SAS | ] D t REIS f. P sect Chryso ü red sink. or uE andrade ff, rel EMEN thick wath: pungent apices and usually with well developed adaxial ribs separated by deep furrows (referred to as type I leaf anatomy in Arriaga 1983). “Short-bristle” group The foll ll share ively a short final segment of the awn (referred to as a bristle; Jacobs et al. 1995) that is 1.5-2 times shorter than the bristles in the “long-bristle” group, of P. sect. Pappostipa. Pappostipa speciosa (Trin. & Rupr.) Romaschenko, comb. nov. Basionym: Stipa speciosa Trin. & Rupr., Sp. Gram. = 5. 1842. Stipa humilis var. speciosa ven niles coe ES ege Gen. Pl. 3(3):371. 1898. Achnatherum speciosum (Trin. & R diea Phytologia 74:13. 1993 ñailillo, Gayana, Bot. 59(1):32. 2002. Der CHILE: H. Se n. (HOLOTYPE: LE-TRIN-1443.011; ISOTYPE: ue, 557434 hana ex LEI). Pappostipa speciosa var. manqueclensis (EA. Roig) o comb. nov. Baam Stipa speciosa var. man- queclensis EA. Roig, Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 12:89, J Roig) Pefiailillo, Contr. U.S. Natl. Herb. 48:407. 2003. Type: ARGENTINA: San GE e de Las A E Roig 5339 ( : ME Pappostipa speciosa var. atuelensis (EA. Roig) Romaschenko, comb. nov. Bastonym: Stipa speciosa f. andinas Eit Roig. Revista Fac. Ci. o Univ. Nac. Cuyo 12(1):91-94, lamina 6. 1-9. 1965. Stipa speciosa var. atuelensis (EA. Roig) EA. Roig, Deserta 2:113. 1 var. atuelensis (EA. Roig) Peñailillo, Contr. U.S. See Herb. 48:407. 2003. Type: ARGENTINA. : MERI wd MENDOZA: Depto. San Rafael, — en márgenes del Rio A ae A. Ruiz Leal 232 Pappo speciosa var. parva (FA. Roi g)R EA. Roig, Revista Fac. Ci. Agrar. Univ. Nac. PA 11(1-2):33-35. ma [1966]. J (FA Roig) Peñailillo, Conte U.S. Natl. Herb. 48:407. 2005. TPE: TINA: Las Heras, P e Uspallata, 2800m, 18- ie 1938, A. Ruiz Leal 4880 (uotorvee: MERL). Pappostipa speciosa var. e as (Parodi dee enko, comb. nov. Basiouvw: Stipa speciosa var. breviglumis rodi, Revista Argent. Agron. 27(3-4):74, f. 1, B, b, 1. 1960. Minis nei var. engen (Parodi) Peñailillo, Contr. U.S. Natl. E 48:407. 2003. Tree: ARGENTINA. San Juan: Depto. Iglesia , Mina Fierro Nuevo, 24 Feb 1950, VR. Perrone 54904 1/2 (uororvpz: BAA). Pappostipa speciosa var. pini (EA. iux R henko, comb. nov. B ciliata EA. Roig, Giorn Bot. Ital. 121(1-2):43, f. 2. 1987 JJ ciliata (EA Roig) Peñailillo, Contr. U.S. 5, Natl geg, 48:407. 2003. Tyre: ARGEN- UAN: Iglesia, entre Tocota y le los H ar 1962, 2180m, A. Ruiz Leal 22093 (HOLOTYPE: MERL; 1sorvrE: SI). Pappostipa na f horrida (EA Roig) lo comb. nov. Basseng Stipa speciosa f. horrida EA. Roig, c. Ci. Agrar. Univ. Nac. Cuyo 11(1-2):36, t. 25a. 1964 [1966]. Jarava speciosa f. horrida (EA. Roig) Peñailillo, Contr. U.S. Gë SCH 48:407. 2003. Tree: ARGENTINA: Malalhue, Cerro Huemul, frecuente, matas aisladas, rígidas, pinchudas, 14 Jan 1958, A. Ruiz Leal & EA. Roig 18784 (HoLoTyPE: MERL?). Pappostipa speciosa f. abscondita (FA. Roig) Romaschenko, comb. nov. Bastonw: Stipa speciosa f. abscondita EA. g, Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 11(1-2):36-37. 1964 [1966]. Jarava speciosa f. abscondita (EA. Roig) Peñailillo, Contr. us Natl. Herb. 48:407. 2003. Tree: ARGENTINA: Las Heras, Uspallata, Cumbre del Paramillo, 11 Feb 1964, FA.. Roig 4929 (Ho- orvre: MERL). moi speciosa var. media (Torres) Romaschenko, comb. nov. Basionwm: Stipa speciosa var. media Torres, Comis. Invest. Ci. tee Aires] 12:44, t.1, C, c; t.4d. 1993. Jarava speciosa ) Peñailillo, Contr. U.S. Natl. Herb. 48:407. RGENTINA. Buenos Ames: Pdo. Villarino, no Chasicó; 19 Apr 1976, J. Frangi 79 (RoLoTYek: LP). Pappostipa mila (Phil.) Romaschenko, comb. nov. Basionvm: Stipa vaginata Phil., Linnaea 33(3-4):281. 1864. Jarava ta (Phil.) E Rojas, Gayana, Bot. 54(2):173. 1997 [1998]. Type: CHILE. ACONCAGUA: 5. Felipe, Nov 1862, Landbeck s.n. (HOLOTYPE:?; ISOTYPES: SGO-45662, US-81900 fragm. ex W!). n L [| Ab el mi | ra H Es: , Dannnctina 183 Sepp ca f. rigida (EA. Roig) Romaschenko, comb. nov. Basion: Stipa vaginata f. rigida EA. Roig, Revista r. Univ. Nac. Cuyo 11(1-2):54-55. 1964 [1966]. Jarava vaginata f. rigida (EA. Roig) Peñailillo, Contr. U.S. Natl. Herb. 48:407. 2003. Tree: ARGENTINA: Las Heras, Paramillo de Uspallata, Cumbre del Paramillo, 2850m, 11 Feb 1964, FA, Roig 4927 (HOLOTYPE: ?; ISOTYPE: Peppe iis cane f. inmersa (EA. Roig) Romaschenko, comb. nov. Basionym: Stipa vaginata f. inmersa EA. Roig, c. Ct. Agrar. Univ. Nac. Cuyo 11(1-2):53-54, t. 10 & 25e. 1964 [1966]. Jarava vaginata f. inmersa (EA. Roig) Peñailillo, Ge U. E Natl. Herb. 48:407. 2003. Tree: ARGENTINA: Las Heras, Cumbre del Paramillo e Uspallata, 2800m, 22 Jan 1960, E 4319 [EA. Roig 4308 cited also in parentheses with 4319] (HoLoryrE: MERL?; 1som Paba vaginata f. laevis (EA. Roig) Romaschenko, comb. nov. Basionm: e vaginata f. laevis FA. Roig, Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 11(1-2): 52, t. 10 & 25g. 196 i «ness, Jarava Reg laevis (EA. Roig) Peñailillo, Contr. U.S. Natl. Herb. 48:407. 2003. Tyre: ARGENTINA: Las Heras, P pallat 00m, 22 Jan 1960, FA. Roig & A. Ruiz Leal 4318 (HOLOTYPE: MERL?). Pappostipa vaginata f. contracta (FA. Roig) Romaschenko, comb. nov. Bastonym: Stipa vaginata f. contracta FA. Roig, Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 11(1-2):53, t. 10 & 25f. 1964 [1966]. Jarava vaginata f. contracta (FA. Roig) Peñailillo, ntr. U.S. Natl. Herb. 48:407. 2003. H ARGENTINA: Las Heras, Los Hornillos, 10 Jan 1959, A. Ruiz Leal 20144 (HOLOTYPE: MERL herb. FA. Roig 3405; IsoTYP Pappostipa vaginata var. argyroidea (EA. Roig) Romaschenko, comb. nov. Basiowvw: Stipa vaginata var. argyroidea EA. Roig, Deserta 2:107, t. 1. 1971) ginata var. EE (EA. Roig) Peñailillo, Contr. U.S. Natl. Herb. 48:407. 2003. TPE: ARGENTINA. MENDOZA: Depto Malalhue, entre C y Fortín Malalhue, 9 Nov 1960, EA. Roig 4514 (HOLOTYPE: MERL?; ISOTYPE: x Herb. Roig 4514). Pappostipa ut var. dilatata (EA. Roig) ee comb nov. B st ta var. dilatata EA. Roig, Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 11(1-2):54. 1964 [1966]. J t dilatata (EA Roig) Peñailil o, Contr. U.S. Natl. Herb. 48:407. 2003. Tre: ARGENTINA: Las Heras, Camino a us por ? Potrerillos, ala vera del mismo, en suelos removidos, arenosos, 22 Dec 1961, A. Ruiz Leal, Roig & Wainstein 4523 (HOLOTYPE: ?; 150 Pappostipa nicorae (FA. Roig) Romaschenko, comb. nov. Bang Geer nicorae FA. Roig, Giorn. Bot. Ital. 121(1-2):41, 987. Jarava nicorae (FA. Roig) Peñailillo, Gayana, Bot. 59(1):32. 2002. Tree: ARGENTINA. San Juan: Depto. Iglesia, Reserva " San Guillermo, Cajoncito de la Brea al N del Infiernillo, 3680m, 20 Feb 1981, E. Nicora, Guaglianone & Ragonese 8201 (HOLOTYPE: SI; E LP). Pappostina a oai (Pilg.) Romaschenko, comb. nov. Basonvm: Stipa pi id Pilg., Bot. Jahrb. Syst. 56 (Beibl. 123):24. ine alas EE (Pig) aaa ea E a ):31. 2002. Trre: ARGEN . La Rioja: Depto. Gral. Sarm- iento, C ja, 26 Feb 1879, G. ec Niederlein 314 (HOLOTYPE: B; isorvres: BAA-3071 fragm. ex B, CORD, US- 866109 fragm!). Pappostipa patagonica (Speg.) Romaschenko, comb. nov. Basionym: Sti , Revista Fac. Agron. Veterin. (Buenos Aires) 3:581. 1897. Jarava patagonica (Speg.) Peñailillo, a Bot. 59(1): 32 2002. Tre: ARGENTINA. SANTA Cruz: Río Santa Cruz, Feb 1882, C. Spegazzini s.n. (Lectotype: LP ex LPS-2433 designated by L.R. Parodi, Revista Argent. Agron. 27(3-4):72. 1300; ISOLECTOTYPES: Lin LPS Pap Roig) R henko, comb. nov. 8 FA. Roig, Bol. Soc. Argent. Bot. 14(4):314, f. 2. 1972. Jarava maeviae (FA. Roig) Peñailillo, Gayana Bot. 59(1):31. 2002. Tar: ARGENTINA. ee Depto. Chos Malal, filo volcán Domuyo, en pedregales, s. d., O. Boelck 11290 (HoLoryrE: BAA; 150 AB). Pappostipa parodiana (FA. Roig) Kumpel comb. nov. BASIONYM: cn parodiana FA. Roig, Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 11(1-2):44-46, t. 8, 26b. 1964 [1966]. Jarava parodiana (EA. Roig) Peñailillo, Gayana, Bot. 59(1):31. 2002. Tyee: SE TINA: m. Ranquil de Lirkai, 1100m, frecuente en las laderas empinadas de los cerros, 14 Dec 1960, A. Ruiz Leal 53 (HoLoTYFE: MER oro dea GE (FA. Roig) ee comb. nov. B i blanquetii FA. Roig, Phytocoe SEN 2(1-2): 19, f 1 1975. Jara al ii (FA. Roig g) Peñailillo, Contr. U.S. Natl Herb 48:407. 2003. Tre: ARGENTINA. za: San Rafael, Valle de Las Leñas, 13 ee? 970, A. Ruiz Leal 27150 da MERL; isoTyPe: LP). Pupposdpa semperiana (FA. Roig) jii M comb. no EA. Roig, Revista Fac. Ci. Agrar. Univ. Nac. e ue Se 37t. 6, a gl (FA. Roig) Peñaili lo, Cana Bot. 59(1):31. 2002. Type: AR- TINA: 1 S Albe rto (La toledo) 27 Dec 1947, s e Leal 11138 (Hororyre: MERL). Pappostipa: rui leal (EA. Row) cdo. Com» nov. B Roig, Revista Fac. Ci. Agrar. Univ Nac HE x ie t. 5, 25c. 1964 [1966]. J lii (EA. Roig) Penailillo, Gayana, Bot. 59(1):31. 2002. Tyre: ARGENTINA: Las Her llata, 2800m, 23 Jan 1960, A. Ruiz Leal & EA. Roig 20774 GaoLotyPeE: MERL-20774). "Long-bristle" group The following f 11 s] d j | teristic oF havine the final tofthe awn (referred to as a brisde: aura et al. 1995) as being i 5-2 times longer than the bristles in a he “short-bristle” group of species of P. sect. Pappostipa. Pappostipa malalhuensis (FA. Roig) Romaschenko, comb. nov. Bastonwu: Stipa malalhuensis EA. Roig, Revista Fac. Ci. I Lo fal Dos » In LF PE fT Safa) RE, 184 Agrar. Univ. Nac. Cuyo 11(1-2):60-62, t, 13, 25d. 1964 [1966]. gena ds ees = cece Peñailillo, Gayana, Bot. 59(1):31. 2002. Tree: ARGENTINA: Malathue, Poti Malal, ne “coirón blanco,” 13 Jan 1960, A. R 938 (HOLOTYPE: MERL-20938). Pappostipa major (Speg.) Romaschenko, comb. et stat. nov. Basionym: Stipa speciosa f. major Speg., Anales Mus. Nac. ntevideo 4(2):58, f. 10a-b. 1901. Stipa speciosa var. major (Speg.) Parodi, Revista Argent. Agron. 27(3-4): 75, f. 1, C, c. 1960. biis speciosa var. major (Speg.) Peñailillo, Contr. U.S. Natl. Herb. 48:407. 2003. Tree: ARGENTINA: lipid prope Lago Nahuel- huapi, Jan 1900, E. Fernádez s.n. (syw1vee: ?). Neuquen: in Valle Trolope, Feb 1900, Otto Asp s.n. (SYNTYPE: Pappostipa major can be separated from P. speciosa by having longer glumes, narrower and convolute leaf blades, longer bristles usually 1.5 to 2 times as long, black (yellow in P. speciosa) bristles, larger anthecia, and a chromosome base number of 2n=60 (2n=66 in P. speciosa) [Parodi 1960; Stebbins & Love 1941; Covas & Bocklet 1945]. Futhermore, these varieties were resolved independently of one another in both the plastid and ITS DNA analyses (Figs. 1, 2 & 3). iode vatroensis (EA. Roig) Romaschenko, comb. nov. Basionym: Stipa vatroensis EA. Roig, Revista Fac. Ci. Agrar. Cuyo 11(1-2):40-42, t. 7, 26a. 1964 [1966]. dc vatroensis es Roig) Peñailillo, Gayana, Bot. 59(1): 31.2002. Tyre: Mm Malalhue, el Vatro, 1100m, en l g [ , en laderas, 16 Dec 1960, A. Ruiz Leal 21501 = EA. Roig 3765 ( : M Pappostipa barrancaensis (EA. Roig) Romaschenko, comb. nov. Basionym: Stipa vatroensis EA. Roig, Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 11(1-2):40-42, t. 7, 26a. 1964 [1966]. Jarava barrancaensis (EA. Roig) Peñailillo, Gavana, Bot. 59(1):31. 2002. Tree: ARGENTINA: Malalhue, el Vatro, 1100m, en la margen izquierda del río Barrancas, en laderas, 16 Dec 1960, A. Ruiz Leal 21501 = EA. Roig 3765 ( RL) D H I halla {BRB TA 3n 1 1 roe V Pappostipa sect. — Romaschenko, sect. nov. Tere: Pappostipa chr) n: 1 REN 1 F. :cY EISE A is et fasciculati on coriaceis, À Pappostipa sect. Pappostipa imis vaginis setaceis (valde acutis, a non nba EE us "oluti O in trans-sectione), et cum non m adaxialibus Pappostipa sect. Chrysovaginatae differs from P. sect. Pappostipa by having pale or stramineous to golden, basal sheaths and basal leaves that are thin and not coriaceous, set (sharply pointed but not pungent), straight, convolute (terete in cross section), and with shallow adaxial furrows. All species have type II leaf anatomy as described by iind (1983). Pappostipa so sorianoi (Parod henko, comb. nov. B i Parodi, Revista Argent. Agron. 27(3-4):94, 960. Jarava sorianoi (Parodi) Peñailillo, Gayana, Bot. 59(1):31. 2002. Tre: ARGENTINA. SANTA Cruz: 20 km al norte rito Moreno, 13 Dec 1954, A. Soriano 4775 (HoLoTyPE: BAA!). Pappostipa atacamensis (Parodi) Romaschenko, comb. nov. Basionym: Sti Parodi, Revista Argent. Agron 27(3-4):85, f. 3. 1960 [1961]. Jarava atacamensis (Parodi) Peñailillo, Gayana, Bot. 50(D: 3]. 2002. Tere: CHILE. Atacama: Depto. Copiapó, Quebrada de Plaza, 3600m, 4 Feb 1949, A. Krapovickas & J. Hunziker 5789 (HoLotyre: BAA-2964; ISOTYPES: CONC-29647, 3 SGO-7325 Pappostipa nana a (S eg.) Romaschenko, comb. nov. B St Speg., Revista Argent. Bot. 1:23. 1925. Jarava nana ( cane ñailillo, Gayana, Bot. 59(1):31. 2002. Tree: ARGENTINA. E CRUZ: 1/2 lega al S de la laguna de Sehuenaiken, 3 Feb 190 Spegazzini s.n. (HOLOTYPE: LP ex LPS-2501). "—— v ibarii (Phil.) Romaschenko, comb. nov. Basionvm: Stipa ibarii Phil., Anales Univ. Chile 93:716. 1896. Jarava ibarii (Phil.) Penailillo, Gayana, Bot. 59(1):31. 2002. Tyre: CHILE: Patagonia ad lacum Pinto, Dec 1877, H. Ibar s.n. (HOLOTYPE: SGO-45658; ISOTYPES: x BAN 2028 Em ex D SE US-1762378 fragm. ex SGO!, US- 141025 SE US-photo ex SGO-45658!). p ,CO omn pa llescens Parodi, Revista Fac. Ci. mes r. Univ. Nac. Eno 111-2): ae 57,4. 1L 1964 abad 1 (Parodi) Peñailill pu U.S. Natl. Herb. 48:407. 2003. Tyre: ARGENTINA: Las Heras, Paramillo de Uspallata, entre 2600- 2800m, 18- 19 Feb 1938, A. Ruiz Leal 4830 (noote: BAA). Pappostipa ibarii var. anomala (Parodi) Romaschenko, EE nov. Basiony: Stipa ibarii var. anomala Parodi, Revista Ar in jd 27(3-4):89. 1960. Jarava ibarii var ñailillo, Contr. U.S. Natl. Herb. 48:407. 2003. Tyre: ARGEN- T A Cruz: Depto. Río Chico, Gdor. Gregores, A. Soriano 5080 (noroTrrE: BAA). Pappostipa aio (Speg.) Romaschenko, comb. nov. Basionym: Sti hi g., Anales Mus. Nac. Montevideo (2):163-165, f. 50. 1901. Jarava eee (Speg.) Peñailillo, o Bot. 59(1): 31. 2002. Tree: ARGENTINA. Santa Cruz: Golfo an Jorge, Aest. 1899, C. Ameghino s.n. (HOLOTYPE: LP-12593 pri ameghinoi var. TEEN (Parodi) Romaschenko, comb. 1 NOV. BASIONYM: is ameghinoi var. digona Parodi, Revista Argent. Agron. 27(3-4):80, f. 2. 1960. J ñailillo, Contr. U.S. Natl. Herb. 48:407. 2003. Tree: ARGENTINA. Santa Cruz: Chicorikaike, 50º S 69º 25 W, Jan 1930, A. Renard 13 (HOLOTYPE: Ge Pappostipa ameghinoi var. precordillerana (FA. Roig) Romaschenko, comb. nov. Bastonym: Stipa ameghinoi var. precordillerana FA. Roig, Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 12(1):79, 80-81, t. 1. 1965. Jarava ameghinoi var. precordillerana Damaes Lan) x Jl DL. £A . es J i Pappostipa 185 (EA. Roig) Peñailillo, Contr. U.S. Natl. Herb. 48:407. 2003. Tre: ARGENTINA. MenDOza: Las Heras, Alto de los Manantiales, 20 Feb 1965, EA. Roig 5284 (HoLoTYPE: MERL; Is LP) Pappostipa chubutensis (Speg. ) Romaschenko, comb. nov. Stipa chubutensis Speg., Anales Mus. Nac. Montevideo 4(2):49-51, f. 7. 1901. J peg.) Peñailillo, Gayana, Bot. 59(1): 31 2002. Tyre: ARGENTINA. Cuusut: Teka Choque, Feb 1900, O. Mauri s.n. (HoLoTYPE: BAA ex LPS-2477; ¡soTYPES: US-1721311!, US-141588)). Pappostipa chubutensis var. hirsutissima (EA. Roig) Romaschenko, comb. nov. Basowvw: Stipa chubutensis var. hirsutissima EA. Roig, Bol. Soc. di d Bot. e 316, » 3. 1972. Jarava chubutensis var. t e Ge E ues U.S. Natl. Herb. 48:407. 2003. T to. Puerto Deseado v 1963, Correa et al. 2638 (HoLorrrE: BAA; ISOTYPES: BAA-BAB, LP). Pappostipa frigida (Phil.) Romaschenko, comb. nov. Basionvw: Stipa frigida Phil., Fl. Atacam. 54. 1860. Jarava frigida (Phil) E ee Gayana, Bot. 54(2):173. 1997 [1998]. Tyre: CHILE: cerro Altos de Puquios, ca. 3810m, Feb. 1864, Philippi s.n. (HOLOTYPE: GO-63153). Pappostipa frigida var. Pore (Parodi) Romaschenko, om nov. qn Stipa frigida var. parvispicula Parodi, Revista Argent. Agron. 27(3-4):9 , €. 1960. Jarava frigida var. ñailillo, Contr. U.S. Natl. Herb. 48:407. 2003. Tyre: Se La Zen SE Sarmiento, Paso de inde Negras 4100m, en E límite Argentino-Chileno, 6 Feb 1949, A. Krapovickas & J. Hunziker 5798 (HoLorYPE: BAA: IsoTYPES: BAB, CORD) SE humilis Geen Romaschenko, comb. nov. B Stipa humilis Cav., Icon. 5:41, t Ge l 1. 1799. Jara ums av.) Peñailillo, Gayana, Bot. 59(1):31. 2002. Tree: CHILE: P D lo, "Habitat in A vulgo Deseado o arido, floretque Decembri, Luis Née s.n.,” Nee s.n. (HoLoTYPE: MA; isotypes: SGO photo, US-866105 fra ail): juo humilis var. decrescens (Kuntze) E EEN comi, NOV. Basionym: Stipa humilis var. decrescens Kuntze, Revis. Gen. Pl. 3(3):371. 1898. Jarava humilis var. d illo, Contr. U.S. Natl. Herb. 48:407. 2003. Tyre: ARGEN- A: Patagonia, Paso Cruz, 2000m, 1 Apr 1882, Moreno & Tonini 285 (syntype: NY!; : US-A866104 ex NY!). ge humilis var. ruiziana (Parodi) Romaschenko, comb. nov. Basin: Stipa humilis var. ruiziana Parodi, Re- vista Argent. Agron. 27(3-4):104. 1960. Jarava iind var. ruiziana (Parodi) Penailillo, Contr. U.S. Natl. Herb. 48:407. 2003. TE: ARGENTINA. MrNpoza: Depto. Las Heras, d llos, abundante, 21 Jan 1944, A. Ruiz Leal 8593 (HoLoYrrE: BAA). Pappostipa chrysophylla (E. Desv.) qe nu, comb. nov. Basionim: Stipa chrysophylla E. Desv., Fl. Chil. 6:278, t. 1854. Stipa humilis f. o (E. Desv) Kuntze, Revis. Gen. Pl. 3(3):371. 1898. Stipa speciosa subsp. sata (E. Desv. is Dusén, Rep. Princeton Univ. Exp. Patagonia, Bot., Suppl. 8(3):30-31. 1914 [1915]. Jarava chrysophylla (E. Desv.) Peñailillo, Gayana, Bot. 59(1):31. 2002. Tee: CHILE: Coquimbo, en el valle de Toro a 3497m, oue un terreno "e — pe Patos, C. Gay 376 (SYNTYPE: P; isosynTYPE: US-866134!). CHILE: Habita en los hel 2003m, y en ps an s.n E: psa chryso ophylla f. a A Roig) Romaschenko, comb. nov. Basiony: Stipa chrysophylla f. minuta EA. Roig, Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 11(1- = 66, t. ndi 1964 [1966]. Jarava chrysophylla 1. minuta (EA. Roig) Peñailillo, Contr. U.S. Natl. Herb. 48:407. 2003. Tree: ARGENT Thue, Los Molles, 1850-1950m, césped pequeño, hojas erectas, 11-22 1960, A. Ruiz Leal 20847 (HoLoTirE: MERL?). ratos chrysophylla var. cordillerarum (Parodi) Romaschenko, comb. nov. Basiowvw: Stipa chrysophylla var. cordillerarum Parodi, Revista Argen SE 27(3-4):99. 1960. Jarava EDO ANE var. Mia cii d img VEN a Bot. iin 31. 2002. Tyre: ARGENTINA. La Rioja: Depto. Sarmiento, Kra kas & J. Hunziker 5855 (HoLoTYPE: BAA: CORD). Pappostipa chrysophylla var. crispula ea Romaschenko, comb. nov. Basionya: Stipa humilis f. crispula Kuntze, n. Pl. 3(3):371. 1898. Stipa chrysophylla var. crispula (Kuntze) Parodi, Revista Argent. Agron. 27(3-4):98. 1960. Jarava dc var. crispula (Kuntze) Peñailillo, Contr. U.S. Natl. Herb. 48:407. 2003. Terr: ARGENTINA: Paso Cruz, Cordillera, 34°N, 1500m, 1 Jan 1892, CEO Kuntze 72 (HoLotYrE: NY!). Pappostipa chrysophylla f. modica (EA. Roig) Romaschenko, comb. nov. Basionwx: Stipa chrysophylla f. modica Roig, Revista Fac. Ci. Agrar. Univ. Nac. Cuyo 11(1-2):66-67. 1964 [1966]. Jarava chrysophylla f. modica (EA. Roig) Peñailillo, Contr. U.S. Natl. Herb. 48:407. 2003. Tree: ARGENTINA: Las Heras, Paramillo de Uspallata, 2800m, 9-11 Feb 1964, A. Ruiz Leal 23249 (HOLOTYPE: MERL?). DISCUSSION Basal lineages.—1n order to focus on Jarava s.l. and the major American lineages of the Stipeae in the present paper we have excluded all except representatives of two extra-American lineages (i.e., one Asian Ptilagrostis and five species of the Australian genus Austrostipa) of Stipeae from our DNA analyses. Several small American genera were excluded from the present paper as they were not considered to be pertinent to the subjects of the present paper (see Methods), but they will be treated in subsequent publications. The phylogenetic patterns we detected with the present data are consistent with those detected with our larger worldwide dataset for Stipeae (preliminary ITS plastid analysis presented by Romaschenko et al. 2007; and 186 Journal of the Botanical h Instit Texas 2( unpublished EE e iind Because our taxon sampling here i posefully curtailed, discussions of the basal radiat estrained in nm paper. Si t] is a larger phylogenetic story in the full dataset, not too much should ie radi into e placement of Austrostipa by itself as sister to the Major American Clade (MAC) in which, Achnatherum, Jarava s.s., Nassella, and Amelichloa are included. Nevertheless, some things can be said about the basal lineages as resolved here. The early diverging or sister group position of the basal lineages (here Piptochaetium, Ptilagrostis, and Anatherostipa) in relation to Achnatherum, Amelichloa, Austrostipa, Jarava s.l., and Nassella [AAAJN] has never been indicated in previous reconstructions of the phylogeny of the Stipeae. An early diverging position of Piptochaetium (as a part of a complex of North American Piptatherum/Oryzopsis plus Piptochaetium) within Stipeae was reported by Jacobs et al. (2000) based on ITS sequence analysis. However, the authors assigned more weight to morphological similarities of this group with Nassella and consequently the early diverging position of Piptochaetium was thought to be an artifact. The early diverging position of Piptochaetium also was not recognized when a larger data set was used (Jacobs et al. 2007). Various morphological characters, such as: short, obovate floret, eccentric straight hairless awns, development of a crown, bulbous-based hairs on the apex of the lemma, silicified walls of fundamental cells ofthe lemma epidermis, and type of epiblast has suggested a close relationship between Piptochaetium and Nassella Thomasson 1978; Barkworth & Everett 1987; Barkworth 1990, 1993; Jacobs et al. 2000; Barkworth & Torres 2001; Cialdella & Giussani 2002). In contrast to these conclusions, our data support a monophyletic origin of Piptochaetium near the base of the American Stipeae, thereby rendering Nassella and Piptochaetium as distant relatives. An independent origin of the above suite of characters in Piptochaetium, well removed from Nassella, is in agreement with the latest revision of Piptochaetium based on phylogenetic analysis of plastid trnL-F and rpl16 sequences and morphological characters (Cialdella et al. 2007). All three genera of our “basal lineages” set (i.e.; Piptochaetium, Ptilagrostis, and Anatherostipa) have a "saw" like pattern on the lemma fundamental cells, where the side walls are usually much longer than end walls, strictly sinuate, lobate or dentate, and fundamental cells do not regularly Ee We SE Ree (Barkworth 1983; Barkworth & Everett 1987). This lemma epidermal pattern is ev idently the p condition in Stipeae and can be assumed to be an unspecialized characteristic within the Pooideae since no other pattern has been ii SO i in Méien to EE fossil stipoid grasses (Thomasson 1978, 1980, 1981, 1982, 1985). There a ¡fiable fossils from the Oligocene of the Stipeae called Berriochloa M.K. Elias with a “saw” like pattes in Xd lemma epidermis. Some early Pliocene fossils referred to Nassella (Thomasson 1978) have short fundamental cells but still have sinuate sidewalls and end walls like the basal lineages identified here, such as Piptochaetium but unlike extant species of Nassella. Our molecular analyses indicate that Ptilagrostis and Achnatherum are also phylogenetically distant. This conclusion is further supported with micromorphological data, as the two genera have different lemma epidermal patterns (Barkworth 1983; Barkworth & Everett 1987). Based on the topology of our trees (Figs. 1-3), the polarization of the epidermal characters from Achnatherum (as a conditionally ancient state) to Ptilagrostis (as a conditionally derived state), as proposed Tzvelev (1977) and Barkworth and Everett (1987), is not parsimonious. We deduce that Ptilagrostis has the ancestral conditions for epidermal characters. The monophyly of the genus Anatherostipa is supported by all our molecular analyses (Figs. 1-3) and Ptilagrostis appears to be the sister group of Anatherostipa plus Piptochaetium (Figs. 2 S » Rr dees of Anatherostipa other than the “saw” like lemma epidermal pattern, are the short glumes relative to the lemt long (often prow-tipped) paleas, and lemmas occasionally with small teeth or lees at the apices (! (Matthei 1965; Barkworth & Everett 1987). These lemma apex characteristics were considered by Tzvelev (1977) to be an ancestral or plesiomorphic state in the Stipeae. Species of Anatherostipa share the lemma surface pattern and apical teeth character with Ptilagrostis. The great variation in morphological forms, particularly the diversity of modifications in shape and size of floret and awn indumentum with apparent adaptive functions, within the basal lineages and MÁC, is a primary reason why these separate groups were difficult to detect based solely on the study of mor- DawsanrekanLba neal Dhal £A H Cs: J Pannactina 187 - phology. This confirms the suggestions of Thomasson (1978, 1985) and Barkworth and Everett (1987) that similar adaptive forms evolved independently, pom SUM Wee Sieg NE JM EE in phylogenetically distant groups of Stipeae. Desp eto or herbivoric adaptations, in addition to sharing the unc (ane E pattern, the basal lineages among themselves have relatively low levels of divergence in DNA sequences. The divergence value for the most variable ITS region was 17.696 for the out-group to ingroup taxa and is only 2.996 for the basal lin- eages. This latter value is not significantly higher than for Nassella where it was 2.696. Relatively high (up to 4%) sequence divergence values between the basal lineages and the Hesperostipa clade indicates a greater evolutionary distance between them even though they share the “saw” like lemma pattern. The “saw” like lemma epidermal pattern found in the basal lineages is also known to occur in Hesperostipa (Thomasson 1978, 1980, 1982, 1985; Barkworth 1990; Cialdella & Giussani 2002). Hesperostipa also shares a long and prow-tipped palea with Anatherostipa (Barkworth & Everett 1987). Hesperostipa is a small phylogenetically and geographically isolated genus of the North American Stipeae. Pappostipa.—The phylogenetic history of the Pappostipa clade supported by our molecular analysis of plastid DNA and nrDNA sequences (Figs. 1-3) provides evidence that Jarava s.l. is polyphyletic and the circumscription of the genus proposed by Peñailillo (2002) cannot be maintained. Consequently, we el- evated all of Spegazzini's Stipa sect. Pappostipa to generic status based on corroborating morphological and anatomical evidence of differences with the remainder of Jarava. The phylogenetic resolution of Austrostipa and Achnatherum clades between Pappostipa clade and more highly derived Jarava s.s. clade clearly indicates that Pappostipa and Jarava s.s. do not share a most recent common ancestor. Pappostipa appears to be inde- pendently derived from the group we have called the Major American Clade (MAC). MAC is comprised of related genera that are geographically ca to Due ro dd vun Jarava s.s. and Nassella constituting the principal part of the clade. Recent phyl on plastid DNA for the South American Stipeae (Cialdella et al. 2007) ee the affinities of Pappostipa with the basal lineages since Pappostipa vaginata was inferred to be the sister group of an Aciachne-Anatherostipa-Piptochaetium clade. The phylogenetic position of Pappostipa speciosa (reported as “Achnatherum speciosum”) within the Achnatherum clade based on ITS sequences by Jacobs et al. idi zn is pone oy our more extensive analysis. Some key morphological features of the Pap; itional characteristics leading to the more advanced states found in MAC, or to analogous states of characteristics independently derived in other genera. Unigeniculate awns, straight and long-hairy columns, and usually straight glabrous bristles are the principal diagnostic characters for Pappostipa (Spegazzini 1901). The same suite of morphological features of the awn is developed in central Asian Stipa sect. Pseudoptilagrostis Tzvel. (Tzvelev 1977) and two Mediterranean genera, Celtica F.M. Vázquez & Barkworth and Macrochloa Kunth (Vázquez & Barkworth 2004). Among awn characteristics, Ptilagrostis differs from Pappostipa by having twice geniculate awns, and Macrochloa ditfers from Pappostipa by having falcate rather than straight bristles. The callus of Pappostipa is narrowly acute to acuminate with a characteristic lanceolate or linear-lanceolate form with a basally open foveola that follows the shape of the peripheral ring a O However, a similar structure is known in Stipa subsessiliflora (Rupr.) Roshev. and Stipa sect. P tis (Freitag 1985; Barkworth & Everett 1987. After complete removal of the cuticle the walls of the leming epidermal cells appear to be sinuate in Pap- postipa (Fig. 4G) and not lobate. This type of lemma epidermal wall can be considered transitional between the “saw” like pattern found in Hesperostipa and other basal lineage genera (Fig. 4G) and the pattern found in MAC. However, the moderate shortening of the fundamental d e their TRAE alternating pattern with equilateral silica cells (*maize" like pattern) enables us to l f Pappostipa with other achnatheroid type species rather than with the pattern encountered in the basal lineages. Our ITS analysis and combined plastid DNA and nrDNA analyses (Figs. 2&3) divides Pappostipa into two clades that are correlated with anatomical types 1 & II of Arriaga (1983). These lineages are here named as Papposti Pa ipa and P. sect. Chysovaginatae. Based on our molecular anal 1 by possessing a Bue of teca and Tea morphological characteristics we placed Pappostipa barrancaensis, P. braun- [| £ al Dos » Im Li dt 188 fTexas 2(1) blanquetii, P. hieronymusii, P. maeviae, P. major, P. malahuensis, P. nicorae, P. parodiana, P. patagonica, P. ruiz- lealii, P. speciosa, P. vaginata, and P. vatroensis in P. sect. Pappostipa; and include P. ameghinoi, P. atacamensis, P. chrysophylla, P. chubutensis, P. frigida, P. humilis, P. ibarii, and P. nana in P. sect. Chysovaginatae. Within P. sect. Pappostipa we also recognize an informal “long-bristle” group that includes P. barrancancaensis, P. major, P. malahuensis, and P. vatroensis. These four species all have long straight bristles, whereas the other spe- cies have “short bristles.” We have recognized P. speciosa var. major as a species, P. major, based on different chromosome number reports (2n=60 in P. major; 2n=66 in P. speciosa; Parodi 1960; Stebbins & Lóve 1941; Covas & Bocklet 1945), distinct differences in the length of the hairs on the lemma and glumes, thickness of the basal leaf blades, and color of the sheaths. Austrostipa.—The monophyly of the Australian Stipeae, i.e., Austrostipa was supported in all of our sepa- rate and combined analyses of plastid DNA and nrDNA regions (Figs 1-3). The Austrostipa clade is clearly positioned as a sister group to the Major American Clade (MAC) composed of Achnatherum, Jarava s.s., and Nassella (with Amelichloa nested inside). This general relationship, but with more Asian genera represented in the clade in which Austrostipa was included, was detected by Romaschenko et al. 2007. The monophyly of the Austrostipa was supported by Jacobs et al. (2000, 2007) based on the ITS analysis. However, the clade was thought to be the most derived group in the Stipeae, preceded by clades or grades containing Nassella, Jarava, and Achnatherum. It has been pointed out by Barkworth and Everett (1987) that the majority of the Austrostipa species do not have shortened fundamental cells, which is a distinctive character in species of Nassella and Jarava s.s. In addition, it was found that Austrostipa species typically have fundamental cells with slightly sinuate walls that sometimes are two or three times longer than the silica cells (Jacobs & Everett 1996). Therefore, we consider Austrostipa to be a somewhat specialized group and transitional to the American Achnatherum. However, the nature of this lineage will be covered in more detail in a separate paper where it will be placed in context of a World wide sample of the Stipeae. Major American Clade.—One of the primary challenges of this study was to verify monophyly and ascer- tain the boundaries of the largest group of American Stipeae. We call it the Major American Clade (MAC) because it contains nearly 72% of the species in the American Stipeae. MAC appears to be monophyletic and is a geographically distinct lineage comprised of Achnatherum (American lineage), Jarava s.s. (as a clade or grade), Amelichloa, and Nassella. MAC has consistent statistical support (Figs. 1-3) as a lineage and ap- pears as a sister group to Austrostipa. The detection of MAC provides a new phylogenetic hypothesis for the evolutionary derivation of the majority of the American Stipeae. Achnatherum.—The monophyly of the subset American species of Achnatherum sampled (4 out of 34 spe- cies) was supported by our plastid and nuclear-derived trees (Figs. 1-3) and this Achnatherum clade was always the sister group to the Jarava s.s. plus the Nassella-Amelichloa complex, in agreement with analyses by Jacobs et al. (2000). Jacobs et al. (2000) suggested that Achnatherum was a close relative of Austrostipa, a hypothesis that is also supported by our analysis. BOUM e exhibits a derived lemma epidermal pattern (with fundamental cells that alternate with silic clear ^maize" pattern), and does not have the “saw” like lemma epidermal pattern found in the basal lineages and Hesperostipa. Jarava s.s-Nassella complex.—The core of MAC is the Jarava s.s. plus Nassella-Amelichloa complex. This group contains ca. 160 species, representing ca 60% of ES American Stipeae. In all combined analyses of plastid DNA and nrDNA this clade letic with high statistical support (Figs. 1-3). Jacobs et al. (2000, 2007) did not detect this complex as a dale: in their ITS tree, whereas recent phyloge- netic inferences based on plastid DNA data (Cialdella et al. 2007) are consistent with our results. However, A coto a topologies yielded by parsimony and Bayesian analyses were not consistent when plastid and nuclear markers. Based on combined data of plastid DNA regions, Jarava ista was placed in a polytomy between the Nassella-Amelichloa clade and the core of Jarava s.s. clade. Jarava s.s was resolved as monophyletic only in combined ee of all pie markers (Fig. Sc Morphologically, the Jarava s.s. plus Nassella-Amelichlo E ts short paleas and short funda- Da L E + | Phal EA H Es [| D , Phylogeny I , Pappostipa 189 mi mental cells. These traits are most highly developed in Nassella where species are characterized by having a “ladder” pattern of the lemma epidermal cells, designated by Barkworth (1990) as the “nasselloid type" (short, silicified fundamental cells not alternating with silica cells, or fundamental cells and short cells that lack silica) that are indistinguishable from each other. Another distinct feature of species with nasselloid type lemma anatomy is the develo] t of the crown (a fleshy, ciliate formation) at the apex of the lemma. Species of Jarava s.s. are characterized here as the “achnatheroid type,” with fundamental cells that alter- nate with silica cells of the same form (Fig. 4H), forming a clear “maize” pattern, and with the absence of a lemma crown. Among other achnatheroid traits common for Jarava s.s. species, but not observed in Nassella or Amelichloa, are the protruding and/or in some cases reduced, lemma lobes. Jarava s.s.—Based on our current set of samples, as seen in Figs. 2&3, the Jarava s.s. clade consists of two distinct groups: 1) Jarava ichu group; and 2) J. plumosula group. In the ITS analysis, the Jarava s.s. clade was resolved to be polyphyletic, with three species constituting J. plumosula group forming a clade that is sister group to Nassella-Amelichloa (Fig. 2). In the plastid analysis (Fig. 1) only J. plumosula resolved in a polytomy between the core of Jarava and Nassella-Amelichloa, whereas J. media and J. polyclada were placed just at the base of the clade of Jarava s.s. (core) plus Nassella-Ameichloa complex. In the analysis of the whole molecular dataset, all species of Jarava s.s. were weakly supported as monophyletic clade (Fig. 3). Jarava media, J. plu- mosula, and J. polyclada all share the same habitat and a habit that includes such characters as culms havin g many branches at the upper nodes, and a zoo-anemochoric floret ie ain ge Our combined plastid and ITS molecular analysis (Fig. 3) ts the s O from species formerly recognized as Stipa capilliseta Hitchc. [= S. ada var. capilliseta (Hitchc. ) modi currently treated as a synonym of J. leptostachya in Soreng et al. 2003] and consequently, labeled on the trees as J. leptostachya*. This separation is supported by morphological differences such as, a flexuous awn and sparsely hairy lemma apices in S. capilliseta verses a geniculate awn and pappose lemma apices in J. 1 dama Nassell luding Amelichloa. —According to the latest revision, Nassella is one of the largest genera in the tribe (Barkworth & Torres 2001) with some e 119 spese listed as accepted by Soreng et al. (2003). A number of subgeneric and sectional names were employed in Stipa s.l. by Spegazzini (1901, 1925) and Roig (1964) to recognize the many forms now placed in Nassella. Despite some striking morphological differences among the species, the majority of the species of Nassella share the same "nasselloid" pattern of the floret defined by Barkworth (1990). This pattern includes short paleas, strongly overlapping lemma margins, presence of crown, and a unique lemma epidermal pattern. Our nuclear and combined plastid-nuclear consensus trees yielded a Suae i ie clade with high statistical support (Figs 2 & 3). The era structure within our Nassella-Amelichloa de i represented by two well- supported branch ting groups of species of varying morphological homogeneity. Within Nassella, the upper clade of Figs. 2 & 3 nel N. brachychaetoides, N. brachyphylla, N. caespitosa, N. dasycarpa, N. depauperata, N. inconspicua, N. meyeniana, N. nardoides, N. pubiflora, and N. rupestris) shares similar morphological features as described for Stipa subg. dinis DER by SE SE This group typically has: oblong or lanceolate, ge truncate florets, hairy | h longer at the apex of the lemma), (rarely inconspicuus. and florets with a short callus. The lower clade within Nassella (bise. 2&3) includes Amelichloa (A. caudata and A. clandestina) and taxa attributed to Stipa subg. Stephanostipa Speg. (incl. N. clarazii, N. filiculmis, N. manicata, N. neesiana, N. nidulans, N. pfisteri, and N. rosengurttii), Stipa subg. Nassella (N. trichotoma, N. tenuissima and N. sanluisensis sensu Spegazzini 1901), and Stipa subg. Microstipa (incl. N. filiculmis sensu Spegazzini 1901; and N. tenuissima sensu Roig 1964). In combined plastid DNA-nrDNA and ITS analyses the Amelichloa subclade was resolved as monophyletic (PP-1.00; BF=100) as a sister group to the lower clade of Nassella. The position of the Amelichloa clade near the oe of this WEE Nassella clade was unexpected. However, all species of Nassella-Amelichloa sl elet ] t the end of the ITS-2 region (not coded for this analysis) not found in other taxa of our EEN The position of Amelichloa within Nassella requires study of additional species prior to proposing taxonomic changes. 190 Jo tani i xas 2(1) CONCLUSIONS Jarava s.l. is polyphyletic and species formerly in Stipa subg. Pappostippa are removed from Jarava s.l. and placed in the new genus Pappostipa. This new genus is phylogenetically intermediate between a group of genera designated as the basal lineages in the American Stipeae and a set of genera designated here as the Major American Clade. The Major American Clade includes: Achnatherum of the New World, Jarava s.s., and Nassella (in which the genus Amelichloa is phylogenetically nested). The American basal lineages include a clade of Anatherostipa and Piptochaetium as sister group to the Asian and American genus Ptilagrostis. Phy- logenetically, the North American genus Hesperostipa stands between these American basal lineages and Pappostipa plus the Major American Clade. ACKNOWLEDGMENTS We wish to thank the following organizations or persons: the Fulbright Scholar Program to KR for a re- search visit to the Smithsonian Institution; the Smithsonian Institutions, Restricted Endowment Fund, the Scholarly Studies Program, Research Opportunities, Atherton Seidell Foundation, and Biodiversity Surveys, Inventories Program, and Laboratory of Analytical Biology for financial support and excellent research facilities; Lee Weigt, Jeffrey Hunt, David Erickson, Kenneth Wurdack, and Andrea Ormon for support and consultation; Roser Villatersana, Teresa Garnatje, and Oriane Hidalgo for help in the laboratory; Asuncion Cano Echevarría, Oscar Tovar Serpa, Dorita Susanibar Cruz, Socorro González Elizondo, Nancy Refulio Rodríguez, Fernando Zuloaga, and Stephan Beck for help with our recent field expeditions to Peru, Bolivia, and Argentina; Samuel Pyke for providing samples; Pedro Acevedo Rodríguez for help with the Spanish resumen: Alain Touwaide for help with the Latin diagnosis; and Jeff Saarela and Emmet Judziewicz for providing helpful comments on the manuscript. 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A acd and; 7 Sect. E name section is enin on one ae nes a Ai 3 in Sec GALA LL L l assi t of the alloploid hiefly key feat f leaf sl , habit, lane oO 1 4 + I d al Drak al A al + J1 1 a E + Za E 1 1 mi + a a g Go má partly i Supe f hybrid origin link the basic diploid species of different secti The lamen ofthe 1 1 M s 1 1-5 ] s 1 1 1i 1 H Pi EE EE tioned aboe. I ticulat ti 1 | FM PDA PAR ER O A "- DÉI O ti d on the diploid entities tl lted from early div da Mme in SCH genus iis tlie later-formed b brid 1 Tea A A E lated possible I D 6) ah r 4 JI proposed by Benth (1848) and for Satureja virgini L., the basiony [P th irgini (I ) Robins & Fernald RESUMEN E SEI 1 Lz H el 1 1 Pë? a UE. A aid "P D +] + tf (1848) 1: ] Ji Ls | t 1. Sect. E 2. Sect. uds 3. a Brachystemum, 4. Sect. me 3 Sect. Macrocephalas 6 Seat Ge y 7. Sect. e Cada sección está Lov en una respecte EE EE 3 en la Sec 1 J 1 ] 1 J k * 1 J 1 E + a L L Pu dela Kops Eé fologia de la inf! j ý sti I t bl ] tud ] d > H a 4 e E d hibrid 1Ó ] t d ] e lei hibrid polinloide V ial t I d i hibrido 1 CH L EI EET L 4 D [ar] 1 : Lael Jenlai IJ. JL F : Ta Ta ce dal : Jani J unen I B I principalmente MM EM en la S de uno u l lica decisi bi ias | ] ] t n un complejo genético ous como este, el den de una clasificación m o aa M odd monofiléticas no es BOUE a secciones 4 11 A ] I se basan en q del género, mient L e r más tarde y fol g te int li d lel ] práctico p ible. Se designan lectonipos para las secciones 1 Cl ad DN C A = r E z T 14 A a A Th. Al a y i prog [ ( YE 4 g : é g (L.) Robins. & Fernald A PROPOSED SECTIONAL ARRANGEMENT OF PYCNANTHEMUM SPECIES Up to the time of George Bentham's treatment of Pycnanthemum for the Prodromus of Alphonse de Candolle (1848), two principal species-groups had MR d been recognized, first as the two genera Pycnanthemum and Brachystemum by Michaux (1803) and tl Tullia and Section Bracl | y Bentham (1834). In his more elaborate treatment of 1848, Bentham formally described six sections, to be discussed below. The much later generic monograph by Grant and Epling (1943) did not recognize formal sections, but rather defined two principal groups, termed the Incanum Phylad, of 8 taxa, and the Virginianum Phylad, of 5 taxa The remaining 8 species were irregularly connected to these groups in a chart (their Figure 1) displaying a network of relationships and suggesting multiple transitional forms and gene exchange between taxa. In their analysis, only P. californicum and P. nudum “present no transitional forms with other species.” n our previous papers and unpublished work (Chambers 1961a, b; Chambers & Chambers 1971; Chambers 1993) no formal sections of Pycnanthemum were recognized, but the taxa were arranged in six informal “species groups.” These paralleled the named sections of Bentham but differed in ts, due J. Bot. Res. Inst. Texas 2(1): 193 — 199. 2008 194 t tani i Texas 2( to the addition of SC Se to ee e after his publication, as well as new information gained from tological and hy! ion studies. We went beyond Grant and Epling (1943) in not only recognizing hee cana and “Virginianum” merde but also in organizing the outlying taxa of their relationship network into arbitrary species groups. Our informal classification is given in Table 1, which lists the known somatic chromosome numbers of each species (Chambers & Chambers 1971). While Grant and Epling (1943) spoke of transitional forms between species and the possibility that some species were themselves of hybrid origin, our later work (Chambers 1961a, b; Chambers & Chambers 1971, Chambers 1993) has defined the genus as a polyploid complex, with 9 extant diploid species (Table 1), apparent autopolyploidy in P. albescens, P. muticum, and P. tenuifolium, and 10 taxa that are limited to the tetraploid level. The latter appear, for the most part, to be morphologically EE between known diploids, such that an alloploid origin is strongly suggested. A pan; ploid evolution in some species has been the development of facultative or obli y, da hologically by flow- ers with short filaments and aborted anthers. Where Mese in sse anthered danis of P. vigina, Pp verticillatum, and polyploid P. muticum, seed-set was normal in both bagged and un-bagged inflorescences (Chambers 1961). Plants with aborted anthers also occur in P. tenuifolium, but although apomictic behavior is suspected, it has not yet been checked experimentally (Chambers 1961). Although it would be desirable to divide the genus entirely into monophyletic sections, we believe that the reticulate nature of evolution in Pycnanthemum, involving, as it does, allopolyploidy, a polyphyletic hybrid origin! for several widespread taxa, and the preservation of sterile hybrids via apomixis, precludes such a classification at present. Our approach is to propose a practical infrageneric classification involving certain arbitrary decisions about the placement of species that putatively are derived from allopolyploidy or E £i or DDR Basic to our Meca is that each section contains one or more of the and y OL POly HI fallo or apomicts). By his plat. we have EH to Opa each taxon de suspected hybrid MM into E section contains the diploid(s) it most i This has involved a choice of emphasis on r certain key features, especially those of habit, leaf morphology, anid details of the inflorescence. An example of this decision process is discussed under P. setosum, below.’ Complementing the cl l and hybridization research mentioned above have been two recent studies using molecular approaches. The ¡Hed by Yetter (1989) examined relationships in the Virginianum Group, using 12 enzyme systems to identify synapomorphic alleles as well as alleles unique to particular diploid taxa. The work of Williams (2005) developed a cladogram of relationships among all but one spe- cies of Pycnanthemum, based on sequence data from the ITS region of nuclear ribosomal DNA. (Technical difficulties prevented the inclusion of P. setosum. Nominally no P. clinopodioides was included in her study, but it may have been represented by a population that she classified as P. torreyi) Our propossu or Ree eee takes account of the molecular EE from the above re- search. It emphasi d species, using PEMEX taxon arbitrarily in iation with che parent species that it more o The pamed sections ate their included species are presented below, with diploid species in bold-face. 1. Sect. Pycnanthemum. Tyre: Clinopodium incanum L., typ. cons. = P incanum (L.) Michx. Syn.: Sect. Tulliae Benth. in DC. Prod. 12:187. 1848. Tyre: Ti d ( , designated here, Amer. J. Sci. Arts, Ser. 1, 20:343, t. 5, 1831, = P. pycnanthemoides (Leavenw.) Fernald). Included species. —P. albescens Torr. & A. Gray, P. curvipes (Greene) E. Grant € Epling, P. loomisii Nutt., P. dinopodioides Torr. & A. Gray, P. incanum (L.) Michx., P. pycnanthemoides (Leavenw.) Fernald Jn. nner H I J 1 Lad Ai M " groups hypothesized in our studies. ry: | ee | L ] I s +l + 1. inf | by Chambers and Chambers. Dir p TAB E I a | li tl I I are partly or wholly diploid are in bold-face. Species Group Included species Chromosome numbers (2n) Incanum Group P. albescens 38, 76 P. loomisii 38 P. curvipes 40 P. pycnanthemoides 72 > incanum 75* P clinopodioides 76 P.floridanum 78 Flexuosum Group P. flexuosum 36 P setosum 76 Virginianum Group P. muticum 40, 80, ca. 108 P. tenuifolium 40, 80 P. pilosum 78 P torreyi 80 P verticillatum ca. 76-78 P virginianum 80 Montanum Group P. montanum 40 P beadlei 76 Nudum Group P. nudum 40 Californicum Group P californicum 40 * The reported count of 2n=78 for P incanum (Chambers 1993) was a misprint for 2n=76. 2. Sect. Aristatae Benth. in DC. Prod. 12:186. 1848. Tre: P aristatum Michx. (Lecrorvrz, designated here, Fl. bor amer. 2:8, pl. 33, 1803, = P setosum Nutt.). Included species —P. fl (Walt.) Britton, St & Poggenb. (syn.: P. hyssopifolium Benth.), P. setosum Nutt. 3. Sect. Brachystemum (Michx.) Benth., Labiat. gen. spec. 329. 1834. Tye: Brachystemum muticum Michx. (LECTOTYFE, effectively designated by Grant & Epling, Univ. Calif. Publ. Bot. 20:202. 1943), = P muticum (Michx.) Pers.). Syn.: Sect. Mutica Benth. in DC. Prod. 12:188. 1848. Tree: Brachyst ti Michx. (LECTOTYPE, designated here, Fl. bor.-amer. 2:6, pl. 32. 1803, = P muticum (Michx.) Pers.). Included species. —P. muticum (Michx.) Pers., P. beadlei (Small) Fernald, P. floridanum E. Grant & Epling. 4. Sect. Capitellatae Benth. in DC. Prod. 12:189. 1848 (as Capitellata). Tire: Brachystemum lanceolatum Willd. (LECTOTYPE, designated here, Willdenow, Enum. pl. 623.1809, = P virginianum (L.) Robins. & Fernald). Included species.—P. tenuifolium Schrad., P. pilosum Nutt., P. torreyi Benth., P. verticillatum (Michx.) Pers., P. virginianum (L.) Robins. & Fernald. 5. Sect. Macrocephalae Benth. in DC. Prod. 12:189. 1848 (as "Macrocephala"). Tee: P montanum Michx. Included species.—P. montanum Michx. 6. Sect. Nudae Benth. in DC. Prod. 12:189. 1848 (as “Nuda”). Tres: P nudum Nutt. Included species.—P. nudum Nutt. 7. Sect. Californicae K.L. Chambers & HL Chambers, sect. nov. Te: P. californicum Durand, J. Acad. Nat. Sci. Phila- delphia Ser. II, 3:99. 1855. Included species.—P. californicum Durand. 196 n n H IMA 2(1) Lectotypes are proposed, above, for the sections of Bentham con taining more than one species. As noted, the genus Brachystemum was earlier typified by Grant and Epling (1943) with B. muticum, so Bentham’s Section Mutica becomes superfluous. For Sect. Capitellatae, of the two species cited by Bentham we have elected Mere S EE B. Geh (= P. virginianum) as the type rather than his B. linifolium (= P. tenuifolium), , and Willdenow's d së tion is at variance with the type and descrip- tion of Satureja virginiana L. ET origin of his cited synonym Thymus virginicus L. (1771). Section Pycnanthemum is the only section containing more than one diploid species. Its members are characterized by loose, well-branched flower clusters, both terminal and (often) in the axils of 1—4 bracte- ate leaf pairs below this. Leaves are broad, petiolate, and often canescent on one or both sides, especially distally on the plant. We hypothesize that this section contributed to the origin of intersectional alloploids showing a tendency toward relatively tight but visibly branched flower clusters, for example P. beadlei, P. floridanum, and P. pilosum, the other putative parents having tight capitula. We dd Bentham dd in ice n EE in Sect. Aristatae, in association with its very likely derivative P. I g of 19 IIsand 18 Is at meiosis in triploid backcross hybrid #75, Chambers 1993). The tlie: putative parent of P. setosum, P. muticum, is in Sect. Brachystemum. An alternative would be to create a monotypic section for P. flexuosum and leave P. setosum alone in an alloploid Sect. Aristatae, but this goes against our plan to have each section contain at least one diploid species. Ad- ditionally, it would create a precedent for other alloploid species like P. floridanum, P. beadlei, P. virginianum, et al., to be seg l into monotypic sections of their own. The resemblance of P. setosum to P. flexuosum is expressed in feat puberulence, distinct | reduced calyx pubescence, and calyx teeth acuminate with an aristate tip up to 1 mm long. EM MES in the northern portion of its range, P. setosum often resembles its other putative parent P. muticum in traits of habit (general absence of leafy, sterile side-branches), leaf shape (more broadly lanceolate leaves), and up- per bracteate leaf pair subtending the capitula (reduced, ovate, canescent, and often drooping). Besides its morphological intermediacy, P. setosum has 2n=76, which is the sum of the diploid numbers of the putative parental species. Section Brachystemum, centered on oe P. muticum, also p being included, by choice, with one of their likely diploid ancestors Á MN ri tetraploid P Mala to P. muticum, as is suggested by the DNA data of Williams (2005), implies that the gametic number n-39 in P. floridanum must include a 20-chromosome genome derived from P. muticum. On morphological and cytological grounds, P. floridanum's other diploid parent is probably a species with n=19 in Sect. Pycnan- themum (as predicted by Grant and Epling 1943, p. 210, referring to their Incanum Phylad). One candidate taxon is P. albescens, but this possible PRU ied not yet Hus tested experimentally. In a further ex- ample, P. beadlei, another tetraploid, has traditionally been associa ically with P. montanum (they are sympatric in the southern Appalachians). ete Williams' DNA data from nuclear ribosomal ITS genes (2005) do not support a close phylogenetic relationship between the two. Instead, P. beadlei appears in a clade with P. muticum and some of its other tetraploid derivates. The chromosome numbers of n=38 in the former species and n=20 in the latter suggest that a second genome of 18 occurs in P. beadlei, perhaps from an ancestral diploid in Sect. Pycnanthemum, where x-18 is known (although no extant species have n-18). Artificial hybrids #12 and #15 between P. beadlei and P. muticum (Chambers 1993) showed up to 20 chromosome pairs and 18 singles at first division of meiosis in pollen-mother cells. Pycnanthemum beadlei is therefore assigned to Sect. Brachystemum. Section Capitellatae contains the core of the earlier Kees ee nn we view as comprising P. tenuifolium plus hybrid derivatives of that taxon with ] ne or more t Pycnanthemum tenuifolium is distinguished by its linear leaves, numerous small and compact floral heads, bractlets of the heads acuminate, pungent, and arcuate basally, with stout, emergent midnerve, and a bushy habit due to numerous vegetative branches arising from the stem nodes. In the derived tetraploid species P. virginianum, osum, P. torreyi and P. verticillatum, the leaves are usually lanceolate, the heads somewhat looser (espe- 11 ] hlet the capitula, >J +1 1 Es asas ume Kaesch P. pi ei L ei L if D | H kr ] lat | t f Pycnanthemum 197 cially P. pilosum), the capitular bractlets less pungent (though still with a notably stout midnerve), and the habit frequently less bushy. There is good molecular evidence that the broader leaves, less-branched habit, and non-pungent bractlets came from the second parent of many tetraploid forms, namely P. muticum (Yet- ter 1989; Williams 2005). From his allozyme studies, Yetter concluded: “Tetraploids traditionally labeled Pycnanthemum virginianum and P. pilosum are thought to be allopolyploid derivatives of diploid P. muticum and P. tenuifolium as indicated by the possession of alleles which are unique to the diploids" (1989, p. 80). In the DNA analyses by Williams (2005), samples of the alloploid taxa P. virginianum, P. pilosum, and P. verticillatum, of Sect. Capitellatae, were always associated in her cladograms with P. tenuifolium or P. muticum, not with species from other sections. Additional evidence comes from the artificial diploid hybrid of P. muticum x P. tenuifolium (#84, Chambers 1993). In the morphology of habit, leaves, and floral capitula, this hybrid can easily be classified as P. virginianum (voucher OSC 192224). Our assignment of P. muticum to the separate Sect. Brachystemum recognizes its involvement in other intersectional hybrids in addition to those involving P. tenuifolium. Finally, we differ from Bentham (1848) in placing P. torreyi in the same section as P. tenuifolium rather than with P. muticum, but this is tentative awaiting further study, since DNA evidence suggests that P. torreyi is itself of complex, polyphyletic hybrid origin (Williams 2005). The monotypic sections Macrocephalae, Nudae, and Californicae are not at this point implicated in the origin of any polyploid taxa. However, intersectional hybrids have been produced artificially (Chambers & Chambers 1971; Chambers 1993), and a natural hybrid of P. nudum x P. flexuosum has been described (Chambers & Chambers 1971). Hybrids with mostly sterile pollen have been produced in crosses of P. muticum x P. montanum, P. muticum x P. californicum, P. muticum x P. tenuifolium, P. flexuosum x P. nudum, P. flexuosum x P. tenuifolium, and P. flexuosum x P. loomisii (Chambers 1993). Puzzling int liate forms will no doubt be found in nature involving these and other species, such that species boundaries, especially at the tetraploid level, will be blurred (Grant & Epling 1943, pp. 233—236). Even if sterile, such hybrids may occasionally persist through seed apomixis or vegetative clonal reproduction. Our revised sectional classification of Pycnanthemum is summarized in Table 2. A taxonomic treat- ment of the genus is in preparation for inclusion in a future volume of Flora of North America (Flora of North America Editorial Committee, 1993 onwards). LECTOTYPIFICATION OF PYCNANTHEMUM VIRGINIANUM L. There are three species named by Linnaeus in Species Plantarum (1753) that today are placed in the genus Pycnanthemum. These are Satureja virginiana (p. 567), Nepeta virginica (p. 571), and Clinopodium incanum (p. 588). Clinopodium incanum L., now Pycnanthemum incanum (L.) Michx., is the conserved type of the genus (Rickett & Stafleu 1960; McNeill et al. 2006). It was lectotypified with a Kalm specimen, Herb. Linn. No. 742.4, by Reveal et al. (1987, see Jarvis 2007, p. 428). We would not hazard a guess as to why Linnaeus cited "Habitat in a parean" Hex this SD North Ámerican species The name Sat Linnaeus 1753, p. 567) has since 1908 or before been treated in all North American lies as ER GEN of Pycnanthemum virginianum (L.) Robins. & Fernald, a common and wide- spread taxon in eastern North America. Its lectotypification has been uncertain, in part because some of the pre-Linnaean polynomials cited in the original description are known to refer to the related species, P. tenuifolium Schrad. (Epling 1929; Grant & Epling 1943; Jarvis 2007). Epling (1929) at first suggested that P. virginianum was synonymous with P. linifolium (Willd.) Pursh, a synonym of P. tenuifolium. However, in Grant and Epling (1943) he changed his mind, saying that Linnaeus had in his herbarium a specimen of what we today know as P. virginianum, and because this usage “has now become well established. ...it seems preferable not to make another change” Epling did not formally propose the Linnaean Herbarium specimen as a lectotype, however. This specimen is Herb. Linn. 744.21. It is annotated in Linnaeus’ script "Sat[ureja] 1 virginiana” and reannotated by Smith “Sp. Pl. 1. Pycnanthemum linifolium Sm. in Rees's Cyclop.” It is filed as sheet number 21 under the genus Thymus (Savage 1945). We have ined the excellent image of this sheet available through the Linnaean Herbarium website [| E fal n.a H In LI PR ET STARK! * E 198 Taste 2. Sections of P tt | their included species, as here proposed. Bold-face indicates species that are partly + 1 or wholly diploid. Section Species 1. Pycnanthemum P. albescens P clinopodioides ipes P. loomisii P. pycnanthemoides 2. Aristatae P. flexuosum P.setosum 3. Brachystemum P. beadle P floridanum 4. Capitellatae P. pilosum P torre P. verticillatum P virginianum 5. Macrocephalae 6. Nudae 7. Californicae P. californicum nds nnus org»), and we find that it is undoubtedly a specimen of Pycnanthemum virginianum as that plied in North America for the last 100 years. There are certain details in Linnaeus’ descrip- tion of Satureja virgini that aptly fit this specimen but that are inappropriate for P. tenuifolium (the other species to which “virginiana,” or the variant “virginica,” have been applied). Firstly, Linnaeus described the leaves as "foliis lanceolatis," and in citing his earlier polynomial from Hortus Cliffortianus (Linnaeus 1737, p. 305), he changed the “foliis linearibus acuminatis" from that work to "foliis lanceolatis acuminatis," E better to fit the specimen he had at hand. Secondly, this specimen has tiny stamens and clearly of P. virginianum which is the common type found in the Atlantic Coast states s (Chambers 19614) E non- ain stamens are well described by Linnaeus in his observation, corolla is effoetis, ut fere posset genere proprio tradi”? This differs from P. tenuifolium, which in this same region ieu uses sexually and has exserted stamens with functional anthers (Chambers 19613). Therefore, based on tl iderations, we here designate Herb. Linn. 744.21 (LINN) as the lectotype of Satureja virginiana L., Sp. Pl. 1:567. 1753. The difficult case of the name Nepeta virginica L., which also lacks lectotypification, is under study by James Reveal and Charlie Jarvis (pers. comm.), and their recommendation will be published in due course. The epithet "virginica" is not available in Pycnanthemum, due to the preexisting P. virginicum (L.) Pers. (1806) based on Thymus virginicus L. (1771). “Stamina in f ACKNOWLEDGMENTS We thank James L. Reveal and Charlie Jarvis for advice on nomenclatural matters, and Philip Cantino for his helpful review comments. Mary Stiffler, Deborah Carroll and Gina Dougias gave valuable assistance in our bibliographic research. "m L IrL I St £ Ze eil Lie ag) | lat | t f Pycnanthemum 199 REFERENCES CHAMBERS, H.L. 1961a. A cytotaxonomic study of the genus Pycnanthemum (Labiatae). Ph.D. thesis, Yale University, New Haven, CT. 233 pp. CHAMBERS, H.L. 1961b. Chromosome numbers and breeding systems in Pycnanthemum (Labiatae). Brittonia 13:116-128. CHAMBERS, H.L. 1993. Chromosome survey and analysis of artificial hybrids in Pycnanthemum. Castanea 58:197-208. CHAMBERS, H.L. and K.L. Chambers. 1971. Artificial and natural hybrids in Pycnanthemum (Labiatae). Brittonia BENTHAM, G. 1834. Pycnanthemum. |n: Labiatarum genera et species. James Ridgeway and Sons, London. Pp. BENTHAM, G. 1848. Pycnanthemum. In: A.L.P. de Candolle, Prodromus systematis naturalis regni vegetabilis 12:186-190. Victoris Masoon, Paris. EPLING, C. 1929. Notes on the Linnean types of American Labiatae. J. Bot. 67:1-12. FLORA Or NORTH America Eprrorial COMMITTEE. 1993 onwards. Flora of North America north of Mexico. Oxford Uni- versity Press, New York. GRANT, E. and C. EPLING, 1943. A study of Pycnanthemum (Labiatae). Univ. Calif. Publ. Bot. 20:195—240. Janvis, C. 2007. Order out of chaos. Linnaean plant names and their types. The Linnaean Society of London. LINNAEUS, C. 1737. Hortus Cliffortianus. Reprint 1968, J. Cramer Verlag, 3301 Lehre. 501 pp, tab. i-xxxvi. LINNAEUS, C. 1753. Satureja virginiana. In: Species plantarum. Laurentii Salvii, Holmiae. P. 567. LINNAEUS, C. 1771. Thymus virginicus. In: Mantissa plantarum altera. Laurentii Salvii, Holmiae. P. 409. McNiLL, J., F.R. Barrie, H.M. Bunoer, V. DemouLin, D.L. HAwKsworTH, K. MarHoLD, DH NicoLson, J. PRADO, PC. Sit vA, J.E. SKOG, J.H. Wiersema, and N.J. TuRLAND. 2006. International code of botanical nomenclature (Vienna code). Regnum Veg. A.R.G. Gantner Verlag, Ruggell, Liechtenstein. 146:i-xviii, 1-568. MicHAUx, A. 1803. Brachystemum and Pycnanthemum. In: Flora boreali-americana. Fratres Levrault, Paris. 2:5-8, t. 31-34 PERSOON, C. H. 1806. Pycnanthemum. In: Synopsis plantarum. Parisiis Lutetiorum, Paris. 2(1):128-129. PursH, F. 1814. Pycnanthemum. In: Flora Americae septentrionalis. White, Cochrane, and Co., London. 2:409-410. REVEAL, J.L, CH Broome, M.L. Brown, and G.F. Frick. 1987. On the identities of Maryland plants mentioned in the first two editions of Linnaeus’ Species plantarum. Huntia 7:209-245 Rickerr, H.W. and FA. Starieu. 1960. Nomina generica conservanda et rejicienda spermatophytorum IV. Taxon 9:07-86. ROBINSON, B.L. and M.L. FernaLD. 1908. Pycnanthemum. In: Gray's new manual of botany. 7th. ed. American Book Co., New York. Pp. 707-708. SAVAGE, S. 1945. A catalogue EES H EE iet ned London. WiLLDENOW, C.L. 1809. Bract io plantarum i regii botanici berolinensis. Taberna Libraria Scholae Realis, Berlin. P. 623. WiLLIAMS, R. 2005. The phylogeny of Pycnanthemum: hybridization and polyploidy in a taxonomically confusing genus. Herbarist 71:35-41. YETER, T.C. 1989. Systematic and phylogenetic relationships in the virginianum species complex of the genus Pycnanthemum (Labiatae). Ph.D. thesis, Miami University, Oxford, OH. Pp. 1—122 200 | t tani i Texas 2( BOOK REVIEWS RICHARD B. Primack. 2002. Essentials of Conservation Biology: Third Edition. (ISBN 0-87893-719-6, hbk.). Sinauer Associates, Inc., 23 Plumtree Road, Sunderland, Massachusetts 01375, U.S.A. (Orders: www. sinauer.com, ordersOsinauer.com, 1-413-549-4300, 1-413-549-1118 fax). $69.95, 698 pp., numerous figures, graphs, images, tables, 7.25" x 9.5”. RicHARD B. PRIMACK. 2006. Essentials of Conservation Biology: Fourth Edition. (ISBN 0-87895-720-X, hbk.). Sinauer Associates, Inc., 23 Plumtree Road, Sunderland, Massachusetts 01375, U.S.A. (Orders: www.sinauer.com, orders@sinauer.com, 1-413-549-4300, 1-413-549-1118 fax) $82.95, 585 pp., nu- merous deem dd images, tables, 8.75" x 11.25". Tr ek 2 r D:n] rard J ath — ties M La: =. Sur TI Al E S, | 1 J nf 1 : +1 The books, VO L DIOL (o ara + catons, E Y > EE d oui + Voz aus 1 1: . x 1: Co: | fi eld eraphs new DU DL 7 i 1 Rs SEN ree DEE ose servation biology, and r T1 f. : 1:[F n +1 E ici Ee nisi e ¿kz ul +1 +; ai 1, Ass ass LLULLU mit only A LN- YY LLLALLOS by ling tl ] it] inf | ithin the field. In the mus Edition, the author i d a demia introduction to A ; x 1 probl i iul Ausus sët or are EA CIO ipii qa pee EN) +l 1 1 114] ] f. T 1 E ls. We Ae +1 el ] 1 explain how the field has evolved in 4 Weiter time. This edition al pus ni a glossary e an instructors Resource CD. Tr 14 1 do | : a FRENIS 1 hol Mée 1 » 1 : " ffinldo ta hal 14 ; h] d create so lutions eps ain size of the Third Edition, ink realized b E new ee dd size of the Fourth Edition was A in ge to include En Ee "enn Rue di a any j te i dd d in de relativ ely newer field and their possible DE — Keri McNew, MS Sm Project Manager, Botanical R h Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S.A. GLENN KEATOR AND ÁLRIE MIDDLEBROOK. 2007. Designing California Native Gardens: the Plant Community Approach to Artful, Ecological Gardens. (ISBN 978-0-520-23978-4 hbk.; 978-0-520-25110-6 pbk.). University of California Press, 2120 Berkeley Way, Berkeley, California 94704, U.S.A. (Orders: www.ucpress.edu, California Princeton Fulfillment Center, 1445 Lower Ferry Rd., Ewing, NJ 08618, U.S.A., 1-609-883-1759; 1-609-883-7413 fax). $65.00 hbk., $27.50 pbk., 352 pp., 364 illustrations, it " 7" x 9 3/4". This book should b ] i to California gardeners and land ji hi whether i pl g gardens or renov ovating Tails BA | LI 4 1 = Ê ] 1 4 iq f. J existing ones. Itis also a practical Ge g I pali mountains, tl i hallen oi TT S aa Nui sr e goes repe E e [-] e [e] iu D o S Es 11 ] 1 x sp c | eT Cf 1 1 : A H Lil: and/or Gees in using ¡Her In e to o the O of individual peus there are ae plans, EY ibi drawings, and photographs of completed gardens. T g g5, including green roofs. For each of the ae communities, , there is a “places to go” SERO hich includes state parks and nes Bie scenic areas, and a few DNE gardens I y be ol d in natural habitat f. RI il; 1 e ] r A $c ties, and a losa of terms.—Joann Karges, Texas Christian vera Libras (retired), Fort Worth, Texas US. A. J. Bot. Res. Inst. Texas 2(1): 200. 2008 NOMENCLATURAL NOTES IN NORTH AMERICAN POTENTILLA (ROSACEAE) Barbara Ertter Curator an Western North American Flora == Jepson Herbaria EN sity of California Berkeley, Es ifornia 94720-2465, USA Research Associate du ge Plains Herbarium State University Boise, in 83725-1515, U.S.A. ertteroberkeley.edu ABSTRACT New Manual of Botany of the Central Rocky Mountains, by J.M. Coulter & A. Nelson (1909) is an earlier source than previously noted for Potentilla sections Supinae, Concinnae, Subjugae, Niveae, and Multifidae. Sections Rubricaules and Leucophylleae are also established, with the latter de Mi sect. Hippianae (Rydb.) Ertter & Reveal. The segregates from P. pensylvanica sensu lato that will be used in a forthcoming volume of Flora of North Ámerica are briefly summarized, with Potentilla jepsonii Ertter established as a new name for P. pensylvanica var. ovium Jepson. The distinction and identity of P. nivea var. dissecta S. Watson and P. concinna var. divisa Rydb. are clarified. Complete lectotypifications are provided for P. litoralis Rydb., P. pensylvanica var. ovium, P. nivea var. dissecta, P. concinna var. divisa, and P. pseudosericea Rydb. RESUMEN New Manual d inii aja del Central id Mountains, jk J.M. Coulter & A. Nelson (1909) es una ag ias a ue tenidas en cuenta nnae, Subjugae, Niveae, y Multifidae. 1 ji hyll ie pi (Rydb.) Ertter & Reveal. I dos de P. lato que serán senelp volumen de la Flora of North America se resumen brevemente, con Potentilla ee Ertter establecida como L alr A 1 A + 1 el se establecen, con | e un nombre nuevo para P. Se var. ovium Jepson: Se clarifican la distinción e identidad de P. nivea var. dissecta S. Watson y P. concinna var. divisa Rydb. Se aporta I P. litoralis Rydb., P. pensylvanica var. ovium, P. nivea var. dissecta, P. concinna Y var. divisa, y P. pseudosericea Rydb. For a forthcoming volume of Flora of North America (FNA), several nomenclatural notes, lectotypifications, and a new name are required for Potentilla (Rosaceae); these are provided herewith. I. Some overlooked sectional names in Potentilla Subsequent to the publication of Potentilla sect. Hippianae (Rydb.) Ertter & Reveal (2007), a previously over- looked source of validly published sectional names has been encountered that supersedes sect. Hippianae and several other sections, at least as far as author attribution. In the key to Potentilla on page 255 of New Manual of Botany of the Central Rocky Mountains (Coulter & Nelson 1909), Nelson adopts several of Rydberg's unranked "groups" and explicitly refers to them as sections. The nomenclatural results are as follows: ge SEH Supinae (Lehm.) A Nelson in J.M. Coult. & A. Nelson, New Man. Bot. Centr. Rocky Mt. Basionym: Potentilla series Supinae Lehm., Revis. Potentill. 8, 191. 1856.—Antedates Johnstons (Phytologia 57:297. SE Em to establish the sectional name, but still later than heterotypic P sect. Rivales Poeverl. in Asch. & Graebn. (Syn. Mit- teleur. Fl. 6(1):669. 1904). o sect. mei (Rydb.) A. Nelson in J.M. Coult. & A. Nelson, New Man. Bot. Centr. Rocky Mt. Made BasionyM: Potentilla [unranked] Concinnae - Bull. Torrey Bot. Club 23:431. 1896 — Antedates Johnston's (Phy- to s 57: 985) effort to establish the sectional nam B E Subjugae ae EZ Nelson in J. M. Coult. & A. Nelson, New Man. Bot. Centr. Rocky Mt. 1] Subjugae Rydb., Bull. Torrey Bot. Club 23:397. 1896.—Antedates Johnston's (Phytologia 51:299. SE effort to establish ili Seet name. J. Bot. Res. Inst. Texas 2(1): 201 — 205. 2008 202 [| [| EA naa D Inm L | PE f Texas 2(1) Porn sect. Niveae diis ) A Nelson ing. M. Coult. & A. Nelson, New Man. Bot. Centr. Rocky Mt. 255. ill Bull. Torrey Bot. Club 23:301. 1896.—Antedates Juzepczuk’s (Fl. SSSR 10:133. trib. Niveae Lehm. (Revis. Potentill. 8, 163. 1856) is an invalid name (Art. md Gr fth tional ranl n McNeill et al. 2006). Potentilla sect. Multifidae (Rydb.) A. Nelson in J.M. Coult. & A. Nelson, New Man. Bot. Centr. Rocky Mt. O. Basionym: Potentilla [unranked] Multifidae Rydb., Bull. Torrey Bot. Club 23:262. 1896.—Antedates de s (Fl. SSSR 10:113. 1941) use of the name at sectional rank, but still later than heterotypic P sect. Pensylvanicae Poever Graebn. (Syn. Mitteleur. Fl. 9(1):669. 1904). Potentilla trib. Multifidae Lehm. (Revis. Potentill. 4, 26. 1856) is an Gren name P 33.9 in McNeill et al SE ges sect. kal rican es (Rydb.) A. Nelson in J.M. Coult. & A. Nelson, New Man. Bot. Centr. Rocky Mt. . 1909. Basion: al funranked] Rubricaules Rydb., Mem. Dept. Bot. Columbia Coll. 2:30. 1898.—Not previously est ES shed asa s Potentilla sect. pa A (Rydb.) A. Nelson in J.M. Coult. & A. Nelson, New Man. Bot. Centr. Rocky Mt. 255. 1909. Basionym: Potentilla [unranked] Leucophyllae Rydb., Mem. Dept. Bot. Columbia Coll. 2:31. 1898.—Antedates homotypic sect. Hippianae (Rydb.) Ertter & Reveal (Novon 17:317. 20 II. Segregates from Potentilla pensylvanica L. sensu lato Several entities frequently included within a broadly defined Potentilla pensylvanica L. (e.g., Hitchcock & Cronquist 1961, McGregor 1986) will be recognized as distinct species in FNA. The critical nomenclature is briefly summarized here, along with a new name and a lectotypification. Potentilla pensylvanica L., Mant. Pl. 76. 1767.1 ification by Fernald (1935: 288) les that of Soják (1987: 298) Excluded from synonymy is P SECH anica var. strigosa Durch, since the | typic P stri (Pursh) Tratt. has been es Ed Soják (1987: 293) on a Pall Sib Sojál identified as th hy 'brid between P rictly Eurasian P sanguisorba Wild. In D.FK. S Potentilla jest mt a Rydb., N. deus Ns E a 1908. An earlier name for P finitima Kohli & Packer (Canad. J. Bot. :714. 1976), as first noted by Soják (19 Potentilla litoralis Rydb., Bull. Torrey Bot ub 23:264. 1896. Tree: CANADA. Prov. QUEBEC. Ge ES sta, mie des Monts, 10:0 iod , J.A. ind s.n. pads ae 2 E In 1988, Soják annotated but ly publish lude P virgulata A. Nelson (Bull. Torrey leg. (Autik. Bot.:164 1840. non Fisch ex Ser. in A.P de Candolle, Prodr. 2:581 vd Bot Club 27:265. 1900) 1P ta Raf., nom. i Er 1825). The earlier name e P diseña Pursh (Fl. Amer. Sept. 1:355. 1813) is probably also a synonym but is proposed for rejection (Ertter et al. 2008) since the name has a long history of application to unrelated species. The homotypic names P pensylvanica var. ai cina de S. Watson, P Geen E ex Ho var. P aii o ) pe Se Packer, and P glabrella Rydb. have bee rm, P sericea L. var. glabrata reum. bel I lated ies P ovina J.M Macoun, as noted by nd (1996: 1 20). Potentilla ¡ jepsonii Ertter, nom. nov. Basionym: P e dp var. ovium Deomm El. cM 2:184. 1936; P litoralis var. ovium (Jepson) Soják, pe 16:49. 2006. Tue vee ovina” ic I n with P ovina, which has an overlapping range. O t I f the tyr tion (Jepson 7307), JEPS 2814 is here designated the wee as the dn itl bund I keted notes. This sheet has been annotated by E. Crum as type of an hed v: using the a eee Potentilla Þipinmatiñda EEN ex Hook., Fl. Bor.-Amer. 1:188. 1832. Soják (2008) | P bipi against two older names: P normalis Bess. ex See (Syst. Veg. ed. 16, 4, Cur. pum “199. 1827) loa Homem. a (Bot. Reg. 17:1412. 1831). RE bimundorum Soják, Cas. Nár. Mus., Odd. Prir. 141:195. 1974. Soják (1974) described P bimundorum to accommodate all New World and some Old World plants previously placed in P multifida L., which as now circumscribed is confined to the Old World. III. Lectotypifications and identities of Potentilla nivea var. dissecta and P. concinna var. divisa Potentilla nivea var. dissecta S. Watson (Proc. Amer. Acad. Arts 8:559. 1873) was described on the basis of four syntypes: “in the Rocky Mts. of British America and Montana (368 Drummond; Douglas; Howard); and in the Uintas, Utah (335 Watson, in part)” When Rydberg (1896b: 431) proposed "P. concinna divisa n. v." he cited P. nivea var. dissecta “at least in part" in synonymy and included the Douglas (“Rocky Mountains”) and Howard (*Montana") syntypes of P. nivea var. dissecta among the specimens representing his concept of P. concinna var. divisa. The other collections cited by Rydberg were "South Dakota: Jenney. 1875; W.H. Forwood, 1887; P.A. Rydberg, nos. 672 and 673. 1892. Assiniboia: John Macoun, no. 10,468, 1895.” Ertter, Al | p [| 4 H AS XL. A H Dus PES f | 203 Rydberg’s citation of Potentilla nivea var. dissecta in synonymy has led to the interpretation that the two names are nomenclatural synonyms, with the epithet divisa superfluous at the varietal level (e.g., Boivin 1952, Hitchcock & Cronquist 1961). This interpretation is not supported, however, by Rydberg’s use of “in part” and his explicit citation of some of Watson’s syntypes to the exclusion of others. Rydberg (1896b) preceded the citation of specimens with the statement that “Dr. Watson included the three first specimens [actually only two, those by Douglas and Howard] cited below in his P. nivea dissecta, but in every respect they are much nearer P. concinna than P. nivea." As proof that Rydberg had two distinct entities in mind, he (1896a: 303) had not only previously cited P. nivea var. dissecta in the synonymy of P. nivea var. quinquefolia Rydb., but in his subsequent 1898 monograph of North American Potentilleae, he cited P. nivea var. dissecta “in part" in the synonymy of both P. nivea var. altaica (Bunge) Rydb. and P. concinna var. divisa. The Drum- mond and Watson syntypes of P. nivea var. dissecta, which were excluded from P. concinna var. divisa, were instead listed among the specimens seen of P. nivea var. altaica, preceded by the statement that *Watson's specimens agree exactly with the figure of P. altiaca [sic] in Ledebour's Illustrations, and this name is the only one available . . ." (Rydberg 1898: 87). This dual citation was repeated in Rydberg's 1908 revision, in which P. divisa (Rydb.) Rydb. was raised to a species and P. nipharga Rydb. replaced P. nivea var. altaica. With the distinction of Potentilla nivea var. dissecta and P. concinna var. divisa established, lectotypifica- tion and identity need to be determined for both entities. Rydberg's removal of the Douglas and Howard syntypes from P. nivea var. dissecta leaves only Drummond 368 and Watson 335 as candidates. Soják has an- notated the NY sheet of Drummond 368 as lectotype of P. nivea var. dissecta; this choice is herewith honored and formalized. The identity of the single small, one-flowered plant is unfortunately ambiguous, with no precise locality to aid identification; because of the subpalmate/subpinnate leaves and short columnar style (ca 1 mm), it is tentatively identified as P. saximontana Rydb. Of the syntypes of Potentilla concinna var. divisa, first stage lectotypification has been already been effec- tively accomplished by Holmgren's (1997: 100) st t that Douglas's coll from the Rocky Mountains was designated lectotype by Rydberg (N. Amer. Fl. 22:330. 1908). In that Rydberg only indicated that the Rocky Mountains were the type locality, without actually citing a specific collection, Holmgren's statement serves as an inadvertent lectotypification, without designating which herbarium houses the type. This would be GH 19535, annotated as holotype by B.C. Johnston and cited as such in his (1980:165) unpublished dis- sertation. Foto nivea L. var. dissecta S. Watson, Proc. Amer. Acad. Arts 8:550. 1873, not P dissecta Pursh. Ter: ocky Mountains of British America,” Drummond 368 (Lecroryre, designated here: NY). - Potentilla saximontana Rydb. Potentilla concinna Richardson var. divisa Rydb., Bull. Torrey Bot. Club 23:431. 1896. Tr: Rocky Mountains, Douglas s.n. (Lecrorvre, 1* stage designated by N. Holmgren, Intermount. Fl. 32:100. 1997; second stage designated here: GH 19535). = Potentilla concinna var. divisa IV. Lectotypification of Potentilla pseudosericea Rydb. Potentilla pseudosericea Rydb. (Mem. Dept. Bot. Columbia Coll. 2:98. 1898) was described on the basis of three syntypes: “Nevada: Shockley, No. 592, 1888. Rocky Mountains: Nuttall; Fremont, No. 218, 1845-7.” Ás noted by Jepson (1936: 184), Shockley 592 is actually from the White Mountains of Mono Co., California, not Nevada. Of the two Shockley collections in GH annotated by Rydberg as "P pseudosericea n. sp.” 19583 is labeled in S. Watson's handwriting as originating in Esmeralda Co., Nevada, but with the caveat “I think—ticket lost.” The other (GH 19584) bears an original label on which a printed “Esmeralda Co., W. Nevada” is supplanted by Shockley's handwritten “White Mts., Mono Co., Calif., Aug. 19, 1888, W.H. Shockley ae ous that the Nevada citation was indeed erroneous. The Shockl ey ies to which tl [ traditionally applied, endemic to the east-central oia of Coa and possibly Nevada (unconfirmed). In contrast, the Nuttall 204 J ourna | fall Doa H In LI dedo nd £T. "€ 2(1) and Frémont syntypes belong to an unrelated species, P. bipinnatifida Do ex Eros Rae (1898) also cited Nuttall's manuscript name “Potentilla holosericea" in synonymy y (Rydberg 1908: 348) specified “Rocky Mountains” as the type locality. This led B.C. Johnston to EE Nuttall's specimen at NY as lectotype, and to annotate Shockley's “Nevada” syntype (GH 19583) as an unpublished variety of P. pensylvanica honoring the collector. However, citation of a locality alone is insufficient for lectotypification (Art. 37.3, Note 2, McNeill et al. 2006), so that effective lectotypification begins with W. Jepson's (1936: 184) citation of “White Mts., Mono Co., Cal., Shockley 592” as type locality. To complete the process, GH 19583 is herewith formally designated the lectotype of P. pseudosericea. Potentilla pseudosericea Rydb., Mem. Dept. Bot. Columbia Coll. 2:98. 1898. Tree: “NEVADA. Fauna na Co.” [actu- ally CALIFORNIA. Mono Co.: White Mountains]: 19 Aug 1888, WH. Shockley 592 (Lecrorvre, 1* stage designated by W. Jepson, Fl. Calif. 2:184; second stage designated here: GH 195831; isoLecrorvres: GH 19584, JEPS, UC ACKNOWLEDGMENTS Special acknowledgments are due the herbaria who have hosted me or provided loans used during the extended period of preparing the treatment for FNA (ALA, BM, BRY, CAS/DS, E, F, GH, K, LE, MICH, MO, NY, PR, RM, SRP, UC/JEPS, US, UTC, WIS, WTU), with special thanks to the curatorial staff of GH, NY, and US for locating critical type specimens. The assistance and advice of Kanchi Gandhi, Jim Reveal, and John McNeill were essential in unraveling nomenclatural snarls. Jiri Sojak and John Packer generously shared their expertise on Potentilla, facilitated by Jan Kirschner, Adolf Ceska, and J. Zazvorka. I am also indebted to Jim Reveal, Reidar Elven, and Dave Murray, my collaborators on FNA Potentilla, who have influenced the conclusions summarized here and served as sharp-eyed reviewers. Support from the Lawrence R. Heckard Fund of the Jepson Herbarium is gratefully acknowledged. REFERENCES Bom, B. 1952. Pugillus Potentillarum. Phytologia 4:89-93. CouLTER JM. and A. NeLson. 1909. New manual of botany of the central Rocky Mountains. American Book Co, New York. ERTER. B. and J.L. Reveat. 2007. New sections, combinations, and varieties in Rosaceae, Potentilleae. Novon 173 15-52). Fern B., R. Even, D. Murray, and J.L. ReveaL. 2008. Proposal to reject the name Potentilla dissecta Pursh (Rosaceae). Taxon 57(3): in press. FernaLo, M.L. 1935. Critical plants of the Upper Great Lakes region of Ontario and Michigan [p.p.]. Rhodora 37:272-301. Hitchcock, C.L. and A. Cronauist. 1961. Rosaceae. Vasc. Pl. Pacific NorthW. 3:89-194. Hoimaren, N.H. 1997. Rosaceae. Intermount. Fl. 3A:64-158. Jepson, W.L. 1936. A flora of California. Vol. Il. Capparidaceae to Cornaceae. Associated Students Store, University of California, Berkeley. JoHNsTON, B.C. 1980. Studies of population variability leading to a new classification of Potentilla sect. Multijugae (Rosaceae). Doctoral dissertation, University of Colorado, Boulder. McGrecor, R. M. 1986. Rosaceae. In: Great Plains Flora Association (eds.), Flora of the Great Plains. University Press of Kansas. Lawrence. Pp. 364—406. McNuLL, J, ER. Barrie, H.M. Buroet, V. Demoutin, D.L. HAwkswonrH, K. MARHOLD, H.H. NICOLSON, J. Prano, PC. Siva, J.E. Skog, J.H. Wiersema and N.J. Turano (eds.). 2006. International code of botanical nomenclature (Vienna Code) adopted by the Seventeenth Internationa! Botanical Congress Vienna, Austria, July 2005. Gantner Verlag, Ruggell, Liechtenstein RypeerG, PA. 1896a. Notes on Potentilla.— Il. Bull. Torrey Bot. Club 23:301-306. Ryppers, PA. 1896b. Notes on Potentilla —V. Bull. Torrey Bot. Club 23:429-435. RYDBERG, PA. 1898. A monograph of the North American Potentilleae. Mem. Dept. Bot. Columbia Coll. 2:1-223 + 112 plates. Ertter, N 205 RYDBERG, PA. 1908. Potentilla. N. Amer. Fl. 22(4):293-352. SOJÁK, J. 1974. Potentilla bimundorum sp. n., nový druh seckce Multifidae. Cas. Nar. Mus., Odd. Přir. 141(3-4): 195-196. SOJAK, J. 1987. Notes on Potentilla. V. Potentilla pensylvanica group in the Old World. Preslia 59:289-305. SOJÁK, J. 1994. Notes on Potentilla (Rosaceae). X. The section Dumosae. XI. The P microphylla and P. steno- phylia groups (sect. Pentaphylloides). XII. Key to the taxa of P sect. Pentaphylloides (Anserina). Bot. Jarhb. Syst. SOJAK, J. 1996. Notes on Potentilla (Rosaceae). XIV. Type specimens in the Lehmann herbarium. Preslia 68:97- 124. SOJAK, J. 2008. Proposais to conserve the name Potentilla bipinnatifida against P normalis and P missourica and the name P stolonifera t P sprengeliana (Rosaceae). Taxon ined. Dei n A Dat iral D h Institut f Texas 2(1) 206 BOOK REVIEWS PauL MARTIN BROWN. 2005. Wild Orchids of Florida. (ISBN 0-8130-2933-3, flexbound). University Press of Florida, 15 Northwest 15* Street, Gainesville, Florida 32611-2079, U.S.A. (Orders: www.upf.com, 1-800-226-3822, 1-800-680-1955 fax). $24.95, 409 pp., numerous line drawings, maps, color images, bs checklist, 6.25" x 9.25". LD 1 AN 1 ] o CALI 4 E lc 1 T Fat Le) Co Is. nt nL tion to Orchids; Which Se de RUE Part Two: Wild Ge of Florida—Native and: Natera deca Gre ond dee SE Introduced, Escaped, and W Literature dd n | Taxa, deeg and Additions to A oe oe of eee Synonyms and Misapplied Names d Hunting in Florida; Pl ty Lines; The Southeastern Atlantic iud Gulf Coastal Plains; Using Luer's Native Orchids of Florida (1972) in the New Century: MM Corrections, and Comments); Glossary; Selected Bibliography; Photo Credits; Index. Wild Mebide af Blavida fee lec xu Ecce: "EN EE MET x ER NEUE. +: ee This book, , ] p o how to use a key, d + J t 323 | ful St ] 1 f EE I ani this book to = a i source for any amateur or orchid expert. de quo; sia both common e gege names in P +1 J LN isi r ree I E EE P e | 1 1 A 1 1 fl 1 a: 4 1 1 as 1 1 ] k | acr. "T T 1 r 1 I CO enecies and a di 1 Inded +] tł 1 ] T The! Jz’ J +] 1 dia] ARS E, f r r Lei taking out in the field or anis on a idi I think the bas , ] d by th hor's incl fd pti ral species that have ned HOM to Florida. d detailed checklist, inf t ] to look, a glossary — Keri McNew, MS ELO) Project Manager, Botanical R h Institut f Texas, 500 , a Fast 4th Street, Fort Worth, Texas 76102-4025, U.S.A. Douc WELSH. 2007. Doug Welsh's Texas Garden Almanac. (ISBN 1-59544-619-3, flexbound with flaps). Texas A&M University Press, 4354 TAMU, College Station, Texas 77843-43 54, U.S.A. (Orders: www. tamu.edu/upress, 1-800-826-8911). $24.95, 492 pp., color maps and images, b/w drawings, index, FA x 10" Contents.—Select Plant Lists; Almanac at a Glance; A » ae FERRE p Spring; 2) — Hints of Spring, Reminders of Winter MEA March: Spring—Get Busy; 4) April Full St ) June: Welcome to Texas Heat; 7) July: D fS 8) August: Hot, Hot, Hot, indi Dry Too! 9) September Beete bd Mie 1 nu o No Need to Stop Cu 1D November: Down Time, Not Exactly; 12) December: D g ources; Doug Mise as Garden Al j y-to-read gardening guide especially for gardening in Texas. Each monthly chapter includes nad: p important garden design and that lated, but not exclusive to that month. Major discussion topics include: 1 E 1 LI 1 ] | J ent r] 41 1 J s 1 1 ] Ea ler for diff ituati d of different col 1 plant; pruning basi d ] Chapters also include a few pages Stas : nm P 4 da z E 4 +if,,] Ad of d tips," imp E g healthy d md Wel sh? Tewrae Cardon ] T; } kf Texas deci A Luckeydoo, Botanical Reseaich Zeg of Texas, 500 Fast 4th MR Fort Worth, Texas 76102- 4025, U.S.A. J. Bot. Res. Inst. Texas 2(1): 206. 2008 A SYNOPSIS OF THE MEXICAN AND CENTRAL AMERICAN SPECIES OF VACCINIUM (ERICACEAE) Robert L. Wilbur James L. Luteyn — of icd Institute of Systematic Botany Duke University The York Botanical Garden Durham, North cid 27708, U.S.A. Bronx, ie York 10458-5126, U.S.A. rwilbur@duke.edu jluteyn@nybg.org ABSTRACT Js TE A us - 1 1 1 1 1 4 felon 97 species of V M o America , together A with Gett ahs my and a citation of presente specimens. Two : are T and illustrated here as new: Vaccinium campanense Wilbur & Luteyn and V. chihuahuense Wilbur & Lute RESUMEN Se presenta una sinopsi ] lescripci de las 27 den e Vaccinium que se tran en México y América Central, también se aa su sinonimia y it les. Se describen y se ilustran aquí dos especies como nuevas: Vaccinium campanense Wilbur & ius y v chihuahuense Wilbur & Luteyn. INLRODUCLION The representatives of the genus Vaccinium in Mexico and Central America were most recently treated by Sleumer (1936) in a study with the deceptively modest title of “Die Arten der Gattung Vaccinium L. in Zentral- und Südamerika." Sleumer' account actually covered the species of both the West Indies and Mexico as well as those occurring in Central and South America. Sleumer recognized 30 species in his treatment, but of these only 12 were known from Mexico and Central America. The present synopsis includes 27 species from Mexico and Central America, which gives some indication of the rich yield resulting from the more intensive field work that has taken place within the region in the past seven decades. More than half of the Mexican and Central American species of Vaccinium have been described in the most recent three decades, while only four species (or about 15%) were named in the first seventy-five years of the twentieth century and nine ML (or 3396) during the nineteenth FERE thing should be said about the generic concept adopted in the present study. The generic limits are ge broad m inclusive like that of ene (1936) in e treatments of the West Indian, Mexican, Central American, and South American members of the genus, as well as his survey of the genus Vaccinium (1941) throughout its total range. In the latter paper Sleumer recognized 33 sections in the genus Vaccinium, which clearly demonS Tate great EE EE E Gre conservative bid of Vaccinium s.l. accepted À Flora (1933), the senior author was exposed to a much narrower concept ol genera. In the case of vaccinoioid genera such as Batodendron Nutt., Polycodium Raf., Herpothamnus Small, Oxycoccus Hill, Hugeria Small, and Cyanococcus Rydb., they still appear to demark natural groups biologically meanmglul as Ja than E into the vast diversity of the genus Vaccinium s.l. However, traditional generic were recently followed by the authors (Luteyn & Wilbur 2005). Molecular data is sparse at SE but it is Steeg to be helptul in sorting out some of the relationships within the tribe Vaccinieae (see Kron et al. 2002). Molecular evidence mE po iode et al. (2002) and Powell and Kron (2003) strongly suggests that the genus Vaccinium is pol tural; vander Kloet (2004) and Stevens et al. (2004) are also of this opinion. Kron et "E (2002) concluded, however, that a much more thorough molecular survey must be completed before any formal nomenclatural changes could be made. Their prognosis for Vaccinium is ominous for those who were content with the heterogenous genus of 33 or so sections. Their molecular survey so far suggests that J. Bot. Res, Inst. Texas 2(1): 207 — 241. 2008 208 t tani ti f Texas 2( “Vaccinium will need to be enlarged to include all species currently recognized in the tribe Vaccinieae, or to be broken up" (Kron et al. 2002). Their first alternative would seem to be biologically unsound, because the resulting "Vaccinium" would contain so many and biologically diverse groups as to be meaningless; so we are left with the second possibility. Fortunately, Kron et al. (2002) agreed that a vast amount of work, both morphological and molecular, needs to be done before anyone should contemplate how the diverse tribe of several hundred species might best be redistributed. It seems certain that some of those groups to be recog- nized may be those identified by Small and others a century or more ago. We also feel that the recognition of the Central American greenish-flowered vacciniums into the genus Symphysia as seen by Vander Kloet et al. SE " ee RACAL A CA na logically-based a Ard EM oy ones e the FÉ Vaccinium s.l. in Mexico 1.: achieve a more natural and Central America may prove a asa a classification. Hence, this synopsis of the Mexican e Conil American Vaccinium is ed in the hope that a new species-oriented base line will prove to be a useful summary of what is now known about the genus Vaccinium in the extensive and biologically interesting region of Mexico and Central America. The newly described taxa strongly suggest that additional collecting is still very much needed. The er- roneous impression exists that most of the thousands of new species that are described as new each year are present in abundance in our herbaria and that diligent general collecting is no longer needed. The species of Vaccinium described in the past half century—and that genus clearly is not exceptional in this regard— certainly demonstrate that all or nearly all of the newly described species are the result of material collected since Sleumer's treatment appeared and definitely are not due to the failure of Sleumer or others to make full use of the specimens available to them. A compelling case can be made that what is still needed is a well-planned, intensive collecting campaign throughout much of the area. VACCINIUM L. Shrubs or rarely trees, often rhizomatous, sometimes arising from lignotubers (=burls). Leaves alternate to pseudo-opposite, deciduous or briefly persistent (i.e., barely through the first unfavorable season) or persistent and evergreen, petiolate, the blades with margin entire to (slightly) serrate, the venation pinnate or plinerved. Inflorescences terminal or axillary, racemose, corymbose, rarely paniculate, or 1-2-flowered; floral bract 1, subtending each pedicel; pedicels articulate or rarely continuous with calyx; bracteoles 2. Flowers 4—5-merous, without odor; calyx synsepalous, the tube usually cylindric to globose, the lobes rarely obsolete; corolla sympetalous, aestivation imbricate, valvate or rarely open, cylindric, urceolate, globose or campanulate, white to pink, greenish or yellowish-green; stamens 8 or 10(-12), equal, included or exserted, the filaments equal, distinct or weakly connate at base, equal to or longer or shorter than anthers, the con- nectives lacking disintegration tissue, spurs on filaments present or lacking, sometimes with apparently vestigial spurs, the anthers equal, lacking awns, the thecae surfaces smooth or papillate, the anther tubules 2, distinct or rarely shortly connate at base, dehiscing introrsely (sometimes obliquely), latrorsely or termi- nally by pores or clefts; pollen grains in tetrahedral tetrads, lacking viscin strands; ovary inferior, 4—5 (or falsely 8 or 10 by partial partitions)-locular; style included or rarely slightly exserted, glabrous, the stigma small, simple or somewhat capitate. Fruit a berry, crowned by the persistent calyx lobes and capped by the conspicuous nectariferous disc; seeds many, sometimes with mucilaginous sheath. Vaccinium is a genus with approximately 450 species distributed in the Northern Hemisphere, Meso- america, the West Indies, South America, E. Africa and Madagascar, Malesia, and Pacifica. Twenty-seven species from Mexico through Panama are recognized here. KEY TO THE MEXICAN AND CENTRAL AMERICAN SPECIES OF VACCINIUM 1. Cal ti ith tl | (ie, lacking a disarticulati ) lla open just prior to anthesis or | — TR I d clos sed. 2. Anthers conspicuously exserted beyond the corolla tube; corolla open prior to anthesis, considerably expanding and enlarging thereafter; anther spurs present 1. V. kunthianum KREE «BEN p " cha a Ve a... A D T] o s 200 2. Anthers clearly included within the corolla tube; corolla closed before anthesis; anther spurs present or white ue pinkish idi mm ong, = Roca ics minutely serrate, deciduous or briefly persis! tent (ie 2. V. barely t g ) geminitorum 3. Creeping, viney, epiphytic shrublet; anthers lacking spurs; lla red, 7-10 mm long, 4-lobed; leaves entire, persistent, evergreen 3. V. dissimile 1. Calyx articulate with the pedicel (i.e., marked by a conspicuous disarticulation g 1 lla closed prior to anthesis. 4. Anthers spurred; leaves serrate or indistinctly crenate-serrate. 2 e in cross-section clearly falsely 10-locular; plants of Mexico and Guatemala. . Inflorescences racemose BE ass (1.52)5-9 cm long, ne DEE E EE or Only with | lla 3-6 mm long, small, «iili ilara | ] V | E fl | Hä lanceolate to lance- Toong to ovate, pa 26 : | D only indistinctly if disk glab 4. V. leucanthum Re dm with flowers either solitary in axils of large, leafy floral d or in racemes with ra- chises (1.5-)3-6 cm long with large, leafy, persistent, floral bracts; corolla 5 m long, cylindric to Uu dc es EE to ae elliptic cao GI Ir mm broad: veins on E upper surface oft hort-pilose ` 5. V. stenophyllum e SE in cross- ación 4- ocular not falsely is ocular plants of Costa Rica and Panama 6. V. consanguineum g Si ; t ly serrate to crenate-serru ulate. E | crenate-serrate or en glandularly serrulate. 8. Leaf- blades basally broadly odis mosi 2-5 cm broad 7. V. cordifolium 8. Leaf-blades basally acute to rounded, mostly less than 2 cm broad. 9. Leaves dn on both surfaces, 8-15 cm long and 4-8 cm broad; leaf venation n reticulat V. breedlovei 9. Leaves cem to densely pilose on both surfaces, (0.4-)1-4,5(-7.5) cm long and usually ee than 2 cm broad; eo indistinct to moderately reticulate. 10. cra densely pilos 11. Leaf-blades elliptic to ovate-elliptic, 1-1.5 cm long and 3-8 mm broad; petioles 1-2 mm tora corolla 3.5-5 mm long. 9. V. chihuahuense 11 o oblong to lanceolate, 3-4.5(-7.5) cm long and 0.8-2(-3) cm broad; Dm m long; corolla 8-10 mm long V. lundellianum 10. Leaf enn glabrous or very nearly so except possibly in the vicinity of the e p junction. 12. Coroila tube densely pubescent externally. rolla aestivation imbricate, the tube 5—7.2(-9) x 3 mm, glabrous to pilose s e internally; leaves lacking basal glands on lower surface; surface of anther minutely papillose; forested slopes from 1600-3200(-3700) m in southern debis southward into Honduras and El Salvador 11. V. selerianum 13. Corolla aestivation valvate, the tube 8-13 x 4-6 i EE pilose extemal mog- ind pilose menda E with 1-2 circular | | | om 12500 m and above on EN cupo Costa ica 12. V. talamancense 12. Corolla tube externally glabrous ot scattered trichomes. 14. Pedicels Ge mm Sege or less; E CS ] Fr mm i une or less. 15. t blades mostly c over 2 cm EEN Le BS. LE, f, +| HI E La ] Rar ama cK a nacemes > IA 4 ing foliage leaves or from subtending leaf scars, mostly (1.5-)5-9 cm long; corolla broadly campanulate to urceolate or cylindric, 3-6 mm long; berry clearly falsely 10 locular 4. V. leucanthum 16. boc 12 ith tl func! | rachi tl tly 0.3-1.2 ong; corolla cylindric, 5-9 mm long; berry 4-5- locular 11. V. selerianum 15. ii dE mostly less than 2 cm lon 17. Corolla cylindric, 5-6.5 mm long; Costa Rica 13. V. floribundum 17. Corolla cylindric to cylindric- me or cylindric-urceolate, 4-6.5 mm long; Mexico, Guatemala, and Hondur 14. V. confertum 14. Pedicels (8-)12-14 mm long; calyx lobes ca. E mm long 15. V. wilburii I 1 fal Das irsi D h Institut f Texas 2(1) P» entire. led n slightly clasped by 2 broad, rounded bracteoles; pedicels Ed NA ML AA 3. V. dissimile alyx basally closely ven ban the flowers; corolla era red _ | Ie | L | | 18. € corolla not red, but rather often yellowish- or creamy- green. 9. Secondary and nera leaf veins strongly th forming ticul 20. Petioles lyx lobes 5-6 mm long, | ly nerved; filaments ca. 3 mm long 16. V. jefense a0, Petioles 24 mm long; o jones 1-1 2 mm long; fi EECH = 2. E mm long _ 1. E furfuraceum S reticulum. . Calyx limb saccate, 5(- 10)-spurred opposi | below each lobe 18. V. luteynii i Calyx limb terete and unlobed. 22. Calyx lobes (1.8-)2-5 mm long. 9. V. bocatorense ti y imonteverdense), the bases Ea I |] A Leg, 23. Leaves flat not not claspi ing, t 24. Corolla internally pd floccose with a à dense tangle of white, E omes enter ese ca 0. V. floccosum | + po gee [on Le dp ther tubules ca. 3-5 mm lon 25, Lower leaf surface glabrous; rm 4-8 mm long, glabrous or nearly so; pedicels and rachises glabrous; calyx tube 4.5-8.5 mm long; calyx lobes 2.5-3.5 mm long; anthe tubules 4.8-5 mm long, pilose along inner monteverdense surface 25. Lower leaf surface Loe Bee ai ERU puse enel petiole m long, minutely | | short mine calyx tube | lyx lobes 4-5 long; anther tubules 3-4.6 mm een along i inner d 22. V. campanense 22. Calyx lobes 1.5 mm long or les 26. Corollas cylindric to ale campanulate, (5-)8-12 mm long, glabrous internally; staminal filaments noticeably shorter than anthers, the anther tubules 3 mm or longer 27. Petioles 1—4 mm dri Tale E E mm us and 6- a mm in diam. 23.V. poasanum no g and d mim in niam EN (III MAIGII 28. Leaf venation plinerved; caly obes 0. 2- 04 mm ong or less; corollas pale 24 4. V. almedae tel Il dl El e + t+ Al NI IÇI MIS LV VEUC; elitti kel J " I IH 28. ti sik or greenish- -white; anther tubules dehiscing by elongate, latrorse 25. V. santafeense | I1 me Corollas cylindric to urceolate, 5-8 mm long, pilose internally; staminal filaments about equaling anthers or slightly shorter, the anther tubules less than NJ ON long. 29, Re basally acutely tapering to rounded; calyx lobes 1.2-1.8(-2) m lon ER v. costaricense 29. E blades basally rounded t typically | E lobes 0.8-1.2 mm lon 27. V. orosiense pases (1818) and Dunal (1838) 1 A +1 1. Vaccinium EE EH po i 56. 1851. cited below. ., Bull. Torrey Bot. Club 39:559. 1912. Vaccinium stamineum L. var. fs Kunth in H.B.K., Nov. Gen. Sp. Pl. 3:267, t. 253. 1818. Tyre: MEXICO. Hiparco: “in sylvis Mexicanis, inter Pachuca et Real il Monte, we 1420 wd boda Ge Bf; LECTOTYPE, ee no P- bó n pen of var. ES (=V. kunthianum), without collection number. There is also at P another sheet ai Se Sleumer in 1958, as an ISOTYPE di Vaccinium MN it has the e: iini of Mi ae 4079. T ing World War II. The Field Museum neg. 9020 of tl 1-written label that gives the | lity a as "in Es Mexicanis prope e del Monte" and the collection ow as "Bonpland 4074, 4079." The pl lso sl lant speci left is 4074" and the one on the right “4079,” but there is no way to tell. The type Gamme at F Be the collection number as “Bonpland 4079.” YAI*IL Id 4 e H ha if " La D V ini Y 211 The Ericaceae types of the Willdenow Herbarium (B-Willd.) include a specimen "America meridionalis, leg. Humboldt & Bonpland s.n." Mode ded SE iue: s à label written by panda ER Kunthianum Kl." This is probably an isotype, but does not have al b d ith it. Vaccinium elevatum Banks & cabar ex Wikst. var. fs Dunal, DC. Prodr. 7 7:568. 1838. Tyre: same as for V stamineum var. 8 above. Terrestrial shrubs (1-)1.5—10 dm tall; branchlets densely and softly pilosulous or puberulent, the trichomes throughout usually slender and hyaline but occasionally some gland-tipped. Leaf-blades deciduous, mem- branous, elliptic to oblong-elliptic or ovate-oblong, 1.5—3.5(-6.5) cm long, 5-12(-2.2) mm broad, apically acute to rounded, basally acutely tapering to rounded, marginally entire and sometimes strongly revolute, densely spreading pilosulose on both surfaces or glabrate, the venation pinnate but only the midrib conspicu- ous above, although the lamina also somewhat pustular above, all nerves raised and conspicuous beneath; petioles ca. 1-2.5 mm long, subterete but narrowly canaliculate above, densely puberulent or pilosulose as branchlets. Inflorescences axillary, of often leafy racemes, 3—8-flowered; rachises sharply angled, 4-34 mm long, pubescent hlets; floral bracts leaf-like to leafy, persistent, EE acute, 5— S mm long, glabrous to pilosulose abaxially; pedicels 5- 5—6 mm long, sometimes nodding, p or near base, linear to oblanceolate, ca. 1.5-3 mm long, hirsutulous. Flowers 5-merous; calyx ca. 2.5-4.5 mm long, continuous with pedicel although rarely appearing articulate as fruit matures, the tube campanulate and x flaring distally, ca. 1-1.5 mm high and 2 mm in diam., glabrous to densely hirsutulous with white, downwardly reflexed, delicate, white trichomes, the limb flaring above, glabrous to pilosulose, the lobes deltoid, acute to obtuse, ca. 1-2 mm long, ciliate or puberulous to pilosulose; corolla with aestivation open, membranaceous, unistratose, campanulate, ca. 4.7-6(-8) mm long, white, glabrous, deeply-lobed, the lobes deltoid to oblong, acute to rounded, ca. 3-4 mm long; stamens 10, conspicuously exserted (rarely included), ca. (4.88.5 mm long, the filaments ca. (2.733 mm long, flattened, densely hirsutulous externally and glabrous internally, the anthers ca. (3-)5-6.5 mm long, bearing dorsally, ascending or horizontally wide- spreading, slender, finely muricate spurs ca. 0.7-2 mm long, the thecae ca. (1.3-)2-2.5 mm long, sharply and finely muricate, the tubules ca. (1.72)3—4.5 mm long, muricate, dehiscing by short, introrse, slightly oblique pores that are marginally irregularly 2-4-dentate (lacinate); styles long-exserted, to 11 mm long. Berry spherical, 5-10 mm in diam., black, sparingly to moderately pilosulose, 5-locular (2). Distribution.—Montane forests, dry pine forests, and oak-pine forests of the Sierra Madre Oriental of Mexico from Nuevo León and Tamaulipas, south through San Luis Potosi, Puebla, Hidalgo, and Vera Cruz, at 1550-1785 m. Flowering: Apr-Jul; fruiting: Jun-Sep. Vaccinium kunthianum is characterized by a calyx that is continuous with the pedicel, a corolla that is open prior to anthesis and enlarges thereafter, by anthers conspicuously exserted beyond the corolla tube, and by anther spurs present. This is the only Mexican representative of the very codium (Raf.) Rehder], a largely eastern North American taxon often Rede separate genus Polycodium Rar e is all too easy for us to discount Rafinesquian genera out-of-hand, but this Ce genus does have lent approbation of Nuttall (1843), one of our historically most critical and acclaimed botanists, who Segen this group of species from Vaccinium as the genus Picrococcus Nutt. The systematics of Poly- codium whether treated as a section, subgenus, or genus is unresolved. As many as 20 species or as few as one species (Baker 1970) have been recognized. EEN Vander Kloet nid Ge a single species in hil the Sierra Madre Occidentale distinctive subgenus Polycodium (Raf.) [= sect. Poly- +l eastern North America, V. staminium L., to synonymy. Too little material is available to bé md but the Mexican plants seem very unlike the common V. stamineum of eastern North America. Therefore, for the present the Vaccinium kunthianum is best treated as a distinct species. Like almost all species in Mexico and Central America much more intensive field observation and collecting is needed. Additional MIS ues epis Minds León: oak-pine forest 8 mi E of aic OR 1550 m, 30 Jun 1948, Meyer & b, La Trinida g 1939, Muller 2843 (GH, LL NY). Taumaulipas: Sierra de tela 23º5-7N, 99º15'W, just W a Tierra Colorado, 19-25 ha 1984. Johnston 12813 (TEX). San Luis Potosi: Cerro El Agujón, cerca de El Zapote, approx. 30 km al SSW de Río Verde, 1750 m, 13 May 1968, Rzedowski 25675 [| | E al n.a D Inm LI EA £T mfa 127 212 (CAS, LL, MICH, MSC, RSA, bs dona ay pue forests near pens Station, 5800 ft, May 1904, Pringle 8899 (CAS, CU, F, GH, K MO, MIN, MSC, NY, PH, US). E g km. 240 on Pan-American Hwy., 2 Jun 1948, Atchinson 512 (CU): Zimapán, Coulter 892 (CGE, GH, K, NY, P H); District Z d 1 sphagnum bogs ca. 3 mi from Zacualtipán on road to Tianguistengo, 2100 m, 4 Jul 1947, Moore 3305 (CU, GH). enema Municipio de Zacualpán, 2500-2600 m, 20º24ºW, 98?26'W, 9 mi SE of Palo Bendito, bosque de piños, 22 Jul 1982, Nee & Diggs 25228 (F). 2. Vaccinium ifl Kunth in H.B.K., Nov. Gen. Sp. Pl. 3:267, t. 252. 1818. Tyee: "Crescit in Regno Mexicano, inter Omitlan et Moran, alt. 1400 hex,” (HOLOTYPE: P- HBK!; isorwes: Bt, B-Willd. 8315-1!). The isotype in the Willdenow Herbarium has on its label “leg. Humboldt & Bonpland s.n.” and “Vaccinium geminiflorum H.B.K.” written by Klotzsch (P. Hiepko 2007 to JLL). Erect, rhizomatous, densely branched shrublets mostly 10-25 cm tall, often forming dense colonies, gla- brous throughout with the exception of the youngest stems and leaves; branchlets angled, at first densely but minutely puberulent but becoming glabrate. Leaf-blades deciduous or briefly persistent, membranous, oblanceolate or narrowly oblong-obovate to somewhat elliptic, 5-20(-32) mm long, 3-8(-12) mm wide, apically obtuse to acute, ne narrowed to a + cuneate base, marginally finely crenate-serrulate with the teeth usually t ially when young with a very short, gland-tipped vein, both surfaces glabrous but the veins (especially "wien young) often very minutely and sparsely puberulous, the venation pinnate, forming a conspicuous, elevated reticulum on both surfaces at least when dried; petioles 1-2 mm long, semicircular in cross-section, flatly grooved above, glabrous. Infl illary, of solitary flowers; floral bracts leaf-like in shape and texture, 4.5-6 mm long; pedicels recurved, 3-5 mm long, slightly and very shortly puberulous to glabrous; bracteoles basal, caducous, oblong, rounded, ca. 4—5 mm long, covering ps and calyx. Flowers 5-merous; calyx 2-3 mm long, continuous with pedicel, the tube campanulate, a. 14-1.8 mm long, glabrous, the limb distally truncate to minutely lobed, the lobes obscure to broadly ovate, dá rounded to EES ca. 0.3-1 mm long, glabrous, occasionally with minute, glandular ) imbricate, | , unistratose, cylindrical to more typically sciatis barrel- "n" or globose-urceolate, 4—6 mm long, 3-4 mm broad, white to pinkish or even roseate, glabrous, the lobes broadly rounded, 0.8—1.5 mm long; stamens 10, included, ca. 3.5—4.5 mm long, the fila- ments ca. (I-)2—2.4 mm long, glabrous, the anthers ca. 2.5 mm long, dorsally bearing 2, slender, antrorsely and minutely barbellate, strongly ascendent spurs ca. 0.6—2.2 mm long, the thecae ca. (0.51—1.4 mm long, papillose, the tubules ca. 0.9—1.7 mm long, 10 or more times as long as wide, dehiscing by oblique or nearly terminal introrse pores. Berry dull blackish, unknown at maturity. Distribution.—Montane pine-oak forests, alpine meadows, boulder fields, and seepage areas, from Chihuahua and Nuevo Leon south throughout much of | Mexico into Guatemala, at 1785-4110 m. Flowering: throughout most of the year but mostly from Feb-Aug; fruiting: Mar-Oct. Vaccinium geminiflorum is characterized by its subshrub, rhizomatous habit, deciduous (or nearly so) leaves, calyx that is continuous with the pedicel, corolla with imbricate aestivation, long-spurred anthers, and comparatively Moab m e Kee The most ext scussion 1 Guat ] ] [the V. cespitosum - geminiflorum complex was that of Camp (1942), based largely on his observations in western Mexico. Vander Kloet (1988) recognized both V. cespitosum Michx. and V. geminiflorum Kunth as occurring in Mexico with V. cespitosum represented by “one disjunct population” although his map shows three widely separated localities in west- ern Mexico. Camp noted that the plants of V. geminiflorum from lower elevations, seldom if ever subjected to frost, had persistent leaves while those from higher elevations had deciduous leaves. Roughly correlated with persistence and lower elevations were more deeply grooved twigs and narrow, relatively longer leaves at the lower elevations, while at higher li E use are more likely to be subterete and the leaves wider. Camp concluded that because of the I tion that it would be best to maintain the Mexican material of this complex or section EN a melena name, V. geminiflorum. He speculated that in the Pleistocene the widespread northern V. cespitosum had had to migrate into Mexico where it encountered V. geminiflorum and hybridized with it resulting in the great variability in the populations encountered in the mountains of central Oaxaca. We are unable to evaluate this suggestion. The focus of vander Kloet’s earlier publications was upon the North American representatives of the Geng i Luteyn, S — ——— — 213 genus, but he also delt with the species of “northern Mexico.” Only Mexican species whose range was largely in the United States and Canada were included in his account: Vaccinium cespitosum Michx. and V. kunthianum Kunth (as V. stamineum L.). Little material was available to him for study. From vander Kloet's keys and descriptions the following key has been extracted. Leaf margins sharply serrate along most its length; blade often sparsely glandular beneath; berry dull black; rest ric cted to Mexico and ds mala V geminimorum | 1. | he 1 +l LAT EA L. | j L. | J fe? | | beneath; berry usually blue and glaucous, or rarely dail blacks ranging from Labrador to Alaska and south into New England, Wisconsin, Colorado and California with three disjunct localities in Mexico V. caespitosa Additional collecti ined: MEXICO. Chihuahua: M pio de B fC on W facing slope in pine-oak forest, ca. 7100 ft, 8 May 1973, Bye 3690 (TEX); higl | de Cob rs 7000 ft, Jun 1892, Hartman 527 E Nuevo León: icd meadow, 12,500 ft, summit of Cerro Potosi, 24?53'30"N, 100º10'30 W, 19 Jun 1966, Gilbert 30 (TEX). Z on road NE fr an Juan Peyotán, Nayarit, towards Aserradero Tepetates, 5—25 km N of Nayarit-Zacatecas boundary, 2400—2600 m, 13 Jan 1975, See 25770 (MICH). Durango: high plateau with Pinus, Quercus, Artostaphylos and Garrya, 55-60 km SW of Durango City on road to La Flor, 2400 m, 16 Sep 1979, Breedlove 44097 (CAS); 8-10 km al S de El Salto, hacia Pueblo Nuevo, Municipio de El aa nan m, 3 M: 1982, ae & Tenorio 7589 ge nd Laguna del Progreso, 34 road mi N of railroad at Coyotes, lumber camp orest fb 00-2600 m, 21 Jul 1955, Maysilles 7887 (MICH). Hidalgo: 1.5 al NNW del ganas Municipio de ica 2850 m, 14 v 1976, Medina 1964 (MICH); Peñas i c cerca de RW Municipio i Real del Monte, bosque de Abies, 2750 m, 21 Apr 1966, Rzedowski 22184 (MSC). Y and thin pine woods, Mt. Orizaba, 10 oe ft, 28 ius pe Ba P» rdi p AAU, K, NY, d in Pe a Cofre eo Perote, 10300 ft, 2 May 1938, Balls 4564 (A, K, NY, US). Puel , 3960 m, 1 Aug 1958, Beaman 1995 (GH, MSC); Si pine forests, MINA Station, 5800 ft, Apr 1904, Pringle 8960 (A, AAU, CAS, CU, F, GH, K, MIN, MO, MSC, NY, PH, US). Distr. Fed , top of Cerro Ajusco, 3937 m, 12 Jul 1959, Beaman 278l (MSC). Mexico: grassy ravine in alpine meadow N side of Nésatio de Toluca; 0.7 mi E of pue where road goes above timberline, 3985 m, 28 Jul 1958, Beaman 1926 (GH, MSC, TEX, US, WIS); Municipio de Amecameca, sl tl, ca. 11,000 ft, 4 Jul 1943, Gily & Dodds 9 (MICH, MSC, iM Distr. Ss EE o ees Sec 2880 m, dÉ? 1933 Hinton 4190 E K, MO, NY); rocky moist tussock grassland on , aw 6 Aug 1960, Iltis, et al. 1039 (MSC, WIS); e; Jel Mente Alta P EG Jit IIC Malinche, 3200 m, 13 Apr 1966, Rzedowski 22130 (DS, MICH, MSC). Jalisco: Nevado de Colima, 12500 ft, 16 May 1893, Pringle 4375 (F, GH, K, MIN, MO, M5C, MU, NY, PH, Sieg Cuai Lo. ide Teotepec, Distr. de Miña, 3200 m, 17 Jul 1939, Hinton et al. 14450 (F, GH, LL, MO, NY, PH, US). O the stone of sacrifice, Zempoaltepetl, 19— ii Ge 1937, Camp 2651 (AAU, CAS, DUKE, MICH, MO, NY, US); crest of Mesican Hwy. 175, 30.5 mi kl pio of? N of Guelatao, open, moist, f mt. ridge, ca. 10,000 ft, 24 Dec 1975, Reznicek & Gregory 328 (MICH). Chiapas: slopes with inus, Quercus and Arbut m N of Mean n 190 on logging ne from FUN a Chamula microwave station, 2400 m, 15 Oct 1976, sb 40753 (DS). GUATEMALA. Chi g line, NW e of Volcán Acatenango, ca. 3760 m 16 Aug 1959, Beaman 3270 (GH, MSC). Huehuet go: Si le los Cucl t ; Tojquia and Caxin bluff, 3700 m, o 1942, Steyermark 50214 (A, F, NY, US). Q It it of Volcan S Maria, 12,400 ft, 26 Jul 1934, Skutch 837 (A). uez: summit of S rim of crater, 3760m m, 31 Jul 1959, B 2925 (GH, MSC, TEX, US) San Marcos: in gravelly soil above Sacatepeq MN M E side of peak, Volcán Tajumulco, ca. 4110 m, 9 Aug 1959, Beaman 3159 (GH, MSC, TEX, US). 3. Vaccinium dissimile S.F Blake, J. Bot. 53:271. 1915. Disterigma dissimile (S.F Blake) S.E Blake, Proc. Biol. Soc. Washington 35:120. 1922. Sphyrospermum dissimile (S.E Blake) Luteyn, Opera Bot. 92:126. 1987. Tyre: COSTA RICA. San José: La Palma, 1500 m, Aug 1898, Tonduz 12546 distributed as J. Donnell Smith 7380 (Lectorvre: K! isotecroTYPES: BM! F! GH! K! MO! NY! USD. Creeping, epiphytic or terrestrial, viney shrublets with stems interwoven amongst bryophytes; branchlets wiry, perhaps to 1 m long, 1-1.5 mm in diam., the young stems densely spreading puberulent, glabrous in age. Leaf-blades persistent, coriaceous, evergreen, ovate, lance-ovate, or narrowly to broadly elliptic, mostly 13(4) cm long, 8-15(-20) mm broad, apically acute, basally rounded to cuneate, marginally entire, glabrous or beneath sparingly to moderately beset with dark, glandular, appressed trichomes 0.1-0.2 mm long, the venation obscure; petioles 1-2 mm long, puberulent. Inflorescences axillary, of solitary flowers; floral bracts broadly reniform, ca. 0.3-0.4 mm long, rounded, glabrous; pedicels 1-2(-4 or —8?) mm long; bracteoles basal, ovate to broadly oblong, opposite, obtuse to broadly rounded, 0.8-1 mm long and nearly as wide, puberulent. Flowers 4-merous; calyx ca. 2.2-2.8 mm long, continuous with pedicel, the tube campanulate, 0.8-1.2 mm long, densely hirsutulous with hyaline trichomes 0.1-0.2 mm long, the lobes broadly deltoid, acute, 0.7-0.8 mm long, moderately to densely hirsutulous; corolla with aestivation valvate, thinly fleshy, bistratose, campanulate to broadly flaringly funnelform, 7-10 mm long, 6-9 mm in diam. at the apex or the [| I E al Dad H ID LI da dg a "f" 214 f Texas LANE broadest point, red, sparingly andi ] ] y bes t witl di gg gland- tipped trichomes 0.1-0.2 mrn +1 C] long, the lobes erect, acutely PECH 1. 5-2 mm long: stamens 8, included, ca. 4.3-5.8 mm long, 1.3-4 mm long, elliptical, tapering both to the base and the apex where ca. 0.5 mm wide, sparsely pilose, the anthers medifixed, 23.3 mm long, lacking spurs, the thecae ca. 1.7 mm long, conspicuously papillose, the tubules 1.6-2.5 mm long, dehiscing by introrse clefts 0.5—0.7 mm long. Berry bluntly 4-angled, oblate, 0.7—1 cm in diam., translucent white, densely hirsutulous, 4-locular. Distribution — Wet mountain slopes dominated by premontane rainforests and elfin thickets of shrubs and small trees in Costa Rica and Panama and also in NW South America (Colombia and Ecuador), at 500-1800 m. Flowering and fruiting: throughout much of the year but perhaps heaviest from Feb—Aug. Vaccinium dissimile is characterized by its lianoid, thin-stemmed habit, short pedicels, and dark red, campanulate corollas. This is another species of Vaccinium that rests not too comfortably with its supposed congeners. S.F. Blake, who orginally named the species as a Vaccinium, later transfered it into the genus Disterigma. Luteyn, currently the leading authority on the neotropical representatives of the family, has concluded that this species should be placed in the genus Sph; Poeppig & Endlicher as shown by the mature fruit: "The fruits of Vaccinium and Disterigma (and most eri blueberries) have a thick and leathery pericarp that dries that way, and the seeds have normal embryos whereas in Sphyrospermum and Themistoclesia the pericarp is very thin and papery ... and the seeds have a green embryo ...” (Luteyn 1987, p. 126). [Note: RLW, first author of this paper, wishes to treat this species in Vaccinium; JLL, second author of this paper, however, ndi and still maintains the species a in ki d pil Du Ad ED 11 J. K^£CIPA DISA TT +1 AA 1 El pl] 1.5 km NE e ca. 8 km ow of = ee 10?18'N, 84°02’ W, 450- ia m, 14 Feb 1986, dicis um no pico d o Mo oLaPalm 8 Volcár Barba, ca. 1500 m, 23 May 1972, Wilbur & Almeda 16898 Gees CR, DUKE, F, GH, LL, MICH, MO, NY, US, PMA, WIS). PANAMA. Bocas del Toro: Cerro Colorado, road along top, 1500-1750 m, 13 Aug 1977, Folsom et al. 4726 (DUKE); forested slopes, Fortuna Dam region, along trails leaving pipeline road, ca. 8°45’N, 82°15’W, ca. 1000 m, 8 Dec 1985, McPherson 786 (DUKE, MO). Chiriquí: cloud forest, Cerro Colorado, ca. 50 Kee N ih San np ca. 1400 m, 17 Aug 1975, Dressler 5086 (DUKE, MO). Veraguas: 11 km from Escuela Agrícola Alto de Piedra, D Atlantic slope, 15 Nov 1974, Mori & Kallunki 3147 (MO). Coclé: Cerro Pajita, hills N “of El Valle, 1100 m, 27 Oct 1946, Alle 3782 (MO, t US); Atl El Cope, 8?40'N, 80°36’W, 750-800 m, 13 Feb 1982, Knapp & Dressler 3494B (DUKE, MO); La Mesa, 5 mi N of El Vallé, 2500 ft, 10 Nov 1965, Tyson & Godfrey 2443 (DUKE, MO). Panamá: Cerro Jefé, new road leading N a summit, Ja ap 1975, Wees? ee (DUKE, MO). Colón: $ approach of Cerro Bruja from Río Escandaloso, 18 May 1978, H l t, Cerro Brewster, 9?18'N, 79?16'W, 800—850 m, 20 Nov 1985, de Nevers et al. 6269 (DUKE, MO). Darién: Sambú Basin, Cerro de Garagara, 500-974 m, 7 Feb 1912, Pittier 5644 (US). 4. aon leiicanthtim edidi , Linnaea 8:524. 1833. Tyre: MEXICO. VERACRUZ: “infra S. Salvdor...Pr. Chiconquiaco” santla and Jalapal, Schiede 174 (HOLOTYPE: BT; LECTOTYPE, designated here: NY! Ki MO! P! WI). m label of the NY sheet (ex M herb.) reads "Infra S. Salvador, Mexico"; the K sheet reads Pu Chiconquiaco" 1 1 Lä defi J and is without ti a imply “Mexico.”] hlechtendalii G. Don, Gen Syst. 3:856. Nov 1834. Type: MEXICO: “near Jalacinga, in woods.” The actual collection and location ) ID | hlecht on) Nutt., Trans. Amer. Phil. Soc. n.s. 8:266. 1843 Vaccinium eidam Dunal, Prodr. 7: aed 1839. Tree: MEXICO: “inter a et Real del Monte,” Berlandier 329 (pl. exs. n. 12) [HOLO- Tyee: G-DC (photos F neg. 33846 NY neg. s.n.); isotypes: BM!, NY frag. ex G-DC!, OXF! P!]. The type specimen in the DeCandolle Herbarium (G), | (JLL) 1 label that gives tl ber *12 ex." in the upper left corner and the number *329" in the upper right corner. There is a fragment of the "i DC: type at NY, uec T ed of ERU A at P!, and one each at the BM and OXF (all isotypes). T na ] hat the pl l by G Don (1834) and by Dunal (1839) are = same collectio + [m] | n exsiccatae sent out by Berlandier. It i Terrestrial shrubs 1—4.5 m to trees to 15 m tall; branchlets somewhat angulate when young but becoming terete, minutely but densely puberulent or glabrate. Leaf-blades persistent, coriaceous, evergreen, lanceolate to lance-oblong to ovate, highly variable in shape and size, mostly 2—5(—7.5) cm long, 1.2-2.6 cm broad, apically usually acute to less commonly Boe or rounded, not mucronate, basally cuneate to rounded, marginally crenate-serrate, when mature g t most very sparingly puberulous (or even densely but often i | idrib above), when young glabrous to ensaia Ge beneath with midrib bubemións above bd Ee beneath, the venation pinnate; petioles 2-4 mm long, LAR Mii c TUER Y VES I fanteal À H Wy mi 215 semicircular in cross-section, "i, broadly and shallowly Ge above, glabrous to densely short-puberulous. Infl ill ising from leafl , racemose, 5-16-flowered; rachises finely ridged n grooved, (1. 55 —9 cm long, 0.5- J mm in diam., densely to sparsely but minutely puberulent; floral bracts caducous, oblong, cuculate, ca. 1.5-4 mm long, 2-2.5 mm broad; pedicels nodding, often somewhat secund, ca. 1.5-7(-12) mm long, finely but densely to sparsely puberulent; bracteoles located medially to just above or below middle, + opposite, caducous, linear, 1.5-2.5 mm long, glabrous to puberulous, caducous. Flowers (4—)5-merous; calyx 2.5-2.7 mm long, conspicuously articulate with pedicel, the tube obconical, 1-1.5 mm long, sparingly to ui densely puberulent, the lobes deltoid, acute, 0.8-1.5 mm long; corolla with aestivation imbricate, , unistratose, broadly campanulate to urceolate, 3-6 mm long, white to pinkish tinged, glabrous externally, the lobes deltoid, acute, 1-2.2 mm long; stamens (8210, included, 2.8-5.4 mm long, the filaments ca. 13.2 mm long, ciliate distally, the anthers 1.73.4 mm long, typically bearing dorsally 2, slender, strongly ascendent and outwardly divergent spurs ca. 0.6—1.2 mm long but these apparently sometimes represented by vestigial remnants 0.1-0.5 mm long, the EE ca. l- a 5 mm long, the tubules ca. 0.7-2 mm long, dehiscing by short, oblique, introrse, oval pores; k glabrous Berry spherical, ca. 5-8 mm in diam., blackish, puberulous to ee Aey 10-locular. Distribution.—Montane pine-oak and both evergreen and ts, thickets and — from Hidalgo south through Vera Cruz into Michoacan, Oaxaca, and Chiapas (Mexico), and mucho Guatemala to Honduras, mostly between 1500-3000 m. Flowering: throughout the year put peaking from Apr-Jul; DE Been ci the year. Vaccinium terized by its broad, crenate-serrate leaves, long-racemose inflorescences, its calyx articulate with the pedicel, spurred to unspurred anthers, and a falsely 10-locular berry. No other species of Vaccinium in either Mexico or Central America has proven as baffling as V. leucanthum. As inter- preted in this treatment, it has the greatest range of floral morphological variability of any species in the area. On numerous occasions it was felt that a set of characters had been detected that would allow the clean separation of a block of disconcerting variability but analysis of more specimens dashed the supposed solu- tion. The most strikingly variable features are in leaf size and shape, number of perianth parts, and length of the anther spurs. The variablility of spur length proved especially vexing for a priori it was felt that spur length was of "fundamental" importance. These spurs ranged from slender, upwardly arching, long, horn- like appendages to the merest protuberance and in some cases could not be detected with certainty and so might well be absent. The presence or absence of spurs has usually been treated as of sectional, subgeneric or even generic significance within the Vaccinieae. Belatedly and no doubt reluctantly, it was impossible to employ anther spurs as a taxonomically useful character in this species. This is not a wholly satisfactory solution; it just happens to be the best that can be done now. Extensive field work throughout the range of V. leucanthum would probably be most helpful in resolving the apparent uncertainties. It would also appear that the problems associated with this species can only be resolved in conjunction with its relatives in the Andes of South America and the West Indies. D 1 [* Alajo A dditi l collect ] ] MEXICO Hidalgo: Dist. Tulanci ingo, dry between Tulancingo and Acaxochitlan, 7000 ft, 13 May 1947, Moore 2820 (CU, GH, MICED: hills ca. Trinidad Iron Works, 5800 ft, 6 ili 1904, Tua (A, SCH CU, F, GH, LL, MIN, MO, MSC, PH, US). Veracruz: Perote, SE 6500 ft, on old lava in crevices ,23 Sep 1938, Balls 5519 (GH, US); Ocotepec, cerca del Cerro, M de Jal go, 1300 m bosque de encino, EN de cerro, 4 Mar 1970, Ventura A. 1006 (DS, F, MICH, MO, NY). Puek in f W of H hinango, 1 Nov 1943, ada 12640 (GH, MICH, MO, NY, US); in Pinus-Quercus-P t t, near hwy. 1 istá Ogden & Gilly 51212 (DUKE, MICH). Michoacán: bosque de pino, San Miquel del Monte, 15 km al SE de Morelia, 2200 m 17 Sep 1967, Rzedowski 25165 (CAS. LL, MICH, M MSC). Oaxaca ] d other trees, along road from Teotitlán del Camino to Huautla de Jiménez, 2250-2300 m, 12 Jul 1968, vou 4739 (DUKE, MICH, NY); top of Zempoaltepetl, 19-27 Feb 1937, Camp 2660 (AAU, CAS, DUKE, F, GH, MO, M Villa-Alta, Reeg Se Rincon, 4-6500 ft, Gaio i [NY; mid in caracasanum Kunth 1818), tl 842)]. Chi 1 NN CAO VILE ly misapplied by ————— mone EN C] laal Zi an C 7800 ft, 20 Jan 1965, B il Raven 8135 (DS, F, LL, MICH, MSC Ds steep wooded +1 1 1 Cal D:. LI J A “RT E Tanta] ] D 1.1 AT Col: AA TE sd J J 00 ft, 31 May ee 1 1 : 1 Breedlove 10178 (DS, F, LI , MICH); loud f t the N dW slope of the C 216 I E ~f al Bas H Ip LI add f Texas 2(1) the road from Huixtla to El Porvenir and Siltepec, Municipio de Pi ee m, pda 1972, Breedlove 25827 (DS, DUKE, MICH, MO, NY, RSA) f evergre ide of Cerro Tres Picos and the ridges near summit, - 2100-2500 m, 11 Dec 1972, Breedlove & Thorne 30138 (DS, DUKE, F, MICH, MO, NY, RSA, TEX); Sierra Madre de Chiapas, Pinus-l bar forest, between Finca Liquidambar and Neuva Colombia, 2500 m, 18 Jun 1985, Luteyn & Lebrón-Luteyn 11574 (MEXU, MO, NY). GUATEMALA. mal Mapa nune areas above Santa Catarina, 5000 ft, 24 di 1945, no) ini c Chiquimula: mixed forest below cloud for 1d] o Brujo, SE il 2000 m, : Nov 1939, rn 31056 (F). bueren e Mun bank, 8100 ft, eae #9 ca. 13,3 mi É of Santa Eulalia, 972, Almeda & Luteyn 1689 (DUKE, F, MICI Soloma, 7400 ft, 21 Aug 193 Skutch 1059 (A, F, NY, US). Progreso: on summit of Volcán Siglo, 3300 m, 21 Jan 1942. Steven 43080 (A, F). El Quiché: in arro Nebaj, 5 Jun 1964, Contreras 4883 (CAS, GH, LL); trail between Nebaj and Chajul, 6 Feb 1946, Sharp 4666 (GH, MO, NY). Zacapa: trail between Santa Rosalia de Marmol and Vegas, 19 Jan 1942, Steyermark 42912 (F). HONDURAS Coma Me: be Don Thomas,” 10 km SW of Gracias, 1850-2050 m, 10 May 1992, D'Arcy 17889 (MO); Cortes National Park, t antiles to Cerro fee 2240 m, 20 Mar 1993, Hawkins 659 (MO); Lempira: montaña de aa 2650 m, 18-22 Nov 1974 (MO); Eni Nacional Celaque, cloud forest on ridge to Cerro Oeste del Río Naranjo, 2300 m, 9 Jul 1991, House 1036 (MO). 5. Vaccinium stenophyllum Steud., Nomencl. Bot. 2:740. 1841. Vaccinium angustifolium Benth., Pl. Hartweg, 45. 1840, non Aiton (1789). Tire: MEXICO. JaLisco: "in regione frigida prope Bolaños,” Hartweg 342 (notorvpe: K-Herb. Benth.! photos Univ. Michigan neg. 694 and NY neg. s.n., frag. NY; isorvres: BM! CGE! K-Herb. Hook.! NY! OXF! PD. Vaccinium angustifolium Benth. var. 8 glaucescens Benth., Pl. Hartweg, 45. 1840. Vaccinium gl (Benth.) Riley, Kew Bull.185. 1922. Tyre: MEXICO. Jalisco: “in regione frigida, prope Bolaños,” Hartweg 342B (HoLotyre: K-Herb. Benth.! photo NY neg. s.n., frag. NY! Vaccinium gonzalezii Riley, Kew Bull. 116. 1923. Tyre: MEXICO. Stnatoa: San Ignacio, Mesa de Bueso, 1250 m, Gonzalez 839 (HOLOTYPE: frag. NY!; oppe K!). Terrestrial shrubs or small trees 1— 34-8) m tall; branchlets slender, glabrous to densely, shortly puberulous, J sometimes glaucous. Leaf: bl t, coriaceous, evergreen, narrowly lanceolate, lance- elliptic to linear- oblong, 1—4(—6.5) cm long, SECH 1215) mm broad, apically acute to acuminate, shortly and deciduously mucronate, basally cuneate, marginally entire to minutely serrulate with the teeth often glandular-tipped, lustrous above, glabrous or very nearly so on both surfaces except often densely puberulous along the midrib above, the venation pinnate, conspicuously reticulate and often raised on both surfaces; petioles 1-2(3) mm long, subterete to broadly and shallowly canaliculate above, glabrous to densely although minutely puberulent. Inflorescences axillary, either leafy racemes of 3-10 flowers each, or flowers solitary in axils of apparently normal or reduced leaves at tips of branchlets and then inflorescences apparently differing little from leafy, vegetative branchlets; rachises (when present) finely ridged and grooved, mostly (1.5-)33-6 cm long, ca. 1 mmin diam., puberulent to glabrous, sometimes glaucous; floral bracts slightly smaller than the foliage leaves and of similar shape and texture, 8-30 mm long, 2.5-9 mm broad, puberulent to glabrate; pedicels nodding or at least strongly divergent, mostly 24—8(-10) mm long, 0.2—0.3 mm in diam., glabrous or sometimes puberulent, sometimes glaucous; bracteoles nearly basal, caducous, subopposite to clearly alternate, oblong to lance-elliptic or linear, 1.2-2.2 mm long. Flowers 5-merous; calyx 3.5-4.6 mm long, conspicuously articulate with pedicel, puberulent to glabrous, sometimes glaucous, the tube obconic to somewhat campanulate, 1.5-2 mm SES ES lobes deltoid, acute to acuminate, ca. 1.2-2 mm long, glabrous to densely ciliolate at tips; corolla with tion imbricate, I , unistratose, cylindric to urceolate, 5-7 mm long, whitish to creamy, glabrous externally, e lobes deltoid, acute, 1.2—2.2 mm long; stamens 10, included, 3.5-4.3 mm long, the filaments ca. 1-1.6 mm long, weakly to densely ciliate, the anthers ca. 2.8-3.5 mm long, dorsally bearing slender, strongly ascendent spurs 0.6-1.4 mm long, the thecae ca. 1-1.7 mm ea the pro slender, 1.2-1.8 mm long, dehiscing by short, introrse, oblique pores; styles included; to shortly and densely white-pilose. Berry spherical, 5-9 mm in diam., blackish, puberulous or geen sometimes glaucous, falsely 10-locular. Distribution.—Mountain slopes of the Mexican Sierra Occidentale from Durango south into Oaxaca (Durango, Sínaloa, Nayarit, Jalisco, and Oaxaca), dominated by open pine-oak forests and rocky slopes and summits, mostly between 1075-2800 m. Flowering: (Jan—)Mar—Aug(—Oct); fruiting: (Jan—)Jun-Oct(- Dec). ITT 31..5 ,$ ` cha ` Ir gel A H y In 217 Vaccinium stenophyllum is characterized by narrowly lanceolate, lance-elliptic to linear-oblong leaves with prominent venation and mucronate tips, inflorescences of leafy racemes or flowers solitary in the axils of apparently normal or reduced leaves, a calyx that is articulate with the pedicel, spurred anthers, an often shortly and densely pilose nectariferous disk, and falsely 10-locular berry. Louis Williams (1965, and in Standley & Williams 1966) included V. stenophyllum in the Flora of Guatemala even though he stated that *the material from Guatemala is hardly, if at all, distinguished from V. leucanthum." For this study, no specimens of V. stenophyllum have been seen south of Jalisco, while V. leucanthum is well-represented in Guatemala. Plants of V. stenophyllum are not at all reminiscent of V. leucanthum. We agree with Sleumer (1936) that V. RR is dd a broader leaf form of V. eee Addit ined. MEXICO. D g ] 19] de I Cl Sta. María Ocotán, Munici ipio de ie e 2400 m, 4 n ri Gonzalez 2582 (NY). Sinaloa: Mesa de B , San nacio, 1250 m, 15 May 1919, iiw & Salazar 83 y 1 Bi M ta Teresa, M N ,1850m, 12 Aug 1980, Breedlove € e: 45509 (CAS, MO, NY). Jalisco: o with Pinus, GE and Arbutus 15 km NW of Los E on road to Talpa de Allende, E de "enam o m, ni Nov Breedlove & Almeda 60617 (CAS, MO, NY); Sierra de Manan- tlán, 25—30 km SE of Autlán, al lled “La Cumbre,” between El Chante and Cuzalapa, 19?35'N, 104°8-15°W, summits of high S- lice cliffs, 2750 m, 20-21 Mar 1965, McVaugh 23131 (MICH); bare, gullied hillside, Hacienda del Cura, Sierra Madre, San Sebastián, 1425 m, 2 Jan 1927, Mexia 1340 (A, CAS, DS, F, GH, MICH, MO, NY, US); Municipio de San Martín de Bolaños, Las Vidrieras, 10 km al NW de El Platanar, 2450 m, 1 Oct 1968, Rzedowski 26156 (MICH). 6. Vaccinium gui Klotzsch, Linnaea 24:64. 1861. Tyre: PANAMA: Volcán Chiriquí, Warszewicz s.n. (HOLOTYPE: Bt, photo F neg. 4613). Vi SI izbl. Bot. Gart. Berlin-Dahlem 12:138. 1934. Vaccini gui Kl | iraz (SI ) Sleumer, Notizbl. Bot. Gart, Berlin-Dahlem 13:126. 1936. Tre: COSTA RICA. CARTAGO: I fielder Irazú, Hoffman 140 (HOLOTYPE: Bt, frag. U Terrestrial shrubs or small trees (0.2-)1-3(-10) m tall, often compact and densely bushy; branchlets stiff, rigid, glabrous to densely puberulent. Leaf-blades persistent, coriaceous, evergreen, mostly narrowly to broadly elliptic, occasionally oblong, (1)1.5-3(-4.6) cm long, 5-12-18) mm broad, usually 2-3 times as long as wide, usually tapering to an acute apex and base, marginally callose-thickened and shallowly serrulate with teeth mostly (1-)2-4 mm apart, each tooth often tipped with a darkish glandular callosity, glabrous or with scattered glandular fimbriae beneath, moderately puberulous along the midrib, the venation pinnate; petioles 1.53 mm long, glabrous or puberulent. Inflorescences allan racemose, 5—10-flowered; rachises sharply angled, mostly 1-4 cm long, puberulent; floral bract ovate to oblong, 2-5 mm long, slightly puberulent or ciliate; pedicels 1-2 mm ne La dp ceo subopposit ovate to DUE scarious, 1.5-3mm long, ciliate but otherwise glabroi larly and minutely toothed. Flowers 4-merous; calyx 3-4 mm lone simply aricilate with pedicel, the orbe campanulate, 1-2.3 mm long, glabrous, the lobes deltoid to broadly ovate, 0.8-1.5 mm long, apically ciliate but otherwise glabrous; corolla with aestivation imbricate, | , unistratose, cylindric, 5-7.5 mm long, 2.7-4.5 mm in diam., creamy white with tinges of red or pink, glabrous, the lobes oblong, 1-1.7 mm long; stamens 8, included, 4.4-5.7 mm long, the filaments 3-4 mm long, ciliate, the anthers 2.8—4 mm long, bearing dorsally 2, slender, horn-like, ascendent spurs 0.2-0.6(-1) mm long originating from near the apex of the thecae, the thecae 1.5—1.8 mm long, granular-papillate, the tubules 1.2-2 mm long, dehiscing by introrse clefts 0.5-0.7 mm long. Berry spherical, 5-6 mm in diam., reddish-to Beien -purple, 4-locular. Distribution.—Mountain meadows, -— | pes, oak forests, and p Rica and western Panama (Bocas del Toro and Chiriquí provinces), between (130021 500-3500(-3900) m. This species was included by Standley (1920-26, pp. 1101-1102) as a member of the woody flora of Mexico, but its presence north of Costa Rica was questioned by Sleumer (1936) and the species was not included by Standley and Williams in the Flora of Guatemala (1966). The species is known only from the mountains of Costa Rica and western Panama. Flowering: throughout the year with the preponderance between Jan—Jul; fruiting: throughout the year. Vaccinium consanguineum is characterized by its pinnately nerved and serrulate-margined leaf blades, ill Costa 218 [| | E ul n.a a ID LI di den ndo £fT at i57 racemose inflorescences with small white to pink flowers, spurred connectives (these rarely vestigial), and glaucous berries. The species varies from small, densely branched shrubs to large trees to 10 m tall. Leaf size also varies greatly with trees often having leaves at the higher end of the range. The small-leaved populations of V. consanguineum look especially like V. floribundum. Both species have spurred (or vestigially-spurred) anthers, although the Costa Rican populations of V. floribundum lack spurs. Nevertheless, V. floribundum would appear to be its closest relative. Additional collections examined: Costa Rica. Alajuela: wooded slopes of Volcán Poá f list 2mid d towards Varablanca, ca. 8200 ft, 28 Jan 1971, Wilbur & Teeri 13647 (DAV, DS, DUKE, F, GH, LL, MICH, MO, NY, PMA, TEX). Heredia: woods around the La Ge del Barba at dud m and on Ge down to ca. 2500 m, 18 Dec 1974, el e id 18375 (DUKE). Car- tago: upper crest of Volcán Irazü covered with many feet of hrubs, 6 May 1971, Wilbur 14199 ee SE M trail to top of Volcán Turrialba es Finca Quemado between 2800-3200 m, 10 Mav 1971, Wilbur 14328 (DUKE); oak 14 km SE of El Empalme, 2600 m, 18 July 1977, Wilbur et al. 22873 (DUKE, NY). San José: roadside ge ca. 3.4 km SE of La rm and 10 km SE of El Asunción, ca. 2750 m, 1 Aug 1977, Wilbur et al. 23689 (DUKE, NY). P ordillera de Talamanca, 13 airline km S of the peak of Cerro Echandi, bid 30”- e 30"N, iid W, dal dM m, 9 Mar 1984, de et al. 25492 (DUKE, MO, NY). Limón: Cordillera de Talam t tern branch of the Río Teribe, between Río Sini and the continental divide at Cerro Bekom. 9?10'45"N, 83?03'30"W, 2500-2600 m, a 27 Mar 1984, Davidse et al. 26142 (CAS, DUKE, MO). PANAMA. Bocas del Toro: Cordillera de Talamanca, pa brega Massif, 9º07'N, 82º52'40”W, 3100-3300 m, 6 and 8 Mar 1984, Davidse et al. 25305 (DUKE, MO). Chiriquí: peña ridges S of Finca Lerida, 6000-7000 ft, 26 Jul 1947, Allen 4768 (F, GH, MO, NY, US); western slope of Volcán Chiriquí on open slopes, ca. 9800 ft, Wilbur et al. 11027 (DS, DUKE, GH, MICH, MO) 7. Vaccinium coro nos (M. Maren i Galeotti) Greg Set Centr. Amer. Bot. 2:274. 1881, where mis- takenly print r CAEN se VM Martens & Galeotti, Bull. Acad. Brux. 9:530. 1842, non Vaccinium cordifolium Stapf (1894). ber MEXICO. Vr- RAC t sur les laves du p de E ML Ge e IAM entre 6 et 7,000 pieds d'élévation,” Galeotti 1795 [HoLorvPE: BR! (ROS NY; € BR-S.P SOS 936), G! ds E neg. odds K! bap MINE 10619)]. T Hol si in 1965, at BR, | field 1 “Gay-lussacia cd en n Galeoti 2s eg ? handwriting of the label the speci f Galeotti He at ds G, and Kay EE EE ae 1 " BR Isotype - :spediesdi de las Visas tiui in = "sur ls aves ne Xalapa a 7000"; K = “on Se s as near Jiichlapa a at no ft.”). Vaccinium matudae Lundell, Phytologia 2:4. 1941 ; i C.La]. Tree: MEXICO. Chiapas: Barranca Honda, Siltepec, g ing ] s, 2600 m, Oct-Nov 1940, Matuda 4074 (noLorvree: MICHI; ISOTYPES: A! photo NY neg. 9630, CAS! NY). EE or terrestrial shrubs, ti ising from lignotubers; | hlets stout, rather irregularly angu- late, slightly | but soon glabrate, diss Leaf-blades persistent, stiffly coriaceous, evergreen, broadly ovate or E 2.5—5(-8) cm long, (1.3-)2-3.5(-5) cm broad, apically rounded to subacute, basally shallowly cordate to rounded, marginally subentire to prominently crenate-serrate, glabrous, lustrous, the venation pinnate, slightly elevated above and somewhat more prominently elevated and reticulate beneath; petioles 2-4.5 mm long, ca. 1.5 mm in diam., glabrous or at most very sparingly pilosulose. Inflorescences racemose or paniculate, 6-40-flowered, when paniculate then short-pedunculate and to ca. 8 cm long; ra- chises 3-6 cm long, 1-1.5 mm in diam., glabrous to weakly short-pilose or hirtelous; floral bracts caducous, lanceolate, ciliate, ca. 1.7-3.2 mm long; pedicels erect to secundly nodding, 5-10 mm long; bracteoles nearly basal, lanceolate, ca. 1-2.2 mm long. Flowers 5-merous; calyx 4.8—5.3 mm long, conspicuously articulate with pedicel, the tube obconic to a € 5mm long, deep rose, glabrous, the lobes deltoid, acute, 1-1.8 mm long, glabrous, non-ciliate; ivation imbricate, | , unistratose, cylindric to urceolate, 5-8 mm long, whitish tinged with deep pink or rose, glabrous externally, the lobes deltoid, subacute to rounded or obtuse, ca. 1-1.3 mm long; stamens 10, included, 6.5-7 mm long, the hlaments ca. 3-4.7 mm long, proximally ciliate, the anthers 3.5-4.2 mm long, lacking spurs, the thecae ca. 1.5-2.2 mm long, pappilose, the tubules ca. 1-2.5 mm long, dehiscing by short, introrse clefts. Berry ca. 5-6 mm in diam., 5-locular Distribution —Pine-oak forests, cloud forests, and montain thickets and forests, from the Mexican states TAF: JT de E H ERA = if A LA H If st 219 of Vera Cruz through Oaxaca and Chiapas, and mountainous northern Guatemala (Huehuetenango and Quiché ld ee 2000-3100 m. Flowering and DE ee the year. Vaccinium cordif I terized by its stiffly coriaceous, broadly with margins sub- entire to crenate, fis racemose (or rarely paniculate) inflorescence, a calyx that is articulate with the pedicel, stamens that lack spurs, and a dark rose to pink corolla. Vaccinium matudae Lundell, a most distinctive species was first described as Gay-Lussacia cordifolia Martens & Galeotti (21842), but was later determined to be a Vaccinium by Hemsley (1881). In attempting to form the new combination based on Martens and Galleotti’s binomial, he instead unfortunately pub- lished Vaccinium cordatum (1881), mistakenly, and carelessly also citing the basionym as Gaylussacia cordata Mart. & Gal., Bull. Acad. Brux. 9:529. 1842. As a result, instead of establishing the intended and needed binomial in Vaccinium, Hemsley created an illegitimate name (Article 52 of the ICBN) that is to be rejected as nomenclaturally superfluous. When Hemsley published Vaccinium cordifolium, there would have been no obstacle to proposing the binomial Vaccinium cordifolium based on Martens and Galeotti’s binomial, but after 1894 Vaccinium cordifolium Stapf had been published (Trans. Linnean Soc. Ser. II. 4:189. 1894) and therefore a combination based on Martens and Galeotti’s ee Re nb EH ina ater a However, when querried on this problem, three very J. McNeill, G. Moore, and D. Nicolson) all agreed that Hemsley's melee in transfering Gage cordifolium M Martens & Galeotti into Vaccinium as V. cordatum OM Martens & Galeotti) Hemsley, while also mistakenly citing the basionym as G. cordatum, should nevertheless be treated as a correctable error rather than as the publication of a later invalid synonym that Lundell corrected by publishing a new name since by that time Vaccinium cordifolium Stapf had been published for a species from Borneo. Consequently, the American species will be known as V. cordifolium (M.Martens & Galeotti) Hemsley as originally intended, while the Borneoan plant is provided with a binomial in the footnote below”. Additional collections examined: MEXICO. Veracruz: Municipio de Las Vigas; en las faldas del Volcancillo, por la carretera federal Jalapa-Las Vigas, 19?38'N, 97°03’W, 2000 m, 16 Aug 1979, Calzada 5483 (F); Las Vegas, Linden 420 (MICH); bosque de piño, 2300 m, El Volcancillo, 2300 m, 21 Jan 1976, Ortega et al. 114 (F, NY). Oaxaca: wet hardwood montane forest, Hwy. 175 ca. 31 mi N of Ixtlán de di 7500 ft, 20 Dec 1972, ee & Em 1675 (DUKE). Chiapas: evergreen cloud forest on the N and W slopes of Cerro Mozotal Huixtla to El Porvenir and Dee 3000 m, 30 Dec 1972, Breedlove & Thorne 31178 oss DUKE, MICH, MO, NY, RSA); Niquivil, Municipio de Motozintla, 2786 m, 15 May 1945, Matuda 5497 (DS, LL, MICH). GUATE- ALA , on slope with Pinus, Ge and Drimys, 6 mi N of Santa Eulalia along road to San Mateo Ixtatán, 9200 D 5 Feb 1965, Breedlove 8593 (DS pofC Chemalito, 3.5 mi W of Santa Eulalia, 3100-3150 m, Steyermark 19946 (F, NY). 8. Vaccinium breedlovei L.O. Williams, Fieldiana Bot. 31:173. 1965. Tre: GUATEMALA. HUEHUETENANGO: Sierra de los Cuchumatanes, 6 mi N of Santa Eulalia along road to Santa Eulalia, 2800 m, Feb 1965 (fl), Breedlove 8594 (HoLoTYrE: F! photo NY ne g. 9247; ISOTYPE: DS!) Ln ] Vaccinium tolbertianum Lundell, Wrightia 5:87. 1975. Tree: GUATEMALA. EL QuicHÉ juerón i y y thickets, 8000-8200 ft, 10 Aug 1964 (im fr), Proctor 25479 (HoLoTYPE: LL! isorvrE: LL!). Epiphytic shrubs; branchlets irregularly angled to + terete, glabrous, reddish-brown. Leaf-blades persistent, coriaceous, evergreen, ovate to ovate-lanceolate, 5-15 cm long, 2.5— oe cm vee pics acuminate to acute or rarely even rounded, basally rounded, marginally very finely but distinctly and minutely glandular crenat with the teeth 6-9 mm apart, glabrous on both surfaces, the venation pinnate, prominent on both RM and pennies) elevated E the ue order of branching forming a conspicuous reticulum, the midrib depressed into tl f channel; petioles stout, 4-11 mm long, puberulous. Inflorescences axillary or arising Fom leafless nodes, racemose or paniculate (2—4 branches), often pronouncedly secund, 10-20-flowered, glabrous throughout; rachises 3-7 cm long; floral bracts scale-like, triangular to narrowly deltoid, 1-1.5 mm long, glabrous; pedicels slender, (527-9 mm long, glabrous; bracteoles medially or more TTL H £ n Stapf (Trans, Linnean Soc. Ser. II. 4:189. 1894). The specific epithet recognizes Professor John H. Beaman, a native of tt tains of western North Carolina, for decades Ei 220 t tani i Texas 2( distally attached, narrowly triangular to linear-lanceolate, 1.5-2 m long. Flowers 5-merous; calyx ca. 3—4 mm long, articulate with pedicel, the tube campanulate, terete, slightly constricted near summit, ca. 1.5-2.2 mm long and 2 mm in diam., glabrous, the limb somewhat flaring, the lobes rounded to somewhat deltoid to triangular, usually abruptly acuminate, ca. 1-1.5 mm long, glabrous; corolla with aestivation weakly im- bricate, membranous, unistratose, broadly funnelform, strongly 5-angulate with pronounced medial nerves, 5-6 mm long, flaring distally to 6-7 mm in diam., glabrous, greenish to greenish-white often suffused with maroon, the lobes triangular, 2-2.5 mm long; stamens 10, included, ca. 5-5.5 mm long, the filaments ca. 3 mm long, glabrous, the anthers ca. 3 mm long, lacking spurs, the thecae ca. 2 mm long, papillate, the tubules ca. 0.8-1.5 mm long and 0.2 mm in diam., dehiscing by terminal, obliquely slanting pores; ovary 10-locular (?); styles ca. 5 mm long. Berry not seen. Distribution. —Known only from four collections on forested slopes of pine, oak, and Cupressus, from Chiapas (Mexico) and Lempira (Honduras), at ca. 2300-2670 m. Flowering: Dec—Feb. Vaccinium breedlovei is characterized by large (to 15 x 8 cm), dida nues B are finely crenate margined and prominently, reticulately and pinnately veined, calyx that is articulate with the pedicel, anthers lacking spurs, a greenish Bares and possibly falsely 10 SE ovaries (mature fruits are needed). Although unable to make better suggestions as to the closest relatives of the very distinctive Vaccinium breedlovei, we do not see the close relationships with V. poasanum and the relatives of that species (sect. Leptothamnia Benth. & Hook.f.) as originally suggested by Williams. The inflorescence pattern would seem to argue against any close relationship with species centered around V. poasanum, although the stamens do suggest such a relationship. Vaccinium tolbertianum Lundell was named from perhaps two incomplete collections made by the same collector at about the same time and place in Guatemala. The specimens (both LL) consisted of attached leaves, stems, and young fruits. The most unusual feature was the presence of coarse, stiff, rigid trichomes on a few centimeters of older stem on two of the six sprigs and on a few nearby leaves. Nothing like those trichomes has been seen in Meso-American vaccinia, but they are frequent in the genus Gaultheria. Other than for the presence of these gland-tipped trichomes, the specimens would fit reasonably well within the current concept of V. haematinum Standley & Steyermark. It seems they can still be best placed there in spite of the presence of those unique trichomes. Obviously further collecting may change our "(— of this little known variant. Specimens with flowers and trichome-bearing stems are unknown and it seems reasonable to conclude that without the presence of those coarse trichomes on a few centimeters of certain of the older branchlets, the species never would have been UM A ates 1 11 ee i J. MAT VIS" rh: 1 Jaa zt ] f AH e qu ,2300m m, 8 Dec 1986, Rreedlo ve & Mally CEE fr^ A CN HONDURAS T pi Ivi tam A Cal | NS of Río now 2670 m, 1,29 May 1991, Davidse 34869 (MO). 9. Vaccinium chihuahuense Wilbur & Luteyn, sp. nov. (Fig. 1). Tree: MEXICO. Cumuanua: Mojarachic, 1 Jun 1939, Knobloch 5791 (HoLorvrE: US! photo NY neg. 13031; sorre: MSU! NY frag. ex US Vaccinium dii Camp, ined. in herb. Frutex. Rami li vel elabrati li pilosuli, tereti. Foli id ist elliptica, 10-15 mm longi et 3- 8mm lati, crenulato- ig pilosis vel pubtisuli "s petiole 1-2 mm longi, silos yakicrescertia racemosa, axilaria, 4-9 mm longa; rhachis ne: praia et Li mate oe ae, 1-2 mm longae; n 1.5-4.5 mm longi, pasce. E pcc articulatus, ca. ] (0.822—2.4 mm lo et 1 5-2 mm diam " | 5, acuti, acuminati, erecti, ca. 1-1. o. mm lone et 0.8 mm lati bask puberulenta; corolla alba vel carnea, 3.5—5 mm longi má 2-2. 5 mm diam., pilosulosi; lobi corollae ca. 1 mm longi, erecti, oblongi. Terrestrial shrubs, 0.8—1.3 m tall; mature branchlets terete, striate, moderately to densely pilosulose with the hyaline trichomes + spreading at right angles to the stem's surface branchlets, to even glabrate, the bark thin, grayish to brownish, splitting longitudinally and NOH UIS twigs sharply and narrowly angled, striate, moderately to densely pilosulose. I y persistent, coriaceous, evergreen, the blades flat to concave, elliptic to ovate-elliptic, 10-15 mm jon 3- 8 mm broad, apically acute to broadly rounded, 221 A OOP ‘re ex S PSOE X ANT IM jj ‘rg j P CZ ES Ec 1. V, showina side. ventral wë F Li A L a D. stamens fruit. with i [| £ ^L Tel r d and immature 222 t tani | Texas 2( basally cuneately tapering to broadly rounded, marginally indistinctly crenulate-serrulate with the teeth terminating in a reddish, gland-tipped vein extension (glandular fimbriae), both surfaces densely puberu- lent or pilosulose with hyaline, spreading trichomes, the lower surface d deciduously glandular-fimbriate, the venation pinnate, inconspicuous to conspicuous above, the midrib impressed above and raised beneath, the secondary and tertiary venation impressed above; petioles somewhat flattened and canaliculate above, E mm long, densely pilosulose. Infl nces axillary, racemose, 2—8-flowered; rachises ca. 4-9 mm long, moderately short-puberulent; floral bracts and bracteoles caducous, oblong, acute to obtuse, 1-2 mm long, + glabrous; pedicels 1.5-4.5 mm long, densely puberulent. Flowers 5-merous; calyx ca. 3.2-3.5 mm long, articulate with pedicel, the tube ca. 1-1.5 mm long, ca. 1 mm in diam., moderately to densely puberulent, the limb (0.8-)2-2.4 mm long, densely puberulent, the lobes erect, narrowly triangular to ovate, acute to shortly acuminate, ca. 1-1.9 mm long, ca. 0.8 mm broad at base, puberulent; sinuses acute; corolla with aestivation imbricate, membranous, unistratose, cylindric, angled opposite the lobes, 3.5-5 mm long, ca. 2-2.5 m in diam., whitish-pink, externally sparingly to moderately pilosulose especially distally along angles, the lobes erect, oblongish, roundedly acute, ca. 1 mm long; stamens 10, included, ca. 2.7 mm long, the filaments ca. 1.7-1.8 mm long, pilosulosely ciliate to densely pilosulose, the anthers ca. 1.5 mm long, lacking spurs, the thecae ca. 0.5-0.7 mm long, + smooth, the tubules distinct, ca. 0. 8—1 mm long, dehiscing by introrse, oblique, tear-shaped pores; styles glabrous, ca. 4-5.5 mm long. Berry not seen. Distribution. —Known only from several collections in forest of Quercus-Pinus with thin, rocky, litosol soils, from the state of Chihuahua, Mexico, at 1950-2200 m. Flowering: Jun-Aug; fruiting: Oct. Vaccinium chihuahuense is characterized by having persistent pubescence in nearly all parts, small leaves that are marginally crenate-serrulate, an articulate calyx, anthers that lack spurs. Morphologically it resembles V. confertum, from which it differs most conspicuously in its persistent, dense pubescence of all vegetative and floral parts. Additional collections examined: MEXICO. Chihuahua: Mojarachic, Jan 1937 (st), Knobloch 5736 (MSU), Parque Nacional Cascada de Basaseachic, Mpio. de Ocampo, 2200 m, 12 Mar 1994 (st), Garcia et al. a iii a m, 2 Oct SE (im fr), Spellenberg et al. 8701 (NMC), 1950 m, 5Aug 1994 (fl), Yen & Estrada 2841 (DUKE, NMC, NY); Divisad eachic, 2100 m, 23 Jun 1987 (fl), T. R. Van Devender et al. 87-160 (NMC); Mpio. de Guachochic, Geen e Cusarare, 28 Jul 1973 (fD, Bye 4400 (NMC). 10. Vaccinium lundellianum L.O. Williams, Fieldiana Bot. 31:173. 1965. (Based on Malea pilosa Lundell, non Vac- cinium pilosum A. Chevalier ex Dop., 1930 Malea Gen Lundell, Amer. Midl. Naturalist 29:484. 1943. Tree: MEXICO. Chiapas: Mt. Malé, near Porvenir, 3200 m, 6-12 Jul 1941, tuda 4614 (HoLorypE: MICH!; ISOTYPES: À! F NY ne 75, CAS! LL! NY! photo NY neg. 9632). Epiphytic shrubs; branchlets terete, + blackish, + unm throughout with whitish trichomes. Leaf-blades persistent, coriaceous, evergreen, oblong to lance-oblong or even lanceolate, 3—4.5(—7.5) cm long, 0.8-2(-3) cm broad, apically obtuse to subacute, basally usually rounded, marginally entire or nearly so, occasionally with a few, remote glandular teeth, somewhat shiny above and dull beneath, persistently pilose beneath and puberulent or pilosulose above at first but glabrescent with age, the venation pinnate, visibly reticu- late on both surfaces with the midrib and primary veins elevated; petioles ridged and grooved, 2.5-8 mm long, densely pilosulose. Inflorescences axillary, solitary, racemose, congested, ca. 2-6-flowered; rachises subsessile, mostly 2-5 mm long, densely pilosulose, bearing small, imbricate bracts at base; floral bracts persistent, deltoid, ca. 1-1.2 mm long, pilosulose; pedicels slender, 3-7 mm long, pilosulose; bracteoles persistent, basal or attached below the middle, opposite to subopposite, lanceolate to linear, 1.5-1.6 mm long. Flowers 5-merous; calyx ca. 3-3.3 mm long, sharply articulate with pedicel, the tube subcampanulate to + globose, ca. 1.5 mm long, densely pilosulose, the lobes narrowly deltoid, acute, ca. 1.3-1.7 mm long; corolla with aestivation imbricate, membranous, unistratose, subcylindric, pentagonal, slightly swelling above, 8-10 mm long, white, pilosulose externally, the lobes erect or nearly so, obtuse, 2-3 mm long; stamens 10, included, 8.5-9.6 mm long, the filaments 6.3-7.2 mm long, pilosulose, at the base weakly attached to corolla, attached to the anthers dorsally near the middle, the anthers ca. 3.5-4 mm long, lacking spurs, the thecae ca. 2-2.3 mm long including the attenuate base, the tubules slender, ca. 1.6-2 mm long, dehiscing by introrse clefts; styles slightly exserted. Berry spherical, at least 4 mm diam. when immature, pilosulose, glabrate, falsely 10-locular. Distribution.—Known only in cloud forests, from a few collections from the Mexican state of Chiapas and the Guatemalan states of Huehuetenango and San Marcos, between 2000-3000 m. Flowering Jul; fruit- ing: immature m Vaccinium li is c] terized by its do to deni entire-margined and pilose leaves to g the rachis, pedicel, calyx, corolla, 1: 4.5(-7.5) cm long, congested racemes, and fruit), relatively long corollas (8— 10 mm long), and a Dos 10 cul berry. Additional co!lections examined: MEXICO. Chiapas: steep slopes with evergreen cloud forest, NW slope of Cerro Malé, 3-4 km W of El Porvenir along road from Huixtla to Siltepec, 19 Sep 1976, Breedlove 40381 REA CAS, MO, NY, RSA, Se sobre SECH e Rodeo-Siltepec, 2000 m, 29 Apr 1964, Matuda 37378 (F, MO). GUATEMALA (Tujimach), across river from San Juan Atitán, 2500-2900 m, upper forested slopes, 8 Sep 1942, Steyermark 52017 (F). San Marcos: montane cloud forest area on outer aa of Geen Kater ca. 8-10 km W of San Marcos, ca. 2300 m, 31 Dec 1964, Williams et al. 26886 (F, NY). HONDURAS N Park, 2435 m, T. Hawkins 920 (MO). 11. Vaccinium selerianum (Loes.) Sleumer, Notizbl. Bot. Gart. Berlin-Dahlem 13(116):124. 1936. Pernettya seleriana Loes., Bull. Herb. Boiss. ser. Il. 3:217. 1903. Tyee: MEXICO. Chiapas: “in distr. del Centro,” in silva montaña inter San Cristóbal Las Casas et Huitzán," 10 Mar 1896, Seler & Seler 2126a (oo oner: Bt, photo F neg. 4597; isorvee: GH!, photo NY neg. 11118). ITATEAJTATA LI Vaccinium RAUM A ANE KR Field Mus. Nat. Hist., Bot. Ser. 23:139. 1944. TYPE: : Cerro Huitz, bet | 500-2600 m, 14 Jul 1942, Steyermark 48572 (HoLoTyrE: F! photo F neg. 52552 and NY neg. 9656). Vaccinium minarum Standley & Steyermark, Field Mus. Nat. Hist., Bot. Ser. 23:219. 1947. Type: GUATEMALA. Zacapa: middle and upper uthern slopes of Volcán Gemelos, Sierra de las Minas, 2100-3200 m, 26 Jan 1942, Steyermark 43295 (HoLorwer: F! photo P neg. 52553 and NY neg. 9657; isorvrE: US! photo NY neg. 11119). MED epi A.C, Smith, Ceiba 3:185. 1953. Tyre: HONDURAS. Morazán: San Juancito Mts., ca. 6 km W of San Juancito, t, 2100 m, 8 Jul 1852 [1952], L.O. Williams 18567 (uororvee: US! photo NY neg. 9443:1 isotypes: DS! F! GH!) Vaccinium quicheanum Lundell, Wrightia 5:86. 1975. Tree: GUATEMALA. EL Quicut: Nebaj, 8000 ft, in pineland, 13 Jun 1964, Contreras 4983 (HOLOTYPE: LL!). Epiphytic or terrestrial, bushy shrubs or rarely small trees (0.5-)1-3(-4.5) m tall; branchlets ridged and grooved and somewhat angular to nearly terete, when young often reddish but becoming brown or even blackish-brown in age, at first glabrous to more typically moderately to even densely puberulent and then later becoming glabrate. Leaves often numerous and overlapping, the blades persistent, coriaceous, ever- green, narrowly to broadly elliptic to oblong-elliptic or even obovate-oblong to even lanceolate, 2—4.5(—6.5) cm long, (0.8-)1.2-2.5(-5) cm broad, apically acute to obtuse or even broadly rounded to tapering to an ultimately obtuse apex, basally rounded to acutely cuneate, marginally remotely and inconspicuously to conspicuously crenate-serrulate, often lustrous especially above, glabrous or nearly so on both surfaces to moderately iu EE and finely puberulent to pilosulose, na venation pinnate, the midrib lly somewhat elevated but mostly somewhat channe led - elevated, the lateral veins above + flush with the surface while beneath either flush or elevated and often comspictiously so and then forming a reticulum; petioles subterete, often canaliculate above, rugose, 1.5— = mm long, iid Or puberulent to short-pilose. Infl illary, racemose, 6—12- NEM ] J 3-12 mm long, sparsely to moderately puberulous; floral bract os narrowly la to oblongish, often concave, (1.5-)2-7 mm long, shortly ciliate, + scarious, EINE Bue or medially ciliate; pedicels reflexed or turned downwards, 1.5-5(-9) mm long, puberulent; | lly to distally placed, alternate, appressed, caducous to deciduous, broadly elliptic to lanceolate, 2-3.5 mm long, sparingly indistinctly ciliate, occasionally externally medially short-pilose. Flowers 5-merous; calyx 3-4.5 mm long, conspicuously articulate with pedicel, the tube spherical to oblong-cylindric, 1.5-2.2 mm long, glabrous to densely puberulent or pilosulose, the lobes deltoid to narrowly triangular, acute, (0.8-)1.2-1.8 mm long, externally and internally glabrous to externally moderately puberulent to pilosulose, marginally glabrous to minutely ciliate and often sparingly and inconspiciously glandular; corolla with aestivation imbricate, [| [| fal nad . In LI E- ef ET 5/43 i7 224 J membranous, unistratose, cylindric, 5—7.2(-9) mm long, ca. 3 mm in diam., deep rose to pinkish or white, glabrous both externally and internally or moderately to densely pilosulose externally, the lobes broadly oblong to deltoid, obtuse to acute, ca. 1-2 mm long; stamens 10(-12), included, 3.7-7(-7.7) mm long, the filaments 2.3-5(-5.7) mm long, attached dorsally to the lower third or half of the anther, flattened, densely ciliate, the anthers 1.8-4 mm long, lacking spurs, the thecae ca. 1.3-2.2 mm long, minutely papillose, the tubules slender 0.7-2.2 mm long, dehiscing by short, oblique pores or short clefts. Berry terete, 5-9 mm diam., reddish during maturation but apparently blackish at maturity, glabrate to sparingly pilosulose, 4-5 -locular. Distribution —Oak forests, evergreen cloud forests, and mossy thickets, in southern Mexico (Oaxaca and Chiapas), south through Guatemala (El Quiché, Huehuetenango, and Zacapa) into Honduras (Lempira and Morazán) and El Salvador (Santa Ana), from 1600-3200(-3700) m. Flowering: (Jan)May—Aug(Dec); fruiting: throughout the year. Vaccinium selerianum is a morphologically variable species characterized by its congested leaves that are elliptic to oblong-elliptic to lanceolate, 2-4.5 cm long, with margins crenate-serrate, short-racemose inflorescences, an articulate calyx, glabrous to pilose corollas, anthers that lack spurs, and a 4—5-locular ovary. Recently, as exemplified by the treatment in the Flora of Guatemala (L.O. Williams in Standley & Wil- liams 1966), two species have been recognized within what is treated here as a single species of Vaccinium. They are separated there by the following straightforward couplet: Flowers glabrous V. haematinum V. minarum V Flowers densely pubescent The brief descriptions there indicate that the calyces of V. haematinum vary from “sparsely and minutely puberulent or glabrate." Specimens have been examined with glabrous or at least glabrate calyces and pilo- sulose corollas. The leaves and vegetative features of both "species" seem to have the same range of variation. It cannot be claimed that a morphological continuum exists between the two extremes, but the variation exhibited by the small sample available suggests that a much larger sample would in all probability further blur the supposed distinctions between the named extremes. It seems that only one, not extremely poly- morphic species is represented. The extremes in pubescence of the calyx seem to be rather widely bridged, but there seems to be no similar blurring in corolla pubescense; those previously assigned to V. minarum have densely pubescent corollas while those attributable to V. haematium are glabrous. The flowers on the type specimens of V. minarum, V. quicheanum, and V. hondurense are e actly similar as to pubescence, although the leaves of V. quicheanum are more elongate, while those of V. minarum are smaller and narrower. As far as the long list of names is concerned, V. selerianum (Loes.) Sleumer (1936) has priority. TL ined: MEXICO. Oaxaca: dry led ding Llano e ca. 12 km ESE of Ixtlán de Juarez, 17º18'N, 96°22’W, 2000-2400 m, 6 Jun 1970, Graham Eron 1101 (MICH); vicinity ltepetl, openings of Cupressus-Pinus woodland, ca. 5 km E of summit, 2800 m, 9 Aug 1950, Hallberg 884 (MICH) | DEDE Ixtlán and Valle Nacional, just below summit at Cerro Pilon, 17º35'N, 96?30"W, ca. 8000 ft, 20 Jun 1969, Webster S Breckon 15350 (GH). Chiapas: steep forested slopes on the SE side of Cerro Tres Picos ang n nip near summit, 2100—2500 m, 28 May 1972, Breedlove 25443 (DS, MICH, MO, NY, RSA); cloud forest g itas to Campo Alegre, 2300 m, 24 Oct 1976, Breedlove 41099 (DS, MICH, MO) ` GUATEMALA. El Quiché: El Boquerón; i loud forest, 8000-8200 ft, Proctor 25435 (MO). ] ll in pine forest between Km 324 = 325 on Ruta Nacional 9N Ee Chemal and a Ixcoy, 3140 m, 4 Aug 1959, Beaman 3051 (MSC) | San Juan Ixcoy, 3700 m, 12-23 Jan 1966, Molina et al. 16438a (E, NY, US). cin on summit between Lema El Picacho aná Cerro de Moños, 2000-2600 m, 16 Jan 1942, a ien (F, US. HONDURAS Celaque o, 2750 m, 18-22 Nov 1974, Hazlett 2262 (F, MO). M t la floresta Humala y nebulosa de Montaña La Tigra, entre El Piliguin y Prado de Fatima, 1800 m, 30 Dec 1962, Molina R. 11217 (F, LL, NY, US). EL SALVADOR. Santa Ana: bosque nebulosa, de Montaña Montecristo, 2300 m, 23 May 1963, Molina R.. 12683 (F, LL, NY, US). 12. Vaccinium talamancense (Wilbur& Luteyn) Luteyn, Brittonia 53:444. 2001. Macleania talamancensis Wilbur TEM) M | "1 1 c E ERA E IA A La H 1 “at 225 & Luteyn, Brittonia 29:263, fig.3. 1977. Type: COSTA RICA. San José: al il from C Chirripó via Los Angeles, N of Río Talari, 3200-3400 m, 19-22 Jan 1970, Burger & Leisner 7401 (HOLOTYPE: M JKE!; isotypes: Fl MO! NY!) Terrestrial shrubs, 0.5-1 m tall; mature branchlets terete, striate, glabrous; twigs subterete, bluntly and broadly compressed, puberulent to densely short-pilose, glabrate. Leaves imbricate, the blades persistent, coriaceous, evergreen, obovate to more typically oblong-elliptic, 2.5—4.7 cm long, 1.2-1.8(-2.3) cm broad, apically acute or rounded but with a 1-2 mm long apiculus, basally rounded, marginally indistinctly and minutely serrulate, the actual margin a lighter color and seemingly a bit thinner than lamina proper, es- sentially glabrous above except finely whitish, appressed-pilosulous near the base along midrib and incon- spicuously ciliate especially proximally, indistinctly and sparingly whitish, appressed-pilosulous beneath basally and also bearing appressed, reddish-brown, glandular-fimbriae «0.2 mm long, provided with 1-2 reddish, circular basal glands, the venation pinnate with 3—4 arcuate-ascending lateral nerves per side, the midrib slightly thickened and raised proximally but impressed distally, lateral nerves impressed above, all veins pene o Sene and of a lighter color than lamina proper; petioles thick, flattened or broad] lightly winged throughout, TUBOS; 2-3 mm long, moderately Pubetuieus to RAL id e short-pilose. Infl illary, racemose, 10—15-flowered; racl gly and sharply angled, 1-2(-4 cm long, pilose with soft, short, white trichomes; floral bracts appressed, linear to lanceolate 3-4 mm long, puberulent; pedicels 6—10 mm long, softly pilosulous or puberulous, bearing distally a series of glandular fimbriae; bracteoles basal, appressed, linear to narrowly lanceolate, 2-3 mm long, puberulent, marginally glandular-fimbriate, without dorsal glands. Flowers 5-merous; calyx 4.5-5 mm long, articulate with pedicel, the tube campanulate to cylindric, ca. 2.5-3 mm long and in diam., longer than limb, moderately puberulous with whitish trichomes +0.1—0.2 mm long, the lobes triangular, acute, 1.2-1.8 mm long, puberulous exter- nally and internally; corolla with aestivation apparently valvate, membranous, weakly bistratose, cylindric, terete, 8-13 mm long, 4-6 mm i uiui pink to red, appressed- to spreading-puberulous externally with moderately abundant hyali - 0.4 mm long, moderately to densely pilose with hyaline trichomes within, the lobes — reflexed, la 1.5-2 mm long, pink to red although sometimes becoming pale pink to whitish, densely pilosulous both externally and internally; stamens 10, included, 7-7.5 mm long, the filaments 3.5-4 mm long, glabrous proximally but marginally with appressed-pilosulous hyaline trichomes distally, the anthers 4.5-5 mm long, lacking spurs, the thecae 1.8-2 mm long, smooth, basally strongly incurved, the tubules 2.5-2.7 mm long, dehiscing by oblique pores ca. 0.6 mm long; styles exserted. Berry spherical, 8-10 mm diam., blue-black, short-pilose, 5-locular. Distribution. —Known only in upper oak forests and disturbed páramo habitats, from the upper slopes of Cerro Chirripó, from ca. 2500-3819 m. Flowering: Jan, Mar-Apr; fruiting: Dec, Mar-Apr. Vaccinium talamancense is characterized by its leaves with ui margins Rasa and nie ser- rulate, red to pin] llas that are ap d- to spreading-pul ternally and moderately to densely pilose internally, anther connectives SCH lack spurs, and a pilose berry Chat is not glaucous. Its interspecific eege are uncertain at this time. Additional coll ined: COSTA RICA. S e é: Cordillera de Talamanca, C Chi if: f just below the param: it] y dead Davidse & Pohl 1594 (DUKE, MO); Parque Nacional Chirripó, Cuenca Térraba- Sierpe, b fC Crestones eee m, 7 Dec 1996 SC Alaro 1011 (MO) Pacific slope, trail to Cerro Urán, 3000 m, 7 Apr 1995 (fD, González 662 (INB); sendero de la , Monte Sin Fe, 2590 m, 18 Mar 1999 (fl, fr), Luteyn & Chaverri 15415 (CR, INB, NY), trail beyond Rio Talari to Valle de Los Leones, 3200-3300 m, 20 Mar 1999 (fl, fr), Luteyn & Chaverril5430 (AAU, CAS, CR, DUKE, F, INB, K, MEXU, MO, NY, TEX, US); Sabana de Los Leones, Cerro Lol 3200 m, 26 Jan 1996 (fI), Morales 5177 (F, INB, NY); between La Piedra and lower refugio, 17-28 Mar 1976 y Weston 10134 (CR). 13. Vaccinium floribundum Kunth in H.B.K., Nov. Gen. Sp. Pl. 3:266, t. 251. 1818. Tyr: PERU. Amazonas: "Crescit in montanis Peruviae, prope Carxamarca, alt. 1700 hex. Floret Augusto,” Humboldt & Bonpland s.n. (notorvee: P-HBK! ISOTYPES: B- Willd. 7353-1! PI). Densely branched, terrestrial shrubs 2-8(-12) dm tall, sometimes ane en EE to 10 cm in diam.; branchlets few, angled to terete, glabrous or puberulent. Leaf-blades p t, coriaceous, evergreen, [| a £ al ns H Im LI P MERCI £T [4 UN 226 highly variable in shape and size from very broadly ovate or nearly orbicular to elliptical or ovate-oblong, 0.4-1.5(-2.5) cm long, 0.3-1.2 cm broad, apically acute to obtuse, basally rounded to somewhat cuneate, marginally thickened and + revolute and shallowly crenate-serrulate, occasionally inconspicuously and sparingly spreading pilosulose above and this pubescence more abundant basally and + glabrate except for the moderate to dense puberulence along the midrib, iie ie SE EE to cdm An pe with E | appressed to somewhat ascendent, glandular 1 drop away, the venation pinnate; petioles flattened above, 2304) mm bue very shortly puberulent. In- florescences axillary, racemose (very rarely weakly paniculate), 6—10- flowered; rachises strongly angulate, 1-2(-4.5) cm long, puberulent to glabrous or very nearly so; floral bracts broadly ovate to oblong, 2.5-4.5 mm long, (223—4.5(-5.5) mm broad, minutely appressed puberulent on the abaxial surface to nearly gla- brous, minutely ciliate and occasionally glandularly ciliate; pedicels 3-4(-6) mm long, usually spreading puberulous but occasionally glabrate; bracteoles medially to basally attached, + ovate to lance-ovate or even narrowly lunear-oblong, 2.5-4.8 mm long, 1.5-2.5 mm broad slightly appressed puberulous externally to glabrate, marginally minutely ciliate and occasionally also Md EE Flowers 4—5-merous; calyx ca. 3—4 mm long, sharply articulate with pedicel, the tu rt-cylindric, 1.5—1.8 mm long, 1.2-1.8 mm in diam., glabrous, the lobes deltoid to ovate- gf acute, ca. 1-1.5 mm long, glabrous, usually ciliate; corolla with aestivation imbricate, membranous, unistratose, cylindric, 5-6.5 mm long, 2-3 mm in diam., white to roseate, glabrous, the lobes + erect, deltoid, acute, ca. 1.5 mm long, glabrous; stamens 8 or 10, included, 3.8-6 mm long, the filaments 2-2.5 mm long, flattened, densely spreading ciliate with hyaline trichomes, the anthers 2.53.5 mm long, lacking spurs, the thecae 1-1.5 mm long, densely papillate or sharply and minutely spinulose, the tubules 1.5-2 mm long, dehiscing by introrse, oblique pores. Berry spherical, ca. 4—6 mm in diam., dark bluish-black, sometimes reddish when immature, 4—5-locular. Distribution.—Disjunct from the South American Andes. Mossy thickets of the highest reaches of the Talamancas of Costa Rica, at 1400-3450 m. Flowering: Feb—Apr, Oct; fruiting: Feb-Mar. Vaccinium floribundum is characterized by its small leaves the blades of which are pinnately nerved and minutely crenate-serrate margined, inflorescences with small white to pues flowers, spurless con- nectives, and glaucous berries. In Costa Rica the stamens lack spurs, wi in South America have at least vestigial spurs. Vaccinium floribundum is most closely related to V —M as mentioned herein. lditional coll ined: COSTA RICA. S C de la Muerte, Cerro Sábila, 80 RUE pd 28 Mar di an Horn 131 (WIS); wet páramo thicket, Cerro de las tala 2700-3000 m, 1 Jan 1926, Standley & Valerio 43697 (1 summit of Cerro Sákira, 3417 m, 29 Aug 1969, Weston 6005 (P); pu Nac. RED. ni de Los Condos 3500-3600 m, 19 Mar 1999, Luteyn & Chaverri 15417 (AAU, CAS, INB, NSW, NY, STE, TEX der, Cerro Echandi, 9°02’, 82º49"W, 3050-3160 m, 22 Aug 1983, Davidse et al. 23856 (MO). Limón: Cordillera de Talamanca, Cerro mile 9*16'N, 83*02/30"W, 3200—3350 m, 24 Mar 1984, Davidse et al. 25979 (MO). 14. Vaccinium confertum Kunth in H.B.K., Nov. Gen. Sp. Pl. 3:265, t. 250. 1818. Ter: MEXICO. "prope Moran et Cerro de Oyamel, alt. 1400 hex," Bonpland 4099 (HoLotyre: P-HBK! frag. F! isotypes: Bt, B-Willd. 8314-1! PD. The isotype in the Willdenow Herbarium has the label as "Humboldt & Bonpland 4099." Vaccinium eriocladum Dunal, DC. Prodr. 7:571. 1839. Vaccinium confertum var. eriocladum (Dunal) Sleumer, Notizbl. Bot. Gart. Berlin . 1936. Tire: MEXICC. Veracruz[?]: inter Tampico et Real ue ee 20 Mid 1827, Sedan 254 n Li G-DC! ne 1 Tas NY s.n., frag. NY!; ISOTYPES: a BMI, FI, ECH Esch wo TI | É V Weeer BEEN both have the “ex. 12” written on the label too. Then there are two Weg ll ER t OXF f which have the “ex. 12" on the label. All sheets are definitely V confertum! Vaccinium adosada Benth. , PI. Duet 65. 1840. Tree: MEXICO. Oaxaca: “in montibus Carmen in jugo la Sierra dicto,” Hartweg 4 TYPE: K-Herb. Peia |, frag. US!, photo NY neg. 11116; isorvees: CGE (x2)! a iq ! NY! pini M.Mart tti, Bull. Acad. Brux. 9:531. 1842. Type: MEXICO: " lalpan, Pelado de San Andres, etc.) , entre 7,500 et 9,500 La ` Galeotti 1818 ee BR!; isoTYPES: BR, 2x! F! NY, 2x! P 2x! Us! photo NY neg 11117, W! photo F neg. Ser Ti mj, E , and US give “Oaxaca, Llano Verde" as the locality. ts at BR, th labeled as “TYPUS” has the locality as OMIM num dici Sierra 7000"; another unannotated isotype has no locality data on its label, and the third isotype has *Oaxaca, Llano Verde.' — XN i p" a "IT e FPE" " IT "m 227 P Fernald, Proc. Amer. Acad. 36:496. 1901. Tre: MEXICO. C tai ER San Cristóbal Las Casas and Huitzilán, 10 Mar 1896, Seler & Seler 2126b (HOLOTYPE: GH! photo NY s.n.). Densely branched, terrestrial shrubs, 2-6(-10) dm tall; branchlets strongly ridged and grooved and hence appearing angulate, very densely puberulent or pilosulose to shortly hirsutulose (or reportedly very rarely glabrous). Leaf-blades PESCE thick-coriaceous, evergreen, oval to oblong-ovate, 6-15(-30) mm long, 4—10(-15) mm broad, apically hat rounded, basally rounded, marginally finely but conspicu- ously crenate-serrulate, the upper surface finely e inconspicuously pilosulose to glabrate, deep green and shining while beneath pale and usually with scattered, inconspicuous, appressed, gland-tipped trichomes, the venation pinnate; petioles semicircular, canaliculate above 1-2(-3) mm long, puberulent to pilosulose. Inflorescenses axillary, densely compact, racemose, 3—7-flowered, almost always shorter than the subtending leaf; rachises mostly 8-12 mm long; floral bracts EE cuculate, oblong, ca. 2-4 mm long; pedicels (1232344 mm long, finely short-pubescent; | t l, caducous, cuculate , oblong, 1.2-1.5 mm long. Flowers 4—5-merous; calyx 3-3.3 mm long, conspicuously iene with pedicel, the tube campanulate or globose, ca. 1-2 mm long and in diam. at anthesis but enlarging to ca. 3(-4) mm in diam. in fruit, glabrous, the lobes deltoid, acute, ca. 1-1.5 mm long; corolla with aestivation imbricate, membranous, unistratose, cylindric to weakly cylindric-campanulate or weakly cylindric-urceolate, 4-6.5 mm long, white to pinkish; stamens 10, included, 4.7-6.4 mm long, the filaments (1.5-)2.4-3.5 mm long, ciliate, the anthers 3.23.6 mm long, lacking spurs (or rarely bearing dorsally 2, minute, vestigial spurs), the thecae ca. (121.42 mm long, the tubules ca. (1-)2-2.3 mm long. Berry 4-6 mm in diam., bluish or blackish with a glaucous bloom, 4—5-locular. Distribution — Mont forested slor f pine-oak, rocky meadows, and clearings, from northern Mexico (Chihuahua, Tamaulipas, San Luis Potosi, Nayarit, Aguascalientes, Queretaro, o: Vera Cruz, Puebla, Mexico, Jalisco, Guerrero, Oaxaca, and Chiapas), south into Guatemala (Chimaltenango, Huehuetenango, Quetzaltenango, San Marcos, Solola, and Totonicapan) and Honduras (Comayagua, Lempira, Olancho, and Ocotopequez), at 1538-4038 m. Flowering: throughout the year but especially in Jan—Jun; fruiting: throughout most of year but especially in Mar-Aug. Vaccinium confertum is characterized by small, more or less concave leaves, 6-15 mm long (rarely to 30 mm long) with finely crenate-serrulate margins, densely bus ae racemose inflorescences of 3-7 flowers that are usually shorter than the sul ling leaves, an articulate calyx, anthers that lack spurs (or are rarely = and a a 4—5-locular Sege Its similarities with V. chihuahuense are mentioned above. A dle: 11 l. 44a CwWir gr CL: Ar” lupe and Calvo, 7200 ft, 12 Oct 1959, niin 6 SE 23048 (LL, NY); Sierra ue 15 mi E ge EE 1-2 Oct 1898, Goldman 183 (GH, US); Lagotera, 5-6000 ft, 21 Jul 1965, Pennington 95 (TEX). T i Gómez Farias, Apr 1960, Duke M3543 (AAU, MO, NY). San Luis P í: San Luis Potosí, 6000- 8000 ft, Parry & Palmer 561 (F, GH, MO, NY, PH, US); Cerro Grande, 8 km al NW de Guadalcazar, 2100 m, 19 Jun 1955, Rzed i 5960 (F, MO, TEX). iu inp Hose en barranca, 1.5 km al S de Santa Teresa e Nayar, Municipio de Jesús Maria, 8 Jun 1978, Diaz L. 9541 (MICH). A del Laurel, near the Jalisco-A ler, ca. 10 mi SE of Calvillo, oal ts, 2500-2700 m, 26-28 Aug 1960, M 18445 (MICH). Queretaro: ladera de cerro con bosque de coniferas, 2 km al SW de Pintal de Amoles, 2400 m, Fernández 2710 (MO, NY). Hidalgo: bosque de pino y encino, alrededores de Zacualtipán, 1 Feb 1964, Gonzales Q. 311 (DUKE, MICH, MSC, TEX); bosque de encino, 1.5 km al SSW de Tezuantla, 2950 m, 29 Jun 1975, Medina 472 (CAS, NY). Veracruz: on old lava, La Joya, Perote, 6500 ft, 23 Sep 1938, Balls 5519 (NY); Peñas-Largas, cerca de Tezoantla, Municipio de Real de Monte, 2750 m, 21 Apr 1966, Rzedowski ee F, E Puebla: pine-oak woodland, 74 kms S of the Puebla-Orizaba Rd. (Mex. Se on Ges to Nicolas Bravo, 2560 m, 15 Apr 1985, Gri 09 (CAS, MO, NY, TEX); bosque de piño y encino, ca. 1 km al NE de Hon Pahuatlán, 2050 m, 13 v 1978, Perino 3336 (CAS, F, MO, WIS); wet cid of river ou below Trinidad ion Works, 5700 ft, 25 Apr 1904, Pringle 8911 (A, CAS, CU, F, GH, LL, MIN, MO, MSC, NY, PH, US). ^ str aaepe 18 Feb 1935, Hinton et al. 7389 (DS, F, LL, MO, NY, US). Jalisco: long] Fl Cl El Guisar i án, 2160 m, 17 Aug 1980, Breedlove & Almeda 45724 (CAS, NY); trail from: Real Alto to San Sel án, 2000 m, 3 Feb 1927, Mexia 1630 (A, CAS, DS, F, GH, MICH, MIN, MO, NY). Guerrero: Vertiente SW del Cerro Teotepec, ca. 17°29’N, 100?12"W, 2900 m, 29 Jan 1965, Rzedowski € dech 231 (CAS, MICH, MSC). Oaxaca: near Tres Cruces S of San José El Pacifico, 10 Feb 1945, Alexander 665 (MICH, NY); top of tl Zepoaltepetl, 19—27 Feb 1937, Camp 2649 (DUKE, NY, US); Distrito de Ixtlán; Municipio de ned M 0.5 A fo Hwy. 175 on road to Cerro Humo Chico, 3000 m, 23 Feb 1981, Martin 341 (CAS, MSC, MO); Cerro P I ltepec, 2950 m, Rzed 28855 (CAS, MICH, TEX, US). Chiapas: 228 t tani i Texas 2( steep slope, evergr loud forest on the N and W slope of Cerro Mozotal below the mi t ] t] df Huixtla to El Porvenir and Siltepec, 30 Dec 1972, Breedlove & Thorne 31182 (DS, LL, MO, RSA); Mount Hueitepec, 3 mi al NW of San Cristóbal de las Casas, 7100-7500 ft, 25 Mar 1949, Carlson 1582 (F, MICH, NY); Mt. Tacana, 2000-4038 m, Aug 1938, Matuda 2363 (A, F, MICH, NY, US). GUATEMALA. Chimaltenango: north side of Volcán de Fuego, ca. 3600 m, 6 Aug 1960, Beaman 4024 (GH, MSC); Volcán dee crater, 13 ids » 7 Feb 1907, Kellerman s.n. iie Buena Vista above Tecpán, 10,000 ft, 25 Mar 1933, Shutch 339 (A, US). Cuchumatanes, | land San Juan Ixoy and kms ~ a 325 on Ruta ae 9 N, ca. 3140 m, 4 Aug 1959, I B 3053 (DUKE, GH, MSC, NY, TEX, US). Quetzal: d 00-3800 m, 22 Jan 1940, wa adiu 34861 ~ San Marcos: trail hen San Sebastian to Tajamolso Ste, cloud forest near summit, 14 Aug 1979, Boeke & Utzschneider 2900 (NY te, S of Volcán Tajumulco, 11 Mar 1971, Plowman 3055 (GH, TEX). Sololá: mountain slopes above Santa María, 10, 500 ft, 14 April 1937, ersten 12363 (CU, F). Totonicapán: ca. 21 km E of Totonicapán at km 153 on Ruta Nacional 1, ca. 3260 m, 8 Aug 1959, Beaman 3123 (GH, MSC, TEX, US); in moist bank of María Tecun, 3000-3600 m, 12-23 Jan 1966, Molina R. et al. 16380a (F, NY, US); summits of Sierra María Tecum between Los E ntros and Totonicapán, 3200 m, Williams 14269 (F, GH, MO, US). HONDURAS. Lempira: Montaña de Celaque, 2650 m, 18-22 Nov 1974, Hazlett 2266 (MO). Ocotepequez: Cerro El Capitán, 10 km E of Belén Gualcho, 2-15 Apr 1977, Nelson et al. 4016 (MO). Olancho: alrededor de la Picucha, 13 km NE de Catacamas en el Parque Nacional de Agalta, 2 June 1992, Mejiá Dario 149 (DUKE). 15. Vaccinium wilburii Almeda € Breedlove, Brittonia 44:50. 1992. Tre: MEXICO. G : ridge with Pinus, Quercus, Clethra and Cleyera, W of Puerto El Gallo along road to Toro Muerto, 2530 m, 9 Oct 1986 (fD, Breedl me 65040 (HOLOTYPE: CAS!; isotypes: DUKE! MEXU, MICH, MO! photo NY neg. 13045, NY). Epiphytic, erect to pendent shrubs with branches 1-2 m long, arising from swollen lignotubers up to 2 dm in diam.; upper branchlets ridged and grooved and there densely spreading puberulent, lower ones becoming terete and glabrate. Leaves densely arranged with most internodes ca. 4—/ mm long, the blades persistent, coriaceous, evergreen, lanceolate to lance-elliptic, mostly 2-3.4 cm long, (5-)7-14 mm broad, apically acute the tip itself often blunt, basally broadly cuneate to obtuse, marginally indistinctly and bluntly crenate- serrate, with a narrow, oni c and E border, the venation pinnate, with all veins conspicuously elevated on both the upp t least when dried; petioles subterete, flattened to canaliculate above, 2-4 mm long, puberulent to glabrate. Infl nces axillary, loosely racemose, sometimes corymbose, 5-10-flowered; rachises sharply angled, 1.5-2.5 cm gës nae Or did so; floral bracts narrowly lan- ceolate to linear, scarious, 2-3 mm long, marginally ciliate; p y angled, ca. (8212-14 mm long, ca. 0.5 mm in diam., glabrous or nearly so; ew m (SHORE lanceolate to linear, scarious, 1.52.2 mm long, distally ciliate. Flowers 5-merous; calyx ca. 5 mm long, articulate with pedicel, glabrous, the tube ca. 1.5-2.5 mm long, ca. 2 mm in diam., the lobes narrowly triangular to deltoid, acute, ca. 22.5 mm long; corolla with aestivation imbricate, | , unistratose, cylindric-urceolate, angulate, 7.6—10 mm long, ca. 4 mm in diam., pink, glabrous externally, the lobes + oblong, obtuse 1-1.5 mm long; stamens 10, included, ca. 6 mm long, the filaments 2.53 mm long, flattened, attached medially to anthers, glabrous to ciliolate, the anthers ca. 4-4.2 mm long, lacking spurs, the thecae ca. 1.52.3 mm long, slightly papillose, the tubules slender, distinct, ca. 1.6-2.2 mm long and 0.1-0.2 mm in diam., dehiscing by introrse, oblique, terminal pores; styles ca. 11 mm long. Mature berry unknown, but at least 5 mm diam. when immature. Distribution. —Known now only from the montane, pine-oak forests of the Sierra Madre del Sur of the Mexican state of Guerrero, at ca. 2500—3000 m. odds and ei Jul, Oct. Vaccinium wilburii is characterized by its lanceolate t p lal calyx with compara- k cylindri olat llas that are angulate and glabrous throughout. tively long lobes, and uniformly p ) Although we have not studied this species in the n the suggestions of Almeda and Breedlove (1992), who stated that the relationships of this species seem to lie with V.selerianum (V. haematinum to them) and to a lesser extent with V. lundellianum, are herein followed. Additional collecti ined: MEXICO. G Distr. Galeana, Piedra Ancha, oak and pine forest, 3000 m, 29 Jul 1939, Hinton et al. 14508 (NY, US); Sierra Sur, Mpio. Metlatónoc, Xatu Yahta, al W de Coicoyán ( (Oaxaca), terrenos de Atzompa, 17?15'N, 98º20'W, 2600 m, 20 Aug 1989, A. de Avila 737 (CAS). 16. Vaccinium jefense Luteyn & Wilbur, Brittonia 29:272. 1977. Symphysia jefensis (Luteyn & Wlbur) vander Kloet, Taxon 53:97. 2004. Tre: PANAMA. PANAMÁ: at the top of Cerro Jefé, 1000 m, 5 Jun 1973 (fl), Luteyn & Kennedy 3955 (HOLOTYPE: DUKE!; sorres: MO! NY! Rasen Mila E "r a CRA a le “Ta D TI E 229 Lianoid epiphytes or arching terrestrial shrubs to 2 m tall; branchlets subterete, glabrous, green becoming reddish-brown, drying brown, the bark thin, cracking into longitudinal strips; twigs subterete, glabrous to densely hirsute distally. Leaf-blad istent, coriaceous, evergreen, elliptic or ovate-elliptic, (4.52)8—12(-20) cm long, (2-2)3-7(-8.5) cm broad, apically acuminate to short-acuminate, basally rounded and cordate to subcordate, ae entire, a EE Ane and pat pensam, e olive to gray Pa DINE glabrous above, sparsely t the principal vein S, with numerous, minute, glandular aba: the venation 5— 9). menea: or sometimes appearing weakly pinnate with 3 pairs of lateral nerves, the veins impressed above and raised beneath (i.e., the leaves somewhat bullate), proximal 1-2 cm of midrib er MR EE above; petioles terete, rugose, 5-9 mm long, 223.5 mm in diam., hirsute to glabrate. Inflore llary (rarely appearing terminal), racemose or subcorymbose, (6-)8-17-flowered; rachises bene 1-1.5 cm long, glabrous or sparsely hirsute, densely glandular-fim- briate; floral bracts linear-lanceolate, conspicuously nerved, (2.523—9 mm long, ca. 1 mm broad, hirsute, glandular-fimbriate; pedicels terete, striate, ca. (6—)14-26 mm long, 1 mm in diam., sparsely hirsute, densely glandular-fimbriate; bracteoles linear-lanceolate, (2.5-)5-8 mm long, ca. 0.5 mm wide, hirsute, glandular- fimbriate. Flowers 5-merous; calyx 6-10 mm long, articulate with pedicel, sparsely to moderately hirsute and pun S the tube cylindric, conspicuously bluntly and roundedly 10-ribbed or fluted, slightly angled opposite the si 2-4 mm long, 2-5 mm in diam., the limb campanulate, 4-6 mm long, the lobes oblong to ovate, acute, often basal slightly imbricate, (2—)4.5-6 mm long, 1.7-3 mm broad, conspicuously nerved; corolla wi tion valvate, thinly fleshy, weakly bistratose, cylindric, 10-13.5 mm long, 5.5-10 mm in diam., green, sparsely hirsute and glandular-fimbriate especially on the surface of the lobes, the lobes triangular, 2-4 mm long and broad; stamens 10, included, ca. 8-10 mm long, the filaments distinct, ca. 2.5—4 mm long, broadest basally, marginally densely sericeous especially on the connective, translucent when fresh, the anthers ca. 7-8 mm long, lacking spurs, orangish-brown when fresh, the thecae ca. 3-3.7 mm long, strongly incurved at the base, papillose, the tubules ca. 3.5—5 mm long, sparsely pilose along in- ner surfaces, dehiscing by latrorse clefts ca. 1.5—2 mm long; styles included, 9-10 mm long, glabrous. Berry spherical, ca. 12 mm in diam., purple, 5-locular. Distribution —Elfin forests, d moss-draped thickets of shrubs and small trees on mountain sum- mits and upper slopes, in P Chiriquí thru Panamá provinces, at 750-1400 m. Flowering: Jan-Jul and sporadically through the rest of the year; fruiting: throughout most of the year. Vaccinium jefense is characterized by its weakly bullate leaf blades that are basally rounded and subcor- date to cordate, conspicuous leaf nervation, glandular-fimbriate (sometimes densely) inflorescences, long and narrow bracteoles, elongate and conspicuously striate calyx limb and lobes, sparsely hirsute corollas (especially on the lobes), and anther tubules that are sparsely long-pilose along the inner surfaces and de- hisce by latrorse clefts. It resembles several other green-flowered species of Vaccinium and its morphological relationships are mentioned below in the discussion of V. furfuraceum. Additional collections examined: PANAMA. Chiriquí: Fortuna, camino de quebrada Bonita, hacia el este del Río Chiriquí, 1150 m, 7 Apr 1987, Valdespino et al. 560 (NY). Veraguas: ridge of Cordillera de Tute, along trail to Cerro Tute, 3-4 km past Escuela Agrícola Alto de Piedra, just W of Santa Fé, premontane and montane rainforest, 8°32’N, 81°07’W, 800--1400 m, 20 Mar 1982, Knapp & Kress 4367 (MO). Coclé: near sawmill above El Copé, ca. 3000 ft, 4 Apr 1978, H 12363 (MO). Panamá: | mi después de La Eneida cerca de Cerro Jefé, + 750 m, 8 Aug 1968, sioe 6 oe 955 (DUKE, MO, PMA); La Eneida, region of Cerro Jefé, 12 Jul 1968, Dressler 3541 (DUKE, MO, PMA); path I ida, ca. 1000 m, 2 Jul 1970, Lutyen & Foster 1124 (DUKE, F, MO, US); top of Cerro Jefé, 9 Jul 1966, bun et al. 4344 (DUKE, MO, SCZ). 17. Vaccinium furfuraceum Mia & E Pida al 1609, fig. 2. 2005. Ter: COSTA RICA. Limón: Cantón de Talamanca, fila d tío i, Muragubishi, 9?22'50"N, 82º56'50"W, 700 m, 14 July 1989 (fl), G. Herrera 3286 (HOLOTYPE: INB!; isotypes: DUKE! F! NY) Much-branched, epiphytic, lianoid shrubs; branchlets + terete, finely ridged and grooved, densely spread- ing hirsutulous, 1.5-4 mm in diam. Leaf-blades persistent, coriaceous, evergreen, ovate to ovate-elliptic, (2.5-)4-6 cm long, (1.2-)1.8-2.5 cm broad, apically acuminate and + tapering abruptly to the narrowly rounded tip, basally rounded, marginally entire, glabrous above or nearly so except moderately pilose along 1 r m.a * Im Ly PEE e PR ” 230 Journal of tl Texas 2( the principal veins, moderately to densely spreading pilose beneath along the principal, secondary and even tertiary veins with hyaline tricl up to 0.7 mm long and also densely beset especially when young with short, thick, glandular, scale-like trichomes ca. 0.1-0.2 mm long on both the veins and the surface, the venation weakly 5-plinerved, the secondary nerves joining together in a series of prominent arches, the lamina + bullate, the midrib and secondary veins impressed above and elevated beneath and the tertiary veins slightly elevated on both surfaces; petioles 2-3(-4) mm long, densely spreading pilose. Inflorescences ti ppearing terminal, racemose but with the rachis often contracted and t] PI g tcorymbose, 3-10(-12)-flowered, 2.5—5 cm long overall; rachises (224-8 mm long, densely glandularly strigillose with thickened trichomes 0.1-0.3 mm long and spar- ingly to moderately pilosulose with hyaline spreading trichomes 0.3-0.6 mm long; floral bracts narrowly lanceolate to linear-lanceolate, 2.5-2.6 mm long, marginally glandular ciliate; pedicels (0.6—)1-2(-2.4) cm long, moderately to densely glandularly strigillose and also sparingly pilosulose with hyaline trichomes; bracteoles located in about proximal third, appressed, linear-lanceolate, glandular ciliate, ca. 2.8 mm long. Flowers 5-merous; calyx 3-4.5 mm long, strongly articulate with pedicel, the tube shallowly obconic, ca. 1.5 mm long, ca. 5-6 mm in diam. distally, pronouncedly flaring from the approximately 3 mm in diam. basal portion, densely and scurfily glandular-squamate throughout, the limb ca. 3 mm long, the lobes deltoidly triangular to broadly oblongish with an abrupt distal tip, ca. 1-1.2 mm long, ca. 2.3 mm broad at base; corolla with aestivation valvate, thinly fleshy when fresh but membranous when dry, bistratose, broadly cylindrical, relatively thin in texture, 8-12 mm long, ca. 8 mm in diam. distally, densely scurfily glandular-squamate externally, glabrous internally, the lobes triangular, acute, 2.5 mm long, ca. 5 mm broad basally; stamens 10, included, ca. 7-9.5 mm long, the filaments 2-5 mm long, conspicuously hyaline ciliate, flattened, the anthers ca. 7 mm long, lacking spurs, the thecae 2-2.2 mm long, basally incurved, papillose, the tubules ca. 3—5 mm long, dehiscing by latrorse, elongate clefts ca. 1-2 mm long; styles about as long as the corolla, straight, glabrous. Berry not seen. Distribution. —Known only from the type collection made in Costa Rica in the Talamanca foothills of Limón Province, at ca. 700 m. Flowering; Jul. Vaccinium furfuraceum is characterized by its generally densely glandular-fimbriate leaf blades and flo- ral parts, thin-petiolate leaves with blades that are bullate, apically short-acuminate, and basally rounded, plinerved venation with the reticulate veinlets conspicuously raised beneath, short calyx lobes, thin and membranous corollas that are cylindric-campanulate, and staminal tubules that are pubescent along the inner surfaces and dehiscent latrorsely. The corollas are proportionally conspicuous with respect to the calyx. Its morphological relationships lie near V. monteverdense and V. jefense with which it has in common pilose inner surfaces of the staminal tubules and latrorse dehiscence. It also shares densely fimbriate leaves and inflorescence parts with V. jefense, perhaps its closest relative. Vaccinium furfuraceum differs most con- spicuously from V. jefense by its possession of an overall shorter calyx (3-4.5 mm long vs. 6-10 mm) and calyx lobes (1-1.2 mm long vs. 4.5-6 mm), inconspicuously apophysate (not 10-ribbed) calyx tube, and non-striate calyx limb and lobes. axillary, con gregated distally, 1 P SH 18. Vaccinium luteynii Wilbur, Sida 21:1612, fig. 3. 2005. Tree: PANAMA. Bocas DEL Toro: Fortuna Dam-Chiriquí Grande R _3 km N of the Continental Divide, ca. 8º48"N, 82°12’W, 700-760 m, disturbed, steep roadside slopes, 15 Oct 1998 (fl), Luteyn et al. 15322 ( PMAL; AAU, CAS, DUKE, E, G, MO, NY, SCZ, TEX, US Terrestrial or epiphytic shrubs to 4 m tall; branchlets terete, tan to dull reddish-brown, glabrous. Leaves subopposite, the blades persistent, coriaceous, evergreen, ovate to oblong-ovate or ovate-elliptic, ca. 5-15 cm long, 3-6 cm broad, apically acute to somewhat acuminate, basally rounded to almost subclasping, marginally entire and slightly thickened and revolute, glabrous or very nearly so on both surfaces, the ve- nation 3-5 (7)-plinerved with lateral nerves arising from proximal 1/4—1/5 of midrib, midrib proximally much thickened and weakly elevated in basal '4 above, then thinner and weakly impressed becoming flush to slightly elevated near apex, lateral nerves weakly impressed to flush proximally above but soon slightly elevated distally, reticulate veinlets elevated above, all venation somewhat elevated beneath throughout; TTHTI IN 8 ; £A . 1 e teni A H Ww Ini 231 “71 £f. E 4 V 153 petioles stout, 1-5 mm long, glabrous. Infl mostly 5-8 cm long overall, 10—13-flowered, with a dieron: to UR dy and xüintatels men peduncles; rachises 1.5-4.5 cm long; floral SE EE ER or lanceolate, scale-like, 2-2.5 mm long, glabrous but often marginally, minutely ciliate; pedi y angulate or ridged, 20-40 mm long, ca. 0.6-1 mm in diam., glabrous; bracteoles at or very near the base persistent, subopposite, lanceolate to narrowly triangular, 1.222 mm long, glabrous or minutely ciliate. Flowers 5-merous; calyx 4—4.2 mm long, articulate with pedicel, the tube subcylindric, deeply and bluntly rugose, ca. 1 mm long, 4-4.5 mm in diam., glabrous, the limb smooth above but broadly and saccately spurred opposite and below each lobe, ca. 3 mm long, the spurs solid (not hollow) and sometimes slightly and broadly bilobed, the lobes triangular, acute, 0.6-1 mm long, glabrous; corolla with aestivation valvate, thick and somewhat fleshy, bistratose, broadly urceolate to urceolate-campanulate, ca. 10 mm long, ca. 8 mm broadest diam. when fresh, but drying 5-8 mm long and ca. 6 mm in diam. just beneath the throat and 3.5-4 mm in diam. just above the calyx, green, externally glabrous and internally densely pilose with sordid, white trichomes especially in the throat and lobes, the lobes acute, ca. 3 mm long; stamens n included, ca. 6.5 mm m long, E Sane 3.5-4 mm long, long-pilose in distal half, attached to the anther dorsa lly slightly al tl 3.5 mm long, lacking spurs, golden, the thecae ca. 2-2.5 mm long, densely and finely pasillóse. the lower third strongly incurved, the tubules ca. 1 mm long, dehiscing by introrse, strikingly oblique pores. Berry not seen. Distribution. —Montane rainforests in Panama, along the continental divide, between Bocas del Toro and Chiriquí, at 700-950 m. All specimens yet seen were ae in flower in Mar. Vaccinium luteynii is characterized by its sul , glabrous leaf blades with founded to almost /7 471 TY 0803783131 Tzu subclasping Bases, Baa 5- ped calyx lib. geck, EE broadly urceo late to urceolate corollas that lly densely pilose at the throat and onto the lobes, and glabrous stamens that dehisce introrsely. The is pede e of V. luteynii are uncertain at this time, although it is morphologi- cally similar to V. bocatorense and V. floccosum. Additional collections examined: PANAMA. Bocas del Toro: along road between Florida and Chiriquí Grande, 1.2 mi N of Conti- nental Divide, 5.3 mi N e Ve over im Dam, dudo N, 82?17"W, 910 m, 12 Mar 1985, Croat & Grayum 60459 (DUKE, MO, NY, M PMA, WIS); f 1, 31.7 km S of Chiriquí Grande and 15.7 km N of Sitio de Presa, ca. 8°49’N, 82°12’W, 1070 m, mee 14831 (BRIT, COL, DUKE, MO, NY, P, PMA, SCZ, W); Fortuna Dam-Chiriquí Grande road, 2 N of the Continental Divide, ca. 848'N, 82°12 W, 700—760 m, 15 d 1998 d Bon et al. p bins MO, NY, PMA, U), 15323 on. CAS, DUKE, MEXU, NY, PMA ). Chiri , Km , ca. 8º48'N, 82?12"W, 890 16 Jan 2003 (bud), Luteyn & Se in dis ne ema de SEN FE 19 Mar 1976, Mendoza et al. 260 (DUKE): ne m Fortuna Lake to Chiriqui Grande, 5 j wet forest, 8°49°N, 82°12’W, 700 m, Hampshire & fro Whiteford 441 (NY, PMA). 19. Vaccinium bocatorense Wilbur, Sida 11:441. 1986 [as oc. Tyee: PANAMA. Bocas DEL Toro: headwaters of Río Colubre, 2400-2550 m (Colubre Camp), 3 Mar 1984, Gómez et al. 223 : DUKE!, isotype: MO! photo NY neg. 13046). Epiphytic, scandent treelets (fide label); branchlets + terete, drying brownish. 1 tly subopposite, the blades persistent, coriaceous, evergreen, ovate to ovate-elliptic, 4-6.5 cm long, 3. 5-5 cm broad, apically + rounded, basally rounded and slightly cordate, marginally entire, glabrous above and moderately but inconspicuously erect-pilosulose beneath with widely spaced, slender, hyaline trichomes 0.2-0.3 mm long, apparently eglandular, the venation 5(—7) plinerved with 2-3 pairs of lateral veins arising within 5-10 mm of the base, the midrib and lateral nerves impressed above and elevated beneath, the reticulate venation not apparent; petioles 2-3 mm long, short-puberulent adaxially, + glabrous abaxially. Inflorescences axil- lary, congregated distally, sometimes appearing terminal, racemose to subcorymbose, 6-12-flowered, 3-7 cm long overall; rachises + terete, 2-5 cm long, moderately and inconspicuously pilosulose; floral bracts lanceolate to lance-ovate, acute to acuminate, 3-6 mm long, sparingly pubesuentt to pilosulose externally; pedicels terete, 10-15(-20) mm long, ca. 1 mm in oo , densely but y puberulent with erect hy aline trichomes, striate, eglandular; | teol | g the proximal third of Ge pedicel, lanceolate to lance-oblong, 4-6 mm long, 0.6-1.5(-1.8) mm broad, aaa glal dal to pilosulose and marginally ciliate. Flowers 5-merous; calyx 7-10 mm long, ca. 6 mm in diam., articulate y spal ingly puber ulent 232 J ti tanical Instit exas 2( with pedicel, inconspicuously and moderately pilosulose with erect trichomes, the tube broadly cylindric, 3.5-4 mm long, pilosulose and glandular-pustulate throughout, the lobes spreading-recurved or reflexed, narrowly triangular to lanceolate, 4—5 mm long, ca. 2-3 mm broad at base; corolla with aestivation valvate, thick and fleshy, bistratose, urceolate to cylindric-urceolate, constricted at throat, 7-9 mm long, 6-7 mm in diam., externally glabrous, internally glabrous or very nearly so for the basal half and increasingly tangled pilose distally, the lobes broadly deltoid, ca. 2-2.5 mm long, 2.5 mm broad at base, exceedingly densely matted tomentose within; stamens 10, included, ca. 5-6.2 mm long, the filaments very weakly connate basally, ca. 2.53 mm long and medially ca. 0.5 mm wide, very much flattened and rather broad, marginally and on both surfaces moderately to densely sericeous especially on the connective abaxially, the anthers ca. 3-4 mm long, lacking spurs, golden, the thecae ca. 2.5 mm long, finely or moderately papillose, strongly incurved basally, the tubules 1-1.5 mm long, about as wide as the thecae, glabrous, dehiscing by introrse, oblique pores; styles included, ca. 7 mm long glabrous. Berry not seen. Distribution —Known only from the type collection, made along the heavily forested | dwaters of the Río Colubre, in the province of Bocas del Toro, Panama, at 2400—2550 m. Flowering: Mar. Vaccinium bocatorense is characterized by its subopposite leaves with stout, rugose petioles, bullate leaf ] ded to sul te and oft mewhat clasping, plinerved blades with the apices rounded and tl venation with the reticulate veinlets obscure beneath, elongate calyx with proportionately long lobes, thick lat d j ly constricted at the throat, and by 11 1 S TA PACATA A and fleshy corolla that is u cylindric I y stamens with introrse dehiscence and glabrous tubules. The corolla is proportionally small and inconspicu- ous with respect to the calyx. Vaccinium bocatorense keys closest to V. floccosum and V. monteverdense because of their long calyces and calyx lobes, but its overall relationships within the genus are uncertain. 20. Vaccinium floccosum (L.O. Williams) Wilbur & Luteyn, Brittonia 29:272. 1977. Hornemannia floccosa L.O. Williams, Brittonia 18:248. 1966. Symphysia floccosa (L.O. Williams) L.O. Williams, Phytologia 24:158. 1972. Type: PANAMA. CHIRIQUÍ: Boquete Distr., Bajo Chorro, rainforest, 6000 ft, 11 Jan 1938, Davidson 104 (HoLoTYPE: F! photo F neg. 52534; ISOTYPES: A! MO! USD. Terrestrial or epiphytic shrubs 1—4 m tall; branchlets slender grayish to olive-brown, glabrous. Leaves alter- te to subopposite, the blad istent, coriaceous, evergreen, ovate to lanceolate or elliptic, 8225-9 C11) cm long, (1.5-)3-5(-6.5) cm broad, apically acute to acuminate, basally cuneate to rounded, rarely short- attenuate, marginally entire and slightly thickened, glabrous to moderately appressed glandular-fimbriate above and especially beneath, the venation indistincly 3—5(-7)-plinerved and also somewhat arcuately veined; petioles 2-4(-6) mm long, glabrous. Inj axillary, loose, umbelliform racemes, mostly 2-4 cm long overall; rachises 2-10(-20) mm long, glabrous to sparingl puberulent; floral bracts persistent, ovate to lance-ovate or lanceolate, 1-3.3 mm long, glabrous to sparingly short-ciliate; pedicels 10-20(-25) mm long, glabrous to sparsely puberulous, but distally often tringed with glandular trichomes 0.1-0.2 mm long; bracteoles approximately medial, subopposite, ovate to lance-ovate, persistent, 1-3.2 mm long, glabrous to inconspicuously ciliate. Flowers 5-merous; calyx 4—7.8 mm long, articulate with pedicel, glabrous, the tube campanulate, 2-3 mm long, 3-5 mm in diam., the limb longer than the tube, commonly striate, the lobes broadly triangular, acute, 1.8-3.5 mm long; corolla with aestivation valvate, fleshy, bistratose, broadly cylindric-campanulate, broadest at mouth, 7-10 mm long, 4-5 mm in diam., pale greenish often suffused with red, externally glabrous to moderately appressed spiculate, internally densely floccose with a tangle of white, villous trichomes, the lobes erect to spreading or reflexed, triangular, acute, 2-2.2 mm long, densely villous-floccose within; stamens 10, included, ca. 5.2 mm long, the filaments slightly adherent to the very base of the corolla tube, ca. 3-3.2 mm long, flattened, slightly ciliate, the anthers 3-3.5 mm long, lacking spurs, golden, attached along the lower 1/3, the thecae ca. 2.2-2.5 mm long, papillose, basally incurved and bluntly tapering, the tubules inconspicuous, stout, ca. 1-1.2 mm long, dehiscing by terminal, truncate pores. Berry spherical, 8-10 mm in diam., dark purplish at maturity, 5-locular. Distribution —Elfin and [ tane cloud forests in the high land along the border of the Panamanian Bocas del Toro and Chiriquí provinces, between 1000-2500 m. Flowering: Jan-Oct; fruiting: May-Dec. Vaccinium floccosum is characterized by its alternate to pseudo-opposite leaves with flat blades, entire margins and paneled’ venation, eouspicucusty striate calyx limbs that are longer than the tubes, greenish corollas that ly densely f] lly, and anther tubules that are short, ca. 1 mm long, glabrous, and dehiscene aso or by perfectly ena pores. Leaf morphology is variable with populations from the Boquete region SE leaves coriaceous jue EE EE SE blades ovate to ovate- P the Fortuna Dam a! lanceolate, with and | or short-attenuate area show Coe more thinly coriaceous and more conspicuously petiolate, EE blades more lance-elliptic, with apices longer acuminate, and bases tapering and cuneate. All leaf blades with rounded bases are also pseudo-opposite, whereas leaf blades with cuneate leaf bases may be either distinctly alternate or pseudo- opposite. The EE of Eus oes as listed ane Eeer come UNE of the extent of its anomalous characteristics. The relationships of this species are as close to the species centere d about Vaccinium poasanum as they are to the largely West salian species usually treated as pL Mes racemosa (Vahl) Stearn (= Horne- mannia racemosa Vahl). The inclusion of all these species within Vaccinium does not significantly expand the limits of that extremely diverse genus and still seems to us to be the best treatment for the complex until a worldwide revision of the generic limits is undertaken. Vander Kloet (1985), whose study was largely based upon the West Indian V. racemosum (Vahl) Wilbur & Luteyn [= Hornemannia racemosa Vahl and Symphysia racemosa (Vahl) Stern], argued against such an extension of the generic limits of Vaccinium claiming that such an expansion “is tantamount to making the genus a dumping ground for any Vaccinieae of uncertain affinity.” In Vander Kloet’s experience “the character unique to the genus “Vaccinium” is the absence of calyx tube enlargement prior to anthesis” (see vander Kloet 1985 and vander Kloet et al. 2004 for an alternative opinion). Additional collecti ined: PANAMA. Bocas del T | il on divid Chiriquí and Bocas del Toro, ca. 8?45'N, 82º15"W, 22 Oct 1985, McPherson 7232 (DUKE, MO, NY, TEX). Chiriquí: trail to Eno Pate Ma aha: headwaters of the Río Palo Alto, above Palo Alto, 8?47"N, 82?22"W, 1700-2100 m, 24 Apr 1982, Knapp & EN 4784 RE MO) Paquetes ia e Mono and Bajo Chorro trails out of Boquete, 1 Jun 1972, Luteyn 3069 (DUKE, MO, NY) o Horqueta, 1700-1900 m, 4 Jan 1975, Luteyn & Wilbur 4600 (DUKE, MO, NY); sl C Horqueta, ca. 6. 6 km NNE of Boquete, 1500-1800 m, 5 Jan 1975, Wilbur & Luteyn 19330 (DUKE, F, GH, LL, MICH, MO, NY, PMA, US). 21. Vaccinium monteverdense Wilbur & Luten. Brittonia 53:442. 2001. Tre: COSTA RICA. Guanacaste: Monteverde area, Hacienda Ira Rosa, 3.7 km above Santa Elena ind ca. 1 km before Río Negro just before the Sky Walk, ca. 1500 m, 10 Mar 1999 (fl), Luteyn et al. 15408 (HoLorYrE: NYI; : CR!) Epiphytic or EE ge ise m tall, ae from Kee E + terete, a DIO. E i errarich sx brous, those o gy Le blades persistent, coriaceous, anca ne inne ise to elliptic, (2. 423 4— 8. cm long, (o. 8). E A cm broad, apically somewhat abruptly acuminate to short-acuminate, the tip itself + bluntly obtuse and ca. 2 mm wide, basally rounded to obtuse, usually short-acuminate and narrowly decurrent onto petioles, marginally entire and revolute causing leaves to be concave, weakly bullate, glabrous on both surfaces, the venation 3(—5)-plinerved with usually only midrib prominent; petioles somewhat angulate and strongly canaliculate, 4-8 mm long, ca. 1.5 mm in diam., glabrous or very nearly so. Infl terminal, racemose, 7-9(-16)-flowered; rachises + terete ca. 2-3(-4.7) cm long, glabrous, N floral bracts inconspicuous, appressed, scale-like, ovate, ca. 0.8—1.5 mm long; pedicels terete, 14-19 mm long, ca. 1-1.2 mm in diam., glabrous, dull reddish; bracteoles nearly basal, subopposite, scale-like, appressed, elliptic to narrowly tri- angular, ca. 0.6-0.8 mm long. Flowers 5-merous; calyx 10-16 mm long, articulate with pedicel, glabrous, green flushed with red, the tube cylindric to obconic, 4.5-8.5 mm long, 2-3 mm in diam. basally and 4—6 mm in diam. proximally and ca. 1 cm in diam. distally, striate to slightly ribbed when dry, the limb spread- ing to campanulate, 5.5-7.5 mm long, conspicuously striate, the lobes deltoid, 2.5-3.5 mm long, 4-5 mm broad basally; corolla with aestivation valvate, thinly fleshy, bistratose, broadly cylindrical to campanulate, 13-18 mm long, 9-15 mm in diam. distally, green suffused with maroon to reddish-maroon or purplish, glabrous, the lobes ca. 3-3.5 mm long, 3-4 mm broad at base; stamens 10, included, ca. 11.5-12 mm long, the filaments distinct, ca. 4.7-5 mm long, flattened, marginally long-pilose, the anthers ca. 8-8.4 mm long, I [| ful Dos H Inm LI PP ET afa\ 234 lacking spurs, golden, the thecae ca. 3.2-3.4 mm long, papillose, bluntly apiculate basally, the tubules ca. 4.8-5 mm long, distinct, long-pilose along inner surfaces, dehiscing by short, slightly latrorse clefts ca. 1.4-1.5 mm long; styles slender, 12-15 mm long, glabrous. Mature berry unknown, but at least 8 mm in diam. and spherical when immature. Distribution — Known only from a few collections in cloud forest, from the Costa Rican Cordillera de Tilaran in the vicinity of Monteverde, where the provinces of Alajuela, Guanacaste, and Puntarenas join one another, and also from the Province of Guanacaste in the Cordillera de Guanacaste on the lower slopes of ea La Ta at ca. id 1600 m. died ee C11 telas circular Elaselz3eis glands 1 by having leaf DIAUESs WILT LW , CONCAVE, ca. 0.2 mm diam., one on either side of midrib on lower surface, ane cies striate calyx limb, cylindre: campanulate corolla that is mostly green but suffused with maroon or reddish-maroon apically, stamens that are adherent to the base of the corolla, relatively long filaments, and anthers with pilose tubules that dehisce latrorsely. Furthermore, its leaves are often slightly bullate and concave. The relationships of V. monteverdense are uncertain at this time. P PRÉ Pu COSTA RICA. G te-Punt Al la: Cordillera de erem Mon es ca. 1mS of , ca. 1460 m, 23 Feb 1986, Almeda et al 5082 (C AS). G ste, lower slopes of Cerro La Giganta ca. 2 km W of Río Naranjo, 2600 ft, 27 Feb 1975, Utley 1885 (DUKE, NY). Punt : cloud forest above the Quaker settlement at Monteverde, 1600-1700 m, 4 May 1975, Utley 2376 (DUKE). 22. Vaccinium campanense Wilbur € Luteyn, sp. nov. (Fig. 2). Tw: PANAMA. Panama: Dist. de Capira, Cerro Campana, trocha desde el mirador a la cima, 800 m, 8°41’N, 79°55’W, bosque nebuloso, Galdames et al. 1840 (noLorwrE: PMA!, isotypes: CAS!, DUKE!, MOL. NY!, SCZ!, USD. Frutex epiphyticus. Petioli 3-4 mm longi, pilosuli et puberuli. Folia coriacea, integra, ovata vel ovato-elliptica, 2-4.7 x 1.2-2.4 cm 3—5 plinervia, venatio reticulata, brochidroma. Inflorescentia umbellate-racemosa, 1-2.5 cm longa; rhachis 4-8 mm longa, NUN strigillosa et pilosula; pediceli 7-15 mm longi; bracteae caducae. Calyx pedicellis articulatus; tubo cylindrico, 23.5 mm longo et 2-3 mm diam., striato, lobato, glandulato; lobi calycis 5, detoidi, 4—5 x 2-2.7 mm, acuti; corolla es verde, ca. 7-15 x 9-10 mm, lobi corollae triangulari, acuti, ca. 3 x 3.5 mm, stamina 10, thecae granulosae, + 3—3.6 m Epiphytic shrubs to 2 m tall; branchlets + terete, BOWIE, grayish or blackish, the youngest 1.5-3 mm in diam., finely ridged, inconspicuously puberulous. Leaf-bl tent, coriaceous, evergreen, ovate to ovate- elliptic, 2-4.7 cm long, 1.2-2.4 cm broad, apically usually Gees into a 4-8 mm long, blunt, tongue-like tip, basally usually rounded, marginally entire, the venation 3-5-plinerved, the midrib and secondary veins somewhat sunken above and elevated beneath, the upper surface glabrous at maturity but initially moderately beset with dark reddish, glandular-appressed trichomes that rarely persist into maturity and the sunken veins proximally usually somewhat puberulous, the lower surface moderately beset with minute, often ap- pressed puberulence on the surface and principal veins and those on the surface apparently glandular and darkening reddish-brown upon drying and perhaps 0.1 mm long, usually disappearing with age; petioles terete or nearly so 3-4 mm long, minutely puberulous. Inflorescences axillary, racemose, 1-3-flowered, ca. 1-2.5 cm long overall; rachises mostly 4-8 mm long or less, sparingly puberulous to short-pilose; floral bracts 1-1.2 mm long; pedicels 7-15 mm long, sparingly puberulous to short-pilose, drying reddish-brown; bracteoles basal, linear, 1-2-3) mm long, glandularly minutely ciliate. Flowers 5-merous; calyx ca. 88.5 mm long, clearly articulate with pedicel, the tube cylindric, 5(-10)-ribbed when fresh, 2-3.5 mm long, 23 mm in diam., striate, moderately minutely glandularly appressed pubescent, the limb ca. 5.5 mm long, the dn s triangular, SHE acute, Së a mm tong, 2-2.7 mm broad, striate, green, minutely reddish-glandular valvate, thinly fleshy, bistratose, broadly campanulate, ca. 7-15 mm pus 9-10 mm in diam., green, the lobes triangular, acute, ca. 3 mm long and 3.5 mm broad at base; stamens 10, included, 9.5-9.6 mm long, the filaments distinct, ca. 33.6 mm long and 1.5 mm wide at base, slender but broadening basally, marginally long-ciliate, dorsifixed, the anthers ca. 7.5-8 mm long, lacking spurs, orangish-yellow, the thecae 33.6 mm long and 1-1.5 mm wide, finely papillose, the tubules somewhat divergent, each ca. 3-4.6 mm long and 0.5 mm wide, dehiscing by latrorse clefts ca. 1.5-2.5 mm long and 0.5 mm wide; styles slightly exserted. Mature berry unknown. 235 2 HESS Marea "ue ODORE S Sia ini Do B. flower in bud be 2. Y F D. mature corolla. E. stamens d E? calvx tube rihs. and bi showina ventral. dorsal F Ei | || E al n H Im | | A ndk 236 t t fTexas 2( T species | EE a RES Distribution.—Epiphyte in the canopy of cloud forest trees. This very d only been collected twice, which is surprising as Cerro Campana is a frequently E and botei area relatively close to Panama City, at ca. 800-850 m. Flowering: Feb. Vaccinium is one of the green-flowered species of Vi by its combination of small, entire-margined leaves, few-flowered inflorescences, calyces densely nda fimbriate with 5-10 bluntly ribbed tubes and prominently striate, proportionately long and strikingly acute calyx lobes, and broadly campanulate corolla. +] pi KN EE 11 E 5 1 ] Additional collection studied: Cerro Campana, topotype, 13 Feb 1996, Luteyn & Galdames 14857 (NY) (flowers from forest floor only). 23. Vaccinium poasanum J D. Smith, Ton as (Crawfordsville) 24:395. e ene psum Y D. Smith) vander Kloet, Taxon 53:97. 2004 CA. AlajuFLa: Volcán Poas, 2600 m, Mar 1 x (LECTOTYPE GH! isoLecTOrYPE: K! syntyPe US! photo NY neg. s.n.). Terrestrial or epiphytic shrubs or small trees (122—577) m tall; branchlets glabrous. Leaf-blades persistent, coriaceous, evergreen, ovate to lanceolate or elliptic, 3-8(-10) cm long, (1-)2.5-4(-6) cm wide, apically acute to acuminate, basally acute, Bee cuneate or acuminate (very rarely eo E entire and slightly revolute, glabrous ly appressed glandular-fimbriate above and especially beneath, rarely E beniein near the base, e aaa erii 3—5- BE and also M arcuately veined; petioles 1-4 mm long, glal irsutul y, umbelliform racemes, mostly 2-4 cm long overall, (4-)7-13-flowered; rachises 0.5-2 cm lona, glabrous to sparingly puberulent; floral bracts persistent, lance-ovate to lanceolate, 1.5-2.5 mm long, ciliate; pedicels slender, 7-15 mm long, sparingly puberulent, often distally with a fringe of glandular trichomes 0.1-0.2 mm long; bracteoles sub- medial, subopposite, lance-ovate to | late, often persistent, 1-1.5 mm long, ciliate. Flowers 5-merous; calyx 3.5-5 mm long, clearly articulate with pedicel, glabrous to densely puberulent, the tube short-campanulate to turbinate, 23.5 mm long, about equal in diam., the limb 1-2 mm long, the lobes broadly and shallowly deltoid, apiculate, 0.2-1 mm long; corolla with aestivation valvate, thinly fleshy, bistratose, broadly cylindric, 8—12 mm long, 6-10 mm in diam., pale yellowish or greenish-white and occasionally tinged with bright rose or pink, externally and internally glabrous or rarely apically sparsely villose, the lobes erect to slightly recurved, broadly oblong, apically acute to obtuse, 2-3.5 mm long; stamens 10, included, 7.5-8.5 mm long, the filaments 2-2.5 mm long, ciliate, flattened, broadened basally wl herent to the very base of the corolla, the anthers 6-7 mm long, lacking spurs, golden, attached slips Hors the middle, the thecae 1.8-2.2 mm long, papillose, basally incurved, the tubules 3.5—5.5 mm long, glabrous, dehiscing by slightly introrse, oblique pores but appearing + terminal; styles exserted, 8-10 mm long. Berry subspherical, 7—10 mm in diam., dark pupe at ae e SE Distribution. t t Guatemala south into central Panama, from 900-3300 m. Flowering: throughout most of year especially from Dec—Mar and Jul-Sep; fruiting: throughout much of the year. 1 ] 1 by its leaves with acute to broadly bladel d acute to te apices, apiculate calyx lobes, broadly cylindric corollas, staminal fil ts that i ] shorter than the overall anthers, and elongate anther tubules. Like many other of the sreen-flowered species of Vaccinium, the inter-specific relationships of V. poasanum are uncertain. It is, however, morphologically similar to V. campanense. Additional collections examined: GUATEMALA. Baja Verapaz: in high e on hill, Unión Barrios, 13 Mar 1972, Contreras 11293 (DUKE, LL). Zacapa: upper reaches of Río Sitio Nuevo, 1500-1800 m, 25 Jan 1942, Steyermark 43200 (A, F, NY, US DURAS. Comayagua: in cloud forest, 2300 m, Montaña Comayagua, 6 Mar 1975, Hades 2499 (MO). El Paraiso: dense cloud forest, Mt. Vol- cán, 1950 m, 16 Mar 1947, Williams & Molina R. 12183 (F, GH, US). M Nacional La Tigra, 22-25 km of Tegucigalpa; cloud forest, 14°12’N, 8707'W, 1850-2125 m, 1 Feb 1987, o 64045 a MO, NY); in Bi Mt Uyca, 2000 m, 20 Aug 1964, Williams & Molina R. 10380 (DS, F, LL, MICH, MO, PH); wind cloud forest area, Montaña La bL dd Panda QE Mts. ca. 15 m NE MEL ME 2000-2200 m, Williams et al. 23273 (CAS, DS, F, LL, MICH, NY, US). Santa ro Santa Bárbara, 2750 m, 5-6 Apr 1951, Allen et al 6066 (DS, F, US). EL SALVADOR. ELE RFOLE X. up pl rocky F Tm bd a! ge D FS D ] e A La a LT] "m 237 Santa Ana: Cerro Montecristo, 6000-6500 ft, 17 Jan 1959, Allen 7153 (DS, F, LL, NY, US); moist cloud forest, Cordillera Miramundo, Mt. Montecristo, 2000-2200 m, d n 1966, Molina R. et al. 16711 & 16825 (F, NY, US). rA: Mataa pa cut-over lai forest area, El Porvenir, C tral, 1700 m, 11 Mar 1967, Molina R. 20515 (F, NY, US). Z C La Pimienta, nortl lope facing La Garrapata, ca ici 84*59"W, 900-1180 m, 16 Mar 1980, Pipoly 6040 (DUKE, MO, NY. COSTA RICA. Alajuel led sl 1 km from li 19 km W ot Vara Blanca, Volcán Poas, 2700 m, 21 Jan 1968, Wilbur & Stone 9843 (DS, DUKE, F, LL, MICH, MO, MSC, NY, US). IHE Volcán een. 8.3 km ia bd Dd in S at Porrosati, 2600-2800 m, 16 Mar 1975, Utley 1959 (CR, DUKE, F, MO, NY, US). Finca Coliblanco, 1-2 km S of Esperanza, 3000-2600 m, 28 Jan 1976, Utley 3813 (CR, DUKE, F, MICH, MO, NY, US); on lower slopes of Volcán Turrialba, 1-2 km S of Finca Central, 2600-2700 m, 15 Mar 1976, Utley 4293 (DUKE, MO, NY). Cartago-San José: bog and ET «el surrounding wooded hills ca. 11 mi SE of El E American Hwy, ca. 9000 ft, 2 Feb 1970, Wilbur & Teeri 13901 (DUKE, F). San José: GEN drin WE ge Sa between Alto La See and e La Hondura, ca. 4200—3800 ft, 4 Jul 1972, Luteyn 3315 (DUKE, F, MO, N ro Danser ca 3-5 km SW of Aserri, 2300-240 Mar 1976, dni 4291 (CR, DUKE, F, MICH, MO, NY, US). Punt R d le, Cerro Negro, 1500-1600 m, Haber & Bello C. 2814 (DUKE, MO). Limón: Cordillera de Talamanca, Atlantic slope, Valle de Silencio along Río Terbi, 0.5-1.5 airline km W of Costa Rican-Panamanian border, 2300-2400 m, 908'N, 82?57"W, 9 Sep 1984, Davidse et al. 28717 (DUKE, MO). PANAMA. Bocas del Toro: upper Río Colubre, 2500—3000 m, Aug 1983, Gómez et al. 21923 (DUKE, MO, NY). Chiriquí: mossy forest ca. 7000 ft, E of Guadeloupe along the Río Chiriquí Viejo ca. 2 mi NE of Cerro Punta, ridge of Cerro Respinga, 13 Jan 1971, Wilbur & Foster 13114 (DAV, DS, DUKE, F, LL, MICH, MO, NY, PMA). Coclé: sawmill N of El Cope along divide, 25 Feb 1979, Hammel 6267 (MO). 24. Vaccinium almedae Wilbur & Luteyn, Sida 21:1607, fig. 1. 2005. Tree: PANAMA. Crriqut: Edwin Fabrega Dam and Reserve in Fortuna, along trail to hydrological station, along Río Hornito, below forestry house along the road in wet forest, 8?45'N, 825'W 1200 m, 20 Jan 1989, Almeda et al. 6369 (noLorvee: PMA!; isorvees: CAS! DUKE! NY!). Coarse, rigid, epiphytic shrubs 1—2 m tall; | hlets and branches + terete, glabrous or nearly so but current season growth not present, thin, brownish or grayish. Leaf-blades pe COriaceous, evergreen, de is 6-18 cm ge 3—7. ei cm dus d d acute to acuminate, basally ] hortly tapering and sometimes d the petiole at the aun as entire, glabrous on both surfaces, the venation 3—5 aime the veins + depressed above elevated for the proximal third to half while elevated beneath through the = 4 order and fannie an indistinct reticulum; petioles somewhat flattened above and there rather broadly and shallowly canaliculate, Kony post mm long, 2-3.5 mm in diam., glabrous. Infl tly ramiflorous, of several to numerous, (2-10-flowered) emerging from small, depressed mounds 1-2 mm in diam. and 1-2 mm long; floral bracts scale-like, 0.6—1.5 mm long, ciliate; pedicels slender, 5-10 mm long, 0.2-0.3 mm in diam., glabrous; brac- teoles deltate, 0.5-0.8 mm long, ciliate. Flowers 5-merous; calyx ca. 3-4 mm long, clearly articulate with pedicel, the tube cylindric-obconic, ca. 1.5-2.2 mm long, 1.6-2 mm in diam., glabrous, the lobes broadly based, minute, varying from barely detectable to perhaps as much as 0.2-0.4 mm long, glabrous; corolla with aestivation apparently valvate, membranaceous, thinly bistratose, cylindric to more typically gradually funnelform, 5-9 mm long, flaring to ca. 5-6 mm in diam. from a 2-2.5 mm diam. base, greenish-white, glabrous both externally and internally, the lobes narrowly triangular to deltoid, acute, ca. 2.5 mm long; stamens 10, included, 7-8 mm long, the filaments 1-2.5 mm long, united in the basal 0.5 mm, glabrous, flattened, the anthers 6-7 mm long, lacking spurs, attached medially, the thecae ca. 1.4-2.2 mm long, finely papillose, basally incurved and apiculate, the tubules ca. 3—4.8 mm long, dehiscing by truncate to slightly flaring, terminal pores; styles slightly exserted, glabrous. Mature berry unknown, but when immature at least 3-4 mm in diam., glabrous. Distribution. —Known only from four collections made in wet forest, near the Fabrega Dam site at Fortuna (Chiriquí Prov.) and Cerro Colorado (Bocas del Toro Prov.), Panama, at 1150-1500 m. Flowering: Jan.; immature fruits: Apr. Vaccinium almedae is characterized by its short ramiflorous, rotate to flaring calyx limb, minute calyx lobes, pale green to white corolla, staminal filaments that are conspicuously shorter than the anthers, and anther tubules that are conspicuously longer than the thecae and that dehisce by truncate to slightly flaring, terminal pores. Its relationships are unclear at this time. i f E T " r1 +l " aus Additiona l collecti ined: PANAMA. Bocas del Toro: Hep of Cerro ea 7 mi ds bine Am: ca. 8°39 NN, 8145 W, ca. 1500m, 12 Apr 1986, A g ito, below forestry house on hwy, 8°45’N, 82º15'W, 1150- 1200 m, 1, 20 Jan 1989, EE 13612 (DI IKE, MO); Fortuna d ite, along stream on white sandy soil, 1200 m, 7 Feb 1985, van der Werff & van Hardeveld 6609 (MO, photo NY neg. 13043). 25. Vaccinium santafeense Wilbur & Luteyn, Ann. Miss. Bot. Gard. 65:140. 1978. Symphysia perardua vander Kloet, Taxon 53:97. 2004. Tres: PANAMA. Veraguas: NW of Santa Fé, 2 km from Escuela Agricola Alto de Piedra, ridge top cloud forest below summit of Cerro Tute, 27 Mar 1975, Mori & Kallunki 5248 ( MO!) Epiphytic shrubs 1-2(-3) m tall, vegetatively glabrous throughout; branchlets and twigs terete, gray to light brown when dry. Leaf -blades persistent, drying subcoriaceous, evergreen, elliptic, 5-8 cm long, 2-3 cm broad, apically acute to acuminate or even tapering into a pronounced drip-tip 8-12 mm long, basally cu- neate, marginally entire, the venation pinnate, the midrib elevated proximally above while sunken distally and beneath elevated throughout; petioles dorsiventrally compressed, 6-11 mm long, 1.5-2 mm broad. Inflorescences racemose to subcorymbose, 2—8-flowered; rachises terete, smooth or weakly striate, 5-9 mm long; floral bracts ovate to Se acute, keeled, 1-1.2 mm long, appressed, irregularly and minutely marginally denticulate; p te, smooth to weakly striate, 8-12 mm long, distally minutely fimbriate; bracteoles basal, slightly smaller and more closely appressed, ca. 1 mm long, minutely denticulate. Flowers 5-merous; calyx ca. 3.5 mm long, markedly articulate with pedicel, the tube terete, ca. 2 mm long, 2.5 mm in diam., the lobes broadly triangular with shallowly indented to almost flat sinuses, ca. 0.6 mm long; co- rolla with aestivation valvate, thin-carnose, bistratose, cylindric, 8—9 mm long, abruptly contracted for the lower 1.5 mm and there 2.2-2.5 mm in diam., 4—4.2 mm in diam. above, greenish-white, glabrous both internally and externally, the lobes erect to somewhat recurved, triangular, acute, 2-2.5 mm long; stamens 10, included, 6—7.7 mm long, the filaments distinct, 2-3.2 mm long, ca. 0.5 mm wide, flattened, ciliate, the anthers ca. 5.5 mm long, lacking spurs, the thecae 2-2.5 mm long, finely to coarsely papillose, the tubules 3-3.6 mm long, glabrous, dehiscing by elongate, latrorse clefts 2-3 mm long; styles slightly exserted, 9—10 mm long. Mature berry unknown. Distribution.—Known only from five collections in cloud forest, on or near Cerro Tute ca. 10 km NW of Santa Fé, Veraguas, Panama, at 900-1000 m. Flowering and fruiting: Jan-May. Additional collections examined: PANAMA. Veraguas: NW of Santa Fé, 2 km from Escuela Agrícola Alto Piedr of Cerro Tute, 27 Mar Pe Mori & n i Me Ke photo NY Wes 13044), 28 Mar 1975, Mori & Kallunki 5280 (MO); Cerro Tute, ca. 10 km NW of Santa Fé, , ca. 1000 m, 19 May 1975, Mori 6254 (MO); along steep trail to summit of Cerro Tute, ca. 3 km above Escuela Agricola Alto Piedra, near Santa Fé, 2800-3000 ft, 4 Jan 1981, Sytsma & Antonio 3029 (NY). 26. Vaccinium costaricense Wilbur & Luteyn, Brittonia 29:270, fig. 5. 1977. Symphysia costaricensis (Wilbur & Luteyn) vander Kloet, Taxon 53:97. 2004. Tyre: COSTA RICA. HEREDIA: vicinity of Cerro Chompipe, 21 Dec 1974, Wilbur & Luteyn 18579 (HOLOTYPE: : CR! F! MO! NY). Terrestrial or more seal epiphytic shrubs, 1-3(-4) m tall; branchlets grayish to reddish-brown, gla- brous. Leaf-blades persistent, coriaceous, evergreen, ovate or lanceolate to elliptic, 2.524-8(-1D) cm long, (1.52.5-4.5(26.2) cm broad, apically acute to acuminate, basally acutely tapering to rounded, marginally entire and slightly thickened, appearing slightly revolute, glabrous to moderately but inconspicuously ap- pressed glandular-strigillose both above and more especially beneath, the venation 3—5(-7)-plinerved and also somewhat weakly arcuately veined; petioles 1-2 mm long, glabrous. Inflorescences axillary but often superficially appearing terminal, of umbelliform racemes, 10—15-flowered; rachises 1-2.53) cm long, glabrous to very sparingly and mainutelyE See floral bracts persistent EE to lanceolate or triangular, scale-like, 1-3.5 mm long, gl to minutely cilia licel y 10-30 mm long, glabrous to inconspicuously puberulous or often sino al ECH with Gett trichomes ca. 0.1 mm long; bracteoles usually in the lower third, + subopposite, lance-ovate to lanceolate, persistent, 0.8-3 mm long, appressed, glabrous to indistinctly ciliate. Flowers 5-merous; calyx 3.5-5(-5.8) mm long, clearly articu- late with pedicel, glabrous to densely, minutely puberulent, the tube campanulate, ca. 2-2.5(-4) mm long, 2.5-3 mm in diam., the lobes triangular, acute, 1.2-1.8(-2) mm long; corolla with aestivati valvate, fleshy, vn I cos E t£ M nul I Eapteal À H y In! 239 bistratose, broadly cylindric to somewhat urceolate, 5-8 mm long, 4—7 mm in diam., pale greenish-yellow, externally glabrous and internally moderately to densely pilose with sordid, white trichomes especially in the throat and on the lobes, the lobes erect to spreading or somewhat reflexed, triangular, acute, 1-2 mm long; stamens 10, included, ca. 6 mm long, the filaments 3-4 mm long, flattened, slightly adherent to the very base of the corolla tube, slightly ciliate, the anthers ca. 3-4 mm long, lacking , golden, attached in the lower quarter, the thecae ca. 1.8-2 mm long, papillose, basally incurved and apena the tubules stout, ca. 1-2 mm long, dehiscing by introrse, slightly oblique or slightly flaring, terminal pores. Berry spherical, 11-16 mm diam., dark maroon to blackish-maroon, or purple, 5-locular. Distribution.—Mountain slopes covered by premontane rainforests, on Atlantic slope of Costa Rica (Heredia and San José) and Continental Divide forests in Panama (Bocas del Toro and Chiriquí), between (700-)1100-2000 m. Flowering and fruiting: throughout much of the year. Vaccinium costaricense is characterized by its combination of acutely tapering to rounded leaf blade bases, a calyx tube that is longer than the limb, corollas that are pilose internally at the throat, and short anther tubules. Vaccinium costaricense has a longer rachis, shorter calyx limb and lobes, smooth, not stri- ate, calyx limb and lobes, and longer anther tubules than its morphologically closest relative V. floccosum, from western Chiriquí Prov., Panama. These differences in the calyx may be influenced by where and how the plants grow, i.e., epiphytes in the closed forest vs. terrestrial in exposed, windswept areas (JLL, pers. observ.). Vaccinium costaricense differs from V. orosiense by acutely tapering to rounded leaf blade bases (not cordate and amplexicaul) and shorter calyx lobes (at least in the Costa Rican populations). The characters that distinguish these three species of Vaccinium are admittedly few and collections are needed to help clarify their morphologies and relationships. Additional collections examined: COSTA RICA. H lia: f t Rio Pejé and Rio Sardinalito, Atlantic slope of Volcan Barva, 10°17.5°N, 84%04.5'W, 700-800 m, 3 Apr 1986, Grayum 6706 (DUKE) Las Vueltas, NE of Volcan Barba, 1950 , 25 Jun 1972, Lent 2645 (DUKE, NY, PMA); forests between Rio os elias de Re nda Medi of Cerro Chompipe, 1700-2000 m, 21 Dec 1974, DA e dio 4438 (DUKE, F, MO, NY); vicinity of Cerro Río Las Vueltas and Río Nuevo, 1800-2000 m, ca. 12 fael, 21 Dec 1974, Wilbur & Luteyn 18576 (DUKE, E MICH, MO, NY, tS San José: CR 216, ca. 3-6 km i» dun ne d in jean of Eet on NW flank of Volcán Irazú, 5700 ft, 10 Dec 1975, Almeda et al. 2626 (CAS, icket at Alto La Palma ca. 15 km in straight line NE of San José, 1480 m, 13 Jul 1976, Wilbur 20339 SC DUKE, E, LL, MICH, MO, NY, PMA, US, WIS). PANAMA. Bocas del Toro: along road between Fortuna and Chiriquí Grande, 1.2 m N of Continental Divide, 5.3 mi N of bridge over Fortuna Dam, 8?44'N, 82º17'W, 910 m, 12 Mar 1985, Croat & Grayum 60459 one NY). pui qae east of es Fortuna main SE 1200-1600 m, 12 Sep 1977, Folsom & Dressler 5300 (DUKE, MO); windswept ridge 10 km N ofI , 1150 m, 18 Jun 1982, Knapp & Vodicka 5603 e MO); forested slope along ridge, vicinity of F Dam, ca 8945 N, 82°15 W, ca. 1250 m, 28 Apr 1986, Weg 9094 (DUKE, MO, NY). 27. Vaccinium orosiense Wilbur & Luteyn, BPO 20: E s OMS sia orosiensis (Wilbur & Luteyn) vander Kloet, Taxon 53:97. 2004. Tree: COSTA RICA. C n property of ICE Hydroelec- tric plant ca. 15-20 km upstream from the suspension M 1300- 1800 m, 20 Dec 1974, "dk et al. 4411 (HoLoTYrE: DUKE!; ISOTYPES: CR! Fl MO! NY! US!). Epiphytic or terrestrial shrubs, 1-2 m tall; branchlets glabrous, olivaceous to purplish. Leaves appearing subopposite, the blades persistent, coriaceous, evergreen, ovate to ovate-lanceolate, (2—)4—9(-14) cm long, (1.5-)3-5(-7) cm broad, apically acute, basally rounded to more typically cordate, clasping to amplexicaul, marginally entire and slightly thickened, glabrous to sparsely and inconspicuously appressed, glandular- strigillose, the venation indistinctly 5-7(-9)-plinerved; petioles ca. 1 mm long, glabrous. Inflorescences axillary but often superficially appearing terminal, of umbelliform racemes mostly 2—4 cm long overall; rachises mostly 1-1.5 cm long, glabrous; floral bracts persistent, triangular to ovate-lanceolate, scale-like, 1-2.2 mm long, glabrous to minutely ciliate; pedicels mostly 15-23 mm long, glabrous; bracteoles usually in the lower quarter, subopposite, deltoid to lance-ovate, persistent, 0.8-1.5 mm long, appressed, glabrous to inconspicuously ciliate. Flowers 5-merous; calyx 2-4 mm long, clearly articulate with pedicel, glabrous, the tube campanulate to urceolate, ca. 2-3 mm long, 2.5-4 mm in diam., the lobes triangular, acute, 0.5-1.2 mm long, glabrous; corolla with aestivation valvate, fleshy, bistratose, cylindric to urceolate, 5-8 mm long, 5-7 mm in diam., yellowish-green often suffused with maroon, externally glabrous and internally moder- 240 ately to densely pilose with white trichomes especially on the lobes and the upper half of the tube, the lobes spreading or reflexed, triangular, acute, 1-1.2 mm long; stamens 10, included, ca. 6 mm long, the filaments ca. 3 mm long, flattened, weakly adherent to the very base of the corolla tube, villous, the anthers ca. 4 mm long, lacking spurs, golden, attached medially, the thecae ca. 1.5-2.3 mm long, papillose, basally incurved and apiculate, the tubules stout, ca. 1.5-2 mm long and 0.5 mm wide, dehiscing by slightly introrse, oblique pores. Berry spherical, ca. 12 mm diam., bunt " 5-locular. Distribution.—Known only from the heavily forested sl | the Río Grande de Orosi near Tapantí (Cartago), Costa Rica, and from the Fortuna area, Panama (Chiriquí), at 1130-1800 m. Flowering and fruit- ing: apparently extends through much of the year. Vaccinium orosiense is characterized by its amplexicaul leaves with rounded or cordate blade bases, short calyx lobes, corollas that are moderately to densely pilose internally at the throat, and by short anther tubules. When fresh, the basal region of the leaf blades is red, as is also the case in V. costaricense, V. floc- cosum, and V. luteynii, but usually not in V. poasanum. The relationships of V. orosiense are mentioned under the discussion of V. costaricense. Additional collections examined: COSTA RICA. Cartago: Hwy 224 on property of ICE Hydroelectric Plant, ca. 11.6-20 is E al the church in Orosi, steep roadside banks, 1500-1700 m, 29 Dec 1973, Almeda et al. 2180 (DUKE, F, GH, MO, NY, US); T project, wet road bank, 13 Dec 1975, Utley 3623 (CR, DUKE, F, MO). PANAMA. Chiri ino de quebrada Bonita ll por el embalse hacia el norte, 1130—1150 m, 8 Apr 1987, Valdespino et al. 631 (DUKE, NY) [JLL nes Va leia 631 may be V. Geh Gualaca-Fortuna Dam site road, 10.1 mi NW of Los Planes de Hornito, 1260 m, 10 Apr 1980 (£D, Antonio 4194 (MO ACKNOWLEDGMENTS Grateful acknowledgment is made to the National Science Foundation whose support over the years made this study possible. Obviously a study such as the present one would be impossible without the cooperation of the curators of the many herbaria who loaned their collections for this investigation. We am very much indebted to the curatorial staff of the following herbaria: A, BM, BR, CAS, CGE, CR, CU, DAV, D5, DUKE, F, G, GH, INB, K, LL, MICH, MIN, MO, MSC, NY, OXF, P, PH, PMA, POM, RSA, SCZ, TEX, US, WIS. We know the care and effort each of those loans take to prepare but without their effort this project could not be undertaken. We also express our sincerest thanks to Bobbi Angell for making the new species come alive through her beautiful illustrations. We thank Drs. J. McNeill, G. Moore, and D. Nicholson for their advice on the nomenclature of V. cordifolium for which we are most appreciative. JLL wishes to thank the Missouri Botanical Garden for providing him facilities to complete this manuscript. REFERENCES ALMEDA, E and DE BreepLove. 1992. A new Vaccinium (Ericaceae: Vaccinieae) from Guerrero, Mexico. Brittonia 44:50-53. Baxer PC. 1970. A systematic study of the genus Vaccinium sect. Polycodium (Raf) Sleumer, in the southeastern United States. Unpublished Ph.D. dissertation, University of North Carolina, Chapel Hill. Camp, W.H. 1942. On the structure of populations in the genus Vaccinium. Brittonia 4:189-247 [discussion of V. caespitosum, pp. 214-216] Kron, KA. W.S. Juno, PF. Stevens, D.M. CRAYN, A.A. ANDERBERG, PA. GADEK, C.J. Quinn, and J.L. Luteyn, 2002. Phylogenetic classification of Ericaceae: Molecular and morphological evidence. Bot. Rev. 68:335-424. Luteyn, J.L. 1976. Notes on neotropical Vaccinieae (Ericaceae). |. Gonocalyx - a genus new to Central America. Brittonia 28:3/-41. LUTEWN, J.L. 1987. New species and notes on neotropical Ericaceae. Opera Bot. 92:109- 130. Luteyn, J.L. and R.L. WiLgur. 2005. Ericaceae. In: Wm. Burger, ed. Flora Costaricensis. Fieldiana, Bot. NS, no. 45:i-vi, 1-107 NurrALL, T. 1843. Description and notices of new or rare plants in tl tural orders Lobeliaceae, Campanulaceae, Vaccinieae, Ericaceae, collected in a journey over the continent -— North America, and during a visit to the Sandwich Islands, and Upper California. Trans. Amer. Philos. Soc. Ser. 2. 8251-272 IT T HIT JI de y A E | a ERA » if de La H A EES, 241 Powel, E.A. and KA. Kron. 2003. Molecular systematics of the northern Andean blueberries (Vaccinieae, Vac- cinioideae, Ericaceae). Int. J. Plant Sci. 164:987-995. SLEUMER, H. 1936. Die Arten der Gattung Vaccinium in Zentral- und Südamerika. Notizbl. Bot. Gart. Berlin-Dahlem 13:111-140. SLEUMER, H. 1941. Vaccinioideen-Studien. Bot. Jahrb. Syst. 71:375-510. SMITH, A.C. 1932. The American species of Thibaudieae. Contr. U.S, Natl. Herb. 28:311-547, p1.1-19. [Gonocalyx pp. 352-355] STANDLEY, PC. 1920-1926. Trees and shrubs of Mexico. Contr. U.S. Natl. Herb. 23:1-1721. [Polycodium and Vac- cinium pp. 1101-1103]. STANDLEY, PC. and L.O. WiLLIAMS. 1966. Ericaceae, Flora of Guatemala. Fieldiana, Bot. 24(pt. 8, no. 2):88-127. STEVENS, P, J. L. LUTEYN, T. Ouver, et al. 2004. Ericaceae. In: K. Kubitski, ed. The families and genera of vascular plants, vol. VI. Springer-Verlag, Berlin & New York. Pp. 145-194. VANDER KLOET, S.P. 1985. On the generic status of Symphysia. Taxon 34:440-447. VANDER KLOET, S.P. 1988. The genus Vaccinium in North America. Research Branch. Agric. Canada Publ. 1828. xi + 1-201. VANDER KLOET, S.P, J.L. Barzen J.H. AppLeBy, R.C. Evans, and D.T. Stewart. 2004. A re-examination of the taxonomic boundaries of Symphysia (Ericaceae). Taxon 53:91-98. WiLBUR R.L. and J.L. Luteyn. 2005. Three previously undescribed species of Vaccinium (Ericaceae) from Costa Rica and Panama. Sida 21:1607-1614. WiLLIAMS, L.O. 1965. Tropical American plants, VII. Fieldiana: Bot. 31:167-174, £ T. TEA! A r 242 BOOK REVIEW Perry K. Peskin. 2006. The Search for Lost Habitats: 30 Years of Exploring for Rare and Endangered Plants—Book 1. (ISBN 1-933197-16-1, pbk.). Orange Frazier Press, PO. Box 214, Wilmington, Ohio 45177, U.S.A. (Orders: www.orangefrazer.com, 1-800-852-9332, Customer Service Department, Orange Frazer Press; or The Three-Hour Press Company, 3559 Strathavon Road, Shaker Heights, Ohio 44122, U.S.A.). $34.95, 288 pp., numerous color images and a 8 7/8" x 8". Want to go EE in captivating habitats? Want to set out on a Í t for rare a l plants? Want to enjoy a 1 J LT Jas Tf. 1 + f.I 42 +1 atu g y y y y MR enj ov the new book eel ird : kel c LC T KL ter (eo) ee gt J30 V, fr H E D J Ei I DI 4 Ranh? D T. Peskin, jor Lost Hap j EX] ch 5 1 +1 de - 241 " 1.1 ] J E tol J gl F. 5 e Pi MIL MARE NK GA Ikki A y AA Ando NS: A 1 TA 1 1 1 à s 1 1 141 "1 C LC 7 Fy Ll: A s Tu: Ce 1 1 1 H Le gag E g 1975 Search jot g curiosity of nat 1 his intriguing hes f d ic pl in both the Midwest and Canada. The book consists of nd + IJ.) s E C E ei 1 PE f 2 | EI 1 tl 7 SC, +1 r r IN d E 1 1 f. Js "r1 1 1 JE Sg) Lä 1 i a | calle pu. 4 T 1 M "p Jp while LE odd bee bae nian nature pan PANE heh SEN deen of an area. A few chapters provide information ] Sage M EAM are focused on plant families like E D El Kaz l.l F-11 1 1 +l Es = F 11 J The author has writter The Search for Lost Habitats t ible t t readers. S basic botanical knowledge would be helpful, but the book is unassuming and readable b un interested pe The author has provided helpful information within Ee text Ge De current Endangered Species A g g e species discussed. Peskin has = ae many colorful r I o i d Additi l I I t L I fl bitat S ol i j l i various ftl Aid 1G La! i Appendix A.A I lix B includ I lant 2 tifi I ith an U. H status under the Endangered u Act A S a. 1 an ind plete the text. Although habitat and specific | are Mondes the See has wisel ly t to allow p ial pl hi |; | iti TT asco cas 1.1] 1.1 ] 1 de 1 A r cmm | E 1 ny: A zl + + a 4 "IP al TI ao} ee A. eke + 1- 1 1 1 1 [a ] a Cl 41 y 21.311 J L1 JA +I [5 2] — AF xx E h of o F ALLA MIL VY AL AL was to the Lakeside Daisy Sut Nature Presa where I saw tl jue limestone habitat these pl inhabit (Lakeside daisies are discussed in Essay 4). Peski h inf ti both text and color photos to visualize and experience the location or plant discussed Ilook lourd to iBook Two of the series.—Lee ebe Botanical Research Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S.A J. Bot. Res. Inst. Texas 2(1): 242. 2008 RE-DISCOVERY AND TYPIFICATION OF THIBAUDIA LAURIFOLIA, MACLEANIA INSIGNIS, AND M. COCCINEA (ERICACEAE: VACCINIEAE), AN ADVENTUROUS HISTORY James L. Luteyn Institute of Systematic Botany The New York Botanical Garden Bronx, New York 10458-5126, U.S.A. jluteyn@nybg.org ABSTRACT Herbarium work has now revealed the true identity of and the ability to typify Thibaudia laurifolia M. Martens & Galeotti, Macleania insignis M. Martens & Galeotti, and Macleania coccinea Decne. Orthaea laurifolia (M. Martens & Galeotti) Luteyn is proposed as a new combination. RESUMEN El trabajo de herbario] lado la identidad verdadera de | ies y | ibilidad de tipificar Thibaudia laurifolia M. Martens € Fa TP! T EE Ys (ha NH + Keen a | PA abajo E H I 1 KA + Ce fal tti Ad T : H Ta C Galeotti, M ia insignis M y Luteyn como combinación nueva. INTRODUCILON In 1967, Yeo wrote a nice little Paper d Tone on some SES of Macleania (Ericaceae).” That paper helped to sort out the t of Macleania, the species numbered 1—15 in Smith's (1932) Geh of the Thibaudieae. That sions was characterized by having a single anther tubule and connate staminal filaments, the subgenus Macleania as defined by Luteyn (1997), several of which are still cultivated. Yeo recognized Smith's taxonomy as “largely satisfactory, but that he [Smith] has misinterpreted a number of type specimens." ie pue 51 i: of the article, Yeo discussed M. insignis with its numerous synonyms. Yeo's attempt to clarify the t of some of these names was also largely satisfactory, but the types of three names discussed b still need further clarification. Henri Galeotti was one of the first botanical explorers of Mexico. His collections were offered for sale and this explains their dispersion over many institutions. However, the herbarium (BR) of the National Botanic Garden of Belgium acquired large quantities of Galeotti material on two separate occasions. A first and the largest part was purchased after Galeotti's death in 1858, as reported in the minutes of the meeting of 14 February 1859 ofthe "Société royale d'Horticulture" [which later became the National Botanic Garden]. This was not a complete set, however, of Galeotti's Mexico collection of ca. 7300 specimens (perhaps only about half). Much later, in 1932, Pierre Martens donated the herbarium of his father Martin Martens to BR. It contained circa 2000 Galeotti specimens that M. Martens himself had examined with Galeotti. Upon his return from Mexico, the horticulturist Galeotti developed a collaboration with M. Martens (himself a professor in chemistry and botany at Louvain). The main output was the “Enumeratio synoptica plantarum phanerogamarum ...," identifying the Mexico collection and published in a long series of articles presented at the meetings of the "Académie royale des Sciences et Belles-Lettres de Bruxelles" (Martens & Galeotti 1842—1845) (pers. comm. 2008 with E. Robbrecht). It is with the aforementioned brief introduction that I set the context for the typifications mentioned in this paper. Thibaudia laurifolia M. Martens & Galeotti In 1842 (page 530), Martens and Galeotti described Thibaudia laurifolia based on Galeotti 1814 (protologue: leaves short-petiolate, acuminate; racemes elongate, 6—8-flowered). In 1932, in his classic "The American J. Bot, Res. Inst. Texas 2(1): 243 — 248. 2008 ae MEAN xas 41) [| [| fal Dos a ID L I Mon ndis f Tex 244 Species of Thibaudieae,” A.C. Smith cited the Galeotti 1814 sheets at B and US as the “type collection of Thibaudia laurifolia." Unfortunately, Galeotti 1814 is now seen to be a mixed collection of at least four elements based on ma- terial seen at BR, G, K, NY, P, US, and W. These include examples of Macleania insignis, Satyria warszewiczit Klotzsch, an unknown species possibly Satyria panurensis (Benth. ex Meisn.) Benth. & Hook.f. ex Nied., and the true Thibaudia laurifolia. Based on my studies, there were at least 11 herbarium sheets numbered as Galeotti 1814 including one destroyed at B (after A.C. Smith had seen it), but not including the fragments at F. All the duplicate collections of Galeotti 1814 seen, except the herein designated lectotype, have original printed Galeotti herbarium labels (Fig. 1A) that cite the collection locality as “Chinantla” and [Oaxaca] “Mexico,” and the collector as "H. Galeotti.” All the duplicates were identified as Macleania insignis, with the Eee of the maa sheet. They are renes by: "vi e (fide AC Smith 1932 de y A dar) g WWID BR Greg S.P 627855) = = Thibaudia laurifoli al ing mod | red tation label “TYPUS,” and an original hand-written speci- d of Martin M that ;“Thibaudial oe pee d D d on T 2-3 in Dorr (1986), r] and pers. comm. 2008 with E. Robbrecht]; =lectotype of Thil BR = M. insignis (recently incorrectly annotated as “TYPUS” by means of a sed felt-tip pen in an — hand; not seen ia Smith; photo NY neg. 12557 gnis (unannotated, bearing only an original printed Galeotti herbarium label that has on it the hand-written “Thibaudia - M. insi laurifolia H. Galeotti") BR - unknown Ericaceae, perhaps Satyri iczii (bearing original printed Galeotti herbarium label that | it the hand-written "Thibaudia laurifolia, nobis") F -Mi ig ' (f g e P correct 1 y TES | AS insignis, f type cetate) F=M.i : o g tex Gi tly tated as i on of T. a G-M.i 1 on two sheets, | ted ti herbarium label that | t the hand-written “Thibaudia,” and EH annotated as “type coll of Thibaudia hanai M. & G.” by Smith in 1932; pe F neg. 26647 P - M. insignis cm annotated as "type coll. of Thibaudia laurifolia M. & G." by Smith in 1931, by Sleumer in 1959, and also by Luteyn in 1978) S (sheet 573239; barcode 00113543) = M. “Thibaudia” (Fig. SECH and i dud annotated as Wee col e Thibaudia E M & ES ? El Se in En bel that has on it the hand-written "Thibaudia laurifolia 1 Galeotti herbarium label that has on it the hand-written W (sheet no. geng g I H. Gal." tation label as Macleania insignis by Cufondontis in bis: W (sheet no. 117231) ] Eri , perhaps Satyri is (1 g W herbarium label that reads “1814. Thibaudia laurifolia Gal.” and tation label as Macleania insignis by MM in 1933) LECTOTYPIFICATION OF THIBAUDIA LAURIFOLIA — pee (M. Martens & Galeotti) Luteyn, comb. nov. Basionym: Thibaudia laurifolia M. Martens & Galeotti, cad. Roy. Sci. Bruxelles 9(2):530. 1842, non Blume, eas E Ned. Ind. 859. 1826. Tyre. MEXICO. Oaxaca: Chinantla, 4000 F s (ID, H Galeotti 1814 (ectorvre, designated her (l 627855), photo NY neg. 12556). Be stipitata Luteyn, Brittonia 28:403, fig. 2. 1976[1977]. Orthaea stipitata (Luteyn) Luteyn, Nord. J. Bot. 7:33. 1987. Tyre. Oaxaca: Comaltepec, Ixtlan, La Esperanza, 1524 m, 19 Apr 1970 (£D, MacDougall 606.5 (HOLOTYPE: NY). The sheets of Galeotti 1814 from B and US that A.C. Smith (1932) cited as the “type collection of Thibaudia laurifolia” in his “The American Species of Thibaudieae" are now both seen to be M. insignis. In that same paper, on page 369, Smith also included Thibaudia laurifolia in the synonymy of Macleania insignis. He did not see (or at least did not annotate) the specimens of either Galeotti 1814 or 1827 at Brussels (BR), but re- lied on sheets at B and US for his concept. Thus he did not know that Galeotti 1814 was a mixed collection; therefore, he synonymized Thibaudia laurifolia under Macleania insignis. Nevertheless, the only specimen that fits the protologue of T. laurifolia is one of the sheets at BR, which Smith did not see, and which bears an original hand-written specimen collection label (Fig. 1B) that perfectly matches the handwriting sample for Martin Martens kept at BR (pers. comm. 2008, E. Robbrecht). That sheet has now been properly an- notated as the lectotype of T. laurifolia and the remaining sheets as Macleania insignis, Satyria warszewiczii, or Satyria cf. panurensis. In 1976, Luteyn described Empedoclesia stipitata based on MacDougall 606.5 (holotype: NY). Later, lutevn. T d F "al, WK A $i Coll il. Gartorn. > 4446 bea a A 2 &, x) SE AM £t Zë ey Ex Pd e E hn * Card k | "e Mexico, RT ely dee LAMA e p nr E 4 d d ER , S ad La PF A DADES : à "1 "AH (AR ^ ff iy e£ | 1 DA “wy y / d É rue Pi “A Le TA € d > ite É FLAG e c to : = f ELA A 5 / / ] ^3 "d " i Ge - “a A = L t d t Es á sá + D T E a v «4 = "xw [ = É Pd É a É 7 ` » 6 e S "J E Er ir; c i e Ag er] iiig rae b= S Coll. H. GasgormI , (Vera-Cruz) h 1840, Vera a ES e E j i * s PAL e Fi E. Fd j E | CE, INN (a, tro 2; r A r3 D e , É : S F Pn fot. wth y Ba / B. y É Act £ £f / Y. Per EI “a E Le i Pd af d? " x y A7 Doo r e d v d A al AA Lé its JE Dr, À d Cé EP Wo | T Cordilie L A Cordi E 7 . ata Ci Coil. 4] rri ANTES UST Fá e i peras Aux) d is Meri, t P 52 d dr ¡Es Ze Ld "ES + d Li L J L | | L L 11 : P [| DAA AA H H H o Li 2 Fic. 1 f , all probably 18405. A, Original printed Galeotti herbarium label in rf.» | E (IT) | | Ey i 404 ATA I SA R PF. . 1L i Pag E] II! "2 | L ge Al L J rad f 3 Pt f E Pn al E eet O44 L | ZE "FL ot F’ I Eis Fr. A len L E | p I al L J £u af i - LI Dre 1827, aL | de A fag i s n Pa 1L I A Hi Im. L T L L J fi fal i E nm L r £u - T bi LI tyr EMS i H F Ft | | B a || L L | L | L L | EII £P I AA E nm L A E ez? PAR Fe bo brd > y yl faa i g n al | yl fas |] PA EL J | . HI AP AL ye Al L I EAR Martanc i, nmm al £I J "^" TE Dall La sor: é JI. J ore IL L| L | | nni 245 246 Jo t tani i Texas 2(1) Luteyn (1987) synonymized the genus Empedoclesia Sleumer under the genus Orthaea Klotzsch and made the new combination Orthaea stipitata (Luteyn) Luteyn. As a result of herbarium studies carried out in the United States (F, GH, MO, NY, US) and in Europe in 1986, 2006, and 2007 (B, BR, CGE, E, G, K, L, MA, OXF, P, S, W), it is now clear that both Thibaudia laurifolia and Orthaea stipitata represent the same species. Therefore, the new name becomes Orthaea laurifolia (M. Martens & Galeotti) Luteyn. Macleania insignis Martens and Galeotti (1842, page 531) also described Macleania insignis in tl ticle as above, based on Galeotti 1827 (protologue: leaves subsessile, ovate, obtuse, 3-nerved; es fasciculate). The protologue stated that the species was found on oaks near to the German colony of Mirador at 4000 feet elevation, and that it was also found in the eastern cordillera of Oaxaca near Talea and Llano-Verde at 5000 and 6500 feet. Based on my studies, there were at least nine herbarium sheets numbered as Galeotti 1827 including one destroyed at B, but not including the fragments at F. The original printed Galeotti herbarium labels of all the duplicates of Galeotti 1827 (Fig. 1C,E) give the general locality as “Cordillera, (Vera Cruz) Mexico,” along with the dates of “Jun.—Oct.” and “Coll. H. Galeotti, 1840,” although the individual labels of some of the duplicates seen at BR2x), G, KQx), NY, and P have three small differences with regards to more specific hand-written information. Two of the sheets at BR mention Llano Verde at 6000 feet (Fig. 1D), while the sheets at G and P give no specific locality, but do say the plants were “parasite” on oaks; the K-Herb. Hook. sheet says “parasite on oaks at 4—6000 ft." One of the BR sheets that was relatively recently designated as “HOLOTYPUS” by P. Bamps (curator at BR) bears two hand-written labels at the bottom of the sheet—one an original hand-written specimen collection label mentioning “Prés Llano Verde 6000" (Fig. 1D), and the other an original printed Galeotti herbarium label that reads “Macleania insignis, nobis” (Fig. 1C). This is the only sheet upon which the original printed Galeotti herbarium label gives the word “nobis”. The two labels match the handwriting of H. Galeotti based on personal communication in 2008 with E. Robbrecht (BR). This I feel means that this sheet was seen by the authors and is appropriate to designate as the lecto- type, which is how I have annotated it. The W sheet reads “Macleania insignis nobis,” but does not have an original printed Galeotti herbarium label nor is it written in a recognized hand. The other Galeotti 1827 sheets have all been annotated by me as syntypes. All of these are identified as Macleania insignis and are represented Së B = Mad e fide A C Smith 1932 (not E A during WWII) BR (barcode S.P. 627888) - M. insignis (etos: annotated as “HOLOTYPUS” by P Bamps, bearing also an original printed Galeotti herbarium label that reads “Macleania insignis, nobis” (Fig. 1C), and a second hand-written specimen label that reads “Prés Llano Verde 6000” (Fig. 1D), both in the hand geck Gase e comm. ii ads EE BR (barcode S.P 627921) = M. insignis (syntype, t leotti herbarium label with hand-written “Macleania coccinea Nobis” in the hand of H. Galeotti (Fig. 1E) chai NY geg 12555; also herein designated as td e M. oa Cd E nu F = M. insignis (syntype tragment ex G; voucher [or r neg. 26 46) G=M. sds e cal Mine *TYPE COLL." by Smith in 1932; photo F neg. 26646) K-Herb. Benth. = M. insignis (syntype, correctly annotated as “TYPE COLL.” by Smith in 1931) K-Herb. Hook. = M. insignis (syntype, correctly annotated as "TYPE COLL." by Smith in 1931) NY - M. insignis (syntype, duplicate ex W) P - M. insignis (syntype, correctly annotated as "TYPE COLL." by Smith in 1931, by Sleumer in 1959, and also by Luteyn in 1978) W (sheet no. 0008497) = M. insignis (syntype, with unrecognized hand-written label reading *Macleania insignis nobis") TYPIFICATION OF MACLEANIA INSIGNIS Macleania insignis M. Martens & Galeotti, Bull. Acad. Roy. Sci. a 902): o ae Tier. MEXICO. VERA Cruz: Cordillera, Llano Verde, 6000 ft, Jun—Oct, 1840, Galeotti 1827pp | 27888), pho NY neg. 12554]. Macleania coccinea The name “Macleania coccinea" was first introduced in the protologue of M. cordata Lem. (Lemaire 1848). In that publication, which included a color illustration of the new M. cordata, Lemaire mentioned that his new Luteyn, Typifi ti | bination in Eri 247 species was cultivated at the establishment of “M. Jacob-Makoy, de Liége,” having been introduced alive in 1842 or 1843 by “M. Ghiesbregt” along e two other "pes “les i coccinea ... et insignis... In 1851, Decaisne formally described Macleania a cultivated plant collected by A.B. Ghies- brecht, presumably from Mexico (see Yeo 1976). The protologue (Decaisne 1851) included a description of the new species (in French), a paragraph about cultivation techniques for species of Macleania, and a beautiful color illustration of M. coccinea that as Yeo (1976) mentioned is *very similar" to the one of M. cordata. It does not, however, give any information about the original collection or its locality; therefore, the actual location of any type material is still uncertain. There are two herbarium specimens at BR bearing the name M. coccinea, the one of Linden 432 (barcode S.P. 627954) from the *Collection Martin Martens," donated to BR by Pierre Martens in 1932, bears the name "Macleania coccinea” on an original hand-written specimen collection label in the hand of Martin Martens (pers. comm. 2008, E. Robbrecht) (Fig. 1F). The second sheet, Galeotti 1827pp (type number for M. insignis), also ex "Collection Martin Martens," donated to BR by Pierre Martens in 1932, has the more explicitly written name "Macleania coccinea Nobis" in the hand of Galeotti on an original printed Galeotti herbarium label that gives the general locality as “Cordillera, (Vera Cruz) Mexico,” along with the dates of “Jun.—Oct.” and “Coll. H Galeotti, 1840" (Fig. EE These are the ony two herbarium sheets that I have seen that bear the name M. coccinea. Since D yI Dt at BR a E Cowan 1976, ti t f Macleania tion number page 607), Lam herewith d l heet yp of which is itself a syntype of M. uus TYPIFICATION OF MACLEANIA COCCINEA Macleania coccinea Decne., Rev. Hort. [Paris], sér. 3, 5:301, t. 16. 1851. Tr. MEXICO. Vera Cruz: SEN “Totutla, 4000 [ft], Llano Verde 4-6000 [ft],” Jun-Oct, 1840, Galeotti En NEOTYPE, designated her 921), oto NY neg. 12555 ACKNOWLEDGMENTS +1 I wee] E + Liz I wish to t] kthe Nati | Science Foundation for their ti ] support me to visit European and American herbaria, and to the herbaria themselves (mentióned above) and their curators for their help and hospitality. I also wish to thank Bruno Wallnófer (W) for checking a type at Vienna; Scott A. Mori (NY) TM Han er PIED dinis for me; Michael Nee (NY) for beneficial discussions; Larry Dorr (US) f final manuscript; Rose Gulledge (US), Nestor Perez- Moliere (NY), and Robbin Moran (NY) for help preparing the figure; and especially to Elmar Robbrecht (BR) for his help to identify the handwriting of Martin Martens and Henri Galeotti and for historical information about the acquisition of Galeotti material at BR. REFERENCES DECAISNE, J. 1851. Macleania coccinea. Rev. Hort. [Paris], sér. 3,5:301, t. 16. Dorr, L.J. 1986. Jean-Baptiste Duerinck (1809-1857) and his collections from the Middle Western United States Bull. Jard. Bot. Nat. Belg. 56(3/4):397-416. [ EMAIRE, CH. 1848. Macleania Midi i m Jard. Eur. 14:312, plus color plate. LUTEYN, J.L. 1976. Notes on Vaccinieae (Ericaceae). Ill. New and noteworthy species from Mexico and Central America. Brittonia 28: 400- 406. LUTEYN, J.L. 1987. Orthaea (Ericaceae-Vaccinieae): new species and redefinition of the genus. Nord. J. Bot. 7:31-37. LUTEYN, J.L. 1997. A review of and taxonomic realignments within the neotropical genus Macleania (Ericaceae: Vaccinieae). BioLlania Ed. Espec. No. 6:455-465 MARTENS, M. and H. Gateotti. 1842. Notice sur les plantes des familles des VACCINIEES et des ERICACÉES, recueillies au Mexique par M. Henri Galeotti, et publiées par MM. Martens et H. Galeotti. Bull. Acad. Roy. Sci. B.-Lettres Bruxelles 9(6):526—544. [reprinted as Martens & Galeotti,"Enum. pl. Galeotti” Part 1, pp. 1-19. 1842, fide Stafleu & Cowan (1981, page 2191 248 Martens, M. and H. Galeom. 1842-1845. Enumeratio synoptica plantarum phanerogamarum ab Henrico Ga- leotti, in regionibus Mexicanis collectarum. Bull. Acad. Roy. Sci. B.-Lettres Bruxelles 9(6):526-544, 9(7):32-47, 9(8):227-249, 9(10):372-393, 10(2):110-134, 10(3):208-224, 10(4):341-360, 10(7):31-52, 10(8):1 78-200, 10(9):302-321, 11(3):121-137, 11(4):227-243, 11(6):355-376, 11(8):61-79, 11(9):185-196, 11(10):319-340, 12(2):129-149, 12(7):15-36 and 12(9):257-278. Sum A.C. 1932. The American species of Thibaudieae. Contr. U.S. Natl. Herb. 28(2):i-xiii, 311-547, plates 1-19. SrAFLEU, FA. and R.S. Cowan. 1976. Taxonomic literature, Ed. 2, Volume |: A-G. Regnum Veg. 94. Bohn, Scheltema € Holkema, Utrecht. SrAFLEU, FA. and R.S. Cowan. 1981. Taxonomic literature, Ed. 2, Volume Ill: Lh-O. Regnum Veg. 105. Bohn, Scheltema & Holkema, Utrecht. Yeo, PF. 1967. Notes on some species of Macleania (Ericaceae). Baileya 15(2):45-59. REVISION OF SIPHONANDRA (ERICACEAE: VACCINIEAE), A GENUS ENDEMIC TO PERU AND BOLIVIA James L. Luteyn Edgardo M. Ortiz Institute of Systematic Botany Herbarium Areqvipense (HUSA) The New York Botanical Garden Universidad Nacional de San Agustín de Arequipa Bronx, New York 10458-5126, U.S.A. PERU ABSTRACT The genus Siphonandra is revised. T ies, Sipl | Lut & E.M. Ortiz and Siphonandra santa-barbarense ] put int ae context e a ge? to the five known species in the genus. An fe Luteyn & E.M. Ortiz, are described, iustrated 1, mapped, m Asas Ta 1 r š E i i o J * JL RESUMEN c H 1 DH Cus J E: ] -1 21 " ] z c+} J T t Er ed M (Art Cial 3 + - ME "TAE : GE santa barbarénse Luteyn n & E.M. Ortiz y E p £ ) F especies, Ci E INTRODUCTION Siphonandra is a small, distinctive, high-elevation genus of five species that are endemic to the geographical range from northern Peru (Amazonas) to northern Bolivia (Cochabamba). It was named by Klotzsch (1851) on the basis of its long and slender staminal tubules, but is also characterized by its articulate calyx, usually connate filaments, and perfectly terminal pores at the tips of the tubules (Smith 1932; Luteyn 1998, 2002b). Siphonandra elliptica, the type species, spans the entire geographical range of the genus from northern Peru (Amazonas) to northern Bolivia (Cochabamba), where it is common. The other four species, however, are only known from one collection site each, thus making it difficult to assess relationships between any of the species in this genus. The description of Siphonandra nervosa and S. santa-barbarense brings to three the number of species found in Peru and to five the number described overall in the genus, which until a few years ago was only known a the bue e S. elliptica (see Luteyn 20022). (1932) ically with Lysiclesia (=Orthaea), Ceratostema, and Orthaea EN A ) on the basis of their hayi ing in common the characte: of articulate calyces and anther dehiscence through apical pores, although he believed that these similarities were “doubtless derived by distinct and somewhat parallel courses.” It is true that the genus Siphonandra is morphologically most similar to those species of Ceratostema that have anther dehiscence by perfectly terminal pores (but there are about as many species in Ceratostema that have dehiscence through oblique or lateral pores); both genera have articulate calyces, large corollas, and equal stamens with very slender anther tubules with apical dehiscence. The major dif- ferences are that Ceratostema has corolla lobes proportionally very long and narrow (usually greater than 4 mm long) and its corolla is also usually basally ventricose, whereas Siphonandra has corolla lobes very short (equal to or less than 2 mm long) and its corolla is cylindric with parallel sides along its entire length. Also the geographical ranges of the two genera do not overlap, with Ceratostema ranging from Venezuela to northern Peru (Piura, Cajamarca), north of the Huancabamba Depression, whereas Siphonandra ranges from northern Peru (Amazonas), south of the Huancabamba Depression, and into northern Bolivia (south to Cochabamba). Orthaea seems more distantly related having alternately unequal stamens. Morphological and molecular studies clarify that Siphonandra is definitely found within the blueberry tribe Vaccinieae (Kron et al. 2002a). The only published molecular phylogeny of Vaccinieae that included Siphonandra (Kron et al. 2002b) used matK s HIE data, which placed Siphonandra within an “Andean clade" sister to a polytomous group containing , Disterigma, and one species of Ceratostema. LA Li + L | Rot Pac Inet Tavac 2(1): 240 — 261. 2008 [| [| fal Dos H In LI PICA EF fa 250 t U A subsequent study within the “Andean clade" did not include Sip! lra (Powell & Kron 2003). Therefore, the overall data that was based on only one sequence from one Geet of Siphonandra p and there is still uncertainty about its affinities. KEY TO THE SPECIES OF SIPHONANDRA . Hlaments distinct. 2. Rachis to 0.5 cm long, 3-6-flowered; calyx ca. 9 mm long, the tube densely bearing sq warts; corolla 35-37 mm lon 3. Si Sonda magnifica 2. Rachis 12-25 mm long, 10-22-flowered; calyx 6-10 mm long, the tube without warts; um: 26 mm lon 5. Siphonandra santa-barbarense 1. Filaments connate. 3. Corolla 43-48 mm long 1. Siphonandra boliviana 3. Corolla ca. 25 mm long. 4. Leaf blades coa: (3.5-)5.5-8.5 x (1-)1.5-3 cm, a | and bluntl inally entire, the lateral nerves 6-8 per side; inflorescences SE owed the rachis’ ^ 0.7 cm bo hob cels to 7 mm long 4. Siphonandra nervosa 4. Leaves oblong, ovate-oblong, or slightly obovate, 3-5.5 x 1-2.5 cm, apically obtuse or subacute, margin- ally entire or faintly crenulate, the lateral nerves 3-5 per side; inflorescences 5-15-flowered, the rachis striate, 1.5-5 cm long; pedicels 7-20 mm long 2. Siphonandra elliptica TAXONOMY Siphonandra Klotzsch, Hanara 24: Së ee Cea ana uus sect. ~o e Hook. f. ex Benth. & Hook. f., Gen. Fl 2:570 p.p. 1876. I ex G. Don) Klot Sip! i Griseb. in Lechl. Berb. Amer. Austr. 58. 1857, nom. nud. Tyre species: Siphonostema myrtifolium Griseb. Tens oro D yu hrubs. I It te. with blad ergreen, marginally recurved and entire or faintly crenulate; pinnately nerved, Ton panie axillary bud GER 2, valvate. Inflorescences axillary, solitary, racemose, with 5-22 pedicellate flowers; floral bract inconspicuous; bracteoles 2. Flowers (4-)5-merous, without odor; aestivation valvate; calyx articulate with pedicel, the tube short-cylindric, terete to narrowly winged, the limb spreading, the lobes 5; corolla cylindric; stamens P equa, pen Sugar longer than corolla, the filaments distinct to connate, equal, shorter than anthers lacking disintegration tissue, the thecae granular, the tubules distinct to Base elongate, slender, flexible. dient half as wide (or less) as EES 4-5 times e Pun nd sayeth d ed dics nd dee pores; pollen without viscin t Fruit a spherical De The current g t follows that of Klotzsch (1 851) and Smith (1 932), but differs by the acceptance D of species with dana staminal filaments (vs. connate in original concept) as first noted by Sleumer (1941). 1. Siphonandra boliviana Luteyn, Sida 20:13-15. 2002. (Fig. 1). Tre. BOLIVIA. La Paz: Prov. Bautista Saavedra: Cha razani, W of Chullina, 3400 m, 1 Aug 1994 (il), B. Herzog H200 ( : NY; 1sotyres: LPB, n.v., LZ, n.v.) oa ee ERN mature bi FUN terete, glabrous, the bark exfoliating in thin strips; immature ] I] angled or ribbed, short-pilose with white hairs; axillary bud scales 4 ca. 2 mm long, slirieilase. Leaves with blades coriaceous, elliptic to oblanceolate, 2.5—5.5 x 1-1.8 cm, basally cuneate, apically broadly acute to nearly obtuse, marginally entire, the lamina essentially glabrous above or sparsely short-pilose proximally along midrib, sparsely pilose beneath especially along midrib, also provided with reddish-brown, basally swollen, glandular fimbriae beneath; pinnately nerved with 4—6 o. veins ee near un dus en ^" eel veins impressed above and raised beneath, t | ly raised beneath; petioles rugose, subterete, broadly RE above 4— 5 mm lone inflorescences ca. 20-flowered, apparently nod- ding; rachis subterete, striate to angled, at least 5 cm long (still in bud, apparently still elongating), densely short-pilose with white hairs; floral bract ovate, acuminate, 3-4 mm long, densely short-pilose; pedicels subterete, striate to angled, 11-13 mm long, densely short-pilose as rachis; bracteoles located in proximal Luteyn and Ortiz, Revision of Siphonandra 251 à d ^ WES. AA NIV (5 GE DUO ae A pira A Ais Toe tart ¡o E | 1. E “gr H :ud CH: xA RU. t. x d ato ee VE G icm. Kë UPS [| E tu i i t i 1 D A E 1 Fic 1 es L J L F I A A Hahit R m PO E Fi L Li L laaf attachmant | ln L J || Te ri L Li Fr [| P d "2 F f d bract. pedicel. hracteoles. calyx. and corolla. D. | itudinal ti f lla «I i iti f ct ES | | | ventral and F | f f d Zi, ki 1 kl F F A asf agl [EP RET | fa D I.E: Fra rj A IT as ` LI = £8 1/3 pedicel, similar to floral bract, 2-3 mm long. Flowers with calyx 7-8 mm long, densely short-pilose as rachis, the tube cylindric, ribbed, 4—5 mm long, rounded at base, the limb spreading-campanulate, 3.8—4.5 mm long, the lobes deltate, acute, 1.5—2 mm long; sinuses obtuse; corolla long-cylindric, 43-48 mm long, 6—7 mm diam., short-pilose throughout with white hairs, the lobes deltate, acute, ca. 2 mm long; stamens ca. 32 mm long, the filaments connate, ca. 7 mm long, glabrous, the anthers ca. 28 mm long, the thecae granular, ca. 6 mm long, incurved at base, the tubules ca. 22 mm long; style about equaling corolla. Berry not seen. 252 Icones.—see Luteyn 1998, 2002 (Fig. 1). Distribution (Fig. 2). —Endemic to Bolivia (La Paz) and known only from the type collection, which was made in a Weinmannia forest at 3000-3500 m. Rare and endangered. Siphonandra boliviana is easily distinguished morphologically from the other two species by the char- acters mentioned in the key and diagnosis. 2. een elliptica (Ruiz & Pav. ex G. Don) Klotzsch, Linnaea 24:24. 1851 (Fig. 3). Thibaudia elliptica Ruiz & Pav., Fl. Peruv. Chil. 4: pl. 384, fig. b. 1802, nom. nud. Thibaudia bo dm Ruiz m m Don, Gen. Hist. 3: 861. 1834 Ceratostema da (Ruiz & Pav.) Benth. & Hook. f., Gen. Pl. 2: 570. q & Pav.) Britton, Bull. Torrey Bot. Club 20:137. 1893. Tree. PERU. Muña, Pillao and Acomayo, Ruiz & NM s.n. (setos, TEN designated by Luteyn: MA! (numbered “15/50” and Gerten MA747445; photo F neg. 29347)]. I suspect that G. Don validated the name in Thibaudia based Illustrations: Hooker. Icones pl 2: t. 108. simply on the plant illustrated by Ruiz and Pavón, 1837. The Hooker illustration is based on *Cuesta of Huanacabra, Andes e Peru. Mathews. (n. 884)” which is at K (1 full sheet and another 1/2 sheet), probably BM, CGE, and OXF also. See also Luteyn 1998. [There is a “884” numbered collection without collector or locality at BM, which also has the hand-written name “Ceratostema corymbosa” written on its label; at OXF too. We do not know of any citation anywhere of the name “C. corymbosa,” which must be unpublished.] (Fig. 3). dci monum bac dd in Lechl., n Amer. ioe eg 1857, nom. nud. Terr collection cited: Geste 2276 (K!). The K yr 1931. The sheet at G S. elliptica by ACS in 1932. | bach in Lech). Berb. Amer. Austr. 58. 1857, nom. nud. Ter collection cited ees (KD. The K sheet is 11 ked f ACS in 1931. The sheet at G S. pilosa by ACS in 1932 Ceratostema ESCH Britton, Bull Torrey Bot. Club 20:137. n nom. ride Britton ie this new name on the Thibaudia UM il- eg in WJ. Hookers, Icon. Pl. t. 108 (1837), R y T cá A O ll ca ana qu c M ee d to match, the plant illustrated by Ruiz and Pavón (1802) and tl f the lectotype itself at MA sl ] ] ly plinerved with 3—5 nerves Get Sg bere the ill ti f Hooker (1837) sl plant with definitely pinnately nerved leaves. Britton Pol vim. Ds weberbauerii Hoerold, Bot. ped Syst. 42: 316. 1909. TPE. PERU. Puno: Sandia, 3100-3300 m, Apr 1902 (fl), Weberbauer 740 (HoLoTYPE: B, destroyed). Siphonandra pilosa A. C. Smith, Contrib. U.S. Natl. Herb. 28(2):355. 1932. Ceratostema pilosum (A.C. Smith) J. E Macbride, Univ. Wyo- ming Publ. 11: 42: 1944. Ty. BOLIVIA. La Paz: Larecaja: Cerro de Tuile, vic. Tacacoma, 3350 m, May or June 1860, Mandon 549 [HoLoTyPE: NY; rsotvrEs: G(4x; photo F neg. 26655), K-Herb. Hook., P(3x)]. Terrestrial or epiphytic shrubs to 2 m tall; mature stems terete, EROS the bark cracking in thin, longi- tudinal strips yellowish to light brown in color; immature stems subterete , striate, puberulous when young, glabrate or persistent; axillary bud scales 2, valvate, ovate, inate to Pilas striate, ca. 3-4 mm long, not pseudo-stipular. Leaves with blades thick-coriaceous, oblong, ovate-oblong, or slightly obovate, 3-5.5 x 1-2.5 cm, basally cuneate, apically obtuse or subacute, marginally entire or faintly crenulate, slightly revolute, the lamina glabrous or sparsely brownish-pilose above, pilose to sparsely pilose beneath with scat- tered, appressed, dark brown hairs ca. 0.4-0.5 mm long, glabrate, often strongly punctuate on both surfaces from deciduous glandular fimbriae; pinnately nerved, the midrib impressed above, prominent beneath, the secondary nerves = 5 pe ue MNA ascending at margin, slightly impressed above, plane to raised beneath, t ioi netimes obscure, plane; petioles subterete, rugose, 3-5 mm long, puberulous or ibus estime 5-15-flowered, deciduously bracteate at base with several, ovate bracts ca. 1.5-2 mm long; rachis striate, 1.5-5 cm long, glabrous or pilose with copious, pale, spreading hairs to 0.3 mm long; floral bract deciduous, ovate, acuminate, ciliate with white hairs to 1 mm long, the lamina glabrous, ca. 3.5-4 mm long; pedicels rugose, 7-20 mm long, pilose or glabrous, bearing several minute, cartilaginous teeth at apex; bracteoles basal, ovate, acuminate, ca. 1.5-3 mm long, ciliate. Flowers with calyx ca. 7-8 mm long, the tube cylindric, sometimes 5-10-ribbed, ca. 3-3.5 mm long, glabrous to pilose with short, pale, spreading hairs, the limb 3.5-4.5 mm long, the lobes to ca. 1.5-3 mm long; sinuses broadly rounded; corolla cylindric, 20-26 mm long, 4-5 mm diam., glabrous to sparsely pilose, red with whitish or pale greenish tips when fresh, the lobes triangular, obtuse to subacute, ca. 1-1.5 mm long; stamens ca. 25 mm long, the filaments connate (supposedly rarely distinct in type of S. weberbauerii), ca. 5-6 mm long, 253 Luteyn and Ortiz, Revision of Siphonandra ==) 05º NO 1 | T BC n sa ut |. N Pa: 4 E - "i Sip ET pw A E | Yy- " Rope = d FL nor ? s ei P" do ee Ed 3d 15* Fig. 2. Distribution of species of Siphonandra. A. O 5. elliptica. B. O. santa-barbarense; A S. nervosa; a S. boliviana; © A. magnifica. £T e TOM) lexas 411) i 254 — oe mm e e o o S * "ac P p ns 3 = ! s per AA DR m: Y ian 9 Zi. ` kee dree Za y pad = E = — —— —N E tie deht ee Ju Eat? Lë 1 f e == PETET T H TA et m "eer E e^ Fi TE 5 * Rr MATE T Capa uU 32. EE sen E. UNE ime pasa vs $ = = D ECK €. Flower E ex G Don) Klotzsch. A. Habit. B. | f 1 showing pedicel. bracteoles. calyx. and corolla. D. | Li LÀ F F a I | E e LN | I P ey a T J La Fic. 3 Si f | ventral, E d wë EA bi al] £ WA LL A LI 7r5,2417.mD f - 15456). F Jn á HUO Loree Pad CI ur PFPE Fas Fa T F Luteyn and Ortiz, Revision of Siphonandra 255 glabrous, the thecae incurved and sometimes setose at an ca. 4-5 mm long, the tubules 13-18 mm long, less than 0.4 mm diam, style al g ll htly exserted, the stigma peltate. Berry cylindric- to elliptic-ovoid, glabrous to sparsely ile to 10-12 mm GC the calyx limb decurrent. Distribution (Fig. 2). —Common from Peru (Dpts. Amazonas, Apurímac, Cusco, Pasco, Puno) into Boli- via (Dpt. Cochabamba: Prov. Chapare and Dpt. La Paz: Provs. Bautista Saavedra, Franz Tamayo, Inquisivi, Larecaja, Murillo, Nor Yungas, Sud Yungas). Found growing in montane cloud forest, shrub vegetation on sandstone, wet puna, bosque siempreverde, bosque húmedo montano de las yungas, ceja de montaña, at elevations of 2200—4000 m. Flowering in Feb, Mar, Apr, May, Jun, Jul, Sep, Oct, Dec; fruiting in Jun, Dec. Common to abundant. Representative specimens examined: PERU. Amazonas: Chachapoyas, Weigend et al. 2000/876 (F, NY); Luya et al. 8843 (USM); near Molinopampa, van der Werff et al. 14894 (MO); Laguna de Pomacochas, van der Werff et al. 15822 (MO). Apurímac: Pinkos river, Weber- a 5861 ^u Cusco: MEG n Weberbauer 6929 (B+, F, US); Cerro de GC SE 13857 dium NY, e La EE NM Ae OTs Tu MO); Paucartambo, Cano 311 West 7047; Balls 6704 (BM. K, NY, US), 6 6705. Canó 3117 (USM), 3174 (USM), 3660 AO 4520 o Cm no & Aguilar 3157 (USM, F); Cano & Baldeón 4967 (USM); Huapalla 136 (USM), 211 (USM); ace & Lebrón- mci 6396 is MO, NY, I Plowman & Davis Si Kee Tupayachi 30 (MO); Challabamba, Cano 4339 rA Lluntuyoc, Woyth ba, Heller 2190 ond C. Vargas 3422, 4295 (both US); F. L. Herrera 3339 (US); Hoogte & Roersch 1397 (NY); Keel & voi 439 (AAU, NY); Funk et al. 3444 (NY, US); N. Salinas et al. 7620 (CUZ, Se Pedraza et al. 1541 (CUZ, NY, USM); Gentry et al. 23475 (MO, NY); Luteyn & Lebrón- Luteyn 6377, 6450. 6464 (all at NY, USM): B. León & al 2333 EN is. R. Foster & Wachter 7545 (CUZ, E NY, USM); Nuñez 7780 (NY, USM); Nuñez et al. 8520 (NY, USM). Juní ler, Boyle et al. 4133 (F, USM), 4138 (F, USM). Pasco: Oxapampa, E. Ortiz 225 (HOXA, MO, USM), 802 (HUSA, USM): D. N. Smith 2441 (MO), 2516 (MO), 8082 (USM, MO). Puno: San Gaván, Lechler 2276 (K); Tabina, Lechler 2053 (S. pilosa form) (K); Sandia, Soukup 223; Mathews 884 (BM, K). IVIA. Eastern Andes, Pearce ddl MS Seege Chapare, i anh e abd 1226 (LPB, BOLV); gege 876 ear Cárdenas 60 (BOLV, US); Bang 2003 ( 1, G, K, NY, US; R (US). La Paz: Mapiri, Rusby 2219 (NY), 2036 (B+, BM, F, G, K, NY, US); Yungas, Bushy 2034; Bautista A Cute 458 (LPB); Cocopunco, Tate 371 (NY); Franz Tamayo, Maldonado et al. 3210 (LPB); Inquisivi, Lara 236 (LPB LPB, MO, NY), 39358 Ea n a 39421 Pond ge 39719 (LPB, MO), 40940 (LPB, MO); Larecaja, Krukoff 11466 (F, NY); Clark 6642 (LPB); Beck ncoma, Tate 862 (NY); Luteyn & Dorr 13584 (AAU, BOLV, LPB, ND. Luteyn et i 15456 dd Goa OE cas CTES, | E, LPB, NY, ENE, TEX); Gutte & Herzog G458 (LPB); Pizarro E. 28 (LPB); Lara D. et al. 236 (LPB); Clark t al. 3210 (LPB) 9 (LPB); Solomon & Moraes 11424 (K); Nor Yungas, Beck 21472 (LPB); Lucis et al. 15477 (LPB, NY); Sol & Moraes 11424 (LPB. F, MO, NY); Sud Yungas, Luna-Pizarro 28 (LPB). Murillo: Beck 21551 (LPB); Luteyn & Dorr 13584 (LPB, BOLV, NY), Luteyn et al. 15431 (LPB, NY); Solomon 5230 (MO, NY). Unknown locality: Pearce s.n. (March 1866) (BM); C. Troll 1643, 2592 (both B, n.v.). Tablas, 3400 m, Herzog 2188 (G). Local name and uses.—willuntuy. Fruits edible, sweet. Siphonandra elliptica and S. pilosa were distinguished only by pubescence persistence of the pedicels and calyces, characters which we do not consider as stable. 3. Sin] ] fica Sleumer, Notizbl. Bot. Gart. Berlin-Dahlem 12(112):132-133. 1934. Bre. BOLIVIA. E PAz: Nor Vigas San Lorenzo-Tola, forest between Tola and San José, 3000 m, 28 Jun 1926 (fl), C. Troll 2593 [HoLorwrE: B, de- stroyed; LECTOTYPE, here designated: US frag. ex B holotype (barcode US00113579)]. Tall shrubs; mature stems thick; immature stems angled, glabrous. Leaves with blades thick-coriaceous, oblong-ovate, basally narrowed, apically rounded, na ec di EEN Dese and minutely denticulate or entire, the lamina glabrous; pinnately nerved (?), t d raised beneath, lateral nerves 4—5, arcuate-ascending, joined at margin, impressed dae and often raised beneath; petioles thick, 5 mm long. Inflorescences 3-6-flowered; rachis short, hardly 5 mm long, or almost none; floral bract unknown; pedicels thick, glabrous; bracteoles basal, ovate-deltate, acute, ca. 3 mm long. Flow- ers with calyx campanulate, ca. 9 mm long, glabrous, the tube densely bearing squamiform, carnose warts, the limb erect-spreading, the lobes broadly deltate, acute, ca. 1.5 mm long; corolla a eee 3.5-3.7 cm long, 7-89) mm diam., glabrous, the lol shorter than corolla, ca. 3 cm long, ds laments distinct, Së puberulent, ca. 6 mm lone the aaa ca. 2.6 cm long, the thecae subpapillose; style thick, glabrous, ca. 3.5 cm long. Berry not seen. Distribution (Fig. 2).—Endemic to Bolivia (La Paz) and known only from the type. Rare and endan- gered. ca 2mm long, reflexe [| [| fal n.a H In LI rr A £T "mia? 256 Siphonandra magnifica is herewith maintained despite the fact that the type and only specimen was destroyed during World War II. Sleumer (1934: 133) mentioned that it “surpasses both species as yet known [S. elliptica and S. pilosa] in all dimensions”, but differed from S. elliptica by the very short, often indistinct ra- chis, the very large corolla, the hairy filaments, and the calyx which was covered with fleshy squamules. NEW SPECIES 4. Siphonandra nervosa Luteyn & E.M. Ortiz, sp. nov. (Fig. 4). Te. PERU. Puno: Carabaya, Dito. Ayapata, near Campamento Chacayage, 13°45.8'S, 70°13.11'W, ca. 2600 m, 10 Mar 2004 (fl), S. Vilca C., K. Arce C., C. Dávalos M, & E. Ortiz V 62 (HOLOTYPE: HUSA; isotypes: HAO, NY) A EE elliptica Fu CNN odd oo we obov: SE mal joribus, 3.5-8.5 cm ddl (non 3—5.5 cm), apicibus foliorum I i 6-8 (non 3-5), inflorescentiis ILALVCVILVLI pauci-floris, 3— 4- floris (non 5- 15- floris), rhachidibus bres ioribus, usque 0. 7 cm longis (dont 1.5-5 cm), pedicellis brevioribus, usque mm longis (non 7-20 mm) differt. Lianoid, epiphytic shrubs with branches to 10 m long; mature stems terete to subterete, ribbed to bluntly angled, short-pilose with white hairs 0.2-0.3 mm long, the bark exfoliating in thin strips; immature stems subterete to bluntly complanate, striate, moderately short-pilose as mature stems; axillary bud scales nar- rowly ovate, long-acuminate, pseudostipular, ca. 3 mm long, short-pilose to glabrate. Leaves with blades coriaceous, lanceolate to elliptic-lanceolate, (3.5-)5.5-9.5 x (1-)1.5-3 cm, basally broadly cuneate, apically short-acuminate, the apex itself blunt, marginally entire, the lamina sparsely short-pilose and ciliate near base otherwise glabrate above, persistently moderately short-pilose along midrib beneath; pinnately nerved with 8-10 secondary veins per side anastomosing near margin, the midrib strongly impressed above and prominently raised beneath, the secondary nerves slightly impressed above and raised beneath causing leaves to appear slightly bullate, the reticulate veinlets obscure on both surfaces; petioles rugose, subterete, canaliculate above, 3-8 mm long, short-pilose. Inflorescences to 5-flowered; rachis sharply angled, ca. 3-5 mm long, sparsely short-white-pilose as stems, circumscribed at base by numerous, ovate, apiculate, ciliate bracts to ca. 2.5 mm long; floral bract ovate, acute, ca. 2.5 mm long, ciliate, marginally also with few glandular fimbriae; pedicels striate to sharply angled, 8-11 mm long, densely short-pilose in bud, glabrate; bracteoles located medially, opposite to subopposite, ovate, long-acuminate, ca. 2-3.5 mm long, marginally glandular-fimbriate, ciliate. Flowers with calyx 7-9 mm long, moderately to densely short-pilose, the tube cylindric, terete, 4. ae 6.5mm long, the pin campanulate, 2-4 mm long, the lobes deltate, short-apiculate, ca. 1.5-2 mm long; si ; corolla (when fresh), cylindric, conspicuously 5-angled (when fresh), 22-26 mm long, ca. ea 6 mm diam. glabrous, ied (when fresh), the lobes erect, deltate, acute, 1.7-2 mm long, yellow to greenish-yellow (when fresh); stamens nearly equaling corolla, 20-21 mm long, the filaments connate, 4—4.5 mm long, glabrous, the anthers 17-18 mm long, the thecae 4.8-5 mm long, strongly papillose, the tubules 12.5-13.5 mm long, the pores 0.2 mm diam.; style slightly exserted, glabrous. Berry not seen. Distribution (Fig. 2).—Endemic to southern Peru (Puno) and known only from the type, collected in heavily-disturbed, mature montane forest along a river margin, at ca. 2600 m elevation. Endangered due to its proximity to gold mining and road building activities. Siphonandra nervosa is characterized by its comparatively large, lanceolate leaf blades with prominent brochidodromous venation, few-flowered inflorescences, short rachis, pedicels and corolla, and connate staminal filaments. When fresh the corolla is 5-angled and red with yellowish lobes. Morphologically it is most similar to S. elliptica, differing primarily in the characters mentioned in the key. 5. Siphonandra santa-barbarense Luteyn & E.M. Ortiz, sp. nov. (Fig. 5). Ter. PERU. Pasco: Oxapampa, Dtto. Huancabamba, Sector Sta. Bárbara, Parque Nacional Yanachaga-Chemillén, source of the Quebrada Cueva Blanca, 10?21'23"S, 75939 20"W, 3420-3510 m, 16 Aug 2005 (fl), E. Ortiz V, S. Vilca C., C. Arias C., 5. Shuña S. & H. Cristóbal E. 807 (noore: HUSA; ISOTYPES: HOXA, MO, NY, USM). A Cal J sf 1 1:.3:1 1 : :1 T3 FR ] : f E ck: zial L corolla breviori, 18-26 mm longa (non 35-37 mm) differt. 257 Luteyn and Ortiz, Revision of Siphonandra d | hracteoles. and D r ada Ortiz A Li L'h vw d to corolla (C). E. St F 258 I I fal Das D In LI FPES f Texas 2(1 Sip honandra- santa -barbarense- Fic. 5. Sipt ira santa-bart Luteyn & E.M. Ortiz. A. Habit showing details of i f | one leaf. B-C. Leaf variation showing lower E Sal [| A... £ Lota L H i] H n EI L J | EN | L IL 4 [| calyx, 1: + ll E r il "1 A SA E ri L a je E eah la || Sal E H EO LÉI je | ALS E | F| GA L H LA l, dorsal, A A | H “al a “1 of obliquely terminal dehiscence pores. (A, D-G d from Ped tal. 1600: B from Ortiz et al. 807; Cfrom Luteyn et al. 15659) Luteyn and Ortiz, Revision of Siphonandra 259 +l 1 1 Terrestrial or ES shrubs to 4 m tall; mature | hes terete to subterete, glabrous, into thin, longitud trips, not exfoliatin tative bracts (at base of new es) membranous, "— lar, 3-5 x 2.5-6 mm, basally articulate, truncate, short-acuminate, fimbriate, glabrous; immature branches subterete to ribbed, glabrous to moderately puberulent, the nodes swollen; axillary bud scales 2, valvate, coriaceous, ovate, 1.5—5 x ca. 0.7-1.5 mm, apparently basally continuous thus persistent, long-acuminate, , Ciliate to glandular-fimbriate, glabrous. Leaves with blades rigidly coriaceous, ovate-oblong, ovate to Geesse 2- 4 x (1-)1.2-1.8 cm, aay rounded to short-attenuate, apically acute to subacute, sometimes shortly acuminate, al ply mucronate, marginally crenate, plane, glabrous above, glandular fimbriate beneath with hairs ca. 0.4mm iene obscurely 3(-5)-plinerved, the midrib impressed above and raised beneath, more prominent in the proximal 1/3 of the lamina, the lateral nerves slightly impressed above and obscure beneath, the reticulate veinlets slightly impressed above; petioles subterete, m flattened dorso- pini EE 3-5 mm long, 1-2 mm diam. ER to moderately puberulent. , 10-22-flowered; infl , ovate, 3 x 2—2.5 mm, basally articulate, "M sounded: entire, glabrous; rachis subterete to ribbed, 12-25 mm long, ca. 2 mm diam., puberulent; floral bract membranaceous, oblong, 5 x 1.5 mm, basally articulate, apically obtuse to rounded, entire, glabrous; pedicels terete, 10-17 mm long, ca. 1 mm diam., puberulent, also with few, scattered, glandular fimbriae distally; bracteoles caducous to persistent, located in proximal 1/3-1/2 of the pedicel, membranaceous, lanceolate, 4 x 1. 2 mm, ay enue apically acuminate, ciliate to marginally glandular-fimbriate, glabrous. Flowers | ; calyx 6-10 mm long, glabrescent, pink when fresh, sometimes obscurely articulate with Pë the iube ak, 5-ribbed (broadly when fresh), 3.5-6 mm long, ca. 4 mm diam., the limb campanulate, ca. 3-4 mm long, glabrous, the lobes deltate, apiculate to acuminate, 1.2-2.5 mm long; sinuses rounded to subacute; corolla tubular, pentagonal, slightly broadest in the middle, carnose especially along the angles, 18-26 mm long, 6-8.5 mm diam., glabrous, rose-red, the lobes erect to reflexed, deltate, acute to subacute, 2-3.5 mm long, glabrous, externally rose-red, internally whitish; stamens 10-20 mm long, the filaments distinct, 3-4 mm long, glabrous, the anthers ca. 17-18 mm long, the thecae tubular with the base shortly incurved and tapering into the tubules distally, 5-6 mm long, slightly papillose, the tubules distinct, 11-13 mm long, dehiscing by an oval, subterminal pore 0.5-1 mm long; nectariferous disc pulvinate, glabrous; style 20-24 mm long, the stigma punctiform. Berry immature, obconic, 5-ribbed, 10 mm long, 6 mm diam., glabrous, greenish-red. Distribution (Fig. 2). —Endemic to the upper limit of montane cloud forest, in elfin forest patches in sheltered areas, alternating with puna grasslands, at elevations of 3400-3600 m. Rare and probably endan- gered. Additional material examined: PERU. Pasco: O , Dtto. Huancabamba, Santa Bárbara, 10?20'35"S, 75939'0"W, 3400-3500 m, 25 Jan 2004 (bud, fl), R. Vásquez et al. 29084 (HOXA, HUT, MO, USM); 10?2124.5"S, 75?39'27.4"W, 3450 m, 9-13 Jun 2006 (fl), Luteyn et al. 15659 (AAU, COL, CUZ, F, HUSA, MO, NY, USM), Pedraza et al. 1600 (COL, CUZ, F, HUSA, K, MO, NY, USM Siphonandra santa-barbarense is characterized by its flat, marginally crenate, obscurely plinerved leaves with sharply mucronate tips, relatively long rachis, many-flowered inflorescences, articulate (sometimes inconspicuously) calyx, and staminal dehiscence by obliquely subterminal, elongate pores. The type was also found growing in the vicinity of S. elliptica. EXCLUDED TAXA Siphonandra mexicana Turcz. =Rondeletia mexicana (Turcz.) Standley [Rubiaceae]. ACKNOWLEDGMENTS JLL thanks NSF for funds for his field work and to support travel to NY for EMO. EMO thanks the Missouri Botanical Garden for funding for his fieldwork in Peru; also to Michael O. Dillon and the Botany Depart- ment, The Field Museum, for support from the Timothy C. Plowman Latin American Research Award. We also thank Wayt Thomas for supplying the GIS layers for the America's Basemap. 260 J tani | Texas 2( REFERENCES Hooker, W.J. 1837. Thibaudia elliptica. Icones Pl. 2: pl. 108. KLOTZSCH, LE 1851. Studien über die natürliche Klasse Bicornes Linné. Linnaea 24:1-88. Kron, KA. W.S. Jupp, PF. Stevens, D.M. Crayn, A.A. ANDERBERG, PA. GADEK, C.J. Quinn, J.L. Luteyn, and R. Futter, 2002a. Phylogenetic classification of Ericaceae: Molecular and morphological evidence. Bot. Rev. 68:335—424. Kron, K.A., E.A. Powe, and J.L. Lutern. 2002b. Phylogenetic relationships within the blueberry tribe (Vaccinieae, Ericaceae) based on sequence data from matK and nuclear ribosomal ITS regions, with comments on the placement of Satyria. Amer. J. Bot. 89:327-336. Luteyn, J.L. 1998. Neotropical blueberries: The plant family Ericaceae. http://nybg.org/bsci/res/lut2. [updated as Luteyn & Pedraza, 2006]. Luteyn, J.L. 2002a. Key to the species of Ericaceae of Bolivia, including two new species. Sida 20:1-20. Luteyn, J.L. 2002b. Diversity, adaptation, and endemism in neotropical Ericaceae: Biogeographical patterns in the Vaccinieae. In: K. Young, C. Ulloa Ulloa, J.L. Luteyn, and S. Knapp, eds. Plant evolution and endemism in Andean South America. Bot. Rev. 68:55-87. Ruiz, H. and J. Pavón. 1802. Flora peruvianae, et chilensis prodromus. Vol. 3. Madrid. SLEUMER, H.O. 1934. Ericaceae americanae novae vel minus cognitae. Notizbl. Bot. Gart. Berlin-Dahlem 12(112):119-140. SiEUMER, H.O. 1941, Vaccinioideen-Studien. Bot. Jahrb. Syst. 71:375—408. SMirH, A.C. 1932. The American species of Thibaudieae. Contr. U.S. Natl. Herb. 28(2):311-547. NUMERICAL LIST OF TAXA 1. Siphonandra boliviana Luteyn 4. Siphonandra nervosa Luteyn & E.M. Ortiz 2. Siphonandra elliptica (Ruiz & Pav. ex G. Don) Sleumer 5. Sip! 1 ta-bart Luteyn & E.M. Ortiz 3. Siphonandra magnifica Sleumer List OF EXSICCATAE (*RECOGNIZED TYPE COLLECTION NUMBERS IN BOLDFACE) Huamantupa, |. & Huamantupa, 4440 (2) (2) Aguilar, B. & Muriel, 1226 (2) Atahuachi, M., 876 (2) Huapalla, N. et al., 136, 211 Balls, E. K., 6704, 6705 (2) Keel, S. & Oqueso, 439 (2) Bang, M., 2003 (2) Krukoff, B.A., 11466 (2) Beck, S., 21472, 21551 (2) Lara, D., 236 (2) Boyle, B. et al., 4133, 4138 (2) Lechler, W., 2276, 2053 (2) Cano, A. 3117, 3174, 3494, 3660, 4330, 4339, 4520 (2) León, B. & Aguilar, 2333 (2) Cano, A. € Aguilar, 5157 (2) Lewis, M,, 38496, 39358, 39421, 39719, 40940 (2) Cano, A. € Baldeón, 4967 (2) Luna-Pizarro, E., 28 (2) Cárdenas, M., 60 (2) Luteyn, J.L. & M. Lebrón-Luteyn, 6377, 6396, 6450, 6464 (2) Clark, J.L., 6642 (2) Luteyn, J.L. et al, 13584, 15431, 15456, 15477 (2); 15659 (5) Croat, T., 78186 (2) Maldonado, C. et al., 3210 (2) Díaz, C. & Peña 8843 (2) Mandon, G., 549 (2) Dudley, T., 11124 (2) Mdb. 884 (2) Ellenberg, H., 4852 (2) ssinger, W., et al. 407 (2) Foster, R. & Wachter, 7545 (2) Metcalf, R.D., 30749, (2) Funk, V. et al., 3444 (2) Nuñez V, P, 7780 (2) Gay, Cl. sn, 709 (2) Nuñez V, P. et al., 8520, 8522 (2) Gentry, A. et al, 23475 (2) Gutte, P, 458 (2 Heller, 2190 (2) Herzog, B., H200 (1) Herzog, Th., 2188 (2) Herrera, F. L., 3339 (2) Hoogte & Roersch, 1397 Q) rr Ortiz, EM. et al 225, 802 (2), 807 (5) Pearce, R.W., 790 (2) Pedraza, P. et al., 1541 (2); 1600 (5) Pennell, EW 13857 (2) Peyton, B. & S. T. Peyton, 318, 771 (2) Ruiz L., H. & J.A. Pavón, s.n. (2) Ri Isby, H.H., 2034, 2036, 2219 (2) Luteyn and Ortiz, Revision of Siphonandra Salinas, N. et al., 7620 (2) Smith, D.N,, 2441, 2516, 8082 (2) Solomon, e 5230, 11424 (2) Solomon, J.C. € M. Moraes, 11424 (2) Soukup, E 223, 383 (2) ) Troll, C., 1643, 2592 (2), 2593 (3) Tupayachi, A., 30 (2) Van der Werff, H. e al., 14894, 15822 (2) Vargas, C., 3422, 4295 (2) Vásquez, R. et al, i (5) Vilca, S. et al., 62 (4) Weberbauer, A., 740, 5861, 6929 (2) Weddell, H. Alg., 4711 (2) Weigend, M. et al., 2000/876 (2) Zimmerer, KS. 114 (2) 261 262 BOOK REVIEW RICHARD K. RABELER. 2007. Gleason's Plants of Michigan. (ISBN 978-0-472-03246-4, pbk.). University of Michigan Press, 839 G e Street, Ann Arbor, Michigan 48104, U.S.A. (Orders: www.pres.umich.edu, robert. pierceOperseusbooks.com, University of Michigan Press, c/o Perseus Distribution, 1094 Flex Drive Jackson, Tennessee 38301, U.S.A., 1-800-343-4499 Ext. 165, 1-877-364-7062 fax). $24.95, 400 pp., b/w drawings, terminology, glossary, indices to plant names and family names, 5" x 7" Contents: Editors Note Preface: Henry A. Gleason: his Michigan period Acknowledgments ee How to use this book Bibliogr Term sii ogy used in plant descriptions List B illustrations Key to Groups Group 1: Woody plants Group 2: Unusual plants Group 3: Monocots Group 4: Dicots Key to the Plants of Michigan (in family order) Monocots Dicots Glossary Subject index Index to plant names ichar abeler has revised the ear s Gleason's Plants o i ised text i llent field guide and designed t Richard K. Rabeler h i arly 1900s Glea e t adi an. g be used as such, f y of the user to view p by sight or with a hand lens, and Į iding Il rul in ees e use. pM isal Ir bp sq ] Ms 1 gy e > 1 1 E fl 1 1 A gl y +l 1 1 F +41 text. T | j fy fi Ê ts, monocots, dicots, and l plants-such as aquatic or ssrasilic plants), then to rd and eventually to > species . Rabeler PHP ECH a ind — famil ly index for when tl has p A H 1 ] T +1 e knowledge Dus np An E I thbl g y of the E guide, as well as 1 | eee x . zt MO ANS: 4 x pu cquo cod "E lel a D ad 1 er +: mall j j g I Overall, Ral 1 3 H 147] sn 4 CRAG. H 3 Es E: 1.1 r 1 H 1 are kien in Sege een puse has many ao aues for the amateur—illustrated terminology eS a E s Je L a 11 1 F+l corrected ME updated from the 1918 ver j T" flora of the region, Edward G. Voss's Michigan Flora B volumes, 1837 pp.!).—Lee SEH Botanical sea Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S.A. J. Bot. Res. Inst. Texas 2(1): 262. 2008 FOLIA TAXONOMICA 4. CONSPECTUS OF MYRIOPUS (HELIOTROPIACEAE: BORAGINALES) IN THE GUIANA SHIELD Christian Feuillet Department of Botany, MRC-166 Smithsonian Institution PO Box 37012 Washington, DC 20013-7012, U.S.A. feuillecsi.edu ABSTRACT "r1 P TS og al f: f 1 " JAM H ad ` H 1 o LE cL:.131f1 = = I y I The synonymy and four new combinations are included. RESUME Le conspectus donne les E taxa du g 5 Myriopus Small (Heliotropi ) présents dans la flore du Bouclier . uo 1: Guy anais.I In the past few years, a better understanding of the relationship between taxa of the Boraginales was the focus of several papers through the re-examination of old and the gathering of new morphological data and the analysis of molecular data (Diane et al. 2002, 2003; Hilger & Diane 2003; Gottschling et al. 2005; Miller & Gottschling 2007). Familial and generic limits have been re-considered. In the preparation of the treatment of the Boraginales for the Flora of the Guianas, I had to decide whether 1 should follow the tra- ditional taxonomy as used for the Boraginaceae treatments in *Flora of Venezuelan Guayana" (Miller et al. 1997) and in "Checklist of the Plants of the Guiana Shield" (Feuillet et al. 2007) or to adapt it to reflect the changes proposed since 1997 in the systematic of this group. As it is often the case with work in progress, some proposals should be adopted, like the era e Varronia P.Br. from Cordia L. (Cordiaceae) and of Myriopus Small from Tournefortia L. (Heliotropiaceae), when otl 1 further work including more species in the data sets, like the split of Heliotropium L. and its (eon with Tournefortia s.s. Currently less than 10% of the species have been analyzed in the whole Heliotropiaceae and it is likely that some major clades of Heliotropium s.l. and Tournefortia s.s. are missing from the data sets. The part of the new classification of the Heliotropiaceae proposed by Diane and her colleagues that is most likely to resist larger sampling is the separation of Myriopus from Tournefortia and its standing as a sister group to Heliotropium s.l. and Tournefortia s.s. It was pointed out by Johnston (1930) that his new Tournefortia sect. Cyphocyema 1.M. Johnst. was distinct from the core of Tournefortia. He suggested on a morphological basis that the two sections of Tournefortia were closer to other genera than to each other. Small (1933) included two species of sect. Cyphocyema in his new genus Myriopus and said there was more Neotropical species. The morphological characters that separate Myriopus apart from Tournefortia s.s. are its elongated corolla lobes, narrow with involute margin, its anthers connate and always hairy at apex, its drupoid fruits distinctly lobed and not dividing into mericarpids, its four 1-seeded endocarpids that are strongly curved, and its curved embryos (Johnston 1930; Diane et al. 2002). The molecular and morpho- logical data sets used by Diane et al. (2002, 2003), Hilger and Diane (2003) confirmed the deep separation between Tournefortia sect. Cyphocyema and most of the Heliotropiaceae. The Cyphocyema/Myriopus branch is EE Së Beete Ee of 100% in all trees. Logically Diane et al. (2002) proposed to use of Tournefortia sect. Cyphocyema. The conspectus presented boues is Tires the Si? of Myriopus present in the Guianas (Guyana, Surinam & French Guiana) and the Venezuelan Guayana (Amazonas, Bolivar & Delta Amacuro), the most part of a region often referred to as J. Bot. Res. Inst. Texas 2(1): 263 — 265. 2008 264 | | .fakn Dad ID Llanta cr the Guiana Shield or Guayana Shield. Four taxa from the Guianas did not have a name in Myriopus and needed a new combination. KEY TO THE GENERA OF HELIOTROPIACEAE IN THE GUIANAS . Herbs. Fruits usually dry when fresh Heliotropium à gie e llanas. a 150 when das t y y | , consisting of 4 similar 7 -seeded nutlets. Emiro: curved. Myriopus 2. Corolla lobes broad and rounded. Fruits obscurely if at all lobed, evidently bicarpellate, breaking up into 1- or 2-seeded irregular nutlets. Embryos straight Tournefortia Myriopus candidulus (Miers) Feuillet, comb. nov. Basionym: Messerschmidia candidula Miers, Ann. Mag. Nat. Hist. ser. 4, 2:202. 1868. Tournefortia candidula (Miers) I.M. Johnst., Contr. Gray Herb. 92:84. 1930. Tree: BRAZIL: Gardner1078 (HOLOTYPE: BM; IsoTYPES: K, NY) = Tournefortia lanuginosa Vaupel, Notizbl. 6:183. 1914. Tree: BRAZIL: Ule 9097 (HoLoTYrE: BD (n.v.); 1soTyPE: K) Myriopus maculatus (Jacq.) Feuillet, comb. nov. Baam: Tourneforti lat Enum. Syst. Pl. 14. 1760; Select. Stirp Amer. Hist. 47. 1763. Tre: COLOMBIA: Cartagena, “Habitat Garer in arbustis Recent? (n.v.). = Tournefortia syringaefolia Vahl, Symb. Bot. 3:23. 1794. Tre: FRENCH GUIANA: von Rohr s.n. (HOLOTYPE: C X : BM) = Tournefortia peruviana Poiret, Encycl. Suppl. 4:425. 1816. T. volubilis sensu Ruiz & Pav. (non L. 1753), Flor. Per. 2:24, tab. 148. 1799. T. scandens Willd. (non Mill. 1768), Enum. Pl. 1:188. 1809. Tree: PERU. Pasco. Pozuzo, Ruiz & Pavón s.n. (HOLOTYP n.v.); IsOTYPE: US). The US emu in the type collection, under T. scandens Willd., | label with handwritten MP. volubilis Fl. Peruv. 2 Zap. 48 ..." and * ex herbario Fl. Peruv. anno 1828,” and off that label, Killips handwriting, "Type of T. volubilis R. & P = T. peruviana Poir." ind “Part of type ex herb. Madrid Ruiz & Pavon.” - ad i diii DC., Prodr. 9:526. 1845. Tree: SURINAM. Hostmann 951 (HoLoTYPE: G-BOISS). x Schomb., Fauna Fl. Brit. Guiana 1151. 1848, nom. nud., based on Hostmann 285 (BD, BM, K, P). Myriopus paniculatus (Chamisso) Feuillet, comb. nov. BASIONYM: Tournefortia paniculata Chamisso, Linnaea 4:468. 1829. Tyre: Sellow (B, extant?). Myriopus paniculatus var. spigeliiflorus (A. DC.) Feuillet, comb. nov. Basionw: T. spigeliaeflora A. DC., Prodr. 9:525. 1845. Tournefortia paniculata Chamisso var. spigeliiflora (A. DC.) I.M. Johnst., J. Arnold. Arbor. 16:49. 1935. Tree: GUYANA: Rob. Schomburgk ser. 2, 427 (HoLotyre: G-BOISS; IsoTYPES: BM, K, P). Myriopus volubilis (L.) SR Man. = E. de E 1933. Tournefortia lubilis L., Sp. PL 140. 1753. LecrorrrE: “Bryonia nigra fruticosa, racemi ramulis i tortis, baccis albis una vel altera nigra macula nd in uude ind mai y t. 143, Í. 2. 1707. = 4 gk, Fauna u. Fl. Brit. Guian. 1084. 1848. non Kunth 1818. KEY TO THE GUIANAN SPECIES — a . Leaves densely white-tomentose on lower surface, grayish green pubescent on upper surface; inflores- cence terminal with short lateral branches, appearing as a very contracted panicle; fruits white-pubescent. uyana) M. candidulus 1. Leaves glabrous or pubescent, never white-tomentose; inflorescence terminal or d paniculate with long latera! branches or a glomerulate with short branches; fruits never white-pubescen 2. Corolla tube 1.5-2.5 mm long, constricted at throat, lobes linear, nearly as long as e tube; fruit white. Sec Surinam, French Guiana) M. volubilis 2. Corolla tube 3-8 mm long, not constricted at throat, lobes broadened below middle, acuminate, half as i. as the tube or Hes men yellowish, pedicel thickemng in fruit. to linear. (Guyana, Surinam, Ten Guiana) M. maculatus ] 11 | J +l L ` I] J L 3. Leaves, y long acuminate. (Guyana, Surinam, French Giana). M. paniculatus v var. spigeliiflorus Feuillet, r p A fa H " al o. eL: 265 ACKNOWLEDGMENTS I would like to thank the reviewers for their suggestions regarding the introduction and the curators of the herbaria CAY, F, NY, P and U for lending the material in their care. This is number 134 in the Smithsonian's Biological Diversity of the Guiana Shield Program publication series. REFERENCES Diane, N., H. FÓRTHER and H.H. HiLcER. 2002. A systematic analysis of Heliotropium, Tournefortia, and allied taxa of the Heliotropiaceae (Boraginales based on ITS1 sequences and morphological data. Amer. J. Bot. 89:287-295. Diane, N., C. Jacos, and H.H. Hier. 2003. Leaf anatomy and foliar trichomes in Heliotropiaceae and their systematic relevance. Flora 198:468-485. FEUILLET, C., J. GAVIRIA, R. Gómez, J.S, Miter, and G. Ropricuez. 2007. ROMS SE 226. In: jus V, T. Hollowell, P. SE Ch; AV) Amazonas, Bolivar, Berry, C. Kelloff, and S.N. Alexander. Checklist of the plants oft Delta Amacuro; Guyana, Surinam, French Guiana). Contr. Us. Natl Herb. 55:1 E GortscHLING, M., J.S. Murs M. WeicenD, and HH Heen. 2005. Congruence of a phylogeny of Cordiaceae (Boragi- nales) inferred from ITS1 sequence data with morphology, ecology, and biogeography. Ann. Missouri Bot. Gard. 92:425—437. HiLGER, H.H. and N. Diane. 2003. A systematic analysis of Heliotropiaceae (Boraginales) based on trnL and ITS1 sequence data. Bot. Jahrb. Syst. 125:19-41 JOHNSTON, LM. 1930. Studies in the Boraginaceae. - VIII. 3. Treatment of Tournefortia. Contr. Gray Herb. 92:66-89. MILLER, J.S., J. Gaviria, R. Gómez, and G. Ropricuez. 1997. Boraginaceae. In: PE. Berry, B.K. Holst, and K. Yatskievych, eds. Flora of the Venezuelan Guayana 3:527-547. Miler, J.S. and M. GOTTSCHLING. 2007. Generic classification in the Cordiaceae (Boraginales): resurrection of the genus Varronia P. Br. Taxon 56:163-169 SMALL, J.K. 1933. Myriopus. In: Manual of the southeastern flora. Hafner, New York. P. 1131. BOOK REVIEWS BETH Jurx 2007. Medicinal Plants of North America: A Flora Delaterre™ Coloring Book. (ISBN 978-0- 9792302-0-2, pbk.). Flora Delaterre™ Productions, PO. Box 8474, Missoula, Montana 59807, U.S.A. (Orders: www.floradelaterre.com, 1-406-728-2977). $12.00, 32 a 33 b/w line drawings, 8.5" x 11". Medicinal Plants of North America: A Flora Delaterre™ Coloring Book is a fun and educati book for any young explorer. The book presents a black and white line pene for oe and pL text E "i MORAN plants Tue author RECH includes a disclaimer that this book is for “knowledge on brief explanation of medicinal value, and fun facts al the plant. Additi i ] bout medici included on the last page. A great activity book for a budding botanist —Lee Luckeydoo, Bovanizal Research Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S.A. Max SCHNEPF AND nate Se (eds.). 2007. Managing Agricultural Land for Envi tal Quality St d i Base. (ISBN 978-0- 9769432-4- 2, pbk.). Se and Water Conservation Societ; 945 SW Ankeny Road, Anken, Iowa, U.S.A. (Orders: www.swcs.org, http://store.swcs.org, 1-515-289- 2331, 1-515-289-1227 fax). $32.00, 196 pp., b/w line drawings, charts, graphs, index, 7" x 10". In October of 2006, the Soil and Water C iety held ing in Kansas City called "Manage Ree oes +l = 1 t a +l x Tas I H P 7 workshop for Environmental Quality". Afterward, name A 1 3 E 1 1 Js 1 1 I ] hat hould il t f i tal effects. Also discussed r 1 1 1 ] ] 1 ] 1 1 1 J 1 ] + e L e e i rr 1 1 1 1 t 3 1 af f, t Ei i D bel L > j À A ix a A À Le S 1 : dos das a f : th A rc th to 1 jd a denlhl + E E e discussions held at the workshop. This book presens em e with an nis and current view of t h d perspectives, including case studies from various peop I i nl regarding c conservation on ind ing scales. SE us "m «1 f ÉS 1 A 1 EN A 1 ] tal valuation, scale, ] h ,and policies can Ir e i mal H — — | quali t1 ali interact. The boo in agricultural [nde —Lee p» Botanical Research Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U S. A. J. Bot. Res. Inst. Texas 2(1): 266. 2008 FOLIA TAXONOMICA 5. A NEW NAME FOR PASSIFLORA HETEROPHYLLA (PASSIFLORACEAE) FROM CUBA AND HAITI Christian Feuillet John M. MacDougal Department of Botany, MRC-166 Division of Arts and Sciences Smithsonian Institution Harris-Stowe State University 7012 3026 Laclede Avenue Washington, D.C. 20013-7012, USA, St. Louis, Missouri 63103, USA feuillec@si.edu threebrane@sigmaxi.net ABSTRACT The name es e ae Lam. does not have priority P ifl het E hy I] Dryand and the taxon does not h tl name available. The new proposed for that species. RESUME disponible Le Le nom -= la Lam. n'a pas puo sur nm ida heterophylla Dryand. et le taxon n'a pas d'autre n nouveau rı E T r r Both Dryander (1789) and Lamarck (1789) published a taxon named Passiflora heterophylla. According to TL-2 (Stafleu & Cowan 1976), Dryander’s name was published between Aug 7 and Oct 1, while Lamarck’s name was from Oct 19 (Stafleu & Cowan 1979), thus giving Dryander’s name priority. Killip (1938) ac- cepted Lamarck's species and name but cited Dryander’s as a synonym under P. suberosa L. as “Passiflora heterophylla Dryand. in Ait. Hort. Kew. 3:309. 1789, not P. heterophylla Lam.” Dryander's species is now con- sidered a synonym of P. pallida L. in subgen. Decaloba supersect. Cieca (Medic.) J.M. MacDougal & Feuillet (Porter-Utley 2003) and P. heterophylla Lam. is a good species without a valid name and without synonym. Passiflora heterophylla “Jacq., Hort. Schoenbr. t. 181. 1797” is Dryander's species because Jacquin credited “Ait. kew. vol. 3. pag. 309.” There is no name available for the taxon currently called P. heterophylla Lam., so we propose the following new name: — insueta Feuillet @ MacDougal, nom. nov. lla Lam., Encycl. 3:41. 19 Oct 1789, non . 7 Aug-1 Oct 1789. (including the Latin diagnosis). Tyee: HAITI: ases à Pitres, omn Icaque, Plumier? s.n. (HOLOTYPE: Pt). Passifl ta belongs in sub Decalol t. Multiflora (Small) J.M. MacDougal & Feuillet (Feuil- let & MacDougal 2003). The leaf blades of this poorly known species are lobed nearly to the base and the lobes themselves are unlobed or lobed in their apical half. Etymology.—The epithet comes from the Latin adjective “insuetus” meaning “unusual” in reference to the leaf shapes. Passiflora insueta is known from Haiti and western Cuba. ACKNOWLEDGMENT Wi ti director d curator wl lO sent E hot ] d all El us to work on the collec- ¥¥L alc Yel y gratel ful t Lu tions in their care at P and MO. The review comments of Kristen E. Porter-Utley are greatly appreciated. REFERENCES DRYANDER, J.C. 1789. In: Aiton W., Hort. Kew. 3:1-547, FEUILLET, C. and J.M. MacDoucaL. 2003 [2004]. A new infrageneric classification of Passiflora L. (Passifloraceae). Passiflora 13(2):34-38. um, E.P. 1938. The American species of Passifloraceae. Publ. Field Mus. Nat. Hist, Bot. Ser. 19:1-613. J. Bot. Res. Inst. Texas 2(1): 267 — 268. 2008 268 Jo tl tanical hi xas 2( LAMARCK, J.B. DE. 1789. Encyclopédie méthodique : botanique 3(1):1-360, Porter-Uriey, K.E. 2003. Revision of Passiflora subgenus Decaloba supersection Cieca (Passifloraceae). Ph.D. dis- sertation, University of Florida, USA. SrAFLEU, FA. and R.S. Cowan. 1976. Taxonomic literature, vol. |: A-G: 1-1136. Grapp, FA. and R.S. Cowan. 1979. Taxonomic literature, vol. Il: H-Le: 1-991. FOLIA TAXONOMICA 6. TWO NEW SPECIES OF BESLERIA (GESNERIACEAE) FROM THE VENEZUELAN GUAYANA Christian Feuillet Department of Botany, MRC-166 ecd Institution, PO. Box 37012 We n, DC 20013-7012, USA. p illec@si.edu ABSTRACT Two species of Besleria (Gesneriaceae) are described from the state of Amazonas in southeastern Venezuela. ida neblinae s B. yatuana are related to B. gibbosa in section Neol il f which tł ley differ mainly by t] gb unequal in a pair RESUME D de e le Besleria (G | ) lécri le l'état d'A ,au sud-est du Venezuela. Besleri bli tB yatuana sont poches de B. gibbosa tion Neobesleri bsection Axillares. dont elles different t ] famil] ——— $ = É A I o I RESUMEN Se describen d ies de Besleria (G l ) del estado de A l sudeste d ee Besleria EEN B. yatu- lan ] B ber aw masses E E EE 1 D Axillares, Asia WE Es 1 iguales en un par. This paper is the first of a series describing new species of Gesneriaceae from the Venezuelan Guayana. This region of Venezuela includes the states of Amazonas, Bolívar, and Delta Amacuro, totaling 454,000 km? that represents 50% of the area of Venezuela. The two new species of the subfamily Gesnerioideae Link described herein belong to Besleria L., tribe Beslerieae Bartl. & H.L. Wendl., and were collected in the state of Amazonas. Besleria neblinae Feuillet, Spo nov. v. (Pig. D Tre: VENEZUELA. Amazonas. Dept. Rio Negro: near Cerro de la Neblina, swamp between Río M Baría, ca. 0°52'N, 66°15'W, 140 m, 8 May 1984, A. Gentry & B. Stein 47253 (Ho- LOTYPE: US; isorvee; MO). j E 1 a | = 1 "E 1 vL re A a «= -l Besleriae gibbosae affinis; lamina angustata, valde inaequali, y g , ang E Į Suffrutescent herb, 0.3-2 m tall, often with spreading, sub-horizontal branches. Stem slender, densely vil- lous. Leaves of a pair mostly unequal: smaller leaves with petioles about 1/5 the length of those of the larger leaves, blades about 1/3 the size of the larger; mature larger leaves with petioles 1.5—4.5 cm, densely villous; blades 9-19 x 3—5.5 cm, lanceolate to elliptic, often asymmetric, base acute to obtuse, occasionally nearly rounded, apex acute to acuminate, margin serrulate and ciliate, both surfaces with appressed trichomes, indumentum loose between veins, dense and spreading on midrib and main veins, 8-12 main veins on each side. Inflorescences sessile, 1—2-flowered; pedicels 3-5 mm long, densely villous. Flowers with calyx hirsute, tube 1.5 mm long, lobes lanceolate, 5.5—6 x 1.2-1.5 mm, attenuate at apex, slightly longer in fruit; corolla white, horizontal in tlie calyx, glandular puberulent outside, prominently spurred, spur 5—5.5 x 2-3 mm, oblong, rounded, prolonging the tube at base, tube 25-35 mm long, 3 mm wide at base and 6 mm at throat, lobes yellow (fide Davidse & Miller 26864), suborbicular, 4 x 4 mm; 4 stamens about 25 mm long, anthers with thecae reniform encircling a round connective; 1 staminode 3 mm long, with a sterile linear anther; J. Bot. Res. Inst. Texas 2(1): 269 — 273. 2008 270 gt e da fTexas 2(1) £539 r> il || Feuillet, New species of Besleria 271 dorsal glands fused into one, semi-annular, tongue-shaped, emarginate; ovary superior, ovoid, glabrous, 3 mm long, style 20 mm long, Enna Young fruits green, narrowly ovate, Jong Ge 9 x3 mm. leri wet forests and on ] tsin the southern Distribution. part of the Municipio Río Negro, Amazonas (Venezuela) near the base of Cerro d la Neblina, at 100-140 m elevation. The Neblina massif is on the border with Brazil and it is possible that populations of this new species are present in nearby Brazil. Phenology.—Blooming probably not seasonal, documented in February-March, May, July, November. Besleria neblinae corollas have a spur at the base that prolongs the tube at a 0-20” angle (cf. Fig. 1 inset) and displaces the attachment to the receptacle in a lateral position. According to Morton (1939), this char- acter places B. neblinae in sect. dcl CV. E Geer, its obsolete inflorescence peduncle and the size of the corolla limb indicate the sp long t. Axillares CV. Morton, one of the two subsections of sect. Neobesleria that remain alter hve were transferred to the reinstated genus Gasteranthus Benth. In the same subsection and from Amazonian Brazil and Venezuelan Guayana, B. gibbosa (Poepp.) Hanst. also has white corollas, but B. neblinae differs by: 1) the leaf blades which are clearly unequal in a pair (versus equal or subequal) and narrower, 2) the calyx lobes that are longer, narrower, and hirsute (versus glabrous), and 3) the dorsal glands that are smaller. In the specimen Liesner 17300, most pairs of leaves are subequal but the smaller petioles are 1/2 as long as the larger ones which is distinctive from B. gibbosa. The collection Thomas 3396 was made in the same locality and the same day as the type collection, Gentry & Stein 47253. Although a different collection (W. Thomas, pers. com.), it is possible that both represent the same population. This species was Besleria “sp. B" in Feuillet and Steyermark (1999). Etymology.—Besleria neblinae is named for Cerro de la Neblina near the base of which all the known collections have been made. PARATYPES: VENEZUELA. Amazonas. Dept. Río Negro: at base of Cerro de la Neblina, swamp forest between Río Marawinuma and Río Baría, 0º50'N, 66°09'W, 8 May 1984, W.W. Thomas 3396 (NY, US); Upper Caño Baría, ca. 0°52'N, 66°15'W, 130 m, 26 Mar 1984, R. L. Liesner 16964 (MO, US, VEN); 1 km E of Cerro de la Neblina b which is on Río Marawinuma, 0°50'N, 66°10'W, 140 m, 25 Feb 1984, R. L. Liesner 16254 (MO, US, VEN), 25-26 Nov 1984, R.L. ester 17300 (MO, US), 30 Nov 1984, R. L. Liesner 17382 (MO, SL Río B , ca. 0°52'N, 66°15'W, 100 m, 2-3 Jul 1984, G. Davidse & J. S. Miller 26864 (MO, US, VEN). Besleria yatuana Feuillet, sp. nov. (Fig. 2). Tre: VENEZUELA. AMAZONAS. DepER Río EE near ncs de b: Neblina, up- permost Río Yatúa, 100-140 m, 7-8 Dec 1953, B. Maguire, J.J. Wurdack & G.S j NY, US sheets 2195722 & 2614591). Besleriae neblinae affinis; lamina longe acuminata, lobis calycis liberis, longioribus, latioribus et minute appressis-pubescentibus dif fert. Suffrutescent herb, 1 m tall, spreading branches. Stem hat succulent, appressed-pubescent. Leaves of a pair mostly unequal; smaller leaves with petioles up to 1/4 the length of those of the larger leaves, blades less than 1/2 the length of the larger; mature larger leaf with petioles E cm, a pubescent blades 11-20 x 4—6 cm, lanceolate to PEO slightly asymmetric, base acute, ay lat adaxially glabrous, abaxially gl t for the midrib and main veins sparsely doped. alas 9-10 secondary veins on each side. EN sessile, 1-2-flowered; pedicels 3-6 mm long, appressed- pubescent. Flowers with calyx minutely appressed-pubescent, lobes free to base, lanceolate-linear, attenuate at apex, 6-10 x 2.5 mm; corolla horizontal in the calyx, white, sparsely to moderately pilose outside, trichomes with spherical glandular tip, prominently spurred, spur 6 x 4 mm, ovoid-oblong, rounded, prolonging the tube at base, tube 25 mm long, 4 mm wide at base and 2 mm at throat, lobes suborbicular, 5—6 x 1.5 mm; ^ stamens about 2.5 cm long, anthers orbicular-reniform; staminode not seen; dorsal glands fused into one; ovary superior, ovoid, glabrous, 3 mm long. Fruits not seen. Distribution.—Ihe type collection of Besleria yatuana in bloom was made in December in the flooded forests along the upper-most Río Yatúa in the southern part of the Municipio Río Negro, Amazonas (Ven- ezuela). Besleria yatuana shares with B. neblinae the characters that places both of them in subsect. Axillares. 272 ff 240 e rd me à Sivas um 3i É RER UNITED STATES NATICNAL HERBARIUM Eme E Y Besleria fje c. [i e. | T n sa? KE ee T m. ¿E A. \ ' E he as T AF ir Maauire E Ki Li Feuillet, New species of Besleria 273 It has white corollas like B. gibbosa and B. neblinae, but differs from the first one in having leaves that are strongly unequal in a pair (versus equal or subequal) and long acuminate (versus acute to acuminate), its longer and appressed-pubescent (versus glabrous) calyx lobes, and its smaller dorsal gland Besleria yatuana differs from B. neblinae in having the leaf blades adaxially glabrous (versus appressed pubescent), the calyx minutely appressed pubescent (versus hirsute), calyx lobes free to the base, and the slightly shorter corolla tube. Besleria yatuana was Besleria “sp. C” in Feuillet and Steyermark (1999). Etymology.—Besleria yatuana is named for the Río Yatúa near which the type specimens have been col- lected. KEY TO BESLERIA SUBSECTION AXILLARES IN THE GUIANA SHIELD 1. Corolla not spurred, but ti little saccate at base sect. Besleria 1. A spur at the base that | | g +l II dl J Pu pl Al ++ | ++ +l + ] | 1 position. sect. Neobesleria 2. Inflorescence peduncle, mues — the pedicel in length subsect. Pendulae 2. Inflorescence peduncle obso subsect. Axillares Leaves in a pair SSES or subequal calyx lobes glabrous B. gibbosa 3. Leaves in a pair, at | tiole, strongly unequal i in a pair; calyx lobes hai 4, Stems, petioles pedicels, and ly pl | pubescent; leaf blades EE SE for the main bes free to base, 6-10 x 2.5 mm, appressed-pubesce B. yatuana 4. M petioles, pedicels, and calyx densely villous with lax soreading m up to 2 mm long; leaf blades with appressed tnichomies: Eet tube 1.5 mm long, lobes 5.5-6 x 1.2-1.5 mm, hirsute B. neblinae ACKNOWLEDGMENTS I am grateful for the suggestions of Larry Skog, Jim Smith and an anonymous reviewer. I want to thank the curators of the herbaria MO and NY for lending the material in their care, and especially Jim Solomon (MO) and Wayt Thomas (NY) for the information about the collections cited. This paper is published as No. 135 in the Smithsonian's Biological Diversity of the Guiana Shield Program publication series. REFERENCES FeuiLLer, C. and J.A. Stevermark. 1999, Gesneriaceae. In: Steyermark, J.A., PF. Berry, K. Yatskievych, and BK. Holst, Flora of the Venezuelan Guayana, vol. 5. Missouri Botanical Garden Press, St. Louis. Pp. 542-573 Morton, CV. 1939, A revision of Besleria. Contr. U.S. Natl. Herb. 26(9):395-474. 274 Journal of t tani tit Texas 2( BOOK REVIEW Roy Morey. 2008. Little Big Bend: Common, Uncommon, and Rare Plants of the Big Bend National Park. (ISBN 978-0-89672-613-0, pbk.). Texas Tech University Press, Box 41037, Lubbock, Texas, 79409-1037, U.S.A. (Orders: www.ttup.ttu.edu, ttup@ttu.edu, 1-800-832-4042, 1-806-742-2982, 1-806-742-979 fax). $34.95, 329 pp., color photographs, 7.5" x 10". Contents: Preface Acknowledgments About This Book Big Bend, the Land of Extremes Appendix A: Critically Imperiled, Imperiled, and Vulnerable Plants Appendix B: ee Plants KE E eene Appendix D: A Note on Botanical Nomenclature Glossary ources References Index An updated l f tl lis] lue. There was the 1951 Park Service Pl ft Bend N LP W.B. 3. McDougall and Omer E. y with 1 190 species ger numerous o eg ane white EE Biol Warnock s Wildflowers o ` hen atthe reainn ~ tarned to the genera ] geben to Trans- Pecos os plants by Michael Powell and others, all of which have been LUE This book fills t at least and ly expertly but handsomely. Agaves and cacti g I p separate sec tions at the Bcc amus of the book, after which other pl follow ( ts and dicots mixed). Each pl bly treated with an excellent photo; non-technical description, and indicati f where in the park the plant might be found d. (Eager to see Texas Thelopody? Follow i it and to it g T.wrightii, too.) TI f the showier plants; more By definition coverage is limited. The book does not pretend to be comprehensive. Rather it reflect ion for ER plants and the desire to share that passion. Need for a comprehensive guide to plants of the Park still exists, but this is one very worthwhile contribution to that end. + + E 1 + 1 1 Té p E 1 1 1- ] Amon g the fou g E y Park location. Joann Karges, Texas "usa University Library (retired), Fort Worth. Texas, U.S. A. J. Bot. Res. Inst. Texas 2(1): 274. 2008 FOLIA TAXONOMICA 7. TWO NEW SPECIES AND A NEW SECTION IN EPISCIA (GESNERIACEAE) FROM THE VENEZUELAN GUAYANA Christian Feuillet Department of Botany, MRC-166 Smithsonian Institution, PO. Box 37012 Washington, D.C. 20013-7012, U.S.A. feuillec@si.edu ABSTRACT (G j ) are described from the state of A in the Venezuelan Guayana. Episcia duidae sp. Two new species of E nov. I UIS E, pian in section Episcia by its flowers, but differs by its ES ee parts and densely appressed-pubescent leaves. E p. nov. resembles E. fimbriata and E. sphalera vegetatively, diff its red corollas. The latter three belong FRI E in subsection T RESUME TA H SR eg : ff^ H x +A m ] Pierre HA qm Vénézuélienne | A A LM | 11] I E ke 4 E É A " A 1 5] ; A: tes E md SE É 1 SS Est ] fleurs à E. ver Episcia, I D 8 pius [ ER mas a 7 1.1 S eee thle GD E , ; Jf. 11 : qués. Ep g fimbriata et E sphalera, I ges. Ces trois espèces appartiennent à la sous-section T tant] 5levée ici d ti RESUMEN — Alar oera da Ja A le SEDEM duidae Es nov. s o ee ls H J Trl : ‘fo parece a E. reptans en la Soe isca por sus flores, pero ppm Episcia rubra sp. nov. parece a E. eege yE. sphalera Vegetativa amente, pero varía DTE sus calis rojas. Los gees la subsección Trematanther [3 The two new species of the subfamily G joideae Link described below were collected in the Venezuelan state of Amazonas. They belong to Eun Mart. in the tribe Episcieae Endl. Since the regional treatment by Leeuwenberg (1958) and the formal infrageneric classification he proposed for Episcia, the genus has been split into four groups. Most authors now follow Wiehler (1973, 1978) in adopting Alsobia Hanst. and Paradrymonia Hanst., as well as the transfer of several Episcia species to Nautilocalyx Linden ex Hanst. The transfer to the other genera affected all the species of six of the seven sections and in section Episcia, three of the five subsections. The current concept of Episcia is the equivalent of two of the subsections of Leeu- wenberg (1958), i.e. section Episcia subsections Episcia and Trematanthera Leeuwenb., and follows the most recent molecular studies (Smith 2000; Zimmer et al. 2003; Clark et al. 2006). Episcia Mart., Nova Gen Sp 3:39 Jan-Jun 1829. Trt: E ptans Mart. (LECTOTYPE: cf. Leeuwenberg, Gesner. Guiana 309. 1958) 1.—Sect. Episcia Anther celis dehiscing throughout by a longitudinal split. Episcia reptans (type), E. andina Wiehl., E. cupreata (Hook.) Hanst., E. lilacina Hanst., E. prancei Wiehl., E. xantha Leeuwenb. & E. duidae sp. nov. Episcia duidae Feuillet, sp. nov. (Fig. 1). Tyee: VENEZUELA. Amazonas. Dept. Alto Orinoco: Cerro Duida, Culebra Creek, 1500-1600 m, 21 Nov 1950 (f), B. Maguire, R.S. Cowan & LI Wurdack 29634 (noLoTYPE: US; isotype: NY). c | ol J 1: Herba stolonib , repens. Basis st ioli I laterales; tricl icellul , api Rene i. Calyx hirsutus, lobi integri, vel dentati. Corolla cen tubus longe villosus. Mat forming stolonif herb; whole plant with long, multicellular, gland-tipped trichomes, old stem with a few short tricl (Berry et al. 4972) or 5-8 mm long, dense trichomes (Cowan & Wurdack 31288). Leaves | Bot Rac Inet Tayag 2(1): 7765 ` 280. 2008 276 Fic 1 FI H "at E DTN Dlant £ H 4 L Li Aal fl fi > La É E +! is T4] and flower huds F fm | ki bk "E a ` Ff ` Kl T Fr (middla) r D laf. I L 51500 (RIV. L Las LJ n Sal Ma fans] Iie | +h 371 A AF "1 I^n£z»5A4 fi A An X FF A FF 7 F i P af por] i JT LE hattam riaht | sall D Fi E [| I i, ar | D 1 aL age A Das I.E on I Anz» fl SA E Es onened rancula a F kl ff 1 LE o | |] (left valve 4.7 mm long), Cowan & Wurdack 31288 (NY). Feuillet, N i ] tion in Episci 277 opposite, equal or subequal in a pair, clearly larger in proximal part of the stem; petiole 0.5—3 cm long, hirsute, trichomes 2-2.5 mm long; blade carnose, suborbicular-ovate to broadly elliptic, 2—4.5 x 14-2.5 cm, base rounded to obtuse, apex rounded to broadly obtuse, margin crenate, with dense, long, appressed, yelow trichomes on both surfaces, except above on s main veins that show their dark green color, 3—5 ide of the midrib, ending i in margin sinuses reduced pair- -flowered cyme, fasciculate, 1-4-flowered, axillary in the apical part of the stem with small leaves still growing; pedicel about 8 mm long. Flowers with sepals free, entire or 1-2-toothed in apical half in the same flower, lanceolate to oblanceolate, 8-9 x 1.5 mm, densely hirsutulous on both sides, the posterior one slightly involute and nar- rower; corolla held transversal in the calyx, crimson, basal gibbosity 3 x 4 mm, uniformly densely pilose, tube 15-18 mm long, densely appressed-pubescent outside, inside papillate in apical third and glabrous below, lobes suborbicular, rounded, serrulate, 7 x 4 mm, glabrous on both sides except at very base where joining the tube; stamens inserted about 2.5 mm above base of corolla tube, filament glabrous, broadened at base, anthers suborbicular, 1.5 x 1.2-1.5 mm; dorsal glands fused in a tongue-like gland, about 1 x 1 mm; ovary superior, ovoid, 3 x 3 mm, densely sericeous—pilose, style 12 mm long, densely pilose. Capsule globose, 4.5-5 mm long. Distribution.—Episcia duidae is known from Cerro Duida (Munic. Alto Orinoco) and Serranía Parú (Munic. Atures) in the state of Amazonas (Venezuela) between 1500 and 2000 m. It was collected at the bases of waterfalls and in wet crevices. Phenology.—1t has been collected in bloom in February-March and November, and in fruit in Febru- Episcia duidae resembles the red-flowered species of subsection Episcia: E. andina Wiehl., E. cupreata (Hook.) Hanst., and E. reptans Mart. Inside the corolla tube, E. duidae has the apical third with large papil- lae, E. cupreata and E. reptans have a ring of glandular trichomes in the throat, E. andina is said to have a corolla glabrous inside (Wiehler 1984; but this could not be checked). The style is densely pilose in E. duidae, unknown in E. andina, and glabrous in the other two, occasionally with scattered trichomes at the apex in E. cupreata. Ge duidae and E. andina differ from the other two by much smaller leaf blades with a long dense app | on both sides rather than hirsute leaf blades found in the other two spe- cies. The corolla of E. tenian is about twice as long as in the other three; it is cylindric and nearly straight in E. duidae and E. reptans, infundibuliform and straight in E. andina, and clearly curved in the middle in E. cupreata. E. duidae differs from E. cupreata by the dense appressed (versus hirsute) indumentum on the leaf blades, the corolla posture transverse (versus oblique) in the calyx, with a tube straight (versus curved near middle), densely appressed-pubescent (versus pilose), smaller corolla lobes, and a style densely pilose (versus glabrous or with a few trichomes near the apex). Episcia duidae was “sp. B” in Feuillet and Steyermark (1999), where it should be noted that numbers 2 and 3 were switched in the first bracket of the “Key to the species of Episcia” in that publication. Etymology.—The new species is named after Mt. Duida at the base of which the type specimens have been collected. ParaTYPES: VENEZUELA. Amazonas. Dep Parú, 2000 m, 7 Feb 1951, R.S. Cowan & J.J. Wurdack 31288 (NY), 1200-1250 m, 5-7 Mar 1991, P.E. Berry, O. Huber a Rosales 4972 (MO, US). 2.—Sect. Trematanthera (Leeuwenb.) Feuillet, sect. nov. Bastonva: Episcia subsect. trematanthera Leeuwenb., Blumea 7:309-310. 1 Anther cells dehiscing only on the basal half by a partial split. Episcia sphalera Leeuwenb. (type), E. fimbriata Fritsch & E. rubra sp. nov. Episcia rubra Feuillet, sp. nov. (Fig. 2). Tyee: VENEZUELA. Amazonas. Dept. Atures: Río Coro-Coro, W of Serranía de Yutajé, 3 km N of settlement of Yutajé, along a small tributary, 5°38'N, 66°07'W, 200 m, 19 Feb 1987, B.K. Holst & R.L. Liesner 3087 (HOLOTYPE: US-3120092; tsotypes: MO, US-3284218) TI 1 "| -1 7 + ] qs $5] E E ol: 1 Tla« p + 1 Le + 11 + 278 t tani i Texas 2( En > Fr. $ i r "Fas DI "I fi 21.1 ELI 1. OS [| A Heal DI! sAog il Lad e FI L Le nfs Ev | Hal Al 3 Lu X d et 21 mm), id (isotype: US), bott th 25 ), id (isotype: MO) PI Herb terrestrial, creeping with short stolons; stem with long, multicellular, fuzzy, dense, yellowish trichomes. Leaves opposite, equal or subequal in a pair, clearly larger in proximal part of the stem; petiole 0.5-2.5 cm long, trichomes longer than on the stem; blade 2.5-9 x 1.5—4.5 cm, ovate to elliptic, base acute to obtuse, apex obtuse, margin crenate, ciliate with a tuft of trichomes at apex of each tooth, above pebbled, with trichomes at apex of each of the bullae, beneath with dense, long, silvery indumentum, 6-9 main veins on each side of the midrib. Inflorescence a reduced pair-flowered cyme, fasciculate, 1—4-flowered, axillary in the apical part of the stem with small leaves still growing; pedicels 8—1.5 mm long, hirsute, with long, fuzzy, dense, yellowish trichomes. Flower: sepals free, oblanceolate, 23.5 x 1 mm, apex acute, tip and 1-2 teeth thickened, hirsute, trichomes 1.8-2.3 mm long, fuzzy, dense, yellowish; corolla oblique in the calyx, crimson, basal gibbosity 1.5—2 mm long, conical, rounded distally, tube 1.2-1.6 cm long, outside with scat- tered, 1 mm long trichomes, lobes suborbicular, 3 x 4 mm, serrulate; stamens inserted 2-3 mm from the base of the tube, filaments 9-12 mm long, anthers reniform, about 1 x 1 mm, cells divergent, dehiscence wide at base, narrow then absent toward apex; nectary comprised of one dorsal tongue-shaped gland, 0.8 Feuillet, N i | tion in Episci 279 x 0.5 mm; ovary superior, ovoid, 2.5 x 1.5 mm, hirsute, trichomes 1.5-2 mm long, style 15 mm long, gla- brous, stigma papillose. Young fruit subspherical, 3.5 mm in diameter, hirsute with yellowish trichomes, 1-1.5 m Distribution and phenology.—Episcia rubra is known only from the type collection from crevices in moist, shaded, rocky canyon along a tributary of Río Coro-Coro, in Amazonas, Venezuela, around 200 m in elevation. It was blooming and had young fruits in February. The anthers dehiscent only in the basal half place Episcia rubra in sect. Trematanthera with E. fimbriata Fritsch and E. sphalera Leeuwenb. among which it differs by its red corollas. Etymology.—The new species is named after the color of its red corolla, unique in Episcia sect. Trema- tanthera. KEY TO THE SPECIES OF EPISCIA FROM THE GUIANA SHIELD . Corolla red or red-orange. 2. Petiole of mature leaves 30-70 mm long; corolla 20-35 mm men 3. Corolla tube cylindric, 3-3.5 cm long, nearly straight, pi t, all 5 lot | ling. Colombia, Venezuela (western and Amazonas, Bolivar), French Guiana, Guyana, Surinam, Brazil (Minas Gerais and northern), Per E. reptans 3. Corolla tube ampliate to twice as wide at throat than at base, < 2.5 cm long, and suddenly curved at middle, yellow in the throat, 2 dorsal E ENER SE America, Folombia, Venezuela (western), Brazil (Amapá), Ecuador, Peru 4 E. cupreata 2. Petiole of mature hiis 10-30 mm n long; coroll tube 12- is mm long. 3. Calyx lobes 7-8 mm long; corolla tube densely appressed ; ant! Ils opening by a long tudinal slit fon base to apex. venezuela (Amazonas) E. duidae 3. Calyx lobes g; sparsely hirsute; anther cells dehiscing only on their basal half. Venezuela TD D E. rubra 1. Corolla yellow, white, or lavender to light blue. 4. Corolla yellow; calyx lobes ao Vi la (A ), Guyana, Surinam, French Guiana E. xantha 4. Corolla tube une: lobes while or | ler; calyx lobes | | spathulate. 5. Corolla lok pale green; calyx lobes | late. Brazil (Amapá, Pará), Surinam, French Guiana E. sphalera 5. Corolla with lavender markings or lobes lavender to blue; leaf blades variously colored; calyx lobes lanceolate or spathulate 6. Corolla white with lobes lavender to blue; calyx lobes spathulate; leaf blades mostly with color markings; anther cells opening by a longitudinal slit from base to apex. Central America, Colombia + Guianas (cultivated, rarely escaped) E. lilacina 6. Epralla white, | dd glos kings; calyx lobes | late; leaf blades mostly green, | || Ils dehiscing only on their basal half. Colombia, Venezuela (Amazonas), Brazil (western and northern), Peru E. fimbriata ACKNOWLEDGMENTS I am grateful to Larry Skog and John L. Clark for their reviews and suggestions. I want to thank the curators of the herbaria MO and NY for lending the material in their care. This paper is published as No. 136 in the Smithsonian's Biological Diversity of the Guiana Shield Program publication series. REFERENCES CLARK, J.L., PS. HERENDEEN, L.E. Skoc, and E.A. Zimmer 2006. P Phylogenetic relati hi | icl laries in tl Episcieae (Gesneriaceae) inferred from nuclear, chloroplast, and aca data. Taxon 54:313-336. FEUILLET, C. and J.A. STEYERMARK. 1999, Gesneriaceae: In: Steyermark, J.A., PE. Berry, K. Yatskievych, and BK. Holst, eds. Flora of the Venezuelan Guayana, vol. 5. Missouri Botanical Garden Press, St. Louis. Pp. 542—573. LEEUWENBERG, A.J.M. 1958. The Gesneriaceae of Guiana. Acta Bot. Neerland. 7:291-444 SMITH, J.F. 2000. Phylogenetic resolution within the tribe Episcieae (Gesneriaceae): congruence of ITS and NDHF sequences from parsimony and maximum-likelihood analyses. Amer. J. Bot. 87:883-897. 280 J loft tanical Insti Texas 2( Wienuer, H. 1973. Seven transfers from Episcia species in cultivation (cesneraceae] Ge 27:307-308. Wiener, H. 1978. The genera Episcia, Alsobia, Nautilocalyx and Paradrymo ). Selbyana 5:11-60. Wiener, H. 1984. Miscellaneous new species in the Gesneriaceae. sim 7 328-347. Zimmer, EA. E.H. RoaLson, LE. Skoc, J.K. Bocca, and A. lonurm. 2002. Phylogenetic relationships in the Gesneri- oideae (Gesneriaceae) based on nrDNA ITS and cpDNA trnl-F and trnE-T spacer region sequences. Amer. J. Bot. 89:296-311, FOLIA TAXONOMICA 8. PASSIFLORA TECTA (PASSIFLORACEAE), A NEW SPECIES IN SUBGENUS PASSIFLORA FROM THE GUIANAS Christian Feuillet Department of Botany, MRC-166 Smithsonian Institution, PO. Box 37012 Washington, D.C. 20013-7012, U.S.A. feuillec@si.edu ABSTRACT Passiflora tecta, a new species with red ES is described from up It Seen in EE Pa supersection Coccinea and resembles Passiflora coccinea and P. miniata. The tl g g t i outer corona filaments of P. tecta are scarlet and held e erect, ns oblique «— like a nar d tl ] In P. coc- cinea the outer corona filaments are white and held oblique inward, while in P. miniata ea are dark purple and ex erect, slightly oblique outward like the rim of a flower pot. RÉSUMÉ D Dn A 4 A 11 >a 1] + As Se us eg 11 t1 + D ATI at F^ a] T Wa - tD Lë Li YI miniata. Ces troi pé t faciles à it 1 té le ] Les filaments I externes de P. t oy H t 1,1 + Il: m AS ai A D ET di 2 st E A £orl-zceme lema + 7 PO HLT SOLIL 1 AS? L CALL UAI comme le rebord d'un pot de fleur. In Feuillet (2007) two new species were described in Passiflora subg. Passiflora supersect. Coccinea Feuillet & J.M. MacDougal, and a key to the species was given. In that paper, one branch in bracket 11 of the key led to “P. sp. (Guyana)", known only from photographs. Since publication of that manuscript, herbarium material has become available, thus allowing a good description to be drawn from actual specimens. This new species is described as Passiflora tecta Passiflora tecta Feuillet, Sp. nov. (Fig. 1-2). Tree: GUYANA. PoTARO—SIPARUNI: Eagle Mountain, around the OMI base camp, 5°15'18"N, 59°6'59"W, 383 m, fl., 10 Sep 2006, K.M. Redden 4132 (HOLOTYPE: K) cy at Bi D 1 a EN tecta in subg. Passiflora i pania inea y iseriata; P. coccineae affinis iffert; P. miniatae affinis, timis rubri | is, intime versus extrinsecus obliquis differt. D Liana climbing on shrubs and small trees at forest edge, probably reaching the canopy. Young stems terete, slightly striate, with short, irregularly curved trichomes. Tendrils long, thin, with short arched trichomes. Vegetative bud in apical position, less than 1 mm away from the rest of the axillary complex (petiole, stipules, pedicel, tendril), prophylls acuminate at apex, densely short-pubescent. Stipules linear, 2.5 x 0.8 mm, short-pubescent, deciduous before the leaf differentiate in petiole and blade, leaving a round scar. Petiole 8-12 mm long, curved at base, with 2 lateral glands at the very base; glands scar-like, glandular surface oval, ca. 1 x 1.5 mm, with 1-2 tufts of long white trichomes, otherwise glabrous, rim minute pubescent. Blade ovate, 5-10 x 2.5—6 cm, bi-serrate, teeth ending in small, stipitate, round glands, acuminate and mucronate at the apex, broadly cordate and briefly cuneate at the base, drying brown adaxially and yellow- brown abaxially, 2-3 main lateral veins on each side of the midrib, the first pair about 2/3 as long as the midrib, tertiary venation scalariform. Inflorescence sessile, 1-flowered. Pedicels erect, 4.5—6.5 cm long, with dense short pubescence. Bracts three, verticillate, orange red, ovate, concave, 3-5 x 2-2.5 cm, free to base, round at the apex, glandular serrate at the margin on the apical two-thirds, with dense short trichomes on both surfaces. Flowers orange red, perianth lobes adaxially whitish at base; hypanthium 7 mm long, 9 mm broad on live material, round at base, inflated with the point of contact with the pedicel invaginated, J. Bot. Res. Inst. Texas 2(1): 281 — 283. 2008 282 tani i Texas 2( Fic. 1. Passiffora tecta in Guyana in Feb 2007 (photo by C. Kel- — Fic. 2. Passiflora tecta in Guyana in Feb 1990 (photo by C. loff). Petal 2.3 cm long. Feuillet). Androgynophore 1.7 cm long. E ss la! a Y | LIANA fal MI Fic. 3. Passiflora coccinea in French Guiana in April 1985 — Fig. 4. P. (Feuillet 2128; photo by C. Feuillet). Outer corona filaments to by C. Feuillet). Out l t g densely short pubescent outside; sepals narrow-oblong, 2—2.5 cm long, plus a subapical awn, 0.7-1.0 cm long, 0.5—0.6 cm broad, short-pubescent in the parts exposed in bud, otherwise glabrous; petals similar to the sepal, slightly narrower, thinner, not awned, glabrous; corona bright red, in 3 rows, filamentose, the first 2 rows composed of laterally compressed, narrowly triangular filaments, erect to 70-80", slightly oblique, leaning against the androgynophore and the ovary, the first row about 12 mm long, the second row about 9 mm long, the third row filiform, about 5-6 mm long; operculum borne at the top of the hypanthium, membranous, dependent to 2/3 down the hypanthium, then recurved inward, laciniate at the margin, 9—10 mm long, glabrous; nectar disc annular at the bottom of the hypanthium; limen membranous, surround- ing the base of the androgynophore, 1-2 mm long, glabrous; androgynophore red throughout, cylindric, ca. 1.7 cm long, glabrous; stamen glabrous, filaments flat, pale green with red dots throughout, 6 x 1 mm, joined in a short membrane at base, anthers dorsifixed, rectangular, 6 x 2.5 mm; gynophore shorter than and covered by the membrane at the base of the androecium; ovary green, 6 mm long, 2-2.5 mm in diam., ellipsoid, densely short-pubescent; spes pus as Ten Hos 9-12 mm "dio Fade pubescent; stigmatic g n elongated pedicel, surface light green, papillate. i | é gtop graph) g hidden by yellowish green bracts. Feuillet, A : £ Daceifl € thn Curt 283 Distribution —The only specimen of Passiflora tecta known to me was collected in flower from Guyana, in Tue Potaro- EE EE near Peer Mountain in a clearing at the edge of the forest in September 2006. phed twice in Guyana, once in February 1990 (C. Feuillet) in the Upper Takutu- oe Essequibo num near Surama, at the forest edge along a dirt road (with bud and young fruit), and again in Rep SE (C. Kelloff) in the Rotator paral region, in Kaieteur National Park (flower), though no herl were made. The three localities are in western centra ] Guyan a and the plants were collected or Bst graphed at the edge e lowland rainforest (less than 500 m elevation Passiflora tecta could be easily confused with P. coccinea, the most common species of Passiflora in the Guianas. Passiflora tecta sp. nov. clearly belongs to P. subgen. Passiflora supersect. Coccinea due to its distinctive leaves with serrate margins, red floral bracts with glandular-serrate margins, red-colored perianth, and straight corona filaments that form a cone angled towards the androgynophore. In this supersection, P. tecta most closely resembles P. coccinea Aubl. (Fig. 3), a species with a large Amazonian and Guianan distribution described from French Guiana (Aublet 1775). However, the Guianan endemic P tecta has narrow-elliptic rather than ovate bracts and red rather than white outer corona filaments. Passiflora tecta also resembles P. miniata Vanderplank (Fig. 4), described from western Amazonia (Colombia, Brazil, Peru, and Bolivia) by John Vanderplank (2006), from which it differs in its outer corona fil ts that are orange-red and slightly oblique inwards, rather than dark purple and slightly oblique outwards. Etymology.—The Latin specific epithet means "covered." This refers to the tl f fil t that restrict pollinator access to the nectar chamber contrary to the condition in P. miniata, where only the innermost row of filaments guard the throat. ACKNOWLEDGMENTS I would like to thank Carol Kelloff for letting me use her excellent photograph and S. Pérez-Cortéz and an anonymous reviewer for their careful reading of the manuscript. This paper is number 137 in the Smithso- nian’s Biological Diversity of the Guiana Shield Program publication series. REFERENCES Auster, J.B. 1775. Histoire des plantes de le Guiane Françoise 2:828. FeuiLter, C. 2007. Folia taxonomica 2. New species of Passiflora subgenus Passiflora (Passifloraceae) from the Guianas. J. Bot. Res. Inst. Texas 1:819-825. VANDERPLANK, J. 2006. 562. Passiflora miniata. Curtis's Bot. Mag., new ed., vol. 23:223-230. fT TEA! VAU £4 dj 284 BOOK REVIEW James H. Everrrr, ROBERT I. LONARD, AND CHRISTOPHER R. LITTLE. 2007. Weeds in South Texas and Northern Mexico: A Guide to Identification. (ISBN 978-089672-614-7, pbk.). Texas Tech University Press, Box 41037, Lubbock, Texas 79409-1037, U.S.A. (Orders: www.ttup.ttu.edu, ttup@ttu.edu, 1-806-742- 2892; 1-806-742-2979 fax). $19.95, 222 pp., 6" x 9". Contents: Preface Introduction Class Polypodiopsida (Ferns) Class Magnoiliopsida (Dicots) Class Liliiopsida (Monocots) Appendix d 1: n Plants and Federally Mg c iia e GE Texas ted Pathogens and I t Texas Appendix 3: Selected Pathogens and Insect Pests of Giant Reed (Arundo donax) in Texas E n 4: Selected Pathogens and 1 t Pests of B ] (Cynodon dactylon) in Texas PE Pathogens and 1 t Pests of À ] Blueg (P ) in Texas A ] Kc] TRES | T r Dat E Tal rx 1 Lil pp Pathogens and d g Appendix 7: Sel ] Chemical and Cultural Controls of S M (M) and Dicot (D) Weeds of South Texas and Mexico Glossary Literature Cited Index Any ] e 1 1 Jaf. i weed q+} A +L t, I g "n 1 € , of course, | of tl df] herish l | by native plant enthusiasts: Cirsium texana, Pluchea purpurascens, Oenothera Lii and DRESS The text hint f plants to be so designated though not establishing full justification : g by family, di ] d, each sy with a color J gu EE 1 1 ] f pl 1 ro 1 af, 1 1 ight 1 which the TS “em 41 A af t +1 s 1 1111 to halaf] Dor} f] +1 + D : i iie Bon g I g ] , the majority of 1 + +1 Kat A mE " A o J Tr 11 1 1 1 E fl E 1 4 + f, 1 e L i I O IL L [e] ning amateur. A tl (viral and f D, nematodes, and i t ts of f the pl ] larly need rr e x: KA e D D E Fl to be AT SEAN, TE 4 e xccl. mem o xem eco" E 1 e TL 1 1 1 [a] 1 +1 1 +1 J. Bot. Res. Inst. Texas 2(1): 284. 2008 state of Texas and adjacent areas. — Joann Karges, Texas Emisión University Library (retired), Fort Worth, Texas, U.S.A. A SYNOPSIS OF PACHYPHYLLUM (ORCHIDACEAE) Eric A. Christenson 4503 21% Ave. West Bradenton, Florida 34209, U.S.A. orchideric@juno.com ABSTRACT ft tropical orchid Pachyphyllum Kuntl ted a are SC for all names including 10 new lectot ifications. T divided int 3 sections, based on th Ord and on n new, P. sect. ge 2 e “L r Capitulum E.A. Christ. Three names are considered synonyms within Pachyphyll d eight lud g Key Wonps: Pachyphyllum, Orchidaceae, Neotropics RESUMEN Se enumeran las 41 especi tadas del gé d id tropical Pachyphyllum Kunth. Se citan los tipos de todos | 1 incluyendo 10 nuevas regen El género se divide en 3 secciones, una basada en el género Orchidotypu nzl. y otr talmente nueva, P. sect. Capitulum E.A. Christ. Tres nombres se consideran sinónimos en Pachyphyllum y EE nombres se a del gén PEE the characteristic growth habit and relative abundance of available herbarium material, the genus D 1 71 £..11 ] 4 ] woetully ied. Concentrated biodiversity studies in the Department of Cusco in s ae Peru (www.andesamazon.org) have recorded many species in the genus, some new to Peru and some, no doubt, new to science. Critical study and dissemination of this information has just begun (Repasky & Christenson, in press) Recent collections of Pachyphyllum ecallosum from the Department of Cusco, Peru, have emphasized the distinct nature of its inflorescence. Unlike the few flowered inflorescences of the type collection from the Department of Huancavelica, more recent collections consistently bear 4—6 flowers arranged in a well defined capitulum. A new section is proposed for P. ecallosum and its sister species, P. tortuosum. In addition, the genus Orchidotypus is given formal status as a section of Pachyphyllum. KEY TO THE SECTIONS o Flowers arranged in a dense capitulum P. sect. Capitulum Geh ENEE in d raceme. | together at the | f g a tube for half their length P. sect. Orchidotypus 3 Sepals and petals free P. sect. Pachyphyllum Pachyphyll section Pachyphyll Tyee species: Pachyphyllum distichum Kunth. ini e breviconnatum Schltr., Repert. Spec. Nov. Regni Veg. Ge e "e Se Repert. Spec. Nov. Regni Veg. Beih. 57: t. 130, nr. 510. 1929. Tee: PERU. Derr w Tres Cruces Inn, 3600-3700 , designated here: US; ISOLECTOTYPES: F S m, Weberbauer 6976 (HOLOTYPE: B, d 4 Comment. —The US specimen is designated lectot tl better fl ilable than on either of the two sheets at F or the JI two sheets at MOL. Pachyphyllum bucarasicae Kraenzl., Pilanzenr. 4, 50:25. 1923. Tree: COLOMBIA. Derr. SANTANDER: Bucasasica, 3300 m, Kalbreyer 918 (nororves: ?W). Pachyphyllum capitatum Kraenzl., Bot. Jahrb. Syst. 37:386. 1906. Tr: PERU. Derr. Amazonas: E of Chachapoyas, between Tambo Ventillas and Piscohuañuna, 2600-2700 m, Weberbauer 4421 (morore: B, destroyed; LecroTYPE, designated by Bennett & Christenson in Icon. Orchid. Peruv. pl. 739, 2001: MOL). i Rat Rac Inct Tavac 2(1): TAE — 289. 2008 286 t tani itute of Texas 2( Pachyphyllum cardenasii Smith & Harris, Contr. Gray e ae pe 1936. Tree: BOLIVIA. Derr. COCHABAMBA: Prov. Ayopapa, Sailapata, 3000 m, Cardenas 3289 (LecroTvrE, designated h G). KT 1 1 ] E +1 dos Comment. VIA VU LIL otype É L 5 11 j (Ames & C. Schweinf.) - - US , Lilloa > oe pie Centropetalum costaricense Ames a E Schweinf., Sched. Orch. an 110. 1930. Tre: COSTA R , above Los Lotes, N of El Copey, 2100-2400 m, 21-22 Dec 1935, Standley 42600 Geen AMES; ISOTYPE: US). Pachyphyllum crystallinum Lindl., Orch. Linden. 18. 1846. Tir: VENEZUELA: Merida, 10,000 ft, J. Linden 686 (Ho- LOTYPE: K) 11 cuencae Rchb.f., Linnaea 41:31. 1877. Tre: ECUADOR: S C , 2900 m, Jul 1864, Jameson ZE s.n. (MR W, ISOTYPE: AMES). iai cipe o o Repert. Spec. Nov. Regni Veg. 15:217. 1918, in textu; Repert. no Regni Veg. B 1922. Basionym: Pachyphyllum falcifolium Schltr., Repert. Spec. Nov. Regni Veg. 10: MA non Rchb.f. Tyre: EEN in moss, near Ingenio, R.S. Williams 1631 (HoLoTYrE: B, destroyed; Lectotype, designated E ) NY; ISOLECTOTYPE: BM Comment.—The NY speci is here d e ted lectotype. Pachyphyll fi Kraenzl. i name based on Schlechters P falci- folium Ye CAI 1 kris) 1 his own new name. GEN SÉ Kraenzl., Pflanzenr. 4, 50:22. 1923. Pachyphyllum dalstroemii Dods., Orquideología 20(3):278. 1997. Tre: ECUADOR: Loja, San Ped 1 from Loja t Vilcabamba, 2500 m, Apr 1984, Dalström & Hoijer 802 (HoLoTYPE: SEL). Pachyphyllum denticulatum (Ruiz & Pav.) Schltr. Ge Spec. Nov. Regni Veg. Beih. 9:180. 1921. Fernande- zia denticulata Ruiz & Pav., Syst. Veg. Fl. Peruv. & Chil. 1:240. 1798. Tre: PERU. Deer. Huanuco: Pillao, Pavon s.n. (HOLOTYPE: MA). I ll distichum Kunth, Nov. Gen. & Sp. 1:339. 1816. Type: ECUADOR: I , Gonzanam, 6420 ft (10701 ] J, “Humboldt & Bonpland s.n. (HOLOTYPE: P). Pachyphyllum falcifolium Rchbf., Linnaea 41:32. 1877. Tez: ECUADOR: Loja, Sep, Jameson s.n. (HOLOTYPE: W; ISOTYPE: Pachyphyllum favosifolium Kraenzl., Pflanzenr. 4, 50:23. 1923. Tree: COLOMBIA. Derr. Antioquia: Sonson, 3300 m, Kalbreyer 1935 (HOLOTYPE: ?W). Pachyphyllum hagsateri Dods., Orquideología 20:104. 1996. Tree: ECUADOR. Sucumbios, El Mirador between Playón and San Gabriel, 3300 m, 8 Jul 1990, Dodson et al. 18501 (HoLoryrE: RPSC; isotypes: MO, QCNE Pachyphyllum hartwegii Rchb.f., Bonplandia 2:219. 1854. Pachyphyllum distichum Lindl. ex Rchb.f., Bonplandia 3:219. 1855, non Kunth. Tree: ECUADOR: Pichincha, Hartweg s.n. (HOLOTYPE: W). Pachyphyllum herzogii Schltr., Meded. Rijks-Herb. Leiden 29:80. 1916. Tw: BOLIVIA: Comarapa, 2600 m, Apr 1911, Herzog 1946 (HOLOTYPE: L; ISOTYPE: Z) mo lycopodioides en , Repert Spec. Nov. Regni Veg. Beih. 9:116. 1921; Repert. Spec. Nov. Regni Veg. Beih. 57: t. 130, nr. 511. 1929. Tre: PERU: Huanuco, SW of Monzón, 3400-3500 m, Weberbauer 3327 (HoLoTYPE: B, d yed; Hu here: MOL) 00 m, M. Madero s.n. (HoLotyre: B, destroyed, drawings aE micrangis SSC a pe Now, Regni d Beih. 7:203. 1920; Repert. Spec. Nov. Regni Veg. Beih. 57: t. 73, ca. 30 AMES; LECTOTYPE, designated "a t; nr. 281, loc. cit.). T1 TI " J : AEN | dica] +] | cd vac s +] tati fal Le JA : + AMEC Comment. AT J 1: Cé Tab WW | + 15 “e 1 + 1 + + I Pachyphyllum micranthum Schltr., Repert. Spec. Nov. Regni Veg. Beih. 7:204. 1920; Repert. Spec. Nov. Regni Veg. Beih. 57: t. 63, nr. 282. 1929. Tre: COLOMBIA: Cauca, M. Madero s.n. (HoLoTYPE: B, destroyed, drawings AMES; LECIOTYPE, designated here: t. 73, nr. 282, loc. cit.). Christenson, Synopsis of Pachyphyllum 287 Re LiL. ] - es NS | n D POPE SR + +] A Co O, E GE o + ANTEC 11 Schlt t. Spec. Nov. Regni Veg. 10:460. 1912; Repert. Spec. Nov. Regni Veg. Beih. EI t. 60, nr. 239. E. Toe: BOLIVIA: Unduavi, ca. 3200 m, Nov. 1910, O. Buchtien s.n. (HoLoTyeE: B, destroyed; LECTOTYPE, designated here: t. 60, nr. 239, loc. c Pachyphyllum nubivagum L.O. Wms., Lilloa 3:483. 1938. Tree: COLOMBIA. Derr. SANTANDER: western slope of Páramo de las Puentes, above La Baja, 3300-3400 m, 25-31 Jan 1927, Killip & Smith 18161 (Hororyre: AMES) Pachyphyllum parvifolium Lindl. in Hook., Ic. Pl. 2: t. 117. 1837. Te: PERU. Derr. Amazonas: Chachapoyas. 1836, Mathews s.n. (HOLOTYPE: K) Pachyphyllum pastii Rchb.f., Bonplandia 3:239. 1855. Tre: ECUADOR: Quito, Jameson 439 (HOLOTYPE: W; ISOTYPE: K). Pachyphyllum pectinatum Rchb.f., Xenia Orchid. 3:22. 1881. Tre: BOLIVIA. Prov. Larecaja: Sorata, 3100-3200 m, May 1860, Mandon 1152 p.p. (HOLOTYPE: W; isoTYPES: MICH, NY). Pachyphyllum peperomioides Kraenzl., Pflanzenr. 4, 50:27. 1923. Tyee: COLOMBIA. Prov. Nariño: Pasto, 3300 m, Jun 78, EC. Lehmann s.n. (HoLoTYPE: W). mo piesiki Szlachetko, Mytnik-Ejsmont & Rutkowski, Orchidee e 58:94. 2007. Tre: UADOR: Prov. CHINCHIPE: near Yangana, Nov 2005, Szlachetho et al. s.n. (HoLoTYPE: UGDA-D 11 lodicl Rchb.f., Xenia Orchid. 3:22. 1878. Tree: BOLIVIA. Prov. LangcAJA: Sorata, 3100-3200 m, "May 1860, TEN 1152 p.p. (HOLOTYPE: w. ISOTYPE: NY). Pachyphyllum serra Rchb.f., Bonplandia 3:219. 1855. Tre: PERU. Derr. Amazonas: Chachapoyas, Mathews s.n. (HOLOTYPE: W) Pachyphyllum squarrosum Lindl., Ann. & Mag. Nat. Hist. 15:107. 1845. Tre: COLOMBIA. Prov. Popayán: 10,500 ft, leg. ? 1409 (HoLoTYrE: K; rsorvee: K). duplice RE par , Repert. Spec. Nov. Regni Veg. Beih. 7:204. 1920; Repert. Spec. Nov. Regni Veg. B eih. 57: t. 73 r. 283. 1929. Tee: COLOMBIA: Tolima, Volcan de Tolima, E 4000 m, Nov 1868, A. Stübel 213 (HoLOTYPE: B, destroyed, di AMES; LECTOTYPE, designated here: t. 73, nr. 283, loc Comment. NT. À dro f conl An n 1 1 (Ae TH Lus 4 : A pa bd 5 x. ZE t Jt 5 [2] FE EDT ein tajacayaense D.E. Benn. & E.A. Christ., Icon. Orchid. Peruv. pl. 739. 2001. Tee: PERU. Derr. ANCAVELICA: Tayacaja, Dist. Huacocolpa, zone Inquilapata, 2900 m, M. León M. ex Bennett 7877 (HoLotyPE: MOL). Pachyphyllum sect. Orchidotypum (Kraenzl.) E.A. Christ., stat. nov. B Orchidotypus Kraenzl., Bot. Jahrb. Syst., 37:383. 1906. Tyre species: Orchidotypus muscoides Kraenzl. 1 La JT yphy with the notable exceptions of Sen- ghas (1995) and EE EE et al. (2007). The fusion e sepal ] ls into a tube, however, appears to be ample reason to maintain them as a section of Pachyehyllüm: Peces of this section are also consistently smaller statured than species of section Pachyphyllum. The genus Orchidotypus has been considered a synonym of P Pachyphyllum aurorae D.E. Benn. & E.A. Christ., Icon. Orchid. and pl. 546. 1928.1 Tree: PERU: Chanchamayo, 00 m, 12 Apr 1996, O. del Castillo ex Bennett 7501 COMM designated here: [e] FI o r Comment.—Bennett and Christenson (1998) neglected to designate the hol in the original publication. The name is lectotypified “41 +l = * ACHT E so. a Schltr., Repert. Spec. Nov. Regni Veg. Beih. 27:181. 1924. Orchidotypus bryophytus (Sch Senghas, Die Orchideen, ed. 3, 1/B(31):1923. 1995. Tyre: COLOMBIA. Derr. CUNDINAMARCA: near Bogota, 2500 m, 1921, H. E s.n. See B, destroyed). Comment. NI A 1; f TI £A 1] ; ] ta | + + Since T L — A AA SE ++ e 1A spi I ka al F 1.51.3523. 1 1 sp | as reme Á Heta land | Pi f the ger in i Colombia is needed. 288 [| [| £a n. - Ip LI dis den ok f Texas 2( Pachyphyllum gracillimum C. Schweinf., Amer. Orchid Soc. Bull. 16:564. 1947. Orchidotypus gracillimus (C. Sch- weinf.) Senghas, Die Orchideen, ed. 3, /B(31):1923. 1995. Tyre: PERU. Dept. Cusco: Prov. Urubamba, Puyupata-Tuncapata, 3200 m, 6 Aug 1942, C. Vargas 2932 (HoLoTyPE: CUZ) Pachyphyllum pi HP (Rchb.f.) Garay & Dunsterv., Venez. Orch. Ill. 3:236. 1965. Aeranthus hispidulus Rchb.f., 6; Campylocentrum hispidulum (Rchb.f.) Rolfe, Orchid Review 11:246. 1903; Miu in hispidulus (Rchb.f.) Senghas, Die quim ed. 3, VB(31):1923. 1995. Tyre: COLOMBIA: Santa Martha, Purdie s.n. (HOLOTYPE: W). Pachyphyll les (K 1.) Schltr., Repert. Spec. Nov. Regni Veg. 15:216. 1918; Orchidotypus muscoides Kraenzl., Bot. Jahrb. Syst. 37 385, 1906. Tree: PERU. Derr. Cajamarca: Prov. Chota, west of Huambos, 2700-3000 m, Weberbauer 4189 (B, destroyed). P. (Kraenzl.) Schltr., Repert. Sp. Nov. Regni Veg. Beih. 19:72. 1923; Dichaea jai is Kraenzl., Pflanzenr. 4(50), 83:36. 1923. Tree: ECUADOR: near Cuenca, ca. 3500 m, 16 Aug 1878, E C. Lehmann s.n. (HoLoTYPE: W) Orchidotypus vareschii Foldats, Acta Biol. Venez. 2(4):28. 1957. Tree: VENEZUELA: Edo. Trujillo, Pmo. El E 3200 m, Vareschi 4527-A (HOLOTYPE: Pachyphyllum mexicanum Dressler & Hágsater, Orquidea(Mex.) 6(3):73. 1976. Orchidotypus mexicanus (Dressler & Hágsater) Senghas, Die Orchideen, ed. 3, 1/B(31):1923. 1995. Tree: MEXICO. GUERRERO: Puerto El Asoleadero, entre Carrizal de Bravos y Cruz de Ocote, 2600 m, Hágsater 2839 (HoLorYPE: MEXU; rsorvrrs: AMES, ENCB, K, MO, SEL). REN ein L.O. Wms., Caldasia 1(3):15. 1941. Orchidotypus schultesii (L.O. Wms.) Senghas, Die Orchideen, 3. 1995. Tyre: COLOMBIA. Páramo de Chipaque, SE of Bogota, 3200 m, 25 Sep 1941, RE. Schultes 1020 (HoLo- TYPE: je Pachyphyllum tenue pa o apec, NOM Regni um E 9:116. 1921; Repert. Spec. Nov. Regni Veg. eih. 512. 1929 Die Orchideen, ed. 3, /B(31):1923. 1995. Tyre: PERU. DEPT. m in dmn Valley, Koehler s.n. me OTYPE: B, dest Eu , designated | t. 131, nr. 512, loc. cit): El H Pachyphyllum vaginatum Schltr., Repert. Spec. Nov. Regni Veg. Beih. 27:182. 1924. Orchidotypus vaginatus (Schltr.) Senghas, Die Orchideen, ed. 3, 1/B(31):1923. 1995. Tree: COLOMBIA: Cundinamarca, near Bogota, 2500 m, 1921, H. Hopf s.n. (HOLOTYPE: B, destroyed). Noa y] E + f LI Iu T] t 1 tal tantr Since 11 th t a tth 4 lA SHAT 1 y I e eo I [e] I 2 ( omment COTE fea. SE, ee A | 1 " NE PES NOE: A detailed study of th Ds ps Tai 3.4 Pachyphyllum sect. Capitulum E.A. Christ., sect. nov. Tyr seecies: P ecallosum D.E. Benn. & E.A. Christ. Flores in capitulum. Pachyphyllum ecallosum D.E. Benn. & E.A. Christ., Icon. Orchid. Peruv. pl. 738. SS Tres: PERU: Dept. Huan- cavelica, Tayacaja, Dist. Huachocolpa, Capcasoro, 3400 m, 15 Jun 1999, M. León M. 3067 (HoLorvre: MO Pachyphyllum tortuosum Foldats, Soc. Venez. Cienc. Nat. 28:115. 1969. Tree: VENEZUELA. Edo Tachira: Rio Quini- mari, Quebrada Las Copas, 2600-2630 m, G. C. K. Dunsterville & E. Dunsterville 90 (HoLotyrE: VEN; ISOTYPE: K). The recent collection of P. tort f southern Peru (Dept. Cusco, Machu Picchu, Wiñay Wayna, Nauray 181, CUZ) strengthens the relationship of these two species. EXCLUDED SPECIES D hypl 1] h; arpon (Sw.) Spreng , Syst Veg. 3:731. 1826. Epidend hi pon Sw., Prodr. 124. 1788. Cymbidium éclinatánmm (Sw) Sw. Nov dem Reg. Soc. Si. Upsal. 6:71. 1799. Dick 1 (Sw.) Lindl., Gen. & Sp. Orch. Pl. 208. 1833, nom. illeg. Tree: JAMAICA: Swartz s.n. (HoLoTYrE: BM). = Dichaea pendula (Aubl.) Do. Pachyphyllum haemathodes (Ruiz & Pav.) Schltr., Repert. Spec. Nov. Regni Veg.Beih. 9:180. 1921. Fernandezia mathodes Ruiz & Pav., Syst. Veg. Fl. Peruv. & Chil. 1:240. 1798. Tree: PERU. Dert. Huanuco: Muña, Pavon s.n. (HOLOTYPE: MA) = Maxillaria haemathodes (Ruiz & Pav.) Garay. Pachyphyllum pamplonense Kraenzl., Pflanzenr. 4. 50:27. 1923. Tre: COLOMBIA: Pamplona to Las Vetas, 3300 m, Funk & Schlim s.n. (HoLoTYPE: W) = Maxillaria nubigena (Rchb.f.) C. Schweinf. Pachyphyllum procumbens Lodd. ex W. Baxt., Loud. Hort. Brit. Suppl. 3:607. 1850, nom. nud. Christenson, Synopsis of Pachyphyllum 289 Pachyphyllum ?scandens Llave & Lex., Nov. Gen. 2:17. 1824. Tree: MEXICO: Huandacareo, Sep (HOLOTYPE: preserved?) = Barkeria scandens (Llave & Lex.) Dressler & Halbinger NT T 1 + 1 f omment. GULF we iei dal (Ruiz & Pav.) Schltr., Repert. Spec. Nov. Regni Veg. Beih. 9:181. 1921. Twe: PERU. Derr a, Pavon s.n. (HcLOTYPE: MA) = Fernandezia subbiflora Ruiz & Pav. Pachyphyllum steyermarkii Foldats, Acta Bot. Venez. 3:369. 1968. Tyre: VENEZUELA. Edo. Tachira: Tama, 2475-2550 m, Steyermark & Dunsterville 98345 (HoLoT:PE: VEN) = Fernandezia sanguinea (Lindl.) Garay & Dunsterv. in um uniflorum Llave & Lex., Nov. Md Descr. 2:25. 1824. Tyee: MEXICO: Irapeo (HOLOTYPE: preserved?). = Barkeria niflora (Llave & Lex.) Dressler & ume "KT T : 7] + 1 Comment SU A ET rl bbe REFERENCES Repasky, R. and E.A. Christenson (In Press). Orchids of Peru: Pachyphyllum crystallinum. J. Orchideentr. Repasky, R. and E.A. CHRISTENSON (In Press). Orchids of Peru: Pachyphyllum gracillimum. Austral. Orchid Rev. Repasky, R. and E.A. Christenson (In Press). Orchids of Peru: Pachyphyllum hispidulum. Orchid Digest. Repasky, R. and E.A. CHRISTENSON (In Press). Orchids of Peru: Pachyphyllum pectinatum. Orchid Review. SENGHAS, K. 1995. Orchidotypus. Schlechter's Die EECH ES 3, no 1):1922-1924. SZLACHETKO, D.L., J.M. EJSMONT, IPR 2007.P i Szlach. spec. nov. (Vandoideae, Pachyphyl- linae), eine neue Art aus Ecuador. Orchidee (Hamburg) 58: 93- 97. 290 BOOK REVIEWS BRIAN L. FISHER AND STEFAN P. COVER. 2007. Ants of North America: A Guide to the Genera. (ISBN 978- 0-520-254220, pbk.). University of California Press, 2120 Berkeley Way, Berkeley, California 94704, U.S.A. (Orders: California Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, New Jersey 08618, U.S.A., www.ucpress.edu, 1-609-883-1759, 1-609-883-7413 fax). $34.95, 194 pp., line draw- ings, color photographs, 4 1/3" x 7". Contents —Preface, 1) Introduction, 2) Key to North American Ant Genera Based on the Worker Caste, 3) Taxonomic Descriptions, 4) Subfamily Descriptions, 5) Genus Descriptions, 6) Ant Genera of North America by Subfamily, 7) Ant Species North of Mexico: A Working List, 8) Terminology. Identification References, General References, Inde the environment, it has been difficult to c ded to lead to their identification. This handy field guide OH that need with an excellent key that provides line drawings , followed by color photographs of a specus iden of each genus, us Miri of the eid Neither the key nor the diagnost for the fledgli or the easily daunted, tial. Still, "-—— TEN PU ee hepa Leda da section can master UM EE Ofir interest even to da MR nnd is the ' rires and ecology" of each genus llth Joann Karges, us EE University Library (retired), Fort Worth Texas, U.S.A a £21) Jay H. BuckLeY 2008. William Clark: Indian Diplomat. (ISBN 978-0-8061-3911-1, pbk.) University of Okla- homa Press, 2800 Venture Drive, Norman, Oklahoma 73069-8218, U.S.A. (Orders: www.oupress com, 1-800-627-7377, 1-800-735-0476 fax). $29.95, 320 pp., b/w illustrations, maps, 6" x Following his return to St. Louis from the inf d 9. Sa? T p! A to the Pacific, William Clark spent three de- in as an Se i s US Government's Indian Policy E aes as an oe i a un as EE dace e Missouri and finally irs, Ironically, Clark's 1; J «1 ] 1 1 1 ] ] f Nati A th I ) Cl d ot] ted in tl E of the American West 5 i ich this thoroughly ae biography of Mn Clark. Jay Buckley d pon Clark’ ive par llasp E killed diplomat. R t li US t leaders, many Indian es would 1 11 endorsed by S M MS Lee Hinds, Rio Brazos Master Naturalist and BRIT volunteer, Fort Worth, Texas, U 5. A. J. Bot. Res. Inst. Texas 2(1): 290. 2008 LECTOTYPIFICATION AND NOTES ON BACCHARIS RIOGRANDENSIS (ASTERACEAE: ASTEREAE) Gustavo Heiden Angelo A. Schneider B. Sc, CNPq-PROTAX Fellow M. Sc, Programa de Pós-Graduação em Botánica Escola Nacional de Botánica lon da Universidade Federal do Rio Grande do Sul de Pesquisas Jardim Botánico ei calves Pacheco Ledo, 915, Jem Botánico 9500, Porto Alegre, RS 91501-970, BRAZIL Rio de Janeiro, RJ 22460-030, BRAZIL angeloschneiderayahoo.com.br gustavo.heidenggmail.com ABSTRACT Baccharis riograndensis Malag. & J.E. Vidal, an endemic species from southern Brazil, was described based on two syntypes and two p sl EE no See lesig 1. Study of original material and j | f other herbari ial fi e 1 LS ypifi ] 3 dde: 1 data, including a A: g rie d if ti , illustra- tions, speci itations, geographi g p and IUCN (2001) status of conservation are presented. RESUMO Baccharis riograndensis Malag. & J.E. Vidal, pé lémica do sul do Brasil, foi descrit I loi I E p Em] pécie tenha sido citad avi balhos, ] ] ipificação foi feita até to. O estudo do material original em conjunto com a revisão de coleções em herbári fi individualidade do táxon. Baccharis riograndensis é lectotipificada e dad lici is. incluind d icáo di g , ilustracóes, distribuica geog ab lod 7 g d da IUCN, sao apresentados. Baccharis riograndensis Malag. & J.E. Vidal, an endemic species from Rio Grande do Sul State, in southern Brazil, was described by Malagarriga (1949, under his monk name Irmáo Teodoro Luis) based on two speci- mens collected by J.E. Vidal (syntypes) and two collections (paratypes) made by Malagarriga himself. After the publication, accounts of this species appeared in the taxonomic literature several times (Malagarriga 1957, 1958, 1977, Barroso Mid Diesel 1987; and Oliveira et al. 2007), but no lectotypification was made. The study of the synt at Herbarium R, Museu Nacional do Rio de Janeiro, along with general collections de various herbaria: confirmed this taxon as distinct and allowed a choice of lectotype. Along with the lectotypification, additional data ted, including a hological description, illustration, specimen citations, geographic distributional map, and IUCN (2001) status a conservation. Baccharis riograndensis Malag. & J.E. Vidal, Bol. Inform. Inst. Geobiol. 1:13. 1949. (Fig. 1). Ts: BRASIL. Rio GRANDE DO SuL: Santa Maria, Boca do Monte, Mar 1939, +, J.E. Vidal 37006 (Lecrorvrz, designated here: R!). Diagnostic description. —Subshrub, 25—65 cm tall. Stems mostly erect, sometimes short reptant and decum- bent, 3-alate, indument of clavate uniseriate hairs in tufts, wings of vegetative branches undulate, 0.5-5 cm long, 0.25-0.75 cm wide, fertile branches distinctly winged, wings narrower, 0.1-0.2 cm wide. Leaves reduced to tdi 0. a O. a cm ong, 0.1-0. o cm wide. Capitulescence a panicle of pseudospikes, 5-25 cm long, lateral to solitary capitula, not arranged in glomerules. Mal itul involucre campanulate ER 5—5.5 mm long, 3.5-4 mm wide, phyllaries in 4-5 series, the outer oblong: the mediums ovate, the inner short lanceolate, apex obtuse, margins narrowly hyaline, entire to short toothed apically. Flowers 18-30 [40], pappus 4-5 mm long, uniseriate, 20-22 bristles, corolla 3-5 mm long, apically 5-lobed, tube 2.8-3.3 mm, throat 0.2-0.3 mm, lobes 1.5-2 mm, style 4-5 mm long, style apex not divided. Female capitulum: clinanthium alveolate, bearing biseriate hairs, involucre cylindrical, 7-10 mm long, 2-3 mm wide, phyllaries in 4—7 series, the outers short ovate becoming more linear onwards the innermost series, apex acute, margins narrowly hyaline, entire to short toothed apically. Flowers 30-40, pappus 7-8 mm J. Bot. Res. Inst. Texas 2(1): 291 — 295. 2008 292 ai g a y VE s E MIT PW va fs e d "Fide, 1 E A | US A já : d " a, t EI K ? 3 Ki a E i. 1 4 Ñ B BUE Si V * é E 1 j MERIT FI ME d E Y EI eram. Teac ux m MM ONE. RO mem nm Ve tnu MT eget NT d aie p ee SE dme ATP me at; ai A a j 1 SH DEE Zi : £z [| || A ,B EI Lay E A G. eg 648 (À, Cand D); G. Heiden 649 (B, Eand F). KP ^ si Fo j ' SL, d E. MEC 1m] A AÑ $ ^ y C | M A "à 5 : d Nef + H E Wd e CN Th d F n Famale flower E Male canitulum. E Male flower long, uniseriate, corolla 5-7 mm long, ligulate with the ligule 5- toothed, style 7-7.5 mm long, cypselas 2.5-4 mm long, cylindrical, 12-20 ribbed, surface few papillose and glabrous. Chromosome number 2n - 18 (Heiden et al. 2006). Etymology.—The name of the species refers to the type localit y gy P yp y Vernacular names.—carqueja, chirca "EM Phenology.—Fertile between December and March; dispersion of cypselas may last until June Geographic De A geen from Ges state al e Mapas Ha e Brazil dea sch found in dry or rodiy ns ds mainly in ES southern half of Geen state tel DH sites in vibe Pampa bone. Fic. 2. G hi f Bnrcharis ri lensis (Ast Act Yin Ria 6 le do Sul State. southern Brazil - LI A F F F and sparsely in the northern half (islands of discontinuous grassland areas on plateau sites inserted in the Atlantic Forest biome). Increasing land use and planned forest projects may disrupt the habitat of the spe- cies. Due to the anthropogenous pressure on the south Brazilian subtropical grasslands (campos) with the increasing of displacement of native vegetation in substitution to large-scale Eucalyptus and Pinus plantations, the species is E as Near Threatened (NT), concerning the IUCN (2001) status of conservation. Comments.—Baccha lensis is assigned to subgen. Molina Heering and sect. Caulopterae DC., to the latter mainly die: to estado of winged stems, clinanthium with glandular hairs, and pappus bristles of female flowers that are enlarged basally and fused into a ring. Seege lo aq pl is SE in its entire limited range to Baccharis crispa Spreng., which has a wider distribut l area in sout South America. Similar characteristics to the both species are the similar habitat, erect branches with undulate wings, involucre of male capitula campanulate and uniseriate pappus in male and female flowers. Baccharis riograndensis differs from B. crispa due to the fertile branches distinctly narrow-winged (vs. not distinctly winged fertile branches), lateral spikes reduced to solitary capitula (vs. lateral spikes fully developed or 294 t tani titute of Texas 2(1) with reduced axis seeming glomerules), involucre of female capitula cylindrical (vs. campanulate) and larger cypselas, 2.5-4 mm long (vs. smaller, 1-1.5 mm long). Another taxon, similar to B. riograndensis, is the alopatric Baccharis genistelloides subsp. lorentzii Joch. Mull., from central Argentina, which shares some characteristics with B. riograndensis, such as the pseudospikes with the lateral spikes generally reduced to one capitulum, cylindrical involucre of female capitula and larger capitula and cypselas, but differs on the basis of its erectopatent branching (vs. erect), usually narrower capitulescences (vs. wider), fertile branches not distinctly winged (vs. distinctly narrow-winged fertile branches), fewer cypsela ribs (9-12 vs. generally 12-20) and multiseriate pappus (vs. uniseriate). Representative specimens: BRAZI Arroio dos R F da Faxinal, 10 Mar 1977, K. e 10967 (ICN); Fazenda Faxinal, 13 Feb 1978, K. eye 12057 (ICN). Bagé: BR-153, km 84, 1 June 1985, M. Sobral & C. Grabauska 3920 (IC Ni Casa de Pedra, 12 Apr 1991, M.R. Ritter 612 (ICN); comeco do Passo do Enforcado, 70 km NE de Bagé, 5 Mar 1981, J.C. Lindeman et al. 6997 eos Ge e EE 21 Mar 2001, V. Marin 20 (CNPO); peus sobre o oe bes RS-153, 28 Mar 1985, O. Bueno 4312 (HAS). 0 Mar 1982, J. 2 Bagé, 10 Mar 1982, J. Mattos Les) E N. Mates 23630 (HAS). Caxias do Sul: Ana Rech, 12 Feb 2000, A. Kegler 678 (HUCS). Cachoeira di Sul: Cerro dos Peixoto, 1 Apr 1985, M. Sobral 3783a (ICN). Canoas: 16 Feb 1949, Irmáo Teodoro Luis s.n. (ICN 17068, 17069). Gees BR-116, km 32, 13 Mar 1993, NI Matzenbacher s.n. (ICN 53681). Osório: Faz. do Arroio, 14 Apr 1950, B. Rambo s.n. (PACA 46 de: ant. Fazenda Capivarita, 26 Feb 1990, G. Grazziotin et al. s.n. (HUCS 6591). Pelotas: Cerrito Alegre, 8 Mar ia O. Pereira s.n. (ICN 1469). Piratini: Fazenda São João, 8 Apr 1991, R. Wasum et al. s.n. (HUCS 7575, 7576). Portão: 17 Mar 1986, S. Diesel s.n. (HUCS 3750). Porto Alegre: Jardim Botánico, 11 Mar 1980, O. Bueno 2193 (HAS); Jardim Ingá, 4 Apr 1992, L.T. Pereira 1 (ICN); Morro das Abertas, 26 Mar. 1980, O. Bueno 2281 (HAS); [idem]. 08 Apr 1980, O.Bueno 2337 (RB); [idem], 26 Mar 1980, O.Bueno 2281 (RB); Morro do Osso, 25 May 1992, L.T Pereira 14 (ICN); Morro Santana, 16 Mar 1967, A.G. Ferreira 139 (ICN). Santa Maria: Pus do Monte, Mar 1939, c, J.E. Vidal 37004 (R): 24 May 1960, Irmão Teodoro Luis s.n. ae 876, 884). Sáo Francisco d do Umbu, 1 km, 12 Mar 1994, L.T. Pereira 96 (ICN); Fazenda Mulita, 9 Apr 2001, R. Wasum 1049 (HUCS). Sao Gabriel: 73Km da saída de São Gabriel, 27 Mar 1985, O. Bueno 4277 (HAS). São José dos Ausentes: Silveira, Fazenda do Tabuleiro, 18 Mar 2002, R. Wasum 1405 (HUCS). São Leopoldo: ad montem Sapucaia, 09 Mar 1949, B. Rambo s.n. (PACA); Quinta, 10 Mar 1937, Ar] Dutra 1536 (ICN). Sáo I co do Sul: Passo dos Baios, 25 Feb 2004, G. Heiden 421 (HECT, PEL); [idem], Mar 2006, G. F 48 (RB); G. Heiden 649 (RB). Sao Sebastiáo do Cai: Vila Rica, 9 Mar 1985, S. Diesel s.n. (PACA 68069). Sapucaia do Sul: Morro Sapucaia, 23 Apr 1979, O. Bueno 1295 (HAS). Taquara: 07 Apr 1957, J. Mattos 6803 (PACA). Tramandaí: Parque Osório, 04 Mar 1994, L.T. Pereira 82 (ICN). Viamao: J. Mattos 2249 (HAS). Selected additional specimens Baccharis crispa Spreng. URUGUAY. Monteviveo: 1821-1822, <7, Sellow d397 (isorYPE: R). Baccharis genistelloides subsp. lorentzii Joch. Mull. ARGENTINA. Buenos AIRES: "Sierras Pampeanas,” Arroyo Pigue, 22 Mar 1881, Y, Lorentz s.n. (SP 25365; 25368 ACKNOWLEDGMENTS Authors acknowledge the herbaria CNPO, HAS, HASU, HECT, HUCS, HURG, ICN, PACA, PEL, R, RB, SMDB, and SP; Guy L. Nesom and Jochen Müller for the helpful comments; and Joáo Ricardo Vieira Iganci for providing the illustration. REFERENCES Barroso, G.M. 1976. Compositae - Subtribo Baccharidinae Hoffman. Estudo das espécies ocorrentes no Brasil. Rodriguésia 28(40):3-273 Diese, S. 1987. Contribuição ao estudo taxonómico do género Baccharis L. (grupo Trimera) no Rio Grande do Sul. Pesquisas, Bot. 38:91- 126. HEIDEN, G., J.R.V. IcaNa, V. Stein, and V.L. Boenowski. 2006, Número « 5mico de Baccharis riog lensis Malag. & J.E. Vidal (Asteraceae). Pesquisas, Bot. 57:121-136. INTERNATIONAL UNION FOR CONSERVATION OF NATURE AND NATURAL Resources (IUCN). 2001. IUCN Red List categories and criteria, Version 3.1. IUCN, Gland, Switzerland and Cambridge, United Kingdom. http://www.iucn.org MALAGARRIGA Heras, R.P [IRMÃO Teopono Luis]. 1949. De*Re-Botánica”. Bol. Inform. inst. Geobiol. La Salle 2:17 MALAGARRIGA Heras, R.P [IRMAO TEoDORO Luis]. 1952. Index Baccharidinarum (Compositae). Contr. Inst. Geobiol. La Salle 2:1 MALAGARRIGA Heras, HP [lamAo Teoporo Luis]. 1957. Para o estudo da flora Sul-Riograndense qual o valor da “Flora Brasiliensis" de Martius?. Contrib. Inst. Geobiol. La Salle 8:1-61. | Schneider, Lectotypification of Baccharis ri lensi 295 ei wë MALAGARRIGA Heras, R.P [IRMAO Teoporo Luis]. 1958. Novum index baccharidinarum. Contr. Inst. Geobiol. La Salle MALAGARRIGA HERAS, R.P. 1977. Nomenclator baccharidinarum omnium. Mem. Soc, Ci. Nat. La Salle 37:129-224. MUüLLER, J. 2006. Systematics of Baccharis (Compositae-Astereae) in Bolivia, including an overview of the genus. Syst. Bot. Monogr. 76:1-341. OLivEIRA, A.S., LP Desle, AA ScHNeIDER, and J.N.C. Marchiori, 2006. Checklist do género Baccharis L. para o Brasil (Asteraceae-Astereae). Balduinia 9:17-27. 296 t tani tit Texas 2( BOOK REVIEW ANTONY Wi, 2005 (American Edition). Coffee: A Dark History. (ISBN 0-393-06071-3, hbk.). WW. Norton & Company, Inc., 500 5^ Avenue, New York, New York 10110, U.S.A. (Orders: www.wwnorton.com, 1-800-233-4830, 1-800-458-6515 fax). $25.95, 323 pp., 8 plates (7 color, 1 b/w), 6 3/8" x 9 1/2”. pia OR dn pn s Wes We Live Now, Ge se 3) Enter the depu 4) Wach piu Trade, 2 ea ie uud e St Helena, 10) Eus Brazil, and Coffee, 11) The Great Exhibition, 12) Harrar and Rimbaud: the Cradl 1 the Crucibl 3) Modern Times, 14) Coffee, Science, and History, ei The Site e the e Hemispheres di Fair Trade, 17) Espresso: the Esperanto i Coffee, 18) The Heart of Darkness. Coda, Appendix: Index Coffee: e: AD Dark JA History begir itl d d hift t tright di ] J tl fi t ł t t Here, author Antony Wild ZE É EE? of MN Third World coffee g E in the greedy clutches of the world's ad vanced, mega lt in tl d chapter, I was delighted when Wild turned 3 ex tlh é r £t, 1 e TIA. | 1 1 1 : ] 1 + mf] ` Amd LLL Tux MES, LLL fa) becht É eer d Pi et d 4 H | brings in some very int ff ] ibilities. His jumping off point fi ] ing is the supposed origins of humans and coffee in Me area now know as Spe Taking bu in a little farther along, | puts forth coff the forbidden fruit of tl Garden of Eden g yt I ibl d lti ] ] y j ] ly as the 15th century. LI B 1 4 E 1/ = “E fe + 1 t ce. 1 "m g the way. The first two-thirds of the book continues and expands on the role of coffee in political struggles, colonialism, imperialism, economics and culture, and social development. Botanical and other scientific tidbits are always present. This book is a great way to read through world history as a very organic, O ud Re last pun of the book resumes with the late 18' century and takes you eres to ps id gh tl y ghout, its evilness, according to Wild, has achieved epic A 1 A oe J +1 Ae. | +1 FE J Tad 1F 1 E E i i Lei oO r A ce T A D Geh 1 ] : 1 ] le wht] sea 1 + Fada A FF, T^ Ite FE Ya | le Dose c r A Ge x F "E 1 ly intriguing treatise — Robert J George, Botanical Research Institute of Texas, 500 East La has written a fascinating, well-researched, 4th Street, Fort Worth, Texas 76102-4025, U.S.A. J. Bot. Res. Inst. Texas 2(1): 296. 2008 NEW SPECIES OF MYRTACEAE FROM ECUADOR Bruce K. Holst Maria Lúcia Kawasaki Marie Selby Botanical Gardens Department of Botany h Palm Avenue ir Museum of Natural History Sarasota, Florida 34236, U.S.A. 00 South Lake Shore Drive bholst@selby.org Trin Illinois 60605-2496, U.S.A. lkawasaki@fieldmuseum.org ABSTRACT Four new species of Myrtaceae from Ecuador are described and illustrated: Calyptranthes websteri, Eugenia concava, Eugenia grossa, and Myrcia subcordifolia RESUMEN Se describen y se ilustran cuatro especies nuevas de Myrtaceae del Ecuador: Calyptranthes websteri, Eugenia concava, Eugenia grossa, y Myrcia subcordifolia. INTRODUCTION In preparation for the treatment of Myrtaceae for the Flora of Ecuador, four new species have been recog- nized and are here described and illustrated: Calyptranthes websteri, Eugenia concava, Eugenia grossa, and Myrcia subcordifolia. Calyptranthes websteri B. Holst & M.L. Kawasaki, sp. nov. (Fig. 1). Tee: ECUADOR. PrcmncHa: Canton Quito, Par- roquia Nanegal, Bosque Protectora Maquipucuna, Montafias de Maquipucuna, rain forest on main ridge of Cerro Sosa, ca. 7 km airline SE of Nanegal, 2150 m, 00?04.5'N, 78°36.5’W, 8 Jul 1992 (fl), G.L. Webster, E Hrusa & C. Zurinsky 29164 (HoLoryrE: DAV; ISOTYPES: AAU, SEL} Arbor, ind t inute dil hiat bello; foliis 13.6-20.2 x 6.1-9.4 cm, llipti i l levit ti lol tis, caudatis, venis mediis supra concavis, venis marginalibus 2-6 mm remotis; petiolis 9-13 mm longis; paniculis multifloris, pedunculis et pedicelis longis et E E Tal ez rm fia 1 "E m longis C LN Tree to lO m tall or UE tree; MONS dibrachiate, minute, with very short arms, reddish, abundant on new growth; | l htly com presse d. Leaf blades g lly elliptic to very slightly ovate or obovate, 13.6-20.2 x 6.1-9.4 cm, a when dry, bearing — trichomes, these deciduous with age, olive green to brown on upper surface when mature, lighter yellow green on lower surface, im- mature leaves with slightly yellowish grey pallor due to waxy EN Je caudate, Lu tip 7-19 mm long. 1. +] margins slightly revolute; base attenuate to te; midvein impressec prominent on lower surface; lateral veins 19-26 pairs, flat on upper and convex on lowe surfaces, princi- pal marginal vein 2-6 mm distant from the edge, slightly less prominent than the lateral veins, secondary marginal vein ca. 1 mm from edge, faintly evident; glands scattered, translucid on both surfaces; petiole 9-13 mm long, ca. 2 mm thick, margins inrolled and meeting to form a tunneled petiole. Inflorescences paniculate, decompound, many-flowered, 2 or 4 borne together on abortive branches, when young, densely covered by a light gray-tan waxy bloom over the small, appressed trichomes; bracts early deciduous, ca. 8 x 1 mm, peduncles 1-5 cm long, the first node bearing 3 or 5(7) branches and upper nodes bearing usually 3 branches, all branches from me m in same plane, the branches terminating with 3 long-pedicellate flowers, the terminal tive; pedicels 18-37 mm long; bracteoles early deciduous, 3-4 mm long. Flowers: buds globose, ca. 5 x 5 mm, with a light gray-tan waxy bloom, particularly on lower 2/3 of bud; calyptra ca. 3 x 5 mm, generally lighter in color than hypanthium, apiculum barely evident; petals lack- ing; stamens numerous; filaments ca. 6 mm long; anthers ca. 0.5 mm long; style ca. 5 mm long; hypanthium ca. 4 mm long, the free portion ca. 2 mm long; ovary 2-locular, ovules 2 per locule. Fruits not known. J. Bot. Res. Inst. Texas 2(1): 297 — 303. 2008 I E .£al n.a » In LI d T den e £Tawae M) Vt 7 298 J Fic. 1. Calyptrantt bsteri B. Holst & M.L. Kawasaki: A. Habit. B. Flower. C. Digital ffi buds. (A—C, Webster et al. 29164). er Distribution.—Known only from Maqui Pichincl ince, in primary rain forests at 2100-2150 L E iu m elevation. Calyptranthes websteri is distinguished from all other Andean species by the elongate peduncles and pedicels of the many-flowered inflorescence. This very distinctive species is named in memory of its dis- coverer, Grady Webster, an astute student of the Euphorbiaceae and respected teacher at the University of California, Davis, who passed away in 2005. A A 1 11 ga : l. ESITTATA AD Tas Us 2 y P ” de Maquipucuna, Cerro Sosa, primary rain forest on main ridge, 2100-2150 1 m, 002051 N, 78037 W, 9 Jul 1991 (fb, G L Webster et al. 28884 (AAU, DAV, MO, QCNE Eugenia concava B. Holst Sr M.L. Kawasaki, sp. nov. (Fig. 2). Tr: ECUADOR. Guavas: Guayaquil, Bosque Protector Cerro RS road Guayaqui il-Salinas, Km 15, 200-300 m, 02°10'S, 80°08'W, Jan 1994 (fl), T. Núñez & A. Hernández 197 (HOLOTYPE: 1 4 x : Js Ê [1 21] r E 1 Frut l arbor; folii | llipticis, chartaceis, glabris, a Se Shrub or tree, 2.5-10 m tall, mostly glabrous; twigs smooth, trunk and older branches with bark peeling in plates or strips. Leaf blades mostly elliptic, less commonly ovate or obovate, chartaceous, 4—7.5 x 2.5-4 cm, drying olive green or greenish brown; upper and lower surfaces glabrous; glands punctiform, plane or slightly salient on both surfaces; midvein convex on the upper surface; lateral veins 10—12 pairs, salient on both surfaces; in ben veins 2, the innermost slightly arched, 13 mm from the margin; base usually obtuse, rarely tly acuminate, the acumen ca. 1 cm long; petioles 2-3 mm long, flattened, glabrous. Infloreseenees — ne fasciculate, with up to 4 flowers, axillary or subterminal, the pedicels 1-1.5 cm long, slender, glabrous; bracteoles ovate, ca. 1 mm long, ciliate, persistent. Flowers 4-merous, buds ca. 5 mm long; hypanthium ca. 2 mm long, glabrous; calyx-lobes ovate to lanceolate, 3-4 mm long, obtuse, concave, glabrous; petals suborbicular, ca. 4 mm diam., ciliate or glabrous; disk ca. 3 mm diam., glabrous; stamens ca. 5 mm long; style ca. 6 mm long; ovary 2-locular, ovules several per locule. Fruits berries, el- lipsoid, 1-1.5 x 0.8-1 cm, glabrous, red, gland-dotted; seed 1, ca. 9 x 6 mm, the seed coat membranous; embryo eugenioid, the cotyledons dd connate, the SE indistinct. Distribution — Eugenia from ince, occurring in dry forests at 200-500 FL ALII E. ce P. 4 Le m elevation. Eugenia concava is characterized by small, elliptic, glabrous leaves, abruptly acuminate at apex, the midvein convex on the upper surface; the inflorescences are fasciculate, with slender pedicels and flowers with concave calyx lobes. The pulp is characterized as "sweet" by Palacios & Rubio (9977). Eugenia concava resembles E. rhombea Krug & Urb., a species from Southern Florida, Mexico, Central America, and West Indies (McVaugh, 1989), but differs by the leaves with distinctly convex midvein (vs. concave), lacking a distinct cartilaginous leaf — (vs. present), and by the ellipsoid fruits (vs. globose). Additional coliections examined: ECUADOR. Guayas: Bosque Protector Cerro Blanco, 350 m, 02°10'S, 79°58'W, 16 Jan 1994 (FD, X. Cornejo & C. Bonifaz 1387 (SEL); Pai Bosque Protector Cerro Blanco, 15 km W of Guayaquil, near Quebrada ME 230 m, 02*10'S, 79*58"W, 27 May 1996 (fr), D. Neill & T. Núñez 10474 (F, MO, SEL); Guayaquil, Cerro Blanco, coastal road, Km 15, 350 m, 02º1'5, 79°58'W, 16 Mar 1992 (fr), W. Palacios & D. Rubio 9944 (MO, SEL); Guayaquil, Cerro Azul, coastal id Km 12, 500 m, 02°10'S, 79°58'W, 18 Mar 1992 (fr), W. Palacios & D. Rubio 9977 (F, MO, QCNE, SEL); Guayaquil, B Bl , road to Salinas, Km 15, 400 m, 02?10'S, 79°58'W, 21-25 Jan 1992 (fl), D. Rubio & G. Tipaz 2321 (MO, SEL); ari Bosque Protector Cerro Blanco, road to Salinas, Km 15, 400 m, 02?10'S, 79°58'W, 21-25 Jan 1992 (fl), D. Rubio & G. Tipaz 2348 (MO, SEL). Eugenia grossa B. Holst & M.L. Kawasaki, sp. nov. (Fig. 3). Tre. ECUADOR. Carcut: Tulcan Canton, Parroquia Tobar Donoso, Reserva Etnica Awa, Sector Sabalera, 650-1000 m, 01°00'N, 78°24'W, 18-28 Jun 1992 (fl, fr), G. Tipaz, J. Zuleta & N. Guanga 1370 (Hororvre: QCNE; isotypes: E MO, SEL Arbor, ind t | foliis 27-45 x 19-25 cm, ellipticis vel ovatis vel obovatis, venis mediis supra concavis, venis margi- nalibus 12-21 mm remotis; racemis 15-21 mm longis, 6-floris, pedicellis 1-4 cm longis, floribus ca. 13 mm lon Tree 6-15 m tall, pubescent to puberulent on new growth. Leaf blades elliptic to broadl y elliptic, chartaceous, 21—45 x 19-25 cm, brown on upper surface, pubescent along veins, the lower an ee evenly pubescent, trichomes short and golden brown; glands indistinct above, dark and numerous below; midvein broadly and shallowly concave on upper surface, all veins convex and prominent on underside (midvein extremely prominent), lateral veins 11-16 pairs, first lateral (or secondary) vein extending to form inframarginal vein and straightening out; marginal veins 3, the inner, most conspicuous vein 12-21 mm from blade margin, the middle marginal vein 2-6 mm from margin, and a 3rd inframarginal vein, very faint, ca. 1 mm from margin, apex acuminate; base rounded to obtuse to very slightly cordate; petioles 13-20 mm long, 4-6 300 t tani i exas 2( | J E.’ fm A ~£ * | a [| LN | £. £l Fic. 2. É g i R Holst £ ME Kawasaki: A. Part of st itl g D. Fruit. E. Embryo. F. Digital ffl t anthesis (A-B, Neill & Núñez 10474; C, Rubio & Tipaz 2348; D-E, Palacios & Rubio 9977; F, Nuñez & Hernández 197). mm diam., densely pubescent with golden trichomes. Inflorescences racemose, the axis 15-21 x 3-4 mmi, pubescent, often at leafless nodes, 3 pairs of pedicels per peduncle; bracts deciduous; bracteoles fused and cupping hypanthium, 10-12 (14-15 hydrated) mm long, pubescent inside and out, deciduous after anthesis, pedicel in flower ca. 1 cm long, lengthening and thickening considerably in fruit, to 4 cm long. Flowers 4-merous, buds roundish, ca. 13 mm long; DEE E pubescent inside and out, ca. 6 mm wide at summit, hypanthium after removal of bracteoles htly longer trichomes, having a hint of red or dark orange, apparent wrinkles or bumpy Geen ome a which corresponding to marks on inner surface of bracteoles); outer calyx lobes suborbicular, ca. 9 x 9 mm, the inner lobes more closely ovate, ca. 10 x 7 mm, the tip acute (with triangular or acuminate shaped discoloration or mark, perhaps where 301 3 E 3 DISSE [ 10 cm Au, Cie, J arca ve Un i E + E-. » ett "e y Ê E eum s aet) , Stamen, petal. D. Digital scan of flower igma . Flower bud, style and st C ts. TIS CS 3 D = te" » E ? P t ? ENT. d tem wer? — a CA neben Tx) os re NS - Ha: E E arian’ EN EU Zn v + + d 3 hs MEA € REPOS: KEE D d » el = phe a, « Branch habit. B. Young fru Kawasak Fic. 3. Eugenia grossa B. Holst & M.L buds. (A-B, Aulestia et al. 830; C— Tipaz et al. 1370). D, 302 mal of t tani ti Texas 2(1) Fic. 4. Mvrcia subcordifolia B. Holst & M.L. Kawasaki: A. | infl B. Flower bud. C. Medial section of D.C tion of E Fruit. F. Digital scan of flower buds. (A-C, Palacios 6723; D—F, Quizhpe et al. 145) trichomes grow in a different direction, possibly where bracteoles lobe rests), the margin more or less entire; petals cream color (live) with plentiful glandular dots, ciliate, rounded eliptical, 16-21 x 11-12 mm, the tip more or less obtuse; stamens numerous (approaching 100), the anthers ca. 1 mm long, the filaments 17-20 mm long. Fruits berries resembling drupes, oblong, 13-28 x 8-16 mm, the hairs clustered in groups, black at maturity; seed 1, mature size unknown, the seed coat membranous; embryo eugenioid, the cotyledons fleshy, connate, the radicle indistinct. Distribution.—Known only from Carchi province (Reserva Etnica Awá), in primary rain forests and premontane forests at 650—1800 m elevation. Eugenia grossa is characterized by the very large leaves (27-45 x 19-25 cm), brown to yellowish brown- TEN lie ki, N E a ra E [ 303 pubescent and large flower buds (ca. 13 mm long), with connate, deciduous bracteoles. Tipaz et al. (1370) describe the fruit as "edible" and note the wood is used to make tool handles and walking canes. Additional collecti ined: ECUADOR. Carchi: Mald lo, Parroquia Tobar Donoso, Reserva Etnica Awá, Sector Sabalera, 900 m, 00°55'N, 78°32'W, 22 Nov 1992 (fr), C. Aulestia et al. 830 (MO, QCNE, SEL); Tulcan, Parroquia Tobar Donoso, Reserva Etnica Awá, Centro El Baboso, 1800 m, 00°53'N, 78°25'W, 17-27 Aug 1992 (fr), C. Aulestia et al. 1817 (MO, QCNE, SEL). Myrcia subcordifolia B. Holst & M.L. Kawasaki, sp. nov. (Fig. 4). Teee: ECUADOR. Zamora-C Nangaritza Cantón, valley of Río Nangaritza, Miazi, 04°18'S, 78°40'W, 1200 m, 10 Dec 1990 (fl), V 672 OCNE; E MO, SEL) Arbor; foliis suborbicularis usque ellipticis, coriaceis, glabris, saepe subcordatis; paniculi ltifloris; alabastris 2-3 mm longis. Tree 4—10 m tall, mostly glabrous, the few trichomes on inflorescences and flowers yellowish to yellowish white. Leaf blades suborbicular to elliptic, or ovate, 4—9.5 x 3-6 cm, coriaceous, plane, glabrous, drying dark brown to brownish above, paler below; apex obtuse to rounded; base rounded or broadly subcordate; midvein impressed above, below; lateral veins 7-10 pairs, slightly convex on both surfaces; marginal veins 2, the innermost 1-2 mm from blade margin, similar to the lateral veins; glands numerous, punctiform, indistinct or impresse d above, salient below; petiole 1-3 mm long, channeled, puberulous to glabrous, black. fullorestenes paniculate, bel multiflorous (approx. 20—60 flowers), 6-12 cm long, puberulous; bracts and bracteoles not seen, early deciduous. Flowers 5-merous, buds obovoid to subglobose, 2-3 mm long; calyx-lobes suborbicular, to 1 mm long, appressed-pubescent to puberulous without, glabrous within; petals suborbicular, ca. 2 mm diam.; stamens numerous, the filaments ca. 4-5 mm long, the anthers ca. 0.5 mm long, style ca. 5 mm long, the stigma punctiform; EE not EE E the ESCH sericeous without; disk ca. 2 mm diam., hirsutulous; ovary 2-locular, 2 I (immatur e) berries, ellipsoid, ca. 5 mm long, glabrous; seeds not seen. Distribution.—This species is known from Zamora-Chinchipe and Morona-Santiago, in the Cordillera del con region i it occurs in i primary, pea wet forests, at 900—1200 m elevation. th) DC., a species known from lowland, white-sand savannas from Venezidla Colombia, one m Brazil (Holst et al., 2003). In M. subcordifolia, however, the leaves are subsessile with petioles 1-3 mm long and plane (vs. petiolate with petioles 5-8 mm long, and the leaves convex-conduplicate), the lateral veins are slightly convex, and the veinlets are indistinct (vs. veins strongly convex and veinlets distinctly reticulate) on both surfaces. Additional collections examined: ECUADOR. Morona-Santiago: Limon Indanza, Cordillera del Céndor, Centro Shuar Yunkumas, Asociación po Cerro e mank Naint, poe d idi ubi) 1150 m, e Dec eid a C. SE et See 1595 Ge MO, QCNE). amora lel Có , W of river, 04°12'22"S, 78º 3916"W, 1100 m, 7 Aug 2002 (fl, yf, W. Quizhpe et ball 145 (F, MO, SEL); Nanos 900 m, 04°17'10"S, 78?38'50"W, 24 Jul 2003 (ED, W. Quizhpe et al. 638 (MO, SEL); El Pangui, Cordillera del NEM , W slopes, Machinaza river basin, 03°37'31"S, 78°31'50"W, 1050 m, 22 Mar 2006 (fl), W. Quizhpe & F. Luisier 2043 (F, LOJA, MO). Fr 4-1] As] Cóndor, ACKNOWLEDGMENTS We thank the herbaria cited in the manuscript for providing the collections used in this study. Fred Bar- rie offered helpful suggestions to improve the manuscript. Illustrations are by Valerie Renard and Lucia Kawasaki. REFERENCES HoLsT, B.K, L. Lanprum, and F. Grito. 2003. Myrtaceae. In: J.A. Steyermark et al., eds. Fl. Venez. Guayana 7:1-99. Mis- souri Botanical Garden Press, St. Louis. McVauH, R. 1989. Myrtaceae. In: RA. Howard, ed. Flora of the Lesser Antilles, Leeward and Windward Islands 5:463-532. 304 BOOK REVIEW Grariro ALANIS FLORES. 2007. El Valor de Nuestras Plantas. (ISBN 978-970-9715-26-2, hbk.). Fondo Edito- rial de Nuevo León, Zaragoza 1290, bann Kalos, Nivel A2, Desp. 249, CP 64000, Monterrey, Nuevo León, Mexico. (Orders: s Inl.gob.mx, (81) 8344 2970 y 71). $150.00, 157 pp., color photographs, b/w line drawings, color paintings, 9" x 9”. Contents: Presentación Nuestras plantas oues del ánima anacahuita ¿Por qué comemos mahuacatas? La inundación de 1909 El sabino gordo de Terán La dominante gobernadora La deliciosa nieve de pitaya El preciado oro verde Cómo plantar un árbol qose d unge des Ferr] -— 1 plants. This TL x heen AE | £..11 1 1 1. £:11 1 1 1 a sa 1 MM comes Dom students in dee M dM ES id the state of ie León: Made en Pus dada native pane and uw i to draw men il I I yp el ro 1 r 1 + ] 41 : a Ti : H A ral + tr AT r I [e] A zd r servation al e H quc] E UM WT 7 D 4 1 + 1 +1 FS 11 Jit to E Pr I d :.1 E A e | qe BET f. +1 1 1 + T ] + a El r o L ful caen: grace the apposita 30 plant Species that are described, gege ith t their local distribution, methods and E for Fus acne and: their current and a uses. ie 11 t the illustrati y of Nuevo León. This enchanting call to Gs conservation of EspruollLs Lt > t k 1 A 3 1 1] r ] SS sciences. s— Tiana Franklin, Botanical Research Institute of Texas, 500 East 4th Street, Fort W orth, Texas 76102-4025, USA. J. Bot. Res. Inst. Texas 2(1): 304. 2008 CLARIFICATION OF BORRERIA GYMNOCEPHALA, DIODIA GYMNOCEPHALA, DIODIA SCHUMANNII, BORRERIA FLAVOVIRENS, AND SPERMACOCE SCHUMANNII (RUBIACEAE) Piero G. Delprete Joseph H. Kirkbride, Jr. Federal University of Goiás, Campus ll USDA-ARS, U.S. National Arboretum Institute of Biological Sciences - ICB- 1 Floral & Nursery Plants Research Unit Department of General Biology/Botany Washington, DC 20002-1958, U.S.A. 74001-970 Goiánia, Goiás, BRAZIL joseph.kirkbridemars.usda.gov pdelprete@hotmail.com ABSTRACT Several erroneous, eee and/or D names, e.g., B [4 phala, Diodia g) phala and Diodia scl ii, have been SE g I y many authors. Tl f all tl is | larifi d, d tl D L Key Wonps: Borreria, Spermacoce, Rubiaceae, Goiás, Tocantins, Brazil, Argentina, Neotropics RESUMEN A f. i ~ 1 Al eg D 1 1 T^ ] 1 1 Ty 1 1 a [o D Qe. É f 1 1 A "| 1 ES A -l i t = d Ania Eri aen ab als si d an À ¢ } ii (Standl. ex Bacigalupo) Delprete. I PALAVRAS CHAvE: Borreria, Spermacoce, Rubiaceae, Goiás, Tocantins, Brasil, Argentina, Neotrópicos INTRODUCTION The present work seeks to clarify the status of the incorrect, confused, and/or misapplied names [e.g., Bor- reria flavovirens Bacigalupo & E.L. Cabral, Borreria gymnocephala DC., Diodia gymnocephala (DC.) K. Schum., Diodia schumannii Standl. ex Bacigalupo] of taxa related to Spermacoce schumannii (Standl. ex Bacigalupo) Delprete (Rubiaceae, Spermacoceae). Candolle (1830, p. 549) described Borreria? gymnocephala DC., and was uncertain deet its generic position. He cited a specimen in his herbarium without giving the collector or tl ollection locality. After the description he stated that this species, collected in Brazil, is so distinct that it could even be treated as a separate genus and is cl terized by 4-angular, smooth stems between the capitate inflorescences and small Ronda with included anthers. Schumann (1888) transferred Candolle’s taxon to the genus Diodia as D. gj hala (DC.) K. Schum. After the description, he stated that it is very similar to Borers latifolia (Aubl) K. Schum. var. siderites (Cham. & Schltdl) K. Schum., from which it differed by the yellowish color of the vegetative parts and the fruit and style morphology. However, both Schumann's description and the specimens he cited apply to a mixture of two species. According to Bacigalupo and Cabral (1998), Se was ME d to notice that Schumann's Diodia gymnocephala suco Md two distinct species. He specimens EEN e to the part of D. gy la that differed from Candolle's B i hal B Bacigalupo (1974) ened this same sf volume 6, part 6 ofi the Flora Ilustrada de Entre Ríos (Argentina), and published Standley's name as “Diodia schumannii Standley ex Bacigalupo, nov. nom.” with Schumann's D. gy hala listed as a synonym and Candolle's B. gymnocephala specifically excluded. Article 69 of the ICBN (Stallen et al. 1972) in force at that time stated that “A name is to be rejected if it is used in different senses and so has become a long-persistent source of error.” Whether Bacigalupo intended to completely | Rat Rac Inet Tayac 2(1): 205 — 308, 2008 306 tanical h Institute of Texas 2( reject D. gymnocephala (DC.) K. Schum. or was simply creating a new name for D. gymnocephala sensu K. Schum. is not entirely clear in this Flora, since no further discussion of nomenclature was provided. Since the type of D. gymnocephala, i.e. Borreria gymnocephala DC., was explicitly excluded by Baciga- lupo (1974), the citation of this earlier name does not make D. schumannii illegitimate under current ICBN Art. 52 (McNeill et al. 2006). But, apart from having the correct Latin form, does this name meet the other requirements under Art. 32.1 for valid publication, namely provision of a Latin description or diagnosis (since 1935) and indication of a type (since 1958)? Bacigalupo (1974) provided only a Spanish description, but her reference to Schumann's name, for which a full Latin description was provided, is an acceptable "reference to a previously and effectively published Latin description" under Art. 36. In addition, under this species Bacigalupo cited only the single specimen “Dpto. La Paz, isla Curuzú Chalí, Burkart 27.103 (SD." When Bacigalupo (1974) published her new species, Art. 37 of the ICBN (Stafleu et al. 1972), then in effect, stated: "Publication on or after 1 Jan. 1958 of the name of a new taxon of the rank of family or below is valid only when the nomenclatural type is indicated ..." thus her citation would not then have satisfied this requirement and her new name was not validly published under the 1972 Code. It was this fact that motivated Bacigalupo and Cabral (1998) to later propose a new name, B. flavovirens Bacigalupo & E.L. Cabral, to replace Diodia schumannii Standley ex Bacigalupo, “nom. illeg.”, which had been published “sin seleccionar el tipo”, although in reality this shortcoming would have affected its valid publication and not its illegitimacy. However, these authors had overlooked the fact that already in the 1988 ICBN (Greuter et al. 1988) Art. 37 had been expanded considerably to include, in Art. 37.3, the following: "for the name of a new species or infraspecific taxon, citation of a single element is acceptable as indication of the holotype.” In the current ICBN (McNeill et al. 2006) this is now stated as: “for the name of a new spe- cies or infraspecific taxon, mention of a single element or gathering ..., even if that element is not explicitly designated as type, is acceptable as indication of the type." Since Bacigalupo's (1974) citation of a single specimen fulfills the requirement for a type indication, D. schumannii has been validly published under all editions of the ICBN since 1988. Thus B. flavovirens was at the outset a nomenclaturally superfluous and illegitimate replacement name for D. schumannii. In their 1998 paper, Bacigalupo and Cabral finally clarified the status of Diodia gymnocephala (DC.) K. Schum., which from their examination of the type of the basionym, B. gymnocephala DC. (1830), at G-DC they determined to be synonymous with Borreria palustris (Cham. & Schltdl.) Bacigalupo & E.L. Cabral, this based on the earlier Diodia palustris Cham. & Schltdl. (1828). A further point should be made with regard to the type of Diodia schumannii Standley ex Bacigalupo. Because this name was *validly published by reference to a previously and effectively published description or diagnosis,” that of Schumann (1888) as has already been noted, Art. 7.7 dictates that it “is to be typified by an element selected from the context of the validating description or diagnosis, unless the validating author has definitely designated a different type.” While Bacigalupo can be said to have satisfied the Art. 37 requirement for "indication" of a type, according to Art. 7.11 “designation” S a a EH the Meuse of Luna a] mat “the term ‘type’ or an equivalent.” So unless the term “Exsiccata” used by B o “type,” which seems unlikely, does this mean that the indication of type ner Art. 37 does not Poss, as Mi LN of a different type under Art. 7.7, such that D. schumannii must be typified on an element cited by Schumann? It is our interpretation that in this case an "indication" in the one case is equivalent to a "designation" in the other, such that the type of D. schumannii is that indicated by Bacigalupo in 1974. In conclusion, the species discussed above is maintained within the genus Spermacoce, as S. schumannii (Standl. ex Bacigalupo) Delprete, following the circumscription adopted by Delprete et al. (2005), Delprete and Cortés-B. (“2006” [2007]), and Delprete (2007). SYSTEMATIC TREATMENT Spermacoce En Ge ex Bacigalupo) Delprete in A. Reis, E Ilustr. pam RUBI 2:754, fig. 130. 2005. D dl. ex Bacigalupo in A. Burkart, Fl. Ilustr. Ent 6(6):15, fig. 5. 1974. Borreria flavovirens o Bacigalupo & E.L. Cabral, Hickenia 2: 29h; fig. 1. 1998, nom. mee n Typ: EH See Rios: o La Paz, Isla Curuzú-Chalí, 10 Apr 1968 (fl, fr), Burkart et al. 27103 ( S at NY! Diodia gymnocephala sensu K. Schum. in Mart., Fl. Bras. 6(6):16. 1888, p.p.; non Borreria gymnocephala DC., Prodr. 4:549. 1830 [- permacoce palustris (Cham. & Schltdl.) Delprete] Selected specimens examined: BRAZIL: Goiás: Serra do Caiapó, ca. 30 km (straight line), S of Caiapónia, 950-1200 m, 29 Apr 1973 (fl, fr), Anderson et al. 9363 (NY, UB); Serra Dourada, 16°3'S, 50?7"W, 920 m, 17 Mar 1989 (fl, fr), Cavalcanti et al. 461 (CEN, NY[(2]); Mun. Alto Paraíso, Chapada dos Mee: 10 km W of Alto Paraíso, 1000 m, 24 Mar 1969 (fl), Irwin et al. 24971 (NY, UB); Mun. Mos- sámendes, Serra Dourada, ca. 6 km NE of Mossámendes, 16%04'S, 50°11'W, 1000 m, 7 Feb 1980 (fl), Kirkbride 3293 (NY, UB); Mun. Niquelandia, A Ge 237 entre che e ana 14º21'S, 48°12'W, 470 m, 13 Apr 1992 (fl, fr), e et al. 1226 ES a ay 1973 (fl, fr), Anderson et al. 9 o m, 4 Lacan camp to mai See Cachimbo rd., km 264, 16 Nov 1967 (£D, Philcox et al. 3068 (NY). Minas Gerais: S do Espinhaco, ca. 15 km NE of Diamantina, on rd to Mendanha, 1300 m, 26 Jan 1969 (fl), Irwin et al. 22605 (NY); Mun. Formoso, Parque Nacional Grande Sertão Veredas, towards Faz. Diamante, 15°20'01"S, 45%56'54"W, 815 m, 29 Nov 1997 (fl), R.C. Mendonça et al. 3308 (IBGE NY); Mun. Diamantina, Fundo do Macaco, 1200 m, 16 May 1931 (fl, fr), Mexia 5849 (NY). Paraná: Mun. Morretes, Estr. da Graciosa, Alto da Serra, 12 Feb 1965 (fl), Hatschbach 12367 (US). Rio de Janeiro: Mun. Caxias, Capivarí, 30 Sep 1958 (fl), E. Pereira 4212 (HBR); Mun. Jacarepaguá, Restinga de Jacarepaguá, 10 Sep 1958 (fl), E. Pereira 4159 (HBR, NY). Santa Catarina: Mun. Lages, Alto da Serra, Encruzilhada, mata secundaria, 900 m, 7 Nov 1946 (fD, Reitz & Klein 12571 (HBR, NY); Mun. Lauro Müller, rd. SC-403, km 12, Novo Horizonte, property of Olavo Mariotti, 28°23'S, 49°28'W, 420 m, 5 Dec 1998 (fl), Delprete et al. 6986 (ABR, MBM, MO, NY, UB, US). São Paulo: Mun. Atibaia, 0.5 km N of Atibaia, 24 Sep 1960 (fl), Mattos & Mattos 8389 (NY); São Paulo, Avenida Paulista, campo, Oct 1913, Brade 6803 (NY). Tocantins: Mun. Couto de Magalhães, rd. Couto de Magalhães-Colinas de Tocantins (TO-335), 5-8 km from Couto de Magalhães, 170—200 m, 08°19'29"S, 49°09'25"W, 4 Nov 2005 (fl, fr), Delprete et al. 9267 (NY, UB, UFG); Mun. Araguaína, 15 km S of Araguaína, 300 m, 15 Mar 1968 (ID, Irwin et al. 21199 (UB). Distribution and ecology.—Widespread from Colombia t thern Brazil, Paraguay, and northern Argentina In Brazil, itis known from the states of Rondonia, Mato Grosso, Goiás, Tocantins, Minas Gerais, Maranhão, Bahia, and southwards throughout Rio Grande do Sul. Mostly found at the margins of and inside gallery forests, semi- EES forests, and slope dcn amd less frequently in seasonally flooded vegetation. Ta bservations.—S very similar to S. latifolia Aubl. in its flexuous, some- what climbing habit, and floral breriches with 5-13 glomerules, but it differs in its cylindrical to cupular, 3-8 mm long stipular sheath [vs. short, triangular, (1-)2-5 mm long in S. latifolia] and commonly yellowish tinge of the vegetative parts (vs. pale to olive green). ACKNOWLEDGMENTS This work was initiated by PD during a fellowship for a Visiting Sci t from the National Counsel of Techno- Development (Conselho N lde D lvimento Científico e Tecnológico - CNPq) ofi the Brazilian Government (grant 309885/2003-5), at the Federal University of Goiás (UFG), under the coordination of Vera Lúcia Gomes-Klein (UFG). We wish to thank Kanchi Gandhi (GH) and Dan Nicolson (US) for information about some of the nomenclatural problems involving this species and John Wiersema (BARC) and Guy Nesom (BRIT) for their helpful reviews of the manuscript. The directors and curators of the following herbaria are kindly acknowledged for loans of material, sending digital images, and/or providing working space during PD's visits: CEN, HBR, HTO, IBGE, NY, NX, PACA, UB, UFG, and UFMT. REFERENCES BAcIGALUPO, N.M. 1974. Diodia schumannii. In: A. Burkart, ed. Flora ilustrada de Entre Rios (Argentina). Tomo VI, parte VI, Dicotiledoneas Metaclamideas. Colección Cientifica BE I.N.T.A, Buenos Aires, Argentina. Pp. 15-16, fig. 5. BAciGALUPO, NM and E.L. CagnAL. 1998. Nota sobre dos especies de Borreria (Rubiaceae-5| ). Hickenia 2(56):261-266. CANDOLLE, A.P. DE. 1830. Borreria ? gymnocephala. Prodromus 4:549. Treuttel & Wurtz, Paris. 308 J i of tl tanical | Texas 2( CHAMISSO, L.A. von and D.FL. von SCHLECHTENDAL. 1828. De plantis in expeciti | vatis, Rubiaceae, Sectio Ili Spermacoceae. Linnaea 3:309-366, Da nere, PG. 2007. New combinations and new synonymies in the genus Spermacoce (Rubiaceae) for the Flora of Goiás and Tocantins (Brazil) and the Flora of the Guianas. J. Bot. Res. Inst. Texas 1:1023-1030 De.preTE, PG. and R. Cortés-B. 2006 [2007]. A synopsis of the Rubiaceae of the states of Mato Grosso and Mato Grosso do Sul, Brazil, with a key to genera, and a preliminary species list. Rev. Biol. Neotrop. 3:13-96. DeLpreTE, PG. L.B. Sura, and R.B. Kreis. 2005. Rubiáceas, Volume 2 - Géneros de G-Z: 20. Gardenia até 46. Tocoye- na. In: A. Reis, ed. Flora Ilustrada Catarinense. Herbário Barbosa Rodrigues, Itajaí, Santa Catarina, Brazil. Pp. 345-843. GREUTER, W, HM Burbet, W.G. CHALONER, V. DEMOULIN, R. Gig D.L. EE Ge bës SN SILVA, F. A. STAFLEU, E. G. Voss, and J. McNeLL. 1988. International code of bot ted by the teenth Internationa Botanical Congress, Berlin, July-August 1987. Koeltz Scientific Books, Konigstein. McNent, J., ER. Barrie, H.M. Bunper, V. Demoun, DI are » Moin dh REC e EEN PC. SEA "m es JH, Wiersema, and NJ. TurLAND, 2006. Int | teenth International Botanical Congress, Vienna, Austria, July 2005. A.R.G. Contner ventas: Ruggell, TANE SCHUMANN, K. 1888. Diodia gymnocephala (DC) K. Schum. In: CFP von Martius, AG. Eichler, and |. Urban, eds. Flora Brasiliensis 6(6):16-17. Fleisher, Leipzig Grapp, FA, C.E.B. Bonner, R. McVaucH, R.D. Mere, R.C. Rons, R. Ross, J.M ScHopr, G.M. ScHuLze, R. DE VILMORIN, and E.G. Voss. 1972. International code of botanical nomenclature adopted by the Eleventh International Botanical Congress, Seattle, August 1969. A. Oosthoek's Uitgeversmaatschappij N.V., Utrecht. ARISTIDAE ELUDENDAE II: A RE-EVALUATION OF THE ARISTIDA GIBBOSA COMPLEX (POACEAE: ARISTIDEAE), INCLUDING A. MARGINALIS, A. ORIZABENSIS, AND A. SORZOGONENSIS Robert T. Strahan and Kelly W. Allred Range Science Herbarium Department of Animal & Range Sciences New Mexico State University es, New Mexico 88003, U.5.A Piscis edu; kallredanmsu. edi ABSTRACT T 1 L: 1 ] Ll : 1 S eR Arictida ocihhnen Arictid d hi Á victid i } d and Aristida Pu L “a [e] a a pa] T bw 31 1 4 71 H D 14 Cal DO A: SKS La SE S 1 Je 4 A cibbnca a e e and. D EE E two taxa are e distinguished b y a a numbe: er getati d spikelet feat Ke A. Ge two taxa M DÉI Qd tod 1 that Do wd E e I + lated bet the A. gibbosa and A. is clust Aristid is f.orizal is (E 5. Fourn ) Strahan & Allred e a ek mb EE | gat Li 1 1 DT 1 E A x | is a new combination. A key to p g plex, I Į provided RESUMEN Las DUE taxonómicas a en fológi ntre Áristida gibbosa, A ee A. dedi A A. Moi cnn Do: s 1 Dr AY T le pl and distintos calacterivados por A. gibbosa y Fi sorzogonensis. m DEE entre estas dos especies o ea incluyen ca características ce Trernnnren taya a nivel SOTZOgOnens "forma" basados en el desarrollo de los pulvínu lis está tada por una sola colección (Typus) y se ubicó en E forma aislada entre los Pup la A. gibbosa yA. SE Aristida zog f. orizabensis (E. Fourn.) Strahan & Allred, E pecies del complejo Aristida gibbosa, descripciones breves y los vegetativas y también diferencias entre ps Dentro ES grup especímenes examinados. INTRODUCTION The genus Aristida I | 250—350 species, nearly worldwide in distribution, growing in warmer, more arid environments (Allred 2003). Members are characterized by a panicle inflorescence, one-flowered spikelets, and relatively large glumes, usually larger than the floret. The florets are characterized by three awns, the two lateral awns absent or much reduced in some species, and a sharp pointed callus with a tuft of hair at the base. In some cases the lemma body is drawn out into an elongated beak below the divergence of the awns. The genus has its share of taxonomically difficult species complexes. Members of such groups often differ only slightly morphologically and commonly overlap in their geographic distribution (Allred 1984a, 1984b; Longhi-Wagner 1990). The Aristida gibbosa complex is one such group. This New World complex is characterized by the fol- lowing two features: 1) floret sulcate, the margins of the lemma slightly overlapping, and 2) callus short, 0.3-0.4 mm long, the attachment scar circular and tiny, 0.2-0.3 mm in diameter (Fig. 1). New World taxa corresponding to these features have received the following names (in chronological order): Chaetaria gibbosa Nees (1829): In his ample description, Nees noted the regular branching of the culms, short lower sheaths exposing the gibbous nodes (whence the specific epithet), and folded or rolled leaf blades. Blade margins are not thickened, the marginal veins being no wider than those inward, and the adaxial surface is lacking pilose hairs except at the corners of the sheath. Panicles lack pulvini and are thus narrow and contracted. This is a South American species, and the name was unused in North and Central American works until Longhi-Wagner (1990) called attention to the sulcate lemma of numerous specimens J. Bot. Res. Inst. Texas 2(1): 309 — 322. 2008. 310 | | tanical h Instit Texas 2(1) at MO and US and annotated them as this. The name was then applied, in an expanded sense, to Meso-American plants by Pohl and Davidse (1994) to include Aristida sorzogon- ensis, A. orizabensis, and A. marginalis. Aristida sorzogonensis J. Presl (1830): Erroneously believing Haenke's specimen to be from Sorsogon on the island of Luzon in the Philippines, Presl applied the specific epithet sorzogonensis to his new species. It is highly probable that the specimen actually came from Central America or Mexico (annotation in 1921 on the type by E.D. Merrill; Henrard 1928). The type collection is represented by at least two sheets, both with a single flowering culm: the holotype at PRO) has three mid- to upper blades, but lacks the base of the plant or any basal blades; an apparent isotype at PR(!) has a culm with three upper blades and two mid- to lower blades. Presl described the blades as convolute-setaceous, which the upper are, but the middle and lower blades are flat (perhaps rolled a bit upon drying) in both specimens. Blade margins are noticeably thickened, with a wide band of sclerenchyma at the margin; this is easily visible even on the upper blades that are rolled. The adaxial surface has prominent pilose hairs upwards from the throat region. A close examination of the panicle branches and pedicels reveals small (perhaps young) pulvini in nearly all the axils, with the resultant slight spreading of the axes, a feature not noted “this species in any previous works. The name Aristida sorzogonensis has been largely ignored or unknown by nearly all North American botanists, even though Henrard recog- el, ue nized it in both his Revision and Monograph of Aristida (Henrard 1929, 1932). Pohl and paesi idse (1994) rightly called attention to the priority of j izabensis and rre “2 ) rightly called attention to the priority of sorzog over orizabensis an marginalis, but we believe they misinterpreted the typification in tentatively applying the name sorzogonensis to plants with rolled, rather than flat, blades Aristida liebmanii E. Fourn. (1886): Fournier's description came from a collection by Liebmann (whence the specific epithet) and noted the following distinctive features: basal blades are flat and the upper blades are rolled (much as in the type of A. sorzogonensis), and first glumes are longer than the second. He did not call attention to the nearly complete absence of an elongated beak on the lemma, the awns immediately diverging from the apex, but this was stressed by Henrard (1927), is easily observ- able in the type, and was re-emphasized by Pohl and Davidse (1994). An isotype at MOO) contains three Se Se and one basal portion. Tue margins of the blades are noticeably thickened, and the lower lat the base hat flexuous. Panicl pulvinate. Pohl and Davidse (1994) called attention to its similarity with A. succedeana Henr., a species of Bolivia and Brazil that also has sulcate lemmas, flat blades, and sub-flexuous panicle branches, with the suggestion that the two may be conspecific. However, because of the scarcity of specimens further analysis and conclusions concerning this species must await the availability of more material. Aristida orizabensis E. Fourn. (1886): Fournier’s description, though short, mentions flat blades (rolled upon drying), somewhat unequal glumes, and a narrow lemma that was slightly shorter or longer than the glumes. Material came from “Valle de Orizaba,” Veracruz, Mexico (whence the specific epithet), with Fournier citing two specimens, which are considered syntypes. Isosyntypes at NY() and USC) have flat, curling blades with thickened margins and with long pilose hairs on the adaxial surface near the ligule, and panicles lacking pulvini. Henrard (1927) contrasted A. orizabensis with A. arizonica Vasey and A. appressa Vasey, two species excluded from this complex that lack sulcate lemmas. Aristida marginalis Ekman (1911): This species was named from material collected along the dry, grassy edges of forests or woods (“in margine silvulae,” whence the specific epithet) in Matto Grosso, south- ern Brazil. The new species is very well described by Ekman. In type material (isotypes G!, US!), sheaths are about half as long as the numerous internodes with gibbous nodes; lower blades are flat, curling, and thick-margined with pilose adaxial surfaces; the panicle is obscurely and incompletely pulvinate Oe. with poorly developed pulvini in the axils of some branchlets and pedicels); and the lemmas are sulcate with a scar (arrow). Strahan and Alired, Re-evaluation of the Arístida gibbosa complex 311 short, circular callus. Longhi-Wagner (1990) subsumed this name under A. gibbosa, using comparisons of spikelet features such as beak length and relative glume length. The name A. marginalis has been applied to numerous Mexican specimens with flat, curling blades and well-developed pulvini. This study investigates the relationships among Aristida gibbosa, A. sorzogonensis, A. orizabensis, and A. marginalis using multivariate analysis of morphological characters scored from herbarium specimens to assess species boundaries and determine the usefulness of certain traits used to distinguish them. MATERIALS AND METHODS For analysis of taxonomic relationships, specimens identified as Aristida gibbosa, A. sorzogonensis, A. oriz- abensis, and A. marginalis were obtained from the following herbaria: F, G, MO, NMCR, US, and WIS. All specimens were examined for a sulcate lemma to distinguish them as members of the A. gibbosa complex. A complete examination of all specimens yielded 90 OTUs (operational taxonomic units) for the analysis, including t fA. gibbosa, A. sorzogonensis, A. orizabensis, and A. marginalis. The specimens used represented the full geographic distribution of the species complex from northern Mexico through Central and South America. A complete data set is available upon request. Roughly two-thirds of the specimens received that were identified in herbaria as A. orizabensis were actually A. appressa, and not included in this study. In addition to not having a sulcate lemma, A. appressa develops a thicker beak (0.2-0.3 mm), usu- ally longer lemma (7-16 mm), and longer callus (0.6-1 mm) with a longer, elongated (rather than circular) attachment scar. Each OTU was scored for 13 characters, seven discrete and six continuous (Table 1). All discrete characters were oes for mature vegetative features as pe or absent, with and without the aid of a dissecting mi di the nature of the character. All spikelet ts were taken using e an ocular comer on a dise cine microscope and were obtained from one mature spikelet from each specimen. Panicle measurements were taken using a standard metric ruler. Principal components analysis (PCA) of the standardized data was conducted using Kovach (2005). RESULIS AND DISCUSSION An initial PCA of all OTUs indicated morphological differences with 83% of the variability among speci- mens occurring along the first three components (Table 2). This is due in part to high variable-to-variable correlations among various discrete features of leaf blade pubescence, type of blade involution and curling, thickness of margins, and shape of nodes (Table 3); the high correlations render these features taxonomi- cally important. Component I accounted for 60.196 of the variability among OTUs, mostly reflecting weak correlations of vegetative features. Component II accounted for 14.996 of the variability, correlating variation in awn and glume lengths, and Component III correlated with pulvini and panicle length (Table 2). Dispersion of OTUs was greatest along the first two components, with resolution bet two morpho- logic groups occurring along component I (Fig. 2). The two groups were distinguished by a combination of vegetative and spikelet features. OTUs corresponding to Group 1 are characterized by leaf blades folded and straight (Fig. 3), gibbous nodes (Fig. 4), lacking pilose hairs on the adaxial surface, margins not thickened (Fig. 5), smaller glumes, shorter awns, and shorter callus to awn lengths (Figs. 6-7). OTUs clustered in Group 2 are characterized by contrasting features: curling leaf blades (Fig. 8), terete nodes (Fig. 9), leaf blades flat with thickened margins and prominent pilose hairs on the adaxial surface (Fig. 10), larger glumes, longer awns, and longer callus to awn lengths (Figs. 11-12). The type of Aristida gibbosa fell within Group 1, and the types of Aristida sorzogonensis and A. orizabensis fell within Group 2. The placement of the type specimen of A. marginalis between Groups 1 and 2 is a result of its having some features of Group 1 (gibbous nodes, branching culms) and some features of Group 2 (flat curling blades, adaxial pubescence). Because members of Group 2 had previously been identified as three different species (marginalis, orizabensis, sorzogonensis), as well as including both pulvinate and non-pulvinate panicles, a second PCA 312 Taste 1. Morphological features scored for principal components analysis, showing discrete and continuous features and how each was scored. Discrete Features Feature Present — 1 Feature Absent = 0 BRANCHING culms branched culms unbranched NODES nodes terete nodes gibbous CURLING leaf blades curling leaf blades straight INVOLUTION leaf blades flat leaf blades folded MARGINS margins thickened margins not thickened PUBESCENCE adaxial surface pilose adaxial surface glabrous PULVINI panicle pulvinate panicle non-pulvinate Continuous Features Feature Measurement panicle length (cm) GIL glume 1 length (mm) GAL giume 2 length (mm) CALAWNL length from callus to awn (mm) CENTAWNL central awn length (mm) LATAWNL lateral awn length (mm) TaBLe 2. Factor loadings for the first three components of the principle component analysis of all OTUs. Loadings less that +0.250 have been replaced by zero. Component 1 Component 2 Component 3 Blade adaxial pubescence 0.335 0 0 Central awn length 0 0.504 0 Blade curling 0337 0 0 Culms branching 0 0 0 Glume I1 length 0.263 0.291 -0.372 Glume 2 length 0.262 0 -0,39 Blade involution 0.337 0 0 Lateral awn length 0 0.476 0 Callus to awn length 0.309 0 0 Blade margins 0.337 0 0 Node shape 0.335 0 0 Pulvini 0 0 0.416 Panicle length 0 0.351 0.663 Total 60.1% 14.9% 8.0% was conducted on all 54 OTUs of Group 2 (Fig. 13). The first three components of the PCA explained 82% of the variability of this second analysis (Table 4). Dispersion of OTUs along component I was a function of variability in awn lengths, glume lengths, and callus to awn length. Component II revealed differences in the central awn, glume two, and panicle lengths, as well as the pulvini development. Component III cor- responded to variation in panicle length and pulvini development (Table 4). Dispersion of Group 2 OTUs reveals no further clear morphological distinctions, nor a clear separation between pulvinate and non-pulvinate forms, in spite of these being visually distinctive. Axillary pulvini cause panicle branches and pedicels to spread outward, giving the panicle a somewhat diffuse appearance, contrasting with the dense contracted appearance of non-pulvinate panicles (Figs. 14—17). In summary, spikelet feat are deceptively similar among the three species analyzed, and may lead one to consider them as conspecific. In contrast, vegetative features, though often thought to be of lesser value in grass taxonomy, serve very well in distinguishing the species. 313 Strahan and Allred, Re-evaluation of the Aristida gibbosa complex 000 L L610 9/00 9900 EELO LSC O 990'0 6r l'0 CCVO 81l0 9900 LIFO 00L'0 NWd 0001 v650 €090 Stevo 9vtO €090 esco CLEO 00€ 0- £090 Voto 6850 INIAINd 0001 ££60 9570 CLEO ££60 $990 Kn 979 D- L£60 cre O vS60 SIJON 0001 DE ZO O8E0 000 L 5950 GE 8£9'0- 000"! "bt ££60 SNIDAVIA 000 L PSSO 6£/0 gez 0 ECLO PLS O 6EZ0 8750 CELO INAAVIVO (OO L 08€0 OLSO £95 0 VvcO- 08£'0 L680 L8€0 JNAWVIV'I 000 L $980 SESO 8e9'0- 0001 SSED L60 NOILMIOANI 000 L 6060 OLE 0- $980 vero LSU KI 000 L Ze O Gu EES'O SHS O TLD 000 1 BE90- Dë EF vS90- | DNIHONVSUS 000 L 8586 H ¿160 INMENSO 000"! 99€ 0 INAAVLNIO 0001 — 3ON32S38fd NVd — INIAINd SIGON SNIDUVW INMVIW INMVIVI NOILMOANI 12) 119 5NIHDNVUg ONTEM ` INMVIND DNDSIINA SNLO IIE JO Wd 104 suoge| KO BIN piipA ^ [| (fm fg: AU VENICE E EE 314 | IL fal Das H im LI bikt £T 2(1) TAXONOMIC TREATMENT Consequent to the results of the PCA, three species are recognized (in addition to A. liebmannii, which was not analyzed in this study): Aristida gibbosa (Group 1 OTUs), Aristida sorzogonensis (Group 2 OTUs), and Aristida marginalis (represented solely by the type specimen). Variation in features is described in Table 5. The recognition of Aristida marginalis (based on the two isotypes from a single collection) is problematic, since so little is known about it. It is distinguished from the other two species by a suite of peculiar vegeta- tive features: taller, more robust culms with numerous nodes and conspicuously short sheaths, about half as long as the internodes, exposing the many gibbous nodes. Other features are a mix of A. gibbosa and A. sorzogonensis: ak during DAMES with adaxial pilose hairs characteristic of A. sorzogonensis, and gibbous nodes and | Ims teristic of A. gibbosa. We leave the name A. marginalis intact until further analyses are Bosse KEY TO THE SPECIES OF THE ARISTIDA GIBBOSA COMPLEX 1. Lemma lacking a beak, the awns almost immediately diverging from the not-twisted apex of the e body; awns 2-3 cm iong 1. A. liebmannii 1. Lemma with a well developed beak, often twisted prior to the divergence of awns; awns (0.5-)1-2 cm lon 2. ade folded or rolled, straight, ti gi t thi ne th Jayial surface alabrous 2.A. gibbosa Blades flat, curling, tl | | tl laxi fi il ith long, Ge hairs E Nodes evidently gibbous bulging on one side; culms Ee above the base, with 7-8 nodes elevated above the bas 3. A. marginalis KI | A HEN + | dass | H 4 | E L na | LGL, EN A ] | + J above the base 4. A. sorzogonensis 1. Aristida liebmannii E. Fourn., Mexic. pl. 2:78. 1886. Tyr: MEXICO: Veracruz: Mirador, Apr 1842, EM. Leibmann 662 (HOLOTYPE: C; sorres: MOL US-207485, US-991666 (fragm. ex LE), US-1389797 We have studied only two specimens: the isotype at MO and an additional specimen (Burch 6143A) from Honduras. A brief diagnosis follows [Pohl & Davidse (1994) pon a bad EE Sheaths glabrous; blades flat (rolled upon drying or at the ends of the blades), with thi 1 throat glabrous; 1 1 E: and naked at the base; Lat panicles narrow, elongate, non-pulvinate, but the lower | , sulcate, a beak absent, glumes inverse, the first 10-12 mm long, the second 8-9 mm long; | the awns diverging immediately at the tip of the lemma; awns 2-3 cm long, + Equal in length. The combination of a sulcate lemma and long awns is immediately distinctive. Distribution —Mexico (Veracruz) and Honduras. 2. a oa o Kunth, Enum. pl. l: ee ee BASIONYM: oa ge Nees, Agrostologia Brasiliensis, EP von Martius 14 (HoLoTyeE: M; CH ISOTYPES: LE-TRIN-1286.01,US- 865702 har ex M! & photo!) Aristida gibbosa is characterized by blades that are folded or rolled, stiffly erect, glabrous, and usually lacking thickened margins. The culms regularly branch above the base and have prominently gibbous nodes, with 2—6(—8) nodes elevated above the base. Panicles are narrow and non-pulvinate. Distribution.—South America: Bolivia, Venezuela, British Guiana, and Brazil (Fig. 18). ta C S Cruz, Nuflo de Chavez, S S 10km W of Norte Le Ge 1986, T. ` Killeen 1979 (US). BRAZIL. Without State: No locality, no date N Glaziou 20104, 20106 (US). Bello H r 1936, M. Barret 4521 (US). Amazonia: ERICA ee Clevelandia, km 50, 31 Aug 1955, G.A. Black 18544 (US). Federal District: e of the Goias-Distrito Federal bord y BR-020-030, 9 Apr 1976, Davidse et al. 12170 (MO). Goias: ca. 15 km (straight line) S of Goias Velho, 10 May 1973, W.R. Anderson 10039 (MO); no locality, no collector, 20 SC SEN ee 7 i ee hillside with campo and wooded uu ca. 10 km S of Alto ke Pariso, 20 Mar me Irwin et al. 24781 (US); ca. 30 S. of Caiaponia, 29 Apr 1973, W.R. Anderson 9424 (M ds ky slopes 8 km N of Veadeiros, 16 Mar 1969, Irwin eed 24524 (US). Maranhao: “Ilha de Balsas” Region, | | Bal d corn 20 us 1962, Fiten et al. 4865 (US). Mato Grosso: in cerrado formation on very sandy soil, Ad 1968, Ratter et al. 2081 (MO). Minas Gerais: Sierra do Cipo, 5 Aug 1936, W.A. Archer 5028 (US); Lagon Santa, serras a 20 km da cidade, 7 Apr 1951, G.A. Black 11695 (US); Estado de Minas Gerais, 1 Jan 1951, Pires et al. 2925 (US); Municipio de Sao Goncalo. BRO40, km 251, open cerrado, 25 Jul 1984, Mori et al. 16988 (MO, US). Roraima: along Boa Vista-BV 8 road (BR 174) km 76 Strahan and Allred, Re-evaluation of the Aristida gibbosa complex 315 0.477 0.38- Group 1 Group 2 0.28- 0.19- Component 2 -0.287 "m | | | | -0.47 | | | | | | | -0.47 -0.38 -0.28 -0.19 -0.09 0.00 0.09 0.19 0.28 0.38 0.47 Component 1 E ) n: E E all ATII F del Au Fora E A | E) [| IG plex. G = type of A. gibbosa. M = type of A. marginalis. $ = type = E of A. sorzogonensis. O = isosuntune of A. orizah is (Müller 2103 US) bi d ei hl E r by Uraricoera river, 8 Oct 1977, Coradin et al. 545 (MO); NE Roraima, ca. 110 km NE of Boa Vista between Bonfim and Guyana border on Tacutu River, 20 Oct 1977, Coradin et al 792 (US). BRITISH GUIANA. Rupununi: no locality 1 Sep 1945, Fraser, Dr. 378 (US); St. Ignatius dry lateritic ridged savanna 350 ft, 25 Jul 1963, R. Goodland 22 (US). VENEZUELA. Anzoategui: Distrito Bolivar: Rocky mountain savanna, just S of El Zamuro, Fila El Purgatorio, i airline km NE of of Bergantin, 24 Nov 1981, nain et al. 19319 (MO). Aragua: Carretera nip d PANE t E enla sabana, 11 Sep 1963, P. taldo 3744 (MO); 12km S of Alto de Chor tal tl 1) oad to Maracay, 14 Nov 1971, G. Davidse 3068 (MO); 10 km SE of Rancho Grande along road to Vs 14 Nov 1971, Dav d G. 3053 go: Bouvet: Distrito Piar, middle part of Rio HEU affluent of Rio Ambutuir, along trail to Uriman, 30 Nov 1982, D J d Pie de la Roca, SEb tepui, 24 Nov 1982, Davidse et al. 22559 (MO). Sucre: 7 km E of the Mochima Hay: intersection along Hwy. 9 Desa Cumana and Puerto La Cruz, 16 Dec 1973, G. Davidse 5027 (MO). 3. Aristida marginalis Ekman, Ark. Bot. 10(17):23, t. 3, f. 2, t. 6, f. 12. 1911. Tee: BRAZIL. Marto Grosso: Cuiba, in margine silvulae (capáo”) loco sicco, graminoso, arenoso-argilloso, 26 Apr 1903, G.0. Malme 3143 (HOLOTYPE S; IsoTYPES Gi, US-702283!, US-81202 fragm. ex S!). Aristida marginalis is known only from the type collection, and is characterized as follows: culms robust, generally about twice as thick as in A. gibbosa or A. sorzogonensis, branching above the base, with numerous (7-8) nodes elevated above the base; sheaths short, about Y the length of the internode, straw-colored and contrasting with the pale greenish internode; blades flat, curling, with thickened margins, pilose on the adaxial surface; panicles narrow, but weakly pulvinate. Distribution —Known only from Brazil: Matto Grosso, Cuiba (Fig. 18). The name Aristida marginalis has been applied to similar plants from North America, many of which 316 A ATI A re LL J Er 4 ATII E [a P | Fics. 3-7 1 Pa igl "E ELIAS Le | L dcs qo £r 10TH & Gl rf | a 0TUc 7 El tafi | 1 01) Scales in mm are conspicuously pulvinate, particularly collections reported as such by Howard Scott Gentry (1942) from the Rio Mayo region in western Chihuahua. All of these North American plants fall well within the A. sor- zogonensis cluster on the PCA, and differ from the true Brazilian A. marginalis in the features given in the key. Further field and herbarium work are needed to fully explain the distribution and variation of Aristida marginalis. Its apparent int liacy between A. gibbosa and A. sorzogonensis seems coincidental, and not a result of hybridization or introgression. As with many species of Aristida, species boundaries are likely to be delicate, yet decipherable, and the name should not be discarded as inconsequential. Strahan and Allred, Re-evaluation of the Aristida gibbosa complex 317 Fics. 8—12. 8. Curli ngl f blades of G p 20TU 9 Terete nodes. 10. F g teristicof G 2 OTUs. 11. Gl p 2 OTUs. 12. Floret of Group 2 OTUs. Scales in mm surface, | 4. Aristida e e in C. Presl, Reliq. Me e 224. e Tree: in Luzonia ad Sorzogon, T. ol, W). Haenke s.n. (HOLOTYPE LE fragm., PRC photo!, US-81256 fragm. ex W! & pho Bistum dus Mexico and Central America, with a few speci [ thern South America (Fig. 18). Aristida sorzogonensis is characterized by blades that are flat and curling, with pilose hairs on the adaxial surface and with thickened margins. The culms do not branch above the base and the nodes are not gibbous, with 0—4 nodes elevated above the base. Panicles may be pulvinate or non-pulvinate, which is a minor feature, but the visual differences are immediately noticeable (Figs. 14—17). As has been done 318 I I £ al n.a D In LI Pay S E ET aia) 0.5 0.44- 034 A A 0.1+ À A N E dm 5 A. 0.0 EA 3 Y ER -0.4+ - vi -0.5-1- ne l "- e T L L | | | H | OMNE E S D | q i | | | -0.6 -0.5 -0.4 -0.3 -0.1 0.0 0.1 0.3 0.4 0.5 Component 1 Ic. 13. Dispersion of OTUs along om ts | and Il of PCA of speci ponding to the Group 2 cluster: S = type of A. sorzogonensis. 0 = isosyntype of A lüller 2103, US). Black triangles = pulvinate form. Gray triangles = non-pulvinate form. Taste 4. Factor loadings for the first three components of the principle component analysis of Group 2 OTUs. Loadings less that +0.250 have been replaced by zero. Component 1 Component 2 Component 3 Blade adaxial pubescence 0 0 0 Central awn length 0,451 0.309 0 Blade curling 0 0 0 Culms branching 0 0 0 Glume 1 length 0.468 0 -0.262 Glume 2 length 0.427 -0.376 Blade involution 0 0 Lateral awn length 0.466 0 0 Callus to awn length 0.407 0 0 Blade margins 0 0 0 Node shape 0 0 0 Pulvini 0 0.516 -0.815 Panicle length 0 0.615 0.406 Total 48.1% 224% 11.5% Strahan and Allred, Re-evaluation of the Aristida gibbosa complex 319 AA P obi FEES -Lr Fics. 14-17 14 0j I E A bal | La af nf H Al ai £l * dal L T Js f| 15 | o P A ai fi | a) A . J ( ) 16 Contracted ? 4l I L2 y tt Al ag E all s 1 L 37 Avile | Le | CLE i within A. purpurea and A. pansa (Allred 19842; Allred ër Valdes-Renya 1997) forma are recognized based on pulvini development: KEY TO THE FORMS OF ARISTIDA SORZGONENSIS L DES "M HI £1 | +l +l doa]: LA D I H | lj up Ep RS and somewhat diffuse 4a. f. sorzogonensis Panicles non-pulvinate, rarely developing pulvini in the axils of branches, branchlets or pedicels, the panicle consistently narrow and contracted 4b. f. orizabensis 23141 lexas Z(1) £T dee eh 320 ayeuiA[nd ÁA Lam ajeuiA[nd ajeuiA[nd-uou 3jeuiA[nd-uou apiued osOI —eso[id Agusuluold aso|id Ápusultuold snoJge|b a2u32saqnd [erxepe peue»plui peuexui peuexplu1 paus!) JOU subiu) Je 1814 "m papjo} uonnjoAul buipino 3 xe] DUIN 9 xe| Duin 9 xel Palo 9 Us səpejg snoaqib 94949) 313191 snoqqio SODON payueIq payuelqun payuelgun peupxuelq sun) SJ932eJeQ5 aS UZI-S6 861-56 STFTOL 0v1-09 LL ECOL 0'vi-09 LL+V8 (WU) y18U3] ume [219287 FSE El oke? ETFSTL SLI-V8 PEPALI 0'81-€8 OCULI (ww) u3bue| ume [eu25 ELO E A ¿£1+S56 Dk GLF/6 Alf vVL+VO (ww) (Dua ume 04 sn e) €799 Keen SEKR SOL-09 EK L'6-0'S OL-T C9 (ww) uibus] z euunjo O8-CL LOL-8'7 9LF+8 OLL-O9 VI-06 e 01-61 LLT69 (uuu) ubue| | euun[o 081-091 S26 06 DE LOL UGC 6v+09l |£-06 VS+92Z1 (wd) y1Sua] aplued obuey obuey AS F UBS abuey (JS F ueaw abuey (JS F ueaA SJ932eJeQq) shonumnuo) usunads ədÁ} a}eulAind ajyeuiA[nd-uou sijeuib1eu "y sisuauobozios "y esoqqib "y SIJDUIDIDLU "y pue 'sisuauobozios “yy “ 1916 DpNsuYy JO sanea JO uosueduuJo» "e 218v | Strahan and Allred, Re-evaluation of the Aristida gibbosa complex 321 120" o 1100 -80" DD —— -40" 20 20" D y -20' -20 Md MD | A -120° -100° -60° -60° do —— P ; a ——É—— ES SEN ti llection. Dashed line = A. sorzogonensis. Solid line = A. gibbosa, M = Type specimen of A. marginalis, 4a. Aristida sorzogonensis f. sorzogonensis Distribution.—Mostly Mexico (Chiapas, Chihuahua, Jalisco, Michoacan, Nayarit, Sinaloa, Tepic) and Central America (Costa Rica, Honduras, Panama), with a few specimens from Columbia and Bolivia. lD: 1 SI Specimens examined. COLOMBIA. Santa Mart locality, H.H. Smith 136 (US). BOLIVIA: bet Aguas Zembrad to ae SanLucas Bon 1400 E 12 e 1944, Drew, W.B. 146 (US). COSTA RICA. Puntarenas: 3 km N of the C I c 1968, Pohl et al. 11602 (F). HONDURAS. El Paraiso: Wuebrada del Muro, road to Yuscaran, open ground, 5: Nov 1951, J.R Swallen 11327 (MO). M S lo largo carretena, Las Mesas, 23 Nov 1948, A. Molina 1707 (US); rocky pine woods, hills around Zamorano valley, 2 Nov 1951, Swallen J R 11259 (US); rocky pine-oak slopes, 15 Oct 1951, LR Swallen 10751 (US); rocky pine woods, hills around Zamorano valley, 2 Nov 1951, J.R. Swallen 11256 (US); colinas y bosque micto entre Las Mesas y Guayabillas 5km empalm dalini yuscaran, 04 Nov 1963, A. Molina 13164 (F). MEXICO. Chiapas: Geen Chicocuceo Belen, 1 km E of Las Sal ,2 km SE of C ud 18 Nov 1984, Davidse et al. 30037 (MO). Chihuahua: Rio Ma flat, Guasaremos, 26 Sep 1955, H.S. Gentry 1873 (ARIZ, MO, NMCR, WIS, US). Jalisco: 20 km Guadalajara-Tepic, 26 Sep 1 1946, Hernandez et al. 2745 (US); ( iente) 3.2 km al W de Las Marias; 15 km por camino al E de Avotitlan, 08 hs 1985, T.S Gees 10987 (WIS). Michoacan: along road from Tsitzio-Tiquicheo-Huetamo- Altamirano, 7 Oct 1953, E.R. Sohns 906 (US). Nayarit: 10 km al E de Tepic por la carretera a Mazatlan, 18 Nov 1983, FJ. Santana 1426 (MO); 13 km NW of TEPIC (Municipio de Tepic) along highway 15, 18 Nov 1983, Solheim et al. 971 (WIS); steep treeless hills 10 mi SE of Toni 6 ii 1952, M R 13384 (US). Sinaloa: Cerro cue 1 Nov 1904, Brandequa (US); Mesa Malqueson, Cerro Colorado; h, 2500 ft, 8 Dec 1939, H.S. Genta 79 (MO). Sonora: Sierra Tecurahui, SE Sonora, 4250 m, 27 Oct 1961, Ger et al. 19380 (US). Tepic: no locality, 6 Feb 1892, Dr. E. MA 1916 (US). PANAMA: between Paso Del Arado and Ola, province of Cocle, in savannas and thickets, 7 Dec 1911, H. Pittier 5019 (US); vicinity of Ola, province of Cocle, 7 Dec 1911, H. Pittier 5047 (US); Picacho de Ola, province of Cocle, 8 Dec 1911, H. Pittier 5066 (US); ca. 10 km SW of San Carlos along the Inter-American Hwy, 18 Nov 1975, Davidse et al. 10128 (MO). 322 4b. cun dE f. orizabensis (E. Fourn.) Strahan & Allred, comb. nov. Basionymn: Aristida orizabensis ., Mexicanas Md 11:78. 1886. Tyee: MEXICO. VERACRUZ: Mp: de mn HH Schaffner 136 POE P; ISOSYNTYPES LE); NY! USD. The name E. Fourn totypified P y 1 by Fournier (1886). The specimens at P and LE were dle 2 a + 71/12 + KATY de AAA LINE an Ta » | e 71 ] (C. sn. 132% 114 Distribution. —Mostly Central America (Guatemala, Honduras, Panama), with fewer specimens from Mexico (Sinaloa, Guerrero) and Columbia. Specimens examined. COLOMBIA. Huila: 3 km SE of Neiva, Huila, 1700 m, 23 Feb 1945, Little et al. 9470 (US); 6 km SE of Altamira along road to Florencia, 9 Jan 1974, G. Davidse 5602 (MO). OREA Jalapa: i e pine: -clad o TO and inis ca. 10 mi S Jalapas, 29 Nov 1939, J. Steyermark 32176 (F). Wit t of oa Guatemal city and Jutiapa, 10 Oct 1944, Goodman et al. 3696 (F). HONDURAS. El paraiso: ed to Yuscaran nen places, about km 2, 5 Nov 1951, qos R Sw sala oe eee ind x in A woods, a ond Rio San Francisco, 1 Nov 1951, J.R. Swallen 11212 (US). Morazan: 20 km N edros t on hills, 4 To 1986, Davidse et al. 31584 (MO); Santa Ines, 4 Nov 1943, J.V. — 1486 em ge panya hills, 7 Nov , 1951, J.R. Swallen 1 ta Inez, 5 km E of El Zamarano, 19 Dec 1978, Pohl et al. 13738 (MO); Colinas y Guayabillas, 5 Pe empalme carretera dalini yuscaran, 04 Nov 1963, A.Molina 13167 (F). Without See in mixed oak-pine forest near Piedra Herrada, 11 Sep 1949, L.O. Williams 15975 (E); in pine woods, near Jicarito, 5 Dec 1948, Williamson et al. 14802 (US). MEXICO. Guerrero: Ixtapan de la Sal, 12 Oct 1952, Matuda et al. 27056 (US). Si- naloa: between Mazatlan and Durango, 6 Jan 1975, Beetle et al. 3695 (WIS). PANAMA. Herrera: 10 mi S Ocu roadside, 21 Jan 1966, Tyson et al. 2866 (MO ACKNOWLEDGMENTS Many thanks to the curators of the following herbaria for loaning the material used in the study: F, G, MO, NMCR, US, and WIS. Patrick Alexander provided pictures of some of the grass parts. David Lee Anderson kindly translated the abstract. We thank Paul Peterson and Stephan Hatch for helpful suggestions on an earlier draft. REFERENCES ALLRED, KW. 2003. Aristida. In: ME Barkworth, KM, Capels, S. Long, and MB, Piep, eds. Magnoliophyta: Com- melinidae (in part): Poaceae, part 2, Flora of North America North of Mexico, vol. 25. Oxford University Press, New York, NY. Pp. 315—342. ALLRED, KW. 1984a. Morphological variation and classification of the North American Aristida pur compl (Gramineae). Brittonia 36:382-395 ALLRED, KW. 1984b. Studies in the genus Aristida (Gramineae) of southeastern United States. |. Spikelet variation in A. purpurescens, A. tenuispica, and A. virgata. Rhodora 86:73-77. ALLRED, KW. and J. Vaibes-ReNvA. 1997. The Aristida pansa complex and a key to the Divaricatae group of North America (Gramineae: Aristideae). Brittonia 49:54-56. Exman, EL. 1911. Neue brasilianische Gráser. Arkiv fór Botanik utgifvet af K. Svenska Vetenskapsakademien | Stockholm 10(17):23. Fournier, E. 1886. Mexicanas piantas 2:78. HeNRARD, J.T. 1927. A critical revision of the genus Aristida. Il. Meded. Rijks-Herb. 54A:221-464, HENRARD, J.T. 1928. A critical revision of the genus Aristida. IIl. Meded. Rijks-Herb. 54B:465- 701. HENRARD, J.T. 1929. A monograph of the genus Aristida. |. Meded. Rijks-Herb. 58:1-137 HENRARO, J.T. 1932. A monograph of the genus Aristida. |. Meded. Rijks-Herb. 58A:157-325 Kovach, W.L. 2005. MVSP — A multivariate statistical package for Windows, ver. 3.1. Kovach Computing Services, Pentraeth, Wales, U.K. LoNcHI-WAcNER H.M. 1990. Diversidade e distribuicao geografica d pecies de Aristida L. (Grami ) ocorrentes no Brasil. Acta. Bot. Bras. 4:105-124 Nees, C.G.D. 1829. Agrostologia Brasiliensis. Stuttgart. Pout, R. and G. Davpse. 1994. Aristida. In: Davidse, G., M. Sousa. S., AO Chater, eds. Flora Mesoamericana, vol. 6. Missouri Botanical Garden. Pp. 253-257. Pres, J.S. 1830. Reliquiae Haenkeanae 1(2-4):224. TAXONOMY OF BROMUS (POACEAE: POOIDEAE: BROMEAE) SECTIONS BROMOPSIS, BROMUS, AND GENEA IN BRITISH COLUMBIA, CANADA Jeffery M. Saarela -o Museum of Nature PO. Box 3443, Station TN E KIP 6P4, CANADA jsaarela@mus-nature.ca ABSTRACT At ic treatment of B tions B is, Bromus and Genea in British Columbia, Canada, is presented. Nineteen taxa are e ara dis six native and 12 introduced ab in one dips two varieties. The ee pone B. Reeg ponies in i a Ts ss ] Ti] p :71 E: o 3 a E Iw 5 eg s 1 T4 El E of 1 E L f th I AJ E NT +1 America n Tli: var. occidentalis Vasey D ES yI [S X y E dst ccs g I I E S f "ue H E NS E 1:11 : Cape n i L Af HB. P ` synonymies, descriptions, ps, re] I E E RESUMEN E de B i B psis, Bromus y Genea en British Ge MW Se reconocen diez y nueve taxa, que inclu; is especi tivas y 12 introducidas, una con dos variedad I riparius, fomentada NT + A a f, 1 eux vum | Im a po pr aaa dg e 1 fl E » m su introducción en Norte América: Y se cita Bromus squarrosus var. vill bia, siendo | ita d la provincia y para Norte América. Se P e: din kalmii var. occidentalis in) ex Beal Se aportan E para identifica las especies, sinonimias, descripciones, E L Bromus L. is a large and t ically complex grass g that includ than 160 species distributed in temperate regions worldwide. The, genus is distinguished from other grass genera by the combination of leat sheaths that are closed for most of their length, awns that are inserted subapically and hairy append- ages on the apices of the ovary (Clayton & Renvoize 1986). Approximately 80 Bromus species are currently recognized in North, Central and South America (Pavlick et al. 2003): 29 of these reportedly occur in Canada (Pavlick & Anderton 2007). Among the 10 Canadian provinces and three territories (Fig. 1), the western- most and topographically diverse British Columbia has the greatest number of Bromus species. Bromus is the second-largest grass genus in British Columbia; only Poa L., with 31 species (Soreng 2007), has greater species richness in the province. In North America most workers recognize five sections in Bromus (reviewed by Saarela et al. 2007), but multiple recent molecular phylogenetic studies indicate that not all are natural groups, particularly the globally widespread B. sect. Bromopsis Dumort., which appears to comprise distinct Sierra Madrean (Mexi- can), North American, South American and Eurasian lineages (e. g., Pillay & Hilu 1990, 1995; Saarela et al. 2007). More work is needed to clarify higher-level phylogenetic relationships in Bromus. Nonetheless, the traditionally recognized Bromus sections can generally be recognized morphologically and learning their key characteristics can greatly aid in species identification in the field and herbarium. There has been considerable progress in clarifying lower-level taxonomic problems in Bromus. Several species that occur in North American have received recent taxonomic study, including; B. ciliatus L. and B. richardsonii Link (Peterson et al. 2002); B. grandis (Shear) Hitchc., B. hallii (Hitchc.) Saarela Er P.M. Peterson and B. orcuttianus Vasey (Saarela et al. 2005); Bromus catharticus var. elata (E. Desv.) Planchuelo (Planchuelo 2006); and B. carinatus Hook. & Arn. (Barkworth et al. 2006). Similar recent taxonomic work has been con- ducted on taxa from South America (Massa et al. 2001, 2004; Saarela et al. 2006) and Eurasia (Sales 1993, 1994; Spalton 2002). Comprehensive taxonomic treatments of Bromus for North America north of Mexico J. Bot. Res. Inst. Texas 2(1): 323 — 372. 2008 324 [| [| £ al Dos H Inm L 1 dad f Texas 2(1 Northwest Territories Nunavut Alberta ; Manitoba Quebec Nova New Brunswick Scotia Fic. 1 Map of Canada, ith the 10 i [+ tarritariac indicated Pritich Columbia ic fl + Lë P . Scale bar=500 km. have also recently been pubis nud 1995; Pavlick & Anderton 2007), but despite these resources, Ln: species of Bromus can ly difficult for many workers. In such large and difficult groups, Es zl 4 + ly £ 8 A "ELT e lh focused t treat detailed information than is normally found in done floras, be they regional or continental in scope. The most recent EE treatment da SE for EH GE in de Flora of British Columbia (Douglas et al. 2001), t roach tunderstand- A +l ing of the genus O e t treatment (s Stewart & Hebda 2000) is siete a to only those species that occur in the Columbia Basin region of British Columbia. The purpose of this work is to provide a detailed and updated taxonomic treatment for Bromus in British Columbia, including a dichotomous key, synonymies, species descriptions, distribution maps and citations of representative specimens. Bromus in British Columbia Bromus in British Columbia includes multiple native and introduced species that are currently classified in four sections. Most of the introduced species are annuals classified in B. sects. Bromus (B. briziformis Fisch. Sr C.A. Mey., B. commutatus L., B. hordeaceus L., B. japonicus Thunb., B. racemosus L., B. secalinus L. and B. squarrosus L.) and Genea Dumort. (B. diandrus Roth, B. sterilis L. and B. tectorum L.). All of these (and other) introduced species in B. sects. Bromus and Genea occur throughout western North America, mostly north of Mexico (see Pavlick & Anderton 2007; Saarela & Peterson in press), and several have caused serious eco- logical disruption. In general, annual species can be distinguished from perennial species based on basal vegetative characteristics. In perennial B species, the dried remains of the previous years’ growth are normally present, whereas there is no previous years’ growth in the annual species. In most cases the annual taxa are generally smaller in stature and have a more delicate appearance (with some exceptions, such as some individuals of the large annual, B. diandrus), compared to the perennial taxa. Bromus sect. Bromus is characterized by 3-5-nerved first glumes, 5-7-nerved second glumes and lemmas rounded over the backs, Saarela, Bromus in British Columbia, Canada 325 whereas B. sect. Genea is characterized by 1(-3)-nerved first glumes, 3(—5)-nerved second glumes and nar- row, elongate lemmas. Species of B. sect. Bromopsis in British Columbia are perennials with 1(-3)-nerved first glumes, 3(-5)-nerved second glumes and lemmas rounded or slightly keeled over the backs (i.e., flat- tened dorsiventrally). Two B. sect. Bromopsis species in British Columbia are introduced: the highly invasive smooth brome (B. inermis Leyss.) and meadow brome (B. riparius Rehm.), which is reported here as part of the British Columbia flora for the first time. The remaining five B. sect. Bromopsis species in the province are native: B. ciliatus L., B. pacificus Shear, B. pumpellianus Scribn., B. richardsonii Link and B. vulgaris (Hook.) Shear. The remainder of the Bromus taxa in British Columbia are classified in B. sect. Ceratochloa (P. Beauv.) Griseb., a group that is widespread throughout the province and easily recognizable on the basis of its strongly laterally flattened and keeled glumes and lemmas. Species limits within B. sect. Ceratochloa are not clear, both in ind Columbia and more o d throughout western North America. Taxa in B. sect. Ceratochloa variously reco gnized at different ranks; in British Columbia these include B. aleutensis Trin. ex Griseb. [-B. sitchensis Bongard var. aleutensis (Trin. ex Griseb.) Hultén], B. carinatus Hook. & Arn., B. marginatus Steud. [=B. carinatus var. marginatus (Nees.) Barkworth & Anderton], B. polyanthus Scribn. ex Shear and B. sitchensis Bongard (e.g., Calder & Taylor 1968; Hitchcock et al. 1969; Hubbard 1969; Pavlick 1995; Douglas et al. 2001; Clayton et al. 2002 onwards; Barkworth et al. 2006; Pavlick & Anderton 2007). Because taxon circumscriptions are not clear in B. section Ceratochloa, the section is not treated here (al- though it is included in the key). MATERIALS AND METHODS This study is based on study of over 1200 herbarium peso is CAN, DAO, UBC, US and V, and ob- servations made in the m from 2003-2007. TI | and are based ui] ] ] r li on study of herbarium sp men llected in British Cola Fora nw of int which there are not a lot of collections, Eurasian specimens at CAN were also used to generate the descriptions. Nomenclatural information was compiled through study of the primary literature and numerous secondary sources, including Pavlick (1995) and Pavlick et al. (2003). I have not attempted to account for all of the names that have been applied to the introduced taxa in their native European ranges (i.e., species of B. sects. Bromus and Genea); these names are rarely used in the North American literature. Information on common names (in English and French) was obtained through internet searches and from information EE by Darbyshire (2003) Data on geograp phical distribution, assadas aa ER 1 WEN label daa and personal field. dbssrs ations. Where not available on specimen labels, geographical NM was obtained from the Geographical Names of Canada website (http://geonames.nrcan.gc.ca/index_e.php, accessed 2007). Maps were produced in ArcView (Environmental Systems Research). All illustrations were prepared by Dr. Cynthia T. Roché, published originally in Flora of North America volume 24 (Barkworth et al. 2007); they are used here with permission. TAXONOMIC TREATMENT Bromus L., Sp. Pl. 1:76. 1753. Tre: Bromus secalinus L. (conserven TYPE: LINN 93.1, see Jarvis, Taxon 41:559. 1992, and Taxon 44:611-612. 1995). Plants annual or perennial; usually caespitose, occasionally rhizomatous. Culms 7-172 cm tall, one to several per plant. Leaf sheaths closed for most of their length, glabrous or variously pubescent; auricles present or absent; ligules membranous, to 7 mm long; blades flat, abaxial and adaxial surfaces glabrous or variously pubescent. Inflorescences panicles, sometimes racemose, edd é EE to a single ID open to dense, erect to nodding. Spikelets 1.3-8 cm long, terete lly to laterally, disarticula- tion above the glumes and below the lemmas. Med necia. shorter than the adjacent lemma, glabrous or variously pubescent, acute, ti ] 1—7(9)-veined; upper glumes 3-9(11)-veined. Lemmas rounded to keeled, glabrous or variously pubes ene with a single +terminal or subterminal awn, occasionally unawned; paleas shorter than lemmas, usually variously ciliate on the keels. Styles 2, inserted laterally on a bilabiate appendage of the ovary. Anthers 2-3. Base chromosome number, x = 7. 326 Etymology.—The name Bromus is derived from the Greek bromos, which means 'oats'; bromos is derived from broma, an ancient Greek word for food (Wagnon 1952; Pavlick & Anderton 2007) KEY TO THE SPECIES OF BROMUS SECTIONS BROMUS, BROMOPSIS AND GENEA IN BRITISH COLUMBIA 1 e. laterally compressed; lemmas laterally compressed and strongly keeled (B. sect. Ceratochloa, not treated here). 1. Spikelets not laterally H flatt | dorsiventrally, not strongly keeled. 2. Lemmas narrow, largest lemmas generally « 2 ide; | pex bidentate, with awn-like 1 inat hyaline teeth 1-5 mm long (B. sect. Genea). 3. Lemma bd 18-35 mm long; awns 30-50 mm long; | glumes 12-26 mm long; upper glumes 18-35 mm long 4. B. diandrus 3. Lemma SE 9-22 mm long; awns < 30 mm long; lower glumes 4-14 mm long; upper glumes 7-21 mm long. 4. Lemmas 13-22 mm long; awns 15-30 mm long; panicles sparse, with 1(-3) spikelets per branch; branches aint usually GE Sch E O Oram in a EITA A 16. B. sterilis ng; a g; panicl r dense, ed some branches » 3 spikelets pet h; | | i longer or r shorter than spikelet 17. B. tectorum T broad, largest lemmas generally > 2 mm wide (except in B. vulgaris) and So URN ae across the back; lemma apex entire or bidentate, teeth t usually not hyaline, 0-3 mm long, not awn-like acuminate. bs 5. Plants perennial, often with remnants of a previous years' vegetative growth present; lower glumes 1-3-nerved; upper glumes 3-5-nerved (B. sect. o 6. Plants ua rhizomes; longest anthers > 4 mm lon 7. Awns (3- in zn mm E EE EE basal sheaths reticulate-fibrillose (ie, thatched i in along upper to upper half; f blad bat Ge and conspicuously Nena a the base = 13. on riparius Fi Awn 5 0 rd. a sd E| Ejs N EE 2 4 t ka keels al | 34 to entire | gt d f blades glal pubescent, marginal hairs (if present) not robust and not | dened tthe b 8. Awns pad or up to 3mm no icm EES to variously scabrous or puberulent on lower 1/6 to 1/4 6. B. inermis Awns 1 HS 5) mm | | ith hairs 0.5-0.8 long along gi | midvein, or moderately to densely villous iliroughout 10. B. pumpellianus 6. Plants without rhizomes, all anthers « 4 mm long. )J 9. Most lower glumes 3-nerved; awns 1.5-3(-4) mm long 9. B. porteri 9. Most lower glumes ed awns (2-)3-11 mm long 10. Glumes pubesce 11. Mature inflorescences T p id cm wog branen, broadly spreading, each with (1-)2-5 ) g -4 mm long; leaf blades not thin and papery in | herbarium ae . Mature inflorescences narrow, 1-5 ide; | , lao . B. pacificus ing, each with Ie E SE nds kon n Mi Geen -)3-6(-7) mm long; leaf a Re | thin no mms Im 18. B. vulgaris 10. Glumes glabrous. 12. guie n SS E GE longa awns UE i Ge mm long; larger lemmas < 2 mm wide; e 8. B. vulgaris 12. Ligules 0.5- 2n mm dong; awns 4- a id mm ud arger lemmas generally > 2 mm v leaf bl p mari im 13. Lemma backs glabrous or sparsely puberulent with 250 0.1 Tn ong) margins pubescent with hairs 0.5-1 nes 6. mes ASE 2 mm ong: Bie 0. a 1 6 mm long al leaf sheaths glabrous or sparsely to J MO, pp! E B. ciliatus to densel ith hai 1 mm long; upper glumes (8— e 11 SE (145) am long lower gumes 7- E (1 a E 34) eh Jf + Ia | | "=| £ upper blades TEA KZI NZ ka 12. B. richardsonii Saarela, Bromus in British Columbia, Canada 327 | E E " pi ml +. | i 2 E. nan we I + H i i , 5. Plants annual, g g : glume upper giumes 5-9-nerved (B. sect Bromide), 14. Lemmas appearing infated, 3-4 mm wide; awns absent or to 1 mm long; spikelets ovate 1.B. briziformis 14, Lemmas not appearing inflated, < 3 mm wide; awns 2-20 long; spikelets | late to ovate lanceolate. 15. Lemmas with s ly protruding | Iry; panicles usually dense 5. B. hordeaceus +l + | J | Lamar "ry, 15. Lemmas Brie t | | 16. ANDS arising 1.5-5 mm 1 below the apex ofthe lemma. 17. sided, the branches usually pu m | ik let: | itt h li | 0 5-0. 9m pl m I wide B. squarrosus (E Inflorescences usually paniculate ano not appearing One “sided, 2H least the qe branches usually bearing > 1 sj g 0.5 mm wee 7. B. japonicus 16. Awns arising « 1.5 mm below the apex of the lemmas. 18. ED leai snenie glabrous erit sparse, s tiffha irs 0.5 0.6 mm | ikelets widening Al E ni Iis al Pa a ible as s oe wraps around tl yopses; caryo| and V-shaped in cross sectio 4. B. secalinus 18. Gen leaf sheaths densely pubescent with long, soft hairs to 1.2 mm long; SS not widening substantially as the fruit forms, lemmas continuing to obscure most t or crescent C E section | | £] | «AC f If 19. Panicles | narrow, < 2 node o r ofthe terminal floret) 1 spikelet per brane most larger d < 22 6.5-9.5 mm long 1. B. racemosus l Paice broad and spreading, > 2.5 cm wide; longest inflorescence e > m (as measured from node to apex of the terminal floret); 1-3 spikelets per be larger spikelets » 22 mm long (including awns); lemmas 7.5-11 mm long 3. B. commutatus Ke 1. Bromus pele Fisch. € CA. ae Index Sem. (St. Petersburg) 3:30. 1837. (Figs. 2A-E, 3A). Tre: IRAN: Tahlysh, in idi pag. Limar, 1830, C. Meyer s.n. (nororvee: LE). Annual; rhizomes absent; plants 17-70 cm tall. Culms 1-2 mm wide at base, smooth; nodes 2, brown-black, pubescent, hairs to 0.2 mm long. Leaf sheaths closed for most of their length, densely pilose with soft hairs to 0.6 mm; auricles absent; ligules 0.5-2 mm long, membranous, glabrous, erose; blades 5.5-10.2 cm long, 1.5-5 mm wide, abaxial surface densely pubescent with short, stiff hairs 0.1-0.3 mm long or densely pilose with long, soft hairs to 1.2 mm long, adaxial surface densely pilose with long, soft hairs to 1.2 mm long, margins smooth or serrulate. Panicles 5-11.5 cm Jens, 2-6 mm wide, open, lax and somewhat sparse, rac- emose, often appearing one-sided (secund); | usually longer than spikelets, scabrous, 1(-2) spikelets per branch, lowest inflorescence node with 1-3 benches Spikelets 1. 8-2. 5 cm ES ovate, rachillas not visible at maturity, florets 7-18. Glumes subequal, glabrous; lower glumes 4.6-6 mm long, 3-5-nerved; upper glumes 5-8 mm long, 5-9-nerved. Lemmas 6-9 mm long, 3-4.5 mm wide, 7—9-nerved, nerves not conspicuous, strongly angled near middle, glabrous or puberulent, hyaline margins 0.5-1.1 mm wide at widest point, apex minutely bifid, the cleft 0.1-0.3 mm deep; awn absent or to 1 mm long. Paleas 6.1-6.6 mm long, glabrous. Stamens 3; anthers 0.7-1 mm long, dark brown. 2n = 14. Distribution and Habitat. — Introduced. Bromus briziformis is native to Eurasia. In British Columbia it is known only from a handful of populations near the Canada/ U.S.A. border (Fig. 3A). It has been collected in the northwestern United States and in a few scattered locations in central and eastern North America (Pavlick & Anderton 2007). Habitats include waste places, roadsides and other disturbed areas. Elevation: 600-914 m. Common Names.—rattlesnake brome, rattlesnake bromegrass, rattlesnake chess, rattlegrass. Notes.—Bromus briziformis is an annual species that superficially resembles the quaking grasses (Briza L.; the introduced B. major and B. minor are reported from British Columbia), but it is easily distinguished 328 | lofti tanical h Insti Texas 2( 1 cm . CIR. Fic. 2. Bromus briziformis. A. Habit. B. Spikelet. C. Lemma. D. Inflorescence. E. Ligule. Bromus ciliatus. F. Inflorescence. G. Spikelet. H. Lemma. I. Glumes. Saarela, Bromus in British Columbia, Canada 329 Bromus Bromus mus briziformis ciliatus , commutatus ibution in British Columbia. A. B. briziformis. B. Bromus ciliatus. C. B tatus, Scale bars = 200 km. Fic. 3. G hie dist from species of Briza by its closed leaf sheath. It is easily distinguished from all other Bromus species on the basis of its wide lemmas. bi I ional Boundary between Kettle and o rivers, ando P SA CANADA. British Col 10 Jun 1902, J.M. M. US); St ] Lake, ca. 2.5 km W of Osoyoos Lake, 12 Jun 1991, Douglas 12535 (UBC, V); 2.3 2 Gunby River, 500 m N o Sandy Creek Forest Service Road, about 5 km due N of Grand Forks, 2700 ft, 15 Jun 1996, F. Lomer 96-60 (UBC, V); 400 meters E of Charbonneau Creek, 200 m N of Pend d'Oreille Road, about 3.5 km E of 7 Mile Dam, 20 km SE of Trail, 12 jun 1996, F. Lomer 96-44 (UBC). 2. Bromus ciliatus L., Sp. PL 1:76-77. 1753. (Figs. 2F-I, 3B). Bromus inermis var. ciliata (L.) Trautv., Acta Horti Petrop. 1877. Bromus purgans var. ciliatus (L.) Kuntze, Revis. Gen. Pl. 2:763. 1891. Forasaccus ciliatus (L.) Lunell, Amer. Midl. Na- turalist 4:225. 1915. Bromus ciliatus var. genuinus Fernald, Rhodora 32:70. 1930. Zerna ciliata Ga Henrard, Blumea 4:498. 1941. Bromopsis ciliata (L.) SE no Geobot. Phytotax. 8:167. 1973. Tre: U.S.A. New York: Essex Co., Huntington Wildlife Forest 600 ft, 23 Jul 1939, H.E Heady 768 Deen DAO-53967, designated by McNeill, Station, Newcomb, Taxon 25:613. 1976; ISONEOTYPE: US). Bromus canadensis Michx., Fl. Bor.-Amer. 1:65. 1803. Bromus hooheri var. canadensis dee E; DU Mexic. Pl. 2:128. 1886. Zerna canadensis (Michx.) Tzvelev, Novosti Sist. Vyss. Rast. 7:54. 1970 [1971]. B ) Holub, Folia Geobot. Phytotax 8:167. 1973. Tere: CANADA: Lac St. Jean, Michaux s.n. (HOLOTYPE: P; ISOTYPE: US-A0865 520, pia ex F photostat Bromus purgans var. pallidus Hook., Fl-Bor. Amer. 2:252. 1840. Bromus richardsonii var. pallidus (Hook.) Shear, Bull. 1 Div. Agrostol., 23:34. 1900. Type: CANADA: Saskatchewan to the Rocky Mountains, Drummond s.n. (HoLOTYPE: K; isorYPE: US-A865461, fragment). Bromus ciliatus Í. denudatus bia Rhodo 24:91. 1922. Bromus ciliatus var. denudatus (Wiegand) Fernald, Rhodora 28:20. 1926. Bromus ciliatis subv. denudatus (Wiegand) Farw., Amer. Midl. Naturalist 10:203. 1927. Tyre: U.S.A. Massachusetts: Ashfield, 3 Aug 1909, E.F Williams s.n. (HOLOTYPE: Es ISOTYPE: US-8655271, fragment). Bromus ciliatus var. intonsus Fernald, Rhodora 32:70. 1930. Bromus ciliatus f. intonsus (Fernald) E Sevm., Fl. N. England 60. 1969. Tyre: A. MAssACHUSETTS: Ashfield, 4 Aug 1909, E.E Williams s.n. (HoLOTYPE: GH; isorvee: US-865526!, fragment ex Bromus s dudleyi Fernald, Rhodora 32:63, pe: ndi 1930. Tyre: ME NEWFOUNDLAND: bushy swale along Deer SES Bonne Bay, 26 Aug 1929, M.L. Fernald, Long & Fogg 1223 (n I; ISOTYE 5487!, fragment ex GH Perennial; rhizomes absent; plants 55-149 cm tall. Culms 1.2-4 mm wide at base, smooth; nodes 4-6, brown, pubescent, hairs 0.2-0.5 mm long. Leaf sheaths closed for most of their length, glabrous or sparsely to densely pubescent with soft, wavy hairs to 1.6 mm long; auricles absent; ligules 0.5-1.6 mm long, membranous, glabrous or minutely pubescent, erose; blades 10-27 cm long, 4-12 mm wide, adaxial surfaces pubescent with soft, wavy hairs 0.3-1.3 mm long, abaxial surfaces glabrous, margins smooth or 330 t tani i f Texas 2( serrulate. Panicles 8-21 cm long, 3.5-10.5 cm wide, open; branches ascending to spreading or drooping, shorter or longer than spikelet, glabrous or scabrous, 1-4 spikelets per branch, lowest inflorescence node with 1-5 branches. Spikelets 2.2-3 cm long (including awns); florets 4-9. Glumes subequal, lanceolate, glabrous, keels + scabrous; lower glumes 5-7.5 mm long, 1(-3)-nerved; upper glumes 6.5-9.5 mm long, 3(-5)-nerved, sometimes mucronate with mucros to 0.2 mm long. Lemmas 8-14 mm long, 1-2.5 mm wide, 7-nerved, pubescent along margins with hairs 0.5-1.3 mm long, hairs sometimes restriced to lower 1/3, backs glabrous or puberulent with hairs to 0.1 mm long, apex entire or minutely bifid, the cleft to 0.2 mm deep; awns (1-)2—6.5 mm long, arising 0-0.2 mm below lemma apex, straight, antrorsely scabrous. Paleas 74—9.2 mm long, shorter than lemmas, keels ciliate, cilia to 0.5 mm long. Stamens 3; anthers 1-1.7 mm long, dark brown. 2n - 14. Distribution and Habitat.—Native. Bromus ciliatus is the widest-ranging native B. sect B psis species in North America. lt occurs across the continent, absent only in south-central and south-eastern U.S.A. (Pavlick & Anderton 2007). Bromus ciliatus occurs throughout British Columbia, mostly east of the coastal mountain ranges (Fig. 3B). Its distribution is P with that of B. richardsonii, but B. ciliatus generally occurs at lower elevations (Peterson et al. 2002). B s ciliatus occurs in a wide range of habitats, including wet ditches, roadsides, boggy areas, dry terraces, moist woodlands, forest openings, stream banks, rocky flats and grassy areas. Elevation: 396-1554 m. Common Names.—English: fringed brome; French: brome cilié. Notes.—Bromus ciliatus and B. richardsonii have been variously treated as the same or r separate taxa (see comments under B. richardsonii). Peterson et al. (2002) d trated that tl lly and L e + genetically distinct and appropriately recognized at the species level. Because B. ciliat in a variety of habitats, there is interest, particularly in western Canada, for developing ecological adavan for re-seeding disturbed areas and clearcuts (see May et al. 1998, 1999; Fu et al. 2005). Fu et al, (2005) used AFLP data to examine molecular variation within and among B. ciliatus populations across Canada and found high among-population variation and low within-population variation. Representative Specimens Examined. CANADA. British Columbia: Yellowhead Pass, Rocky Mtns, 20 Jul 1848, W.S. Spreadborough 21012 Ka "i e SE Kootenay Valley, 23 Jul 1883, Dawson s.n. ño e Creek, Selkirk Mtns, 8 Aug 1885, Se Macoun 72994 Six-mile Creek, 8 Aug 1885, J. Macoun 72997 ( 3 Jul 1889, J. Macoun 30013 AN); McLennnan River headwaters of Columbia River, 27 Jul 1898, W.S. Spreadborough s.n. (CAN): Zeiten 27 Aug 1904, E. Es 415 (UBC); Arm- strong, 31 Aug 1904, E. Wilson 421 (UBC); Salmon Arm, 15 Aug 1911, MO Malte s.n. (CAN); Salmon Arm, 15 Aug 1911, MO Malte s.n. (DAO); McBride, 2360 ft, Jun-Jul 1914, J.A. Walker s.n. (UBC); Hazelton, 17 Jul 1917, J.M. Macoun 93974 (CAN, DAO, V); Vanderhoof, 5 Aug 1919, J.M. Macoun s.n. (CAN); Vanderhoof, 6 Aug 1919, J.M. Macoun s.n. (CAN, V); Pouce Coupe, 15 Jul 1921, M.O. Malte s.n. (CAN, DAO); Fraser Lake, 2222 ft, Aug 1922, Mackenzie s.n. (V); Vanderhoof, 18 Jul 1929, S.G. Preston s.n. (UBC); Dawson Creek, 3 Sep 1934, H. Groh s.n. (DAO); Smithers, Jul 1938, J.D. Menzies s.n. (UBC, V); Tetana Lake, 28 Jul 1938, T. Fletcher s.n. (V); Wardner, Aug 1939, WR Foster s.n. eats Nelson, Apex, 10 Jul 1940, J.W. Eastham 7766 (DAO, UBC); near Kinbasket Lake, 70 mi W of Golden, 21 Jul 1941, JW. E 96 (DAO); Kinbasket Lake, Big Bend Highway, 21 Jul 1941, J.W. Eastham s.n. (DAO, UBC, V); near Griffin Lake, between DEEN and Sicamous, 23 Jul 1941, J.W. Eastham 9353 (DAO, UBC); Griffin Lake, 27 Jul 1941, J.W. Eastham s.n. (V); Alaska GE Minaker River, mile 157, 19 Jun 1943, C H.D. Clarke 4 (CAN); old Cranbrook-Moyie road, 6 Aug 1943, J.W. Eastham s.n. (UB : vi- cinity of Buckinghorse River, 31 Aug 1943, H.M. Raup & D.S. Correll 11598 (UBC); S of EE 12 Jul 1944, J.W. Eastham s.n. SEO Woodpecker at Meadow Creek, 18 Jul 1944, J.W. Eastham s.n. (V); Cariboo, Horsefly district, 31 Jul 1944, J.W. Eastham s.n. (UBC, V); Anahim Lake, Chilcotin, Aug 1944, C.F. Cornwall 12362 (CAN, DAO, UBC, V); Smithers, 26 Jul 1946, J.W. Eastham s.n. ae Smithers, 26 Jul 1946 JP W. Bonum 14838 EN Fort St. Jenn, 4 Se 1946, H. Groh 3049 (DAO); Liard Hot Springs, 1 Aug 1946, J.P. Anderson 10362 (CAN), vicinity g d River, 25 Aug 1948, H.M. Raup & D.S. Correll 11519 GE UBC); Deka Lake, 4 Sep 1953, V.C. Brink s.n T BO): 1 i S of Hi I Q 1, 10 Aug 1954, J.A. Calder, D.B.O. Savile & J.M. Ferguson 14341 (DAO); 7 mi SW of Telkwa deeg Goathorn Creek, shilihers area, 31 Aug 1954, J.A. Calder, D.B.O. Savile & J.M. Ferguson 15271 (DAO); 1 mi W of Fairy Creek, Fernie, 3500 ft, 31 Jul 1956, FJ. Hermann 13014 (US); ee National Park, McCleod Meadows, 2 mi S along Nixon Creek, 3700 ft, 9 Aug 1956, FJ. Hermann 13227 (US); 5.5 mi E of Bridge Lake P.O. on roa ad to Littlefort, 11 Aug 1956, J.A. Calder, J. A. Parmelee & R.L. Taylor20007 (DAO), 1 mi W of Me bei oe — E Kleene and Anahim Lake, W of Williams Lake, 15 Aug 1956, J.A. Calder, J. A. ~ & RL. Te 201 ge, Flathead, 5100 ft, 22 Jul 1957, J. Davidson s.n. (DAO, UBC, V); Help Lake area 24 m cf Denald, 8 Aug 1958, R. L M & D.H. Ferguson 3654 Pe Eum Hot EE ca. pid ft, Se 1960, AE icu & R.T. iz 21954 odo Liard REA A 1960, J.A. Calder & | Park, id eek Ranger Station, 5 Pee LE y k + D; 470.1] dins Noam o ai? 1961, L. Ahti & T. Ahti 6877 (DAO); point of land looking P Valley, 25 pape St. John, last Saarela, Bromus in British Columbia, Canada 331 week of Jul 1961, D. GC & M. = Dhaliwal 7 (DAO); Willow River, 20 mi S of Giscome, 10 ins 1963, S. Eis ps Auclair e ete 4 mi N of Davie Lake, ca. 50 mi N of P George, 16 Aug 1963, A Auclair 484 (D 5 CUL TOT V Lake, 2000-2400 ft, 17 y 1963, A. Auclair & S. Eis s.n. ( (DAO); Port Mtn dam, 25 Jul 1965, A. F. p 7/65 (V): N of Williams Lake, 11 Jul 1967, C. Beil 67-7-11-14 (UBC); al Range NE di Pine Pass, Cariboo District, 19 Jul 1971, R. Revel s.n. (UBC); Glacier National Park, ^ € 29 Jul 1972, E. Haber & M.J. Shchepanek 1698 (CAN); Manning Park, Orchid meadow. mi E of Ranger Station, 31 Jul 1973, C. C. Chuang 870, 880 (CAN); Taspai Creek, ca. 27 mi ESE of Prince George, 2550 ft, 5 Aug 1974, VJ. Krajina, J. Pojar & C. Parsons s.n. (UBC); Jim Meadow, Chilcotin, 24 Jul 1979, C. Selby & A. Roberts s.n. (UBC); near Cooper Creek, 1340 m, 20 Aug 1980, L.E. Pavlick 80-560 (V); Cariboo District, near Ratdam, N of Horsefly, 16 Jul 1982, S.G. Aiken & S.J. Darbyshire 2350 (DAO); Upper Teslin River, 8 Aug 1983, C. McEwen 6207 (DAO); Chilcotin area, Cooper Creek, junction of road to Lake Mons and Sky ranch, 14 Sep 1993, J. Cayouette 7615 (DAO); Peace River Land District, 6 km N of Chetwind, 10 Sep 1998, J. Cayouette 8494 (DAO); Cariboo Land District, Perry Road, S of Shelley, 11 Sep 1998, J. Cayouette 8495 (DAO); Cariboo Land District, Dome Creek, road 16, 11 Sep 1998, J. Cayouette 8497 (DAO); Mount Robson Prov. Park, Redpass, close to Fraser River, 11 Sep 1998, J. Cayouette 8499 (DAO); Willow Creek Rest Area on Hwy 16, S of Prince George, 6 Jul 2004, P.M. Peterson, J.M. Saarela & S. Smith 18446 (CAN, US); Liard River Hot Springs Provincial Park, 20 Jul 2004, P.M. Peterson, J.M. Saarela & S. Smith 18659 (CAN, UBC) 3. Bromus commutatus pus e Germ. bd 1806. n 3€ hae Brachypodium commutatum (Schrad.) P Beauv., Ess. Agrostogr. 101, 155 W. Schultz, Flora 32:234. 1849. Serrafulcus com- mutatus (Schrad.) Bab., Man. Brit. Bot. 1:374. 1843 tatus (Schrad.) Coss. & T. Durand, Expl. Sci. Algérie 2:165. 1855. Bromus mollis var. commutatus (Schrad.) Sanio, Verh. Bot. Vereins Prov. Brandenburg 23:Abh. (3)31. 1882. Serrafulcus racemosus var. commutatus (Schrad.) Husn., Graminees 72. 1899. Forasaccus commutatus (Schrad.) Bubani, Fl. Pyren. 4:387. 1901. Bromus racemosus subsp. commutatus (Schrad.) Tourlet, pos n Vasc. Indre-et-Loire 588. 1908. Bromus hordeaceus var. commutatus (Schrad.) Fiori, Nuov. Fl. Italia 1:149. 1923. B (Schrad.) Lloret, Collect. Bot. (Barcelona) 22:151. 1993. Tyre: GERMANY: Inter segetes, ad vias, sepes, alibique. Bromus secalinus var. gladewitzii Farw., Amer. Midl. Naturalist 10:24. 1926. Type: U.S.A. Micuican: Edgewater Park, Farwell & Gladewitz 7434 (isorvre: MICH-1108616). Annual; rhizomes absent; plants 20-100 cm tall. Culms 1.5-3 mm WE at base, smooth; nodes 3—5, yellow- brown, densely pubescent with stiff hairs 0.2-1 mm long. Leaf sl losed for most of their length, densely pubescent, hairs soft and wavy to 1.2 mm long; auricles absent; ligules 1-3 mm long, membranous, glabrous or pubescent, erose; blades 8-16 cm long, 2-4 mm wide, adaxial surface densely pubescent with long, stiff hairs to 2.5 mm long, abaxial surface pubescent with dense hairs 0.1—0.5 mm long, margins smooth or ser- rulate. Panicles 5.5-17 cm long, 2.5—8 cm wide, open, longest lower branch > 4 cm n dc (as measured from node to apex of terminal floret including awn); branches erect to ascen ding, secund, scabrous to pubescent, 13 spikelets per branch, lowest inflorescence node with 1-3 diramdhies. Spikelets 2-3 cm long (including awns), ovate- Mn de ao sometimes visible at maturity, florets 6-12. Glumes slightly subequal, smooth or scabrous, | distally; lower glumes 5.5-8.2 mm long, 5-nerved; upper glumes 6.9-8.5 mm long: 7(9)-nerved. Lemmas 7-11 mm long, 1.5-2 mm wide, 7-9-nerved, nerves visible but not conspicuously raised, bacl scabrous to puberulent, margins often bluntly angled, apex entire or minutely bifid, the cleft 0. E 0.2 mm deep; awns 5-8.5 mm long, arising < 1.5 mm below lemma apex, straight or minutely divaricate, antrorsely scabrous, awn of lowest floret shorter than others. Paleas 6-10 mm long, shorter than lemmas, backs glabrous, keels ciliate, cilia to 0.3 mm long. Stamens 2; anthers 0.5—2.5 mm long, dark brown. 2n = 14, 28, 56. Distribution and Habitat. — Introduced. Bromus tatus is native to Europe. It is common in eastern and western North America and occurs in scattered locations across the United States Great Plains (Pavlick & Anderton 2007). In British Columbia, B. commutatus occurs in the southern portion of the province (Fig. 3C). Habitats include waste places, fields and grasslands. Elevation: 0-967 m. Common Names.—English: hairy brome, hairy chess, meadow brome, upright chess; French: brome variable, brome confondu. Notes.—Bromus commutatus and B. hologicall il ies t] f difficult to distinguish. They are both native to Europe, bur now SE) more del distributed. Hess & Chase (1951:49) noted that these taxa are "differentiated only by arbitrary characters.” In his classifica- tion of subgenus Bromus (=B. sect. Bromus), Scholz (1970) placed B. commutatus and B. racemosus in different series, and many authors recognize them as separate species (e.g., Clapham et al. 1987; Conert 1997; Smith 332 | | E al A * | LJ FIF D f Texas A Fic. 4. Bromus commutatus. A. Inflorescence. B. Lemma. C. Spikelet. D. Ligule. Bromus diandrus. E. Inflorescence. F. Floret. G. Spikelet. H. Glumes. I. Ligule. Saarela, Bromus in British Columbia, Canada 333 1980, 1985; Stace 1997; Douglas et al. 2001; Pavlick & Anderton 2007; Saarela & Peterson in press), but others have treated B. commutatus as a subspecies of B. racemosus (e.g., Shear 1900; Acedo & Llamas 1999; Lauber & Wagner 1996; Meijden 1996) or at least suggested that this treatment might be appropriate (e.g., Pavlick 1995; Planchuelo & Peterson 2000). Some workers lump these taxa and recognize only a single, variable species (e.g., Liang et al. 2006). Acedo & Llamas (1999) noted substantial phological variation among these taxa in the Iberian Peninsula, and indicated that environmental conditions influence plant and inflorescence size. In most treatments, B. commutatus is circumscribed as having longer lemmas (8-11 mm vs. 6.5-8 mm in B. racemosus), broadly angled lemma margins (vs. rounded) and shorter anthers (but measurements in earlier keys vary substantially) [reviewed by Spalton 2002]. Additionally, B. commutatus is generally con- sidered to have more open panicles with longer branches that sometimes have more than one spikelet (vs. less open panicles with shorter branches usually with a single spikelet), but these characters have not often been used in keys. Spalton (2002) conducted a morphological analysis of B. racemosus, B. commutatus and E Rap a a B. secalinus, jdn on material from throughout their native ranges. He overlap i liy used to distinguish B. racemosus and B. commutatus (e.g., lemma length) and found that the single best diagnostic character to distinguish the taxa is the length of the lowest panicle branches (including the terminal spikelet) (i.e., > 4 cm long in B. commutatus vs. < 4 cm long in B. racemosus). Consequently, Spalton’s (2002) circumscriptions of B. commutatus and B. racemosus are slightly different from those employed by earlier authors, but his work is also the most comprehensive study of variation in the complex. Spalton (2002) conc luded that B. commutatus is distinguished by its larger, more open and often branched inflorescences and larger spikelets, and B. racemosus is distinguished by its narrower, unbranched inflorescences and smaller spikelets. Using these characters, most British Columbia individuals can be readily placed into one of these two taxa. My observations of material from British Co- lly more robust, usually with some lumbia indicate that plants with long branches that have more than one spikelet and larger spikelets. 1 M Spalton’s circumscription here and recognize B. commutatus and B. racemosus at the species level, as in most recent treatments of these taxa in North America and Europe. I have ol 1 mixed populations of B. commutatus and B. racemosus in British Columbia near Princeton [Saarela, Sears & Maze 586A (CAN, UBC), 586B (CAN)] and Elko [Saarela 405A (CAN, UBC), 405B (CAN)]. A comprehensive genetic study of this complex, ideally based on material from throughout their native ranges, would contribute to e the natural boundaries between these taxa. Isl l, 16 Jun 1887, Macoun 29982 (CAN); Representative Specimens Examined. CANADA. British Columbia: Cedar vicinity of Victoria, 6 Jun 1908, J. Macoun 77852 (CAN, US); Saltspring "iem 24 May 1914, J.M. Macoun 90094 (CAN, US); Comox, Vancouver Island, 19 Jun 1915, J.M. Macoun 93968 (CAN); Vernon, 27 Jun 1917, E.M. Warren s.n. (UBC); Langley Prairie, 30 May 1931, H. Groh s.n. (DAO); S bank of Fraser River, N Ladner, 7.5.1932, E.P. Hume s.n. (UBC); Sumas Recreation, 13 May 1935, D.W. Thompson s.n. PER Victoria, 2 Jun 1938, J.W. Eastham s.n. (UBC); Creston, 21 Jun 1938, J.W. Eastham s.n. (UBC); The Gorge, dee 11 Jun 1939, J.W. Eastham s.n. (DAO, UBC); Sproat River Falls, 1 Jul 1939, J.W. Eastham s.n. (UBC); Arrowhead, 19 Jul 1939, H. Groh 21 ne S of Kamloops, 24 Jul 1939, H. Groh 29.5 (DAO); Christine Lake, 16 Jun 1940, J.W. Eastham s.n. (UBC); Thetis Lake, 27 May 1941, Fastham s.n. ee mE District, Alkali Lake, 640 m, 18 Jul 1982, S.G. Aiken, "SJ Rana GA. KE See (PAI a ng ramp, 29 Jun 1988, S.J. Dar! ive, Vancouver, 22 Jun 1994, F. Lome 94-251 a Kettle River Campground, 5 km N of A 3 - 33 along Hwy 33, 25 Jun 2003, J.M. Saarela & A. Roe 254 (CAN, UBC); Rocky Mountain Forest District, along Hwy 3, 1.5 km E of Elko, 23 May 2006, J.M. Saarela 405B ae 10 km W of Princeton on Hwy 3, 19 Jun 2006, J.M E C. Sears & J. Maze 586B (CAN); NE 3 km going up just before town lookout and just beyond the first switchback, 20 Jun 2006 JM. hara. C. Sears & J. Maze 644 (CAN, UBC); W of Osoyoos, 1.31 I Road, 19 Jun 2006, J.M ula C. Sears & J. Maze 631 (CAN, UBC); F Road, ca. 2 km NE ith Lowe Road, Blind C E of Cawston, 19 Jun 2006 aiid gedeien C. Sears & J. Maze 619 (CAN, UBC); Spotted Lake, ca. 5km W of Osoyoos, S of Hwy 3, o 2006, J.M. Saarela, C. Sears & J. ^ 79 (CAN) 4. Bromus diandrus Roth, Bot. Abh. Beobacht. 44. 1787. (Figs. 4E-I, 5A). Anisantha diandra (Roth) Tutin, Fl. British Isles, ed. 2. 1149. 1962. Anisantha diandra (Roth) Tutin ex Tzvelev, Bot. Mater. Gerb. Bot. Inst. Acad. Nauk SSSR 22:4. 1963. Tyre: Gr. Bromoides, l , lanuginosum, Italicum, Hist. Nat. 261. no. 444 (neoryrE: OXF-Scheuchzer herb., designated by Sales, Edinb. J. Bot. 50:8. 1993). 334 A A D Ph ol fTexas 2( Bromus diandrus e Bromus Bromus hordeaceus inermis e Fic. 5. G hic distribution in British Columbia. A. Bromus diandrus. B. B C. Bromus inermis. Scale bars = 200 km. wé Mitteleur. Fl. 2:595. 1901. Anisantha rigida (Roth) Hyl. Uppsala Univ. Arsskr. 7:32. 1945. Bromus diandrus subsp. rigidus (Roth) Lainz, Aanl. Inst. Forest. Invest. Exper. 12:49. 1967. Anisantha diandra Roth subsp. rigida (Roth) Tzvel., Zlaki SSSR. 1:223. 1976. Bromus diandrus subsp. rigidus (Roth) O. Bolòs, Masalles & Vigo, Collect. Bot. (Barcelona) 17: Al 1987 nire Bromus diandrus var. rigidus (Roth) Sales, Edinb. J. Bot. 50:9. 1993. Tyre: Habitat in Europa australi (neoryeE: B-Will gnated by Sales, Edinb. J. Bot. 50:9-10. 1993 Annual: rhizomes absent; 15-120 cm tall. Culms 0.8-3 mm wide at base, smooth, sometimes pubescent below panicles; nodes 23, blackish brown, glabrous. Leaf sheaths closed for most of their length, pubescent, with short stiff hairs to 0.6 mm long or soft hairs to 2.5 mm long; auricles absent; ligules 2.2-5 mm long, membranous, glabrous, erose-lacerate; blades 3-18 cm long, 2-7 mm wide; adaxial and abaxial surfaces pubescent with long, soft hairs to 1.5 mm long, margins smooth or hairy. Panicles 6-30 cm long, 2-25 cm wide, erect when young, sometimes becoming spreading or nodding at maturity; branches usually shorter than spikelets, scabrous to densely pubescent, usually with one spikelet per branch; lowest inflorescence node with 1-3 branches. Spikelets 6.5-8 cm long (including awns), cuneate, -compressed dorsiventrally, sometimes spreading at maturity with visible rachillas; florets 5-8. Glumes subequal, linear-lanceolate, glabrous, keels glabrous or serrulate, particularly distally, margins hyaline; lower glumes 12-26 mm long, 1(-3)-nerved; upper glumes 18-35 mm long, 3(-5)-nerved. Lemmas 18-35 mm long, 1-2 mm wide, /-nerved, lanceolate, backs sparsely to densely scabrous, sometimes with hairs to 1 mm long at apex or along margins, occasionally with a dense tuft of hairs 0.1-0.2 mm long at base, margins hyaline (except at base), smooth or minutely serrulate, keels serrulate, apex bifid, the cleft 2.2-5.2 mm deep; awns 30-50 mm long, arising 3.5-6 mm below lemma apex, straight, antrorsely scabrous. Paleas 13-16 mm long, shorter than lemmas, backs glabrous, keels ciliate, cilia to 0.5 mm long. Stamens 3; anthers 0.5-1.2 mm long, dark brown. 2n - 42,00. Distribution and Habitat. —Introduced. Bromus diandrus is native to Europe. In North Geer itis com- mon in the west from Washington to California and Texas, with ttered localities in tl United States (Pavlick & Anderton 2007). In British Columbia the species occurs on southern Vancouver Island and in the Vancouver area (Fig. 5A), where it is distributed in waste places, roadsides, thickets, river banks, sand bars, rocky outcrops, grassy banks and disturbed ground. Elevation: 0—290 m. Saarela, Bromus in British Columbia, Canada 335 Common Names.—ripgut brome, ripgut grass. Notes.—In British Columbia (Douglas et al. 2001) and elsewhere in western North America, this intro- duced taxon has previously been treated as B. rigidus Roth. Sales (1993) conducted a comprehensive survey of morphological, geographical and ecological variation in the Bromus diandrus/ rigidus complex based on material collected throughout its native European range, and recognized these taxa as varieties: B. EE var. diandrus and B. diandrus var. rigidus (Roth) Sales. However, Sales (1993) noted that the diff her varieties are very subtle and that it is often very difficult to one variety or the other. I therefore treat all British Columbia material as B. diandrus s. L witha no recognition of intraspecific taxa, as in the recent Bromus treatment for North America by Pavlick and Anderton (2007) and that forthcoming for California (Saarela & Peterson in press). Bromus diandrus has the largest spikelet characters (glumes, lem- mas, awns) among the three B. sect. Genea species in British Columbia; consequently, it is straightforward to identify. Representative Specimens Examined. CANADA. British Columbia: near Victoria, Jun 1876, Dawson 30040 (CAN); Nanaimo, Van- couver Island, 12 Jul 1893, Macoun 30039 (CAN); Nanaimo, Vancouver Island, 17 Jul 1908, J. Macoun 76832 (CAN, US); Victoria, 17 Aug 1912, MO Malte 108314 (CAN, DAO); vicinity of Sidney, 20 Jun 1913, J. Macoun 88538 (CAN); Victoria, 1 Jun 1938, J.W. Eastham 419c (UBC); C. N. railway yards, Nanaimo, Vancouver Island, 31 May 1939, J.W. Eastham s.n. (DAO); Vancouver Island, Anderson Hill, near Victoria, 65 m, 17 May 1950, V. Krajina & R.H. Spilsbury 3859 (DAO); Victoria, above McNeil Bay, 60 m, 21 May 1950, V. Krajina & R.H. Spilsbury 3950 (UBC); Nanaimo, Vancouver Island, 10 m, 12 Jun 1950, V. Krajina, R.H. Spilsbury & A. Szczawinski 9360 (DAO, UBC); Esquimalt Lagoon, Victoria, 3 May 1962, J. Spraggs s.n. (DAO); Esquimalt Lagoon near Victoria, 29 May 1962, J. Hett s.n. (DAO [3 sheets]); Victoria and vicinity, Vancouver Island, 10 Jul 1964, H.J. Scoggan 15431 (CAN); Gonzales Hill, Vancouver Island, 300 ft, 14 May 1965, B.W. Davies & R.A. Keller 365 (DAO); Smugglers Cove, Ten Mile Point, 10 ft, 19 May 1965, B.W. Davies 446 (DAO); Goldstream Park, Victoria, 20 ui 1980, R. Hainault 8244 (DAO); V 1, S Saanich District, Saanich Peninsula, Central Saanich Munici- pality, Island View Beach, Cordova Spit, 2 Jul 1982, S.G. Aiken, S.J. Daibysire & L.E. Pavlick 2127 (DAO); C han District, Saltspring Island, Mount Tuam above Cape Keppel, 950 ft, 3 Jul 1982, S.C. Aiken, SJ. E & L.E. Pavlick 2154 (DAO); Goose Spit, Comox, 4 May 1984, V.C. Brink s.n. (UBC); Sangster Island, 20 May 1985, A. Ceska, O. Ceska & M. Ceska s.n. (UBC); Holland Point, Victoria, 30 Jun 1988, SJ. add di va p Peri Bay, Victoria, 1 Jul 1988, S.J. pedcs s (CAN, DAO); University of British Columbia, Mus king lot, 15 May 2006, J.M. Saarela 332 (CAN, UBC); Horseshoe Bay, Whytecliff Park, 27 May 2006, J.M. S la 509 (CAN, UBO): S end of Tsawassen Drive S, Tsawassen Indian Reserve #0, 28 May 2006, J.M. Saarela & Sears 544 (CAN, UBC); Vancouver Island, Island View Park, S of Hwy 17 between Sidney and Victoria, 4 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 847 (CAN, UBC 5. Bromus hordeaceus L., Sp. Pl. 1:77. 1753. (Figs. 5B, 6). Bromus caló var. hordeaceus (L.) L., Flora Suecica, ed. 2. llis var. hordeaceus (L.) Fries, Novit. Fl. Suec. 16. 1814-23 bsp. hordeaceus (L.) Hiitonen, Suom Kasvio 219. 1933. Tyre: EUROPE: in collibus aridissimis sabulosis. (Lecrotyre: [icon] “Gramen avenaceum pratense, gluma breviore squamosa et villosa" in Morison, Pl. Hist. Univ. 3: s. 8, pl. 7, £ 18. 1699, designated by Smith in Cafferty et al., Taxon 49:248. 2000; EPITYPE: LINN-93.7, designated by Smith in Cafferty et al., Taxon 49:248). Bromus mollis L., Sp. Pl., ed. 2. 112. 1762. Serrafulcus mollis (L.) Parl., Pl. Rar. Sic. 2:11. 1840. Forasaccus mollis (L.) Bubani, Fl. Pyr. 4:386. 1901. Bromus hordeaceus var. mollis (L.) Fiori, Nuov. Fl. Italia. 1:149. 1923. Bromus hordeaceus subsp. mollis (L.) Maire, Cat. P]. Maroc. 4:943. 1941. Tree: EUROPE: in australioris siccis (LecroryrE: LINN-93.6, designated by Smith in Cafferty et al., Taxon 49:248. 2000). Annual; rhizomes absent; plants 7-110 cm tall. Culms 0.8-5 mm wide at base, smooth; nodes 2-4, yellow- brown, minutely to densely pubescent, hairs soft or stiff, to 0.6 mm long. I losed for most of their length, moderately to densely pilose, hairs soft and wavy to 1.2 mm long; auricles absent; ligules 12.6 mm long, membranous, glabrous or pubescent, erose; blades 2.2-18 cm long, 1-5.3 mm wide, adaxial surface P punc WEE ong; stiff E to 1.2 mm long, abaxial surface pubescent with dense hairs to 0.3 mm long or like adaxial surface th or serrulate. Panicles 2.5—14 cm long, 1-4 cm wide, dense, 1 ew reduced to a single spikele? branc! t to ascending t lax, scabrous to pubescent, most shorter than spikelets, 1 spikelet per branch; lowest en node with 1-3 branches. Spikelets 1.73 cm long (including awns), ovate-lanceolate, rachillas sometimes visible at maturity, florets 6-11. Glumes subequal, minutely to densely pubescent, keels serrulate; lower glumes 5.2-7 mm long, 3-5-nerved; upper glumes 6-8.5 mm long, 7-9-nerved. Lemmas 7.5-9 mm long, 1.9-2.5 mm wide, 7—9-nerved, nerves visible and conspicuously raised, particularly apically, backs densely pubescent with hairs to 0.3 mm long, 336 Fis. 6. Bromus hordeaceus. A, B, C. Habit. D. Spikelet. E. Lemmas. Saarela, Bromus in British Columbia, Canada 337 hyaline margins 0.3-0.6 mm wide, often bluntly angled; apex bifid, the cleft 0.3-0.7 mm deep; awns 4-7.6 mm long, arising 0.4-1.2 mm below lemma apex, straight or slightly divaricate, antrorsely scabrous. Paleas shorter than lemmas, backs glabrous, keels ciliate, cilia to 0.3 mm long. Stamens 2; anthers 0.3-1.3 mm long, dark brown. 2n - 28 Distribution and Habitat. —Introduced. Bromus hordeaceus is thought to be native to the Meditteranean basin (Smith 1986). It is now distributed widely in North America (Pavlick & Anderton 2007), Europe, Africa and Australia. In British Columbia B. hordeaceus occurs throughout the southern portion of the province and on the Queen Charlotte Islands (Fig. 5B). It is distributed in waste places, roadsides, fields, grassy areas and sandy beaches. Elevation: 0-1110 m. Common Names.—English: soft chess, soft brome-grass, soft cheat, tender brome-grass, soft-brome, soft brome; French: brome mou, brome doux, brome mollet. Notes.—Bromus hordeaceus can be very difficult to distinguish from B. racemosus, even with considerable experience. These taxa have been lumped E some workers, reflecting Hus EE e B^ Wilken & Painter 1995), but most workers recognize both taxa. Their major di degree of lemma nerve protrusion. In B. hordeaceus, the lemma nerves are id and M (Fig. 7B), whereas in B. racemosus the lemmas are smooth, with nerves raised Ld or not at A Fig 7A). This dnd is usually most easily observed near the apex of the lemma. B tend d compared with B. racemosus, but there is variation in this character and it is not always reliable. Bromus hordeaceus is a morphologically variable species (Fig. 6). In Europe, most workers recognize several intraspecific taxa (as subspecies) in the complex, differentiated by lemma length, awn width at its base, awn curvature, culm height and spikelet pubescence (e.g., Smith 1968, 1980, 1985; Zajac 1996). These subspecies are generally difficult to tell apart, but they have been recognized in recent treatments for North America (e.g., Pavlick 1995; Pavlick & Anderton 2007). Two subspecies [subsp. hordeaceus and subsp. thominei (Hardham ex Nyman) Braun-Blanquet] were recognized in British Columbia by Douglas et al. 2001). Douglas et al. (2001) also indicated that the putative hybrid B. x pseudothominei P.M. Sm. (B. hordeaceus subsp. hordeaceus x B. lepidus Holmberg) is infrequent on southern Vancouver Island. In material of the B. hordeaceus plex from British Columbia, I have observed that many individuals have a combination of the morphological characters typically used in keys to distinguish the subspecies, and they cannot be reliably differentiated. Specimens of the putative hybrid are also indistinguishable from other B. Nee EE SE et al. (1996, oai found no Moda of DNA sequence or al- 1] d from throughout ¡A ] f rhe putative B | es native eastern Mean ranges, and des noted substantial MM in morphological characters traditionally used to segregate intraspecific taxa among populations they examined. Intraspecific taxa in this group in North America are not recognized here. Representative Specimens Examined. ee pop Columbia: Victoria, 6 Jun 1893, Wm. Scott s.n. (DAO); Cascade Station, near International Boundary between Kettl ivers, 26 Jun 1902, J.M. Macoun 63,330 (US); Armstrong, 29 Jun 1907, E. Wilson s.n. R] (UBC); vicinity of Victoria, 13 Jun 1908, J. Macoun 76853 (US); Vancouver Island, Robertson River, 5 Jul 1911, W. Spreadborough s.n. (CAN); Victoria, 13 Sep 1909, A.S. Hitchcoch 4863 (US); Spence's Bridge, 16 Jul 1913, J. Davidson s.n. (UBC); Mayne Island, 20 May 1914, J.M. Macoun 90,077 (US); White Rock, 4 Jun 1916, J. Davidson s.n. (UBC); Vancouver, 10th Ave. and Willow St., Jun 1918, J. David- son s.n. (UBC); Lulu Island, 15 Jun 1924, T.R. Ashlee s.n. (UBC); White Rock, 25 May 1924, T.R. Ashlee s.n. (UBC); Mante [?] Creek, June 1927, Bostock s.n. (DAO); Agassiz, 3 Sep 1930, H. Groh s.n. (DAO); Langley Prairie, 4 Sep 1930, H. Groh s.n. (DÃO); Abbotsford, 12 Sep 1930, H Groh s.n. (DAO), Sumas Prairie, 15 Sep 1930, H. Groh s.n. (DAO); Sooke, 6 Oct 1930, H Groh s.n. (DAO); Mt Lehman, 12 May 1931, H. Groh s.n. (DAO); New Westminster, 19 May 1931, H Groh s.n. (DAO); New Westminster, 19 May 1931, H Groh s.n. (DAO), Horseshoe Bay, 23 May 1931, H. Groh s.n. (DAO); Saanichton, 27 May 1931, H. Groh s.n. (DAO); Langley Prairie, 30 May 1931, H. Groh s.n. (DAO); Agassiz, 2 Jun 1931, H. Groh s.n. (DAO); S bank of Fraser River, N Ladner, 7.5.1932, E.P. Hume 602 (UBC); Saanichton, Van- couver Island, 24 Jun 1935, A.M. Eastham s.n. (UBC); Lower Fraser, 18 Jul 1935, P.P. Henson s.n. (DAO), Coal Harbour, 25 Jun 1936, L.E. Taylor 2414 (UBC); Crescent Beach, 31 May 1937, LW Eastham 469 (UBC); Summerland, May 1936, V.C. Brink s.n. (UBC); W Point Grey, Vancouver, a 1937, a Ee um s.n. (UBC); Nanaimo, 31 May 1939 SE Eastham s.n. (UBC); Saanichton, Jun 1939, W.R. Foster s.n. na nichton, 5 Jul 1939, I. Mounce s.n. (DAO); Duncan, 7 Jul 1939, W. Newton s.n. Md Langley Prairie, (Cy ? Aug 26 Eu 1939, H. Groh 322 eds Otter, 1 Áug 1939, H. Groh 381 iod Vancouver Island, Shelter Point, 5 m 1939, I. Mounce s.n. (DAO); James Island, off Saanichton, 5 Aug 1939, H. Groh 438 (DAO); Kamloops, Brigade a 2800 ft, 24 Jul 1940, 338 tani ti Texas 2(1) A n A Is B. B. hord [M Calla 8256 (UBC 61258)] l f i h; i | trudi 4 Scale bars = 1 mm. Note that the E.W. Tisdale 40-7 (US); Nelson, 20 Jun 1944, W.G. McCalla 8256 (DAO); N of F River, Mission, 500 ft, 7 Aug 1949, G.F. Ledingham 49-513 (DAO); Duncan, 19 Jul 1950, E. G. Anderson 972 (DAO); Old W Saanich Road near Victoria, drainage of Colquitz Creek, 390 ft, 22 May 1963, A. Young & W. Hubbard s.n. (DAO); Gorden Head, 10 ft, 18 May 1965, B.W. Davies 397 (DAO); Gillespie Road, East Sooke, 30 May 1965, B.W. Davies s.n. (DAO); Government House, Victoria, 200 ft, 7 Jun 1965, R. A. Keller & B.W. Davies 822 (DAO); above Har- rison River, 1 mi from Harrison Mills, 400 ft, 12 Jun 1974, K.1. Beamish, J. Pinder-Moss, C. Selby, J. Pojar & E. Perkins 74063 (unes Long Lake Road, S of Kamloops, 24 Jul 1939, J. Davidson s.n. (UBC); Ladner, 14 Jul 1943, HD Falls s.n. (UBC), Snake Island, W Nanaimo, 30 May 1944, LE Taylor 2296 (UBC); Locarno Park, 11 Jun 1944, J.W. Eastham s.n. (DAO); Nelson, 20 Jun 1944, W. C. McCalla 8256 m Nelson, 21 Jun 1944, W.C. McCalla 8267 (UBC); Hastings Park, Vancouver, 8 Jun 1923, T.R. Ashlee s.n. (UBC); 43 Hwy W of Nelson, 23 Jun 1944, W.C. McCalla 8280 (UBC); edge of Hwy 3, 6 mi W of Nelson, 20 Jun 1944, W.C. McCalla 8256 (DAO); Gray Creek, 27 Jun 1944, W.C. McCalla 8308 (UBC): Fairfield Island, Chilliwack, 10 Jun 1946, J.W. Eastham s.n. (UBC); Victoria, above McNeil Bay, 60 m, 21 May 1950, V. Krajina & R.H. Spilsbury 3946 (UBC); Bear-Cowichan Lakes, dde m, e M p V Mie e R.H. Spilsbury 4096 (UBC), Quinsam Lake, Comox, 1 Jun 1951, J. Bendell s.n. (UBC); Gully at W S f Okanag , 27 Jun 1953, J.A. Calder & D.B.O. Savile 9711 (DAO); along highway on W side of Skaha Lake between Penticton deeg EE 28 Se 1953 i de A. Gg id D.B. id Savile 9784 (DAO); Goose Lake about 3 mi N of Vernon, 6 Jul 1953, J.A. Calder & D the highway, 3100 ft, 29 Aug 1953, VJ. Krajina s.n. (UBC); Port Alberni, 17 May 1955, D.R. Lindsay & W. W bou 658 (DAO); Gieres Lake, 31 May 1954, W.G. Dore & M. Smith s.n. (DAO); SE Bar, 30 May 1955, DR Lindsay & W. Woodbury 862 (DAO); Saltspring Island, islet off Beaver Point, 9 Jul 1955, T.R. Ashlee s.n. (UBC); D d Creek area, Nanaimo River Valley, 840 ft, 22 Jun 1956, D. Mueller- Dombois 90-3 (UBC); Queen Charlotte Islands, Graham Island, Queen Charlotte City, 29 Jul 1957, J.A. Calder, D.B.O. Savile & R.L. Taylor 23000 (DAO, UBC); Moresby Island, just S of Sandspit along road to Copper Bay, 5 Aug 1957, J.A. Calder, D.B.O. Savile & R.L. Taylor 23207 (DAO); W Vancouver, sea level, 27 May 1960, K. Beamish & F. Vrugtman 60501 (UBC); Java Rocks, Saturna Island, 8 Jun 1961, T.R. Ashlee s.n. (DAO); 100 yds W of entrance to Johnstone Creek campsite W of Rock Creek on Hwy 3, 15 Jun 1961, G. Beke 61-81 (DAO); Happy Valley near Victoria, 30 May 1962, J. Hett s.n. (DAO); Queen Charlotte Islands, Graham Island, S end of Richardson Ranch at Tlell, 4 Jun 1964, LA. Calder, R.L. d & L.C. Sherk 34658 (DAO, 2 VECI 2 mi E of Queen Charlotte City, Graham Island, 8 Jun 1964, JA. Calder, R.L. Taylor & L C. Sherk 34872 = AO); S i tead, Tlell, Medis Island, 14 Jul 1964, J.A. Calder & R.L. Taylor 35920 (DAO); Moresby Island, just FS o Copper Bay, 18 Jun 1964, J.A. Calder & R.L. Taylor 35166 (DAO); Lawn Point N of Skidegate Mission, Gralim Island, 24 4 Jun 1: 1964, JA A. Calder & R. L Taylor 35445 (DAO); Smugglers Cove, Ten Mile Point, 10 ft, 19 May 1965, B.W. Davies 448 (DAO); Esquimalt Lagoon, 1 Jun 1965, B.W. Davies & R.A. Keller 699 (DAO); Rocky Point, 100 ft, 8 Jun 1965, B.W. Davies & RA. Ke 844 PAO): Ten a Point area, 20 n Sr May 1965, B.W. Davies 482 (DAO); Lock Bay, Gabriola Island, 4 Jul 1965, K. Wade 65-57 (UBC); Gonzales Hill, l, Victoria, 15 Jul 1966, A.S. Harrison 111.1 (DAO); Ross Lake, Skagit River rid near Canada: U.S.A ela ca. 1000 ft, Eun 1974, K.1. Beamish, J. idi C. Selby, J. Pojar & E. Perkins 74040 (DAO, UBC); Mount Warburton-Pike, Saturna Island, 14 Jun 1974, G. Krause s.n. (UBC); UBC campus, S of Asian Centre, 29 Jun 1975, A. Guppy s.n. o 1 mi NE of Kaleden, 1800 ft, 23 May 1977, B. Wikeem & S. Gale s.n. (UBC); SW coast of Van- couver Island, China Beach, 2 Aug 1980, H.L. Dickson et al. 3804 (DAO); Harrison Hot Springs, along road which goes along Harrison Lake to Sasquatch Provincial Park, 6 Aug 1980, H.L. Dickson 3856 (DAO); W end of Manning Provincial Park, Hwy 3 at 2 Mile Creek, 2 km E of Hope, 29 Jun 1982, S.G. Aiken & S.J. Darbyshire 2081 (DAO); Cowichan District, Salt Spring Island, Mount Taum, above Cape Keppel, 950 ft, 3 Jul 1982, S.G. Aiken, S.J. Darbyshire & L.E. Pavlick 2140 (DAO); Pacific Rim National Park, Long Beach, Schooner Trail, Saarela, Bromus in British Columbia, Canada 339 4 Jul 1982, S.G. Aiken & S.J. Darbyshire 2171 (DAO); Pacific Rim National Park, Long Beach Unit, Wickaninnish Beach, Quisitis Point, 5 Jul 1982, S.G. Aiken & S.J. Darbyshire 2173 (DAO); Vancouver, UBC, along Chancellor Blvd., near University Hill Elementary School, Univ. Endowment HE e in ds 1986, s B. nid 3502 UEM Galiano Island, W of Stockade Hill, overlooking Montague Harbour, 17 Aug 1986, W s 73 S gs, N of Pemberton, Lillooet District, 150 m, “ue 1987, A.A. Rose s.n. (UBC); Abbotsford, intersection of Wess 1 aud 11 along nd 29 jun 1988, S.J. Darbyshire 3729 (DAO) , summit, 3 Jul 1988, SJ Darbyshire & J A. Darybshire 3777 (DAO); 0.25 km E of Vaseux See 1110 m, 13 Jun 1991, G. W. dis 12541 (UBC); Vaseux Lake, S end of Mud Lake, W side, 13 Jun 1991, T. Goward 91-435 (I otsford Dist. Munic., Mt, 25 May 1995, H. Ken- nedy, F.R. Ganders, M. Marshall, D. Justice & L. Goertzen s.n. (UBC); Sooke hills, Ragged Mountain ERP, 380 m, 27 Jun 1996, J.L. Penny & H Roemer 82 (UBC); along Hwy 33, 33.5 km S of Beav E 25 Tun 2003 J M. Sanda & A. Roe 255 en A OBESOS parana lot at UBC Museum of Anthropology, 25 May 2005, J.M. Saarela 300 (CAN, UBC); K y, 24 May 2006, J.M. inn 452 (CAN, UBC); Heede Bay, Whytecliff Park, 27 is 2006, J.M. Saarela 507 (CÁN, UBC); Horseshoe Se SEN Drive, just S of Hwy 1 (Trans. Canada Highway), 27 May 2006, J.M. Saarela 513 (CAN, UBC); Ladner, S side of Canoe Passage, W termi nus of River Road, 28 May 2006, J.M. Saarela & C. J. Sears 537, 538 (CAN, UBC); W of Osoyoos, 1.3 km up Richter Mountain Road, 19 Jun 2006, J.M. Saarela, C.J. Sears & J. Maze 632 (CAN, UBC); 0.5 km up Grizzly Road, S of Hwy 3, ca. 5 km E of Osoyoos, 20 Jun 2006, JM. Saarela, CJ. Sears & J. Maze 646 (CAN, UBC); roadside pullout on W side of Hwy 1 ee at Hells Gate, 22 Jun 2006, J.M. Saarela, C.J. Sears & J. Maze 734 (CAN, UBC); Richmond, London Farm, Gilbert Beach, just E ling, 28 May 2006, J.M. Saarela 6 C. J. Sears 523a (CAN, in eg Gees Park, " May 2006, J.M. Saarela & C. J. Sears 531 (CAN, UBC); Mayne Island, cliff edge of Mount Parke, ca. 1 km V | is, 3 Jun 2007, J. M. Saarela, DM Percy, Y. Chang, & Q.C.E. Cronk 803 (CAN, UBC); Vancouver gek doc View Park, S of Hwy 17 between Sidney and Victoria, along coast, 4 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 851 (CAN, UBC); Vancouver Island, Thetis Lake Regional Park, just N of Hwy 1, N of Langford, 4 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 859 (CAN, UBC); Vancouver Island, just S of Chemanus along Hwy 1A, 4 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 906 (CAN, UBC); Vancouver Island, just off Hwy 19A along Crane Road, N of Qualicum Beach, at confluence of Nile Creek and the Straight of Georgia, 7 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 934a (CAN, UBC); Queen Charlotte Islands: Moresby Island, Moresby Camp, Gillett Arm of Cumshewa Inlet, 11 km from Y junction of the loop road, 25 Jul 2007, J.M. Saarela & D. M. Percy 1271 (CAN, UBC). 6. noue inermis EE EL Mene 16. 1761. (Figs. 5C, SA—E). Dies inermis (Leyss.) DC. & Lam., Fl. Franc. 5 Ess. Agrost. 99:177. 1812. F yss.) Lunell, Amer. Midl. Naturalist 4:225. 1915. Zerna inermis (Leyss.) Lindm., venal came 101. 1918. ria a inermis An a NE Folia Geobot. Phytotax., Praha 8:167. 1973. We: EUROPE. G Amstgarten ad Belberg Crollwitz et alibi frequens, Leysser s.n. a LINN). Bromus pumpellianus var. melicoides Shear, Bull. Div. Agrostol., U.S.D.A. 23: = De i U.S.A. Coronapo: Beaver Ge O alt. ca. 3400 m, 8 Jul 1896, L. H. ua ee US-81589!) a synonym of B pumpellianus, but ly is B. inermis, based on th Bromus inopinatus Brues, Trans. Wis. Acad. Sci. 17: 73. 1911. Tyre: U.S.A. Wisconsin: Milwaukee, McKinley Park, along shores of Lake Michigan, 21 [no month] 1904, B.B. Brues 78 (isotype: US-3168443!, fragment). Bromus inermis f. proliferus Louis-Marie, Rev. Oka Agron. Inst. Agric. 14:14. 1940. Tree: Canada, Quesec: La Trappe, 24 Aug 1931, P Louis-Marie 318 (HOLOTYPE: QFA). Bromus inermis f. bulbiferus Moore, Rhodora 43:76. 1941. Tree: U.S.A. À R C Cl land Ave., 4 mi N of the campus of Minnesota College of Agriculture, 11 Oct 1938, Kaufman s.n. (HOLOTYPE: MINN). L Perennial; st ly rhi tous; basal l ] ing into straight fibres, though not readily, plants 21-150 Pi cm tall. Cas 2. 9- 6 mm wide at base, smooth; nodes 3-4, brown, glabrous or minutely pubescent, hairs to 0.2 mm long. Leaf sheaths closed for most of their length, glabrous or pubescent with soft hairs to 1.2 mm long or stiff hairs to 0.6 mm long; auricles absent or aa ligules 1.5-3.5 mm long, membranous, glabrous, erose; blades 10-21.5 cm long, 3.8-11.5 mm wide, adaxial surfaces glal h occasional long, soft hairs to 1.2 mm long or with dense soft hairs to 0.8 mm long, abaxial surfaces glabrous or pubescent with short hairs to 0.3 mm distributed along blade center, margins serrulate. Panicles 6.5-22 cm long, 3.5~14 cm wide, open; branches erect to ascending, often nodding at maturity, scabrous, usually longer than spikelet, 1-5 spikelets; did inflorescence node with 3-7 branches. Spikelets 1.7-3.2 cm long (including awns), florets 7-10. G I l, lanceolate, glabrous, ves scabrous or with hairs to 0.2 mm long: lower glumes 4.1-7 mm long, 13) -nerved; upper glumes 6-9.5 mm long, 3-nerved. Lemmas 10-13.5 mm long, 1.5-2.5 mm wide, glabrous or variously scabrous or puberulent on lower 1/6 to 1⁄4, nerves scabrous; awns absent or to 3 mm long, arising 0-0.5 mm SUM lemma apex, straight, antrorsely scabrous. Paleas 8—10.5 mm long, shorter than lemmas, keels g t, the cilia ca. 0.2 mm long. Stamens 3; anthers 4—5.5 mm long, yellow. 2n = 28, 56. 340 m T A ee =, SÉ a ) dm: mu Es == m A a [FA d i A if iy d Fic. 8. Bromus inermis. A. Rhizome. B. Inflorescence. C. Lemmas. D. Spikelet. E. Ligule. Bromus japonicus. F. Inflorescence. G. Ligule. H. Habit. |. Lemma. l J, Glumes. K. Spikelet. Saarela, Bromus in British Columbia, Canada 341 Distribution and Habitat.—Introduced. Bromus inermis is native across central Eurasia, and introduced in Africa, Australia, North America and South America. It is distributed widely across North America and it is extremely common (Pavlick & Anderton 2007). Bromus inermis has been collected throughout British Columbia, mostly on southern Vancouver Island, in the Vancouver area and east of the coastal ranges (Fig. 5C; also see Othnowski et al. 2007:188). Habitats include roadsides, fields and pastures, wet ] and open woods. Elevation: 0-1256 m. Common Names.—English: Austrian brome, Austrian brome grass, awnless brome, awnless brome grass, Hungarian brome, Hungarian brome grass, Hungarian fodder grass, Russian brome, smooth brome, smooth brome grass; French: brome inerme, brome de HORES brome sans arétes Notes.—The introduced B. inermis and the native B. pum] ly (e.g., Elliott 1949; Wagnon 1952; Scoggan 1978) or ani species (e.g., Pavlick 1995; Pavlick & Ander- ton 2007). Elliott (1949) demonstrated that the taxa are partially interfertile and presented morphological evidence (i.e., intermediate specimens) indicating that hybridization has likely occurred between the taxa. Nevertheless, the taxa have different morphological characteristics and native distributions and they occur in different habitats, suggesting that they have independent evolutionary histories. Bromus inermis is also morphologically similar to the relatively recently introduced B. riparius (see discussion under that taxon). Bromus inermis was introduced into North America in 1884 by the California Agricultural Experi- mental Station. In Canada it is widely cultivated for hay and pasture and used in revegetation projects, but although economically beneficial as forage, B. inermis is extremely invasive and poses serious threats to Canadian ecosystems (Otfinowski et al. 2007). It was recently ranked as the 8'^ most serious invasive alien plant threatening natural habitats in Canada (Catling & Mitrow 2005). Otfinowski et al. (2007) provide a detailed review of the biology of Bromus inermis. ew icu EA Representative Specimens Examined. CANADA. British Columbia: Bridge Creek, 21 Sep 1896, W. Allan s.n. (UBC); Goldstream, Oka- nagan, 11 Jul 1904, J. R. Anderson s.n. (UBC); Kamloops Lake, 4 Jul 1906, E. Wilson 678 (UBC); Coombe, Surrey, 27 Jun 1910, C. E. Britton s.n. (UBC); E of Field, 1-2 Aug 1914, A.S. Hitchcock 11543 (US); Big Creek, Jun 1915, M. Bertrand s.n. (UBC); Vernon, 20 June 1917, E.M. Warren s.n. (UBC); Quesnel Dam, 1 Nov 1917, W. Newcombe 317 (UBC); Merritt, 28 Aug 1920, G.W. Copley 10-9 (UBC); SE 3 Jul 1937, J.W. Eastham s.n. (UBC); Procter, Nelson, GE 1937 JM. BUE . (UBC); Ymir Rd, Nelson, 10 Jul 1937, J.W. Eastham s. Mirror Lake, Kaslo, 12 Jul 1937, J.W. Eastham s.n. (UB Mts, Hat Creek Valley, 26 Jun 1938, J.W. Thompson & E.M. ees (CAN); Kamloops, D tal Stati 12h Jun 1940, J.W. Eastham s.n. e Manning Park, Hope-Princeton, 9 Aug 1945, G.A. Hardy s. n. (UBC); Cache Creek, 29 Jul 1946,] A Rattenbury s.n. (UBC) ge, 20 Jul 1955, S. Lenaelou 16 (UBC); 2nd Trapp Lake, Kamloops, 19 Aug 1950, VJ. Krajina s.n. (UBC); Kamloops EE Trap Lake, 27 Aug 1950, VJ. Krajina s.n. (UBC); Edgewood, Arrow Lake, 1500 ft, 9 Jun 1957, K. Beamish & A. Gilmartin 7288 UBC: Keene Creek, Kaslo, Kootenay, 16 Jul 1960, M. Bell s.n. (UBC); Tranquille, 6 Oct 1962, V.C. Brink s.n. (UBC); 1 , Aug 1963, H. H. Ross s.n. (UBC); Chilcotin, plot no. 110, Aug 1968, C. Beil 68-8-14 (UBC); Fort Nelson near Mile 300 the Alaska a Highway, 3 Jul 1970, R.M. Annas, VJ. Krajina & R.G. McMinn s.n. (UBC): Maxhamish Lake, 7 Jul 1970, R.M. Annas, V.J. Krajina & R.G. McMinn s.n. (UBC); Cariboo Parklands, near confluence of Chilcotin and Fraser Rivers, 20 Jun 1972, J.F. Hyne 156 (UBC); Lakit Mountain lookout, 18 Aug 1974, F. Fodor 940 (UBC); Callanan Lake, Chilcotin, 8 Jul 1975, D. Jenkins 10122 (UBC); 3 mi W of Red Pass Se Kee 17 Jul 1975, C.C. Chuang 75/5 (UBC); Sundance Lake, Peace River District, 2200 ft, 3 Jul 1978, A.A. Rose AA delta, 6 Jun 1979, G.E. Bradfield 91 (UBC); Ray Farm, 11 km S of Clearwater Lake, Wells Gray Park, 21 Jun 1979, T. Goward 81-326 (UBC); along Rt. 12, ca. 47 = W of jct with Rt. 97, E e a Creek, 21 Jun 1983, G.B. Straley 2691 (UBC); New Westminster, 18 May 1989, F Lomer 89-015 (UB border crossing, W bank Similkameen River near border (Indian Reserve), 2 Jun 1990, T. Goward 90. 804 (UBC) along road to Tumbler Ridge, a few km S of junction with Rt. 97 at Arras, near Dawson Creek, 27 DE A G.B. paced & K.W. Nicholls 6866 (UBC); Spatsizi, Cold Fish Lake, 16 Aug 1992, V.C. ge n. (UBC); n Grand Pod 1, E of Grand Forks, 23 Jun 1993, L.E. Pavlick 93-141 (UBC); Kootenay, Wi B out Camp “Woodsmoke”, 27 Aug 1998, H. Stewart 98091808 (UBC); 18 mi N of Dease Lake (Brady Gee on Has 37 towards Watson M 20 Jul 2004, P.M. Peterson, J.M. Saarela €» S. Smith 18668 (CAN, US); 1 mi S of Iskut on Hwy 37 towards Meziadin Junction, along Coyote Creek, 20 Jul 2004, P.M. Peterson, J.M. Saarela & S. Smith 18673 (CAN, US); 18 mi N of Dease Lake (Brady Ge on Hwy 37 towards Watson See e 2004, P.M. Peterson, J.M. Saarela & S. Smith 18668 (CAN, US); 1 mi S of Iskut on Hwy 37 Junction, al k, 20 Jul 2004, P.M. Peterson, J.M. Saarela & S. Smith 186 s Ge eg binds Se 3 at E bd vacant ju beside river across from “Gold Pan Café and Store", 18 May 2006, J. M. Saarcla 3 Vardner, W of Jaffray, 23 May 2006, J.M. Saarela 412 (CAN, UBC); ca. 2 km E a Efirkbon along Hwy 3, 24 i 2006, J. M. es 439 (CAN, ne Osoyoos, Haynes Provincial Park, 25 May 2006, J.M. Saarela 481, 485 (CAN, UBC); Ladner, S di of Canoe e W terminus ons River Road, 28 May 2006, J.M. Saarela & C. J. Sears 540 (CAN, UBC); Manning Provincial Park, 0.9 p lodge that g lpi , 19 Jun S 1: 342 | tani i Texas 2(1) 2006, J.M. Saarela, CJ. Sears & J. Maze 566 (CA N); Princeton, f Princet Him SERO 19 Jun isi M aie Cd. dh ii J. Maze 597 (CAN, UBC); EE Finmore Road, W of Prince G S ofS River P 1 Park, al b Lake and N of Beverly 1 Road, 18 Jul 2007, IM. Saarela & D. M. Percy 1033 (CAN, UBC); ge gravel mining area, just 5 of Hwy 16, N of Bulkley River, just E of Ross LE 19 Jul 2007, J.M. Saarela & D.M. Percy 1088 (CAN, UBC); S of 100 Mile House, down 83 Mile Road to Green Lake Prov. Park, 28 Jul 2007, D.M. Percy 1366 (CAN, UBC). 7. Bromus japonicus Thunb. in Murray, Syst. Veg., ed. 14. 119. 1784. Thunb. Fl. Jap. 52. 1784, serius. (Fi F-K, 9A). Bromus arvensis var. japonicus, Nuov. Fl. Italia 1:149. 1923. Serrafalcus japonicus (Thunb.) Willmott., Man. Brit. Bot. ege, ed. 10. 510. 1922. Tyre: JAPAN: Thunberg s.n. (HoLoTYPE: UPS). Annual; rhizomes absent; plants 17-85 cm tall. Culms 1-2.2 mm wide at base, smooth; nodes 3-4, black- brown. Leaf sheaths closed for most of their length, densely pilose with soft, wavy hairs to 1.2 mm long; auricles absent; ligules 1-2.2 mm long, membranous, glabrous, apex erose; blades 3.5-13 cm long, 1-6 mm wide, adaxial and abaxial surfaces densely pubescent with short, stiff hairs to 0.6 mm long, margins EE or SE eres 3-26 cm long, 6-16 cm wide, open; branches spreading to ascending (lower ling), flexuous, smooth to scabrous, usually longer than spikelets, 1—6 spikelets per branch, lowest eege mode with 1-6 branches. Spikelets 1.8-4.2 cm long, broadly oblong to ovate- lanceolate, terete to moderately compressed, rachillas sometimes visible at maturity, florets 7-15. Glumes subequal, smooth, margins sometimes hyaline; lower glumes 4—7 mm long, 3—5(-7)-nerved; upper glumes 5-8 mm long, 7—9-nerved. Lemmas 7-9.2 mm long, 1.2-2.2 mm wide, 7—9-nerved, nerves not conspicu- ous, backs glabrous or scabridulous, scabrules ca. 0.1 mm long, hyaline margins 0—0.5 mm wide at widest point, apex minutely bifid, the cleft to 1 mm deep; awns 4.2-13 mm long (awn on lowest lemma usually shorter), arising 1.5-2.6 mm below lemma apex, straight to strongly divergent, widest at base, antrorsely scabrous. Paleas 6—7 mm long, backs glabrous, keels ciliate, cilia 0.3-0.5 mm long. Anthers 2; stamens 0.6-1.1 mm long, brown. 2n = 14. Distribution and Habitat.—Introduced. Bromus japonicus is native to Europe. In North America, B. japonicus is distributed widely throughout the U.S.A. and in southern Canada (Pavlick & Anderton 2007). Bromus japonicus has been collected in various places throughout British Columbia; it is most common in the interior of the province (Fig. 9A). Habitats include roadsides, grassy areas, hillsides and meadows. El- evation: 152-1050 m. Whisenant (1990) suggested that B. japonicus has become problematic in areas that have reduced occurrence of fire and grazing by large animals. Common Names.—English: Japanese chess, Japanese bromegrass, Japanese brome, spreading brome; French: brome du Japon, brome japonais. Notes.—The authority for this name has been variously attributed to ‘Thunberg’, ‘Thunberg ex Mur- ray’, ‘Thunberg in Murray’, and ‘Murray’, because it was published in both Flora Japonica (Thunberg, 1784, August) and Systema Vegetabilum, ed. 14 (Murray 1784, May-June), being attributed to Thunberg in the latter. Because of this confusion, Bartholomew et al. (Taxon 46:311-314. 1997) considered the author citations of the many new names published in these works and concluded that their authorship should be attributed to ‘Thunberg in Murray’ when a complete bibliographic citation is given (as above), or as ‘Thunberg’ when only me aon is E CES T CE ished on the basis of their Kee lemma EE vith and E degree of ade branching (see Savoie Key). Some individuals wit be difficult to determine, but the taxa have consistently been — a as dista species in eine native and introduced ranges. Molecular (Ainouche & Bayer 1997, Ainouche et al. 1999) and serological data (Smith 1972) indicate that they are closely related. Based on al- lozyme and morphometric data, Oja et al. (2003) suggested that the two taxa might be more appropriately recognized as intraspecific taxa of a single polymorphic species. Oja et al. (2003) did not, however, propose a formal taxonomic revision of the group. I recognize these taxa here at the species level. R Speci E ined. CANADA. British Col A trong, 31 May 1928, J.B. Munro s.n. (V); near Kamloops, 2000 ft, 27 May 1934, GV Copley 7 (US); Kamloops, Mission Flats, 1200 ft, 22 Jun 1937, E.W. Tisdale 40-10 (US); Tranquille Ranges, Tranquille, 21 Jul 1939, H. Groh 244 (DAO); 2 mi NW of Roosville, R Jul 1947, H.K. DeBeck s.n. (DAO, UBC, V); Goose Lake e Saarela, Bromus in British Columbia, Canada 343 Bromus Bromus Bromus japonicus pacificus porteri Crala hare nf Leen be 9 G hic distribution in British Columbia A. Bramus ianonicus R. Bromus pacificus. C. Bromus porteri wë d a F i about 3 mi N of Vernon, 6 Jul 1953, J.A Calder & D.B.O. Savile 10145A (DAO); 4.5 mi WSW of Kamloops, 2300 ft, 10 Jul 1953, J.A.Calder & D.B.O. Savile 10297 (DAO, US); two mi W of Kamloops, 26 Jul 1955, G.A. Mulligan & W. Woodbury 1817 (DAO); Stikine River, between Taya and Tahltan Rivers, 500 ft, 10 Sep 1977, A.A. Rose 77-778 (UBC); Lake Koocanusa, Kikomun Creek Provincial Campground, 15 m 1993, F. Lomer SH IN a 7.6 km W a ee arene Hwy 3, 25 May 2006, J.M. Saarela 487 (CAN, UBC); 5.6 km S Ge ee 3 on 1 Area, 20 Jun 2006, J.M. Saarela, C.J. Sears & J. Maze 673 (CAN, UBC). 8. Bromus pacificus Shear, Bull. Div. Agrostol., U.S.D.A. 23:38, f. 21. 1900. (Figs. 9B, 10A—E). Bromopsis pacifica (Shear) Holub, Folia Geobot. Phytotax. 8:168. 1973. Tyre: U.S.A. OREGON. ae Co.: in moist thickets near the seashore, S of Seaside, 11 Aug 1899, C.L. Shear & FL S Scribner 1703 € HOLOTYPE: US-] 15850). Bromus magnificus Elmer, Bot. Gaz. 36:53. 1903. Tyre: U.S.A. WasHINGToN. Clallam Co.: Aug 1900, A.D.E. Elmer 1957 (HoLorvrE: DS; ISOTYPE: US-401765!, with additional locality information: “Olympic Mtns."). Perennial; rhizomes absent; plants 45—172 cm tall. Culms 4-5 mm wide at base, smooth; nodes 5-9, brown, finely to densely retrorsely pubescent, hairs to 0.7(-1) mm long, occasionally glabrous with hairs above and below nodes. Leaf sheaths closed for most of their length, open for 14-35 mm in length, glabrous or sparsely to densely pilose with hairs to 2.1 mm long; auricles absent; ligules 2-4 mm long, membranous, glabrous, erose; blades to 3237) cm long, 7-16 mm wide, adaxial surface pilose, hairs to 1.5 mm long, soft and wavy, abaxial surface glabrous, margins smooth or serrulate. Panicles 17-28 cm long, 12.5-19 cm wide, open; branches mostly spreading and/or nodding, scabrous to pubescent, longer than spikelets, 1—6 spikelets per branch; lowest mee node wien l- É Branenes Spikelets 2.1—3.8 cm long, ovate-lanceolate, terete to slightly compressed, rachi visible at maturity, particularly distally; florets (4)6—8(10). Glumes slightly subequal, pubescent, hairs to 0.6 mm long, longer hairs sometimes restricted to keel; lower glumes 8.2-10.5 mm long, 1-nerved, lanceolate; upper glumes 9.7-12 mm long, 3-nerved, lanceolate to strongly acuminate with mucros 0—1.3 mm long. Lemmas 10-13.5 mm long, 1.8-2.3 mm wide, 7-nerved, sparsely to densely pubescent with appressed hairs to 1.5 mm long, longest hairs sometimes along margins, hairs sometimes denser on lower 1, apex entire or minutely bifid, the cleft 0-0.3 mm deep; awns 3—6.5(-7) mm long, arising 0—0.1 mm below lemma apex, straight, antrorsely scabrous. Paleas 9-10.5 mm long, usually shorter than lemmas, keels densely ciliate, cilia to 0.5 mm long. Stamens 3; anthers 2-3 mm long, dark brown. 2n - 28. Distribution and Habitat.—Native. Bromus pacificus occurs along or near the Pacific coast from south- Fic. 10. Bromus pacificus. A. Ligule. B. Inflorescence. C. Glumes. D. Lemma. E. Spikelet. Bromus porteri. F. Habit. G. Collar. H. Spikelet. 1. Lemma. J. Inflorescence. K. Ligule. Saarela, Bromus in British Columbia, Canada 345 eastern Alaska to central Oregon (Pavlick & Anderton 2007). Pavlik & Anderton (2007) indicated that B. pacificus also occurs in northern California, based on two specimen s from HSC identified as B. pacificus (M. Barkworth, pers. comm.). Examination of these specimens indicates that they were incorrectly determined [Barker 1035 (HSC-44691!) from Del Norte Co. is B. vulgaris; and Anderson 5120 (HSC-22840!) from Hum- boldt Co. is B. carinatus Hook. & Arn. var. marginatus (Nees) Barkworth & Anderton]. Bromus pacificus is not known from California (Saarela & Peterson, in press). In British Columbia, B. pacificus occurs in coastal regions of the province (Fig. 9B); absence of collections along much of the coastline likely reflects a paucity of collecting activities in difficult-to-access areas, rather than the absence of the species. Habitats include moist ravines, shaded woods, grassy swards, saline beaches, ditches and along roadsides. Elevation: 0—50 m. Common Names.—Pacific brome, Shear Pacific brome. Notes.—Many B. pacificus specimens from British Columbia have previously been determined incor- rectly as species of B. sect. Ceratochloa (i.e., B. carinatus or B. sitchensis). With large, open panicles, B. pacificus is superficially similar to these taxa, but it can be easily distinguished from them by its lemmas that are rounded or slightly keeled over the back (vs. lemmas that are strongly laterally flattened and keeled in spe- cies of B. sect. Ceratochloa), and 1- and 3-nerved lower and upper glumes (vs. 3-5- and 5-9-nerved lower and upper glumes, respectively). Additionally, B. pacificus occurs only on or near the coast, whereas plants in B. sect. Ceratochloa are distributed much more widely. Representative Specimens Examined. CANADA. British Columbia: Qualicum, Vancouver Island, 29 Jul 1887, Macoun 30008 (CAN, US); Renfrew, 18 Jul 1902, C.O. Rosendahl 776 (CAN, US); Sidney, Vancouver Island, 30 May 1914, M.O. Malte s.n.(DAO); Courtenay River, Comox, 10 Jul 1915, J. Macoun s.n. (V); Courtenay, 23 Jul M» M.O. oe 108338 (CAN, DAO); ios Judo 1916, W. R. Carter s.n. (V); Prince Rupert, 1 Aug 1916, M.O. Malte 106910 (CAN, ARO land, Port Alberni, head 18 Aug 1922, J. Davidson s.n. (UBC); Triangle Island, 28 Jun 1949, G. Hardy 21645 (Y; Q Charlotte Islands, Moresby Island, eege Inlet, 13 Sep 1951, R. Pillsbury s.n. E Hore of Duncan pay ca. 6 mi WNW V of Prince Rupert, 24 Aug 1954, J.A. Calder, D.B.O. Savile & J.M. Ferguson 14973 (DAO); India y, Langara Island off NW tip of Graham Island, 16 Jul 1957, J.A. Calder, D.B.O. Savile & R.L. Taylor22539 Du near Juskatla on. jock ths Inlet, Graham Island, 8 Aug 1957, D.B.O. Savile 3520 (DAO); Yakoun River Delta S dal Port enu. Masset Ge cin sland, 15 Aug 1957, J.A. Calder & R.L. Taylor 23507 (DAO); Queen Charlotte Islands, Lr Ma 1957, J.A. Calder & R.L. E 23 des — Bs Markale, Kyuquot, 1 Jul 1958, Taylor 143 (UBC): Kyuquot, Malay cove, 8 Jul 1958, Taylor 383 (UBC) by Island, W end of S arm of Kootenay Inlet, 22 Jul 1964, J.A. Calder & R.L. a geg (D40, UBC): Kaisum ancient Haida village, Moresby Island, 4 Aug 1964, LA Calder & R.L. Taylor 36523 (DAO); Tee Island, E sid ] , Graham Island, 12 Aug 1964, J.A. Calder & RI Taylor 36848 (DAO); Long Island, Vancouver Island, 19 Aug 1965, Taylor 6740 (UBC); W of Truck Narrows, Truck Inlet, ca. 7 mi N of Prince Rupert, range 5, Coast District, 31 Jul 1973, V. Krajina s.n. (UBC); Vancouver Island, Tofino, Meares lelana, nén "iw: creek S di Lone cd I Jul 1981, A. Ceska, O. Oldriska, & R.T. Ogilvie s.n. (UBC); Vancouver Island, Nitinat C P, J Nitinat Lake Road, 21 Aug 1989, L.E. Pavlick 89-158 (UBC), ` land, Port Albe erni, th of S River, 11 Sep 1993, : See i. Re DA is EE Rocky Point Park at head of Pu ine SEH 14 km E of Vancouver, 5 Augut 1996, F. Lomer 128 (UB ls, Moresby Island, 11 km SE of S it I , Om, 18 Jul 1997, F. Lomer & N. Grove 97407 (V); Sep by Mission Creek, W edge of Kincolith (Ginglox), 22 Jul 2004, P.M. Peterson, J.M. Saarela & S. Smith 18719 (CAN, US); entrance to Ferry at Prince Rupert, 23 jul 2004, P.M. Peterson, J.M. Saarela & S. Smith 18729 (CAN, US); Vancouver Island, Rupert Inlet, 4 Sep 2006 J M. auda 760 o eee nae Charlotte o bonds E Wie Sound Recreation Site, at end of Rennell Sound F m NW , 23 Jul 2007, J.M. Saarela & D.M. Percy 1202 (CAN, UBC); Rain! Pass, C M ue iet f Prince R 26 Jul 2007, J.M. Saarela & D.M. Percy 1308 (CAN, UBC); along Hwy 16, adj t to SI River between Tyl d SI ,26 Jul 2007, J.M. Saarela & D.M. Percy 1311 (CAN, UBC). 9. Bromus porteri (J.M. Coult.) Nash, Bull. Torrey Bot. Club 22:512. 1895. (Figs. 9C, 10F-K). Bromus kalmii var. porteri J. M. Coult., Man. Bot. Rocky Mt. 425. 1885. Bromus ciliatus var. porteri (J. M. Coult.) Rydb., Contr. U.S. Natl. Herb. 3:192. 1895. Bromopsis porteri J. M. Coult.) Holub, Folia Geobot. Phytotax. 8:168. 1973. Tree: U.S.A. Corogapo. Lake Co.: Twin Lakes, 25 Jul 1872, T.C. Porter s.n (Lectoryee: NY, designated by Wagnon, Brittonia 7:471. 1952). Bromus ciliatus var. montanus Vasey ex Beal, Grass. N. Amer. 2:619. 1896. Tree: U.S.A. Coronapo: 1892, H.N. Patterson 264 (LECTOTYPE: MSC, designated by Beal, Grass. N. Amer. 2:619. 189 00 Bromus ciliatus var. scariosus Scribn., Bull. Div. Agrostol., U. s. DA 13:46. 1898. Tree: U.S.A. WyominG. Albany Co.: Sheep Mountain, 3 Jul 1897, A. Nelson 3305 (HoLorvre: US-1008288!) Bromus kalmii var. occidentalis Vasey ex Beal, Grass. N. Amer. 2:624. 1896. Tree: U.S.A. MONTANA: Anderson 11 (LecrorrrE: MSC-00913261, designated here; sorme: US-865498!, fragment and photostat ex. MSC). Mul wd 1008608!) D 346 t tani tit Texas 2( ] for B. kalmii var. occidentalis, NOTES: Beal (1896:624) lists tl (1) Montana, 1883, WM. Canby & EL. Scrib- ner 384; Q) Monten Anderson 11; m Colorado, Cassidy s.n. e isa sheer in US (US- 1008517; digital photograph!) labeled “B. EL. Scribner, 1883, Northern Pacific R. R. [rail mA Ge or B. kalmii var. occidentalis” tated in A.S. Hitchcocks hand “Probably Vaseys typ Herb. “Mich. Agr. Coll.” TI Iso | l tation "Montana 2 nd ? Hitchcock 1935 811) indicated that dee e geng ge is the iu pe. The Botanical Type Sr R I J yhy pes, accessed 12 jun 2007), | itl Ee name, EE Pdl number (384), nor tl hi fl Į : it is therefore not absolutely clear is tl ited ] Palais et al. (2003:172) ] li 1 that the C 1 Scril is housed at both MSC and US; ne are no mee of this collection ir in MSC (ie LA. nins a The un two ganze) indi been located at MSC ipd 009 ad e MSC-0091327!), E US-86549 tion UE llex MSC” Pavlick et EL Quos a oy indicate al ES J 1 + 1 +1 e 1 E C DI 2] A ja] " g E grar y N (Vienna Code), II:n] 1 1 2 oe o Da 3215-44] c zl 1 1 + a pen Da sl + £c als APA RA ZE r O , the protologue. Among the And | Cassidy collections, the Anderson collection i t plete. It is most appropriate to deciaonate rhe renn 1] t1 | 1 } if 7 } t 1 Jar? to] flier rb rl tal [9] k Perennial; rhizomes absent; plants 32-100 cm tall. Culms 0. ins 8 mm bon at n PE sometimes pubescent below intl ; nodes 3-8, brown, glabrous, sometimes with nodes; lower nodes covered by sheaths. Leaf sheatl ] | for most of their length, gla! icles al t; ligules 0.5-1.9 mm long, membranous, glabrous, erose; blades 7-28 cm long, 1.9— 5 mm "ide adaxial surface glabrous, abaxial surface glabrous or with sparse, soft hairs to 1.5 mm, margins smooth or serrulate. Panicles 5-12 cm long, 1.5-7 cm wide, open; branches ascending to drooping, shorter or longer than spikelet, pubescent, 1-2 spikelets per branch, lowest inflorescence node with 1-2 branches. Spikelets 1.7-3.1 cm long, ovate- lanceolate, terete to slightly compressed, rachillas sometimes visible at maturity, particularly distally; florets 4—9. Glumes lanceolate, pubescent, hairs 0.2-0.3 mm long; lower glumes 5.2-7.5 mm long, (1-)3-nerved; upper glumes 6.2-9.5 mm long, 3-nerved. Lemmas 8-10 mm long, 1.5-2.5 mm wide, 7-nerved, densely pubescent with hairs 0.2-1.5 mm long, hairs sometimes denser on lower 14, apex entire or minutely bifid, the cleft 0-0.2 mm deep; awns 0.8-3.1 mm long, arising 0-0.3 mm below lemma apex, straight, antrorsely scabrous. Paleas 7-8.2 mm long, shorter than lemmas, keels ciliate, cilia to 0.3 mm long. Stamens 3; anthers 1.9-2.5 mm long, dark brown. 2n = 14. Distribution and Habitat. — Native. B teri in sout hern Alberta, Saskatcl and Manitoba and south to western Texas and California (Pavlick & Anderton 2007). In British Columbia B. porteri occurs mostly in the interior of the province. Habitat includes dry talus slopes, meadows, open ground, woodland flats and grasslands. Elevation: 152-1310 m. Common Name. — nodding brome. Notes.—Bromus porteri has been variously treated as a distinct species (e.g., Gould & Moran 1981; Pavlick 1995; Pavlick & Anderton 2007; Scoggan 1978; Soderstrom & Beaman 1968) or as a synonym of Bromus anomalus Rupr. ex E. Fourn. (e.g., Douglas et al. 2001; Wilken & Painter 1993). Bromus anomalus and B. porteri are morphologically and geographically distinct taxa. Bromus anomalus is easily separated from B. porteri by its 1-nerved lower glumes [vs. (1)3-nerved in B. porteri] and conspicuously narrowed midvein of the leaf below the collar (Wagnon 1952; Pavlick 1995; Pavlick & Anderton 2007, J. M. Saarela, personal observation). The two species are largely allopatric. B teri has a wide distribution in western North America where it is distributed from Manitoba to British Columbia and south to California, New Mexico and northeastern Mexico (Wagnon 1952; Pavlick 1995; Herrera Arrieta 2001; Pavlick & Anderton 2007; Saarela & Peterson in press), whereas B. anomalus is distributed in Texas and throughout northern and cen- tral Mexico (Soderstrom & Beaman 1968; Pavlick et al. 2003; Herrera Arrieta 2001; J. M. Saarela, personal observation). Plants in British Columbia are clearly referable to Bromus porteri. Representative Specimens Examined. CANADA. British Columbia: Klinaklini Valley, 3000 ft, 17 Sep 1920, T. MacKenzie 3-1 (UBC); junction of Banbricker and Big Creeks, Chilcotin, 16 Jul 1929, J. Davi ison 63 (UBC); Chilcotin (E), Riske Creek, 13 Aug 1938, E.W. Tis- dale 40-1 (US); Hanceville, Chilcotin, 24 Jul 1944, J.W. Eastham 12236 (CAN, UBC), Francois Lake, 6 Jul 1944, J.W. Eastham s.n. (UBC); Canoe Creek, Gang Ranch road, 13 Jul 1945, J.W. Eastham s.n. (DAO, UBC), above Pavilion Lake, 30 Jul 1946, V.C. Brink s.n. (UBC); Old Saarela, Bromus in British Columbia, Canada 347 "iid Lookout, Ootsa Lake, 8 Jul 1944, J.W. Eastham s.n. aes Hanceville, Chilcotin, 24 Jul 1944, J.W. Eastham s.n. (V); Cariboo zone, mis pas e. 20 Jul 1967, C. Beil 67-07-20 (UBC); C , plot no. 105, 3500 ft, Aug 1968, C. Beil 68-8-21 (UBC); Farwell a 19 Jul 1972, C.E. Beil 282 (UBC); W side of Relay Creek Basin trail ca. 1.5 mi from Relay Creek cabin, 27 Jul 1977, C. Selby plot 78 #627 ps on Creek Basin, 12 Aug 1977, C. Selby s.n. (UBC); Stikine River (Ranch), 500 ft, 9 Sep 1977, A.A. Rose 77-771 (UBC); of Burn Creek, 0.5 km from Stikine River, 744-824 m, 31 Jul 1979, W. Gorman 1349 (UBC); 10 km W of Tatla Lake Village, 5 Jul 1980, LE Pavlick 80-14 (V); Cariboo District, Riske Creek, E pasture of Bald Mountain, 12 Jul 1982, S.G. Aiken, S.J. Darbyshiren & A. Roberts 2287 (DAO); Cariboo District, Alexis Creek, 1 km N of settlement, 17 Jul 1982, S.G. Aiken & S.J. Darbyshire 2369 (DAO), Thompson-Okanagan, Cornwall Hills, along Cornwall Hills Road, 13 Sep 1989, LE Pavlick 89-212 (UBC, V); Thompson-Okanagan, Hat Creek, along Hat Creek Road, 14 Sep 1989, L.E. Pavlick 89-263 (UBC); Chasm, 26 Jul 1989, F. Lomer 89-14 (UBC); Cariboo area, NW of Clinton, White Lake, 13 Sep 1993, J. Cayouette C7601 (DAO); Cariboo-Chilcotin, Dog Creek area, 7 Jul 1994, T. Goward & C. Clement 94-513 (UBC); Cariboo-Chilcotin, Dog Creek area, 21 Jul 1994, T. Goward & C. Clement 94-594 (UBC); Peace River Land District, W of Fort St. John, close to road 29, 10 Sep 1998, J. Cayouette 8491 (DAO). 10. Bromus pumpellianus Scribn., Bull. Torrey Bot. Club 15: 9. 1509. (Figs. HE pa Forasaccus pumpel- lianus (Scribn.) Lunell, Amer. Midl. Naturalist 4:225. 1915. B ibn.) Wagnon, Rhodora 52:211. 1950. Zerna pumpelliana (Scribn.) Tzvelev, Fl. Arct. U.R.S.S. 2:225. 1964. DEN irpta ie Holub, Folia Geobot. Phytotax. 8:168. 1973. Tree: U.S.A. Montana. Meagher Co.: Belt Mountains, Meagher County, Montana, 1883, E Lamson-Scribner 418 (HOLOTYPE: US-81590!). Bromus purgans var. purpurascens Hook., Fl.-Bor. Amer. 2:252. 1840. Bromus inermis subsp. pumpellianus var. purpurascens (Hook.) Wagnon, Rhodora 52:211. 1950. Tre: CANADA: Bear Lake to Arctic seacoast, Richardson s.n. (HOLOTYPE: K; sorre; US-8654551, fragment ex K). Bromus pns var. coloradensis Vasey ex Beal, Grass. N. Amer. 2:619. 1896. Tyre: U.S.A. COLORADO: near Grays Peak, Wolf 1158 (LECTOTYPE: 007969!, designated by Hitchcock, Man. Grass. U.S. 816. 1935). Bromus E var. tweedyi Scribn. ex Beal, Grass. N. Amer. 2:622. 1896. Type: U.S.A. Montana: Yellowstone Park, Slough Creek, 1885, E Tweedy 587 (sore: US-815921). Bromus arcticus Shear in Scribn. & Merr., Contr. U.S. Natl. Herb. 13:83. 1910. B lli . arcticus (Shear) Porsild, Rhodora 41:182. 1939. Tyre: U.S.A. Alaska: 9 Sep 1901, EA. Walpole 2036 (HOLOTYPE: US-3791571, with the following additional location information: “Vicinity of Port Clarence. Tundra banks near buildings at Teller Reindeer Station”. Perennial; rhizomes present; basal leaves readily decaying into straight fibres. Culms 3.2-4.5 mm wide at base, + densely pubescent below nodes; nodes 2-3, light brown-yellow, moderately to densely pubescent, hairs to 1 mm long. Leaf sheaths closed for most of their length, glabrous or pilose with long, soft hairs to 2 mm long; auricles present or absent; ligules 1-4 mm long, membranous, glabrous, erose; blades 9-17 cm long, 2.1-10 mm wide, adaxial surfaces glabrous or pubescent with dense, short hairs to 0.5 mm long, abaxial surfaces glabrous or variously pubescent with hairs 0.5-1.2 mm long, hairs sometimes present only marginally, margins serrulate. Panicles 10—29.5 cm long, 1.5—14.5 cm wide, open, branches erect to ascending, shorter or longer than spikelet, scabrous, 1(-2) spikelets per branch, lowest inflorescence node with 4 branches. Spikelets 2.3-3.3 cm long (including awns); florets 7-14. Glumes subequal, lanceolate, glabrous, midnerves scabrous; lower glumes 6.5-10 mm long, 1-nerved; upper glumes 9-12.8 mm long, 3-nerved. Lemmas 10-13 mm long, 1.5-2.6 mm wide, variously pubescent with hairs 0.5-0.8 mm long along margins and midvein or moderately to densely villous throughout; awns 1-4.5(-5) mm long, arising 0—0.5 mm below lemma apex, straight, antrorsely scabrous. Paleas 9.2-11 mm long, shorter than lemmas, cilia present on keels along lower % to entire length, cilia ca. 0.2 mm long. Stamens 3; anthers 4.1-6.5 mm long, yellow to dark brown. 2n = 56 Distribution and Habitat.—Native. Bromus pumpellianus occurs in western North America from Alaska to New Mexico and west through south-central Alberta, Sasakatchewan and Manitoba to northern Ontario (Pavlick & Anderton 2007). In British Columbia B. pumpellianus occurs in the Rocky Mountains (Fig. 12A). Habitat includes grassy banks, grassy meadows, wet banks, sand and gravel bars along rivers and roadsides. Elevation: 597-2000 m. Common Name. EE SE Notes.—B pump phologi us ¡able t In North America, some authors have treated it | variety of tl hologi similar and putatively closely-related taxon Bromus inermis (e.g., Elliott 1949; Wagnon 1952; CAMS 1978). Several intraspecific taxa have been recognized 348 A Phizoma R Node € Snikelet D. lemma. E Inflorescence Fic 11 E Inflorescence. G Snikelet H, | lemmas Bromus richardsonii. J. Inflorescence. K. Spikelet. L, M. Lemmas. N. Glumes. Bromus riparius. O. Spikelet. P. Lemma. Saarela, Bromus in British Columbia, Canada 349 Bromus rons pumpellianus PSCBIOSUS “Uv ORARE Fig 12 € phi IS au! lae ' PCI Fal |" A D Hz PR E P E ech i EE: C vl ege 200 km. DN 1] t pubescence le.g., var. arcticus (Shear) Porsild, var. purpura- within B. pumpelli | 1 on degree Scens (Böök ) Wagnon] but there is substantial intergradation among these types and recent authors do not recognize them (e.g., Mitchell 1967; Pavlick 1995; Douglas et al. 2001; Pavlick & Anderton 2007). Bromus pumpellianus is also morphologically similar to the introduced B. riparius (see discussion under that taxon). Representative Specimens Examined. CANADA. British Columbia: Moberly, 14 Jul 1930, R. Graham s.n. (UBC), vicinity of Sikanni River, 21 Jun 1943, H.M. Raup & D.S. Correll 10223 (UBC), Perry Creek, Cranbrook (Selkirks), 8 Aug 1943, J.W. Eastham 11035 (UBC); vicinity of AT River, 1 Sep 1943, H.M. Raup & D.S. Correll 11619 (CAN, UBC); Fort St. John, 23 Jun 1948, n.c. (DAO); Ft. St. John, 29 Jun 1949, V.C. Brink s.n. (UBC); Mile 98, Haines Road, 7 Jul 1956, T M.C. Taylor, A.F. Szczawinski & M. Bell 1015 (DAO, UBC), 984 (DAO, SCH 1035 (DAO); Mile 14, Flathead Rd, 15 Jul 1957, M. Bell & J. Davidson s.n. (UBC); Sage Creek Lodge, 21 Jul 1957, M. Bell & J. Davidson s.n. Eon U.S. Sun adc ipe ft, a Jul 1957, M. Bell & J. Davidson 599 (UBC); U.S. Border, Flathead, 4600 ft, 27 Jul 1957, k and Elk River, 4200 ft, 30 Jun 1958, R.L. Taylor & D.H. Ferguson 2264 (DAO); w of Kee Geng at Mile 160, 5 Jul Sasa J A. Calder & J.M. Gillett 26541 (DAO); 4 mi NW of Dawson Creek along highway to Ft. St. John, 12 Jul 1960, J.A. Calder & I. Kukkonen 26820 (DAO); Nevis Creek, 17 Jul 1969, A. Luchhurst s.n. (UBC); Nevis Creek, 24 Jul 1969, A Luckhurst s.n (UBC); Mt Turnbull, Fording River, Creek 4, 29 Sep 1 970, D.C. Morrison s.n (UBC); Mt Beie oe 2 wis River, 28 Jul 1971, D.C. Morrison s.n. (UBC); Emkay, Bleasdell Mt, 10 Aug 1972, n.c. (UBC); intersection of Ew nola River, 9.8.1975, R. Hainault 7713 (DAO); Robb Lake, slopes on N side of lake, 3900 ft, 29 Jul 1977, G.W. Argus & E. pads 10197 pu it of Pinl in, 10 Aug 1992, F. Lomer 92-160 (UBC). 11. Bromus racemosus L., Sp. Pl., ed. 2. 1:114. 1762. (Fig. 11F-I, 12B). serrafalcus racemosus (L.) Parl., Rar. PL Sic. 2:14. 1840. Bromus arvensis var. racemosus (L.) Neilreich, Fl. Nieder-Oesterr. 81. 1859. B. squarrosus var. racemosus (L.) Regel, Act. Hort. Petrop. 7:602. 1881. Bromus mollis var. racemosus (L.) Bubani, Fl. Pyr 4:387. 1901. Bromus hordeaceus var. racemosus (L.) Fiori, Nuova Fl. Analitica Ital. 1:149. 1923. Type: Anon (LecroTYPE: LINN 93.31, designated by Smith, Notes Roy. Bot. Gard. Edinb. 42:499. 1985). Annual; rhizomes absent; plants 17-60 tall. Culms 1-2.1 mm wide at base, smooth; nodes 2-3, brown, minutely to densely pubescent, hairs to 0.6 mm long. Leaf sheaths closed for most of their length, densely pubescent, hairs soft and wavy to 1.2 mm long; auricles absent; ligules 0.5-2 mm long, membranous, gla- brous, erose; blades 2.7-12 cm long, 1-3 mm wide, adaxial surface densely pubescent with stiff hairs to O. 8 mm long, abaxial surface densely pubescent with shorter, stiff hairs to 0.6 mm long or like adaxial sur- face, margins smooth or serrulate. Panicles 3-8.5 cm long (or less when reduced to a single spikelet), 1-2.5 cm wide, longest lower branch « 4 cm dns (as measured from node to apex of terminal floret, including awn); branches erect to ascen ding, und, scabrous to pubescent, 1 spikelet per branch, lowest 350 Journal of t tani itute of Texas 2( inflorescence node with 1-4 branches. Spikelets 1.5-2.2(-2.6) cm long (including awns), ovate-lanceolate, rachilla sometimes visible at maturity, florets 6-9. Glumes slightly subequal, smooth or scabrous, keels oc- casionally serrulate distally; lower glumes 5.2-6.7 mm long, 3—5-nerved; upper glumes 6.2-8.5 mm long, 7-9-nerved. Lemmas 6.5-9.5 mm long, 1.5-2 mm wide, 7—9-nerved, nerves visible but not conspicuously raised, backs glabrous or scabrous to puberulent, hairs to 0.5 mm long, margins + bluntly angled, apex entire or minutely bifid, the cleft 0.1-0.2 mm deep; awns 3.4-8.5 mm long, arising < 1.5 mm below lemma apex, straight or minutely divaricate, antrorsely scabrous, awns of lowest florets shorter than others. Paleas 5.5-8.5 mm long, shorter than lemmas, backs glabrous, keels ciliate, cilia to 0.3 mm long. Stamens 2; anthers (0.5)1-1.6 mm long, dark brown. 2n = 28 Distribution and Habitat—Introduced. In North America, B. racemosus occurs in scattered locations across the S.A. e southern Canada (Pavlick & EE ZOON In British Columbia B. racemosus occurs mostly in t tern and southcentral parts of the province (Fig. 12B). Habitats include waste places, field, roadsides and gravelly hillsides. Elevation: 0-967 m Common Names.—bald brome, meadow brome, smooth brome, smooth chess. Notes. —Bromus racemosus is often confused with Bromus hordeaceus because both taxa have relatively narrow inflorescences, and the two have sometimes been treated as synonyms (e.g., Wilken and Painter 1993). Bromus hordeaceus is distinguished by its lemmas with conspicuously raised nerves and generally denser inflorescences. See additional comments under B. commutatus and B. hordeaceus. R tat E ined. CANADA. British Col Victoria, 7 May 1875, Macoun 29980 (CAN); near Victoria, Áug 1876, D 29979 Sup Nanaimo, tjun 1884 4) Macoun 29982 M Pitt River, Aug 1884, Macoun 29981 (CAN); Vancouver, 23 Jun 1887, J. Fowler s.n. (US); Y facoun idge, 28 May 1889, Macoun (92) (CAN); Victoria area, 27 May 1893, Macoun 116 (CAN); Victoria, SS 1893, Wm. Sege n. (DAO): Nanaimo, 20 Jun 1907, C.O. Rosendahl 1887 (US); vicinity of Victoria, 6 Jun 1908, J. Macoun 76851 E Ne See River, Vancouver Island, 5 Jul 1911, W. Spreadborough 87602 (US); vicinity of Sidney, 16 Jun 1913, J. M 885 , 24 May 1914, J.M. Macoun 90094 (CAN); 10th Ave and Willow St., Vancouver, Jun 1918, J. Pone n. (UBC); Vernon, 16 ne 1918, A.H. ERR n. (UBC); Vernon, 25 Jun 1917, E.M. Warren s.n. (UBC); Vernon, 17 Jun 1918, A.H. Hutchinson s.n. (UBC); Victoria, Jun 1920, G.V. Copley 1-15 (UBC); Courtenay, 3 Oct 1930, H. Groh s.n. (DAO); Steveston, 20 May 1931, H. Groh s.n. (DAO); Saanichton, 27 May 1931, H. Groh s.n. (DÀO); Ganges, 28 May 1931, H. m men W Point Grey, 14 Jun 1934, J.W. Eastham s.n. (UBC); W Point Grey, Vancouver, 20 Jun 1937, J.W. Eastham s.n. (UBC); Fai ] Chilliwack, 10 Jun 1946, J.W. Eastham s.n. (UBC); Fairfield Island, GE SC 1946, J.W. Eastham 15133 (DAO), Christina I P 29 May 1953, G.H. en bap n. slags along highway on W side of Skaha Lal d Kaleden, 28 Jun 1953, JA. Calder & D.B.O. Savile 9784 ul 1953, J.M. Smith s.n. (DAO); Christina Lake, 31 May 1954, W.G. Dore & M. Smith s.n. pe Christina Lake, near "-— Forks, 31 May 1954, W.G. Dore 15053 (DAO), Parksville, 13 Jul 1954, T.M.C. Taylor 3086 (UBC); 7 mi N of Duncan, 18 May 1955, D.R. Lindsay & W. Woodbury 696 (DAO); S Pender Island, 11 Jul 1955, S. Buchanan s.n. (UBC); Saltspring bud 12 Jun 1955, T.R. Ashlee s.n. (UBC); Summerland Research Station, 1300 ft, 14 Jul 1960, A. Mclean 60-83 (DAO); John Dean Park, Saanichton, 1000 ft, 14 Jun 1962, J. Hett s.n. (DAO); Henderson Point, W coast of Saanich Peninsula, 14 e? Ms , J. Hett s.n. (DAO), Uplands Park, Victoria, 15 Jun 1962, J. Hett 62 (DAO); Gordon Head, Victoria, 19 May 1962, M. Bell s.n. (DAO); B y Head, 10 ft, 4 Jun 1965, A. Fisk & B.W. Davies 803 o ~ Head near Victoria, 30 May 1962, J. Hett s.n. (DAO); North Hill, Saanich, 350 ft, 2 Jun 1965, B.W. Davies & R.A. Keller , 14 Jun 1970, S. Kojima s.n. (UBC) ison River, 400 ft, 12 Jun 1974, K.I. Beamish, J. Pinder-Moss, C. Selby, J. Pojar& 5 E. Perhins e DAO) Patricia mae pales one-half mile N of Royal Oak Corner, 20 May 1955, D.R. Lindsay & W. Woodbury Jul 1988, S.J. Darbyshire & J.A. Darbyshire 3777 (CAN); alongside parking lot at UBC Museum of pales? 25 May 05 3 M. E 305 (CAN, UBC); Rock Forest District, along Hwy 3, 1.5 km E of Elko, 23 May 2006, J.M. Saarela 405A (CAN, UBC); Osoyoos, Haynes Provincial Park, 25 May 2006 Ji M. Suena 484 (CAN, UBC); 10 km W of Princeton on Hwy 3, 19 Jun 2006, J.M. Saarela, C. Sears & J. Maze 586A (CAN, UBC); ca. 2] t Lake, 14.6 km N on Garnet Valley Road from Hwy 97 at Summerland, 21 Jun 2006, J.M. Saarela, C. Sears & J. Maze 707 (CAN, UBC). 12. Bromus richardsonii Link, Hort. Berol. 2:281. 1833. (Fig. LIM, 126) Zera richardsonii (Link) Nevski, Trudy Se Aziatsk. Gosud. Univ., Ser, 8b, Bot. 17:17. 1934. Bromus ciliatus var. ri ) B. Boivin, Naturaliste Canad. (Que.) 94:521. 1967. Zerna canadensis subsp. richardsonii mpi Tzvelev, uen RE s has de 54. 1970 [1971;. Bromopsis richardsonii (Link) Holub, Folia Geobot. Phytotax. 8(2):1 1973 (Link) Tzvelev, Zlaki SSSR 214. 1976. Tyee: Hab. in A ntali. Semina mist cl.: Grown from seed in Berlin, PD cun (HOLOTYPE: B, destroyed fide J. Cayouette & PM. Peterson 2007). Bromus purgans var. longispicatus aon a Bor. nid 2: an PAM Tree: CANADA: Rocky Mountains, pd sin: E OH "ms 1 US-A865460!, fragment). N Tu" B. richardsonii, the bases al duced 41 4 Saarela, Bromus in British Columbia, Canada 351 Perennial; rhizomes absent; plants 47-110 cm tall. Culms 1.5-3.5 mm wide at base, smooth; nodes 3-4, brown, glabrous, lower nodes sometimes minutely pubescent. Leaf sheaths closed for most of their length, glabrous or sparsely to densely pilose with stiff hairs 0.2-1.2 mm long; auricles absent; ligules 1-2.5 mm long, membranous, glabrous or minutely pubescent, erose; blades 18.5—34 cm long, 3-9(-13) mm wide, adaxial and abaxial surfaces glabrous, margins smooth or serrulate. Panicles 13-23.5 cm long, 2-17.5 cm wide, open, branches ascending to drooping, shorter or longer than spikelet, smooth-scabrous, 1—5 spikelets per branch, lowest inflorescence node with 1—4 branches. Spikelets 2.4—3.5 cm long, rachilla sometimes visible at maturity; florets 3-9. Glumes subequal, lanceolate, glabrous or minutely pubescent, hairs some- times restricted to margins, keels glabrous or scabrous; lower glumes 7-9.5(-12) mm long, 1(-3)-nerved, lanceolate; upper glumes (8-)9.5-11.5(-14.5) mm long, 3-nerved, sometimes acuminate or mucronate with mucros to 0.7(-1) mm long. Lemmas 9-13.5(-15) mm long, Ze = 8 mm EE 7-nerved, pubescent, hairs along margins 0.2-1 mm long, hai ly or densely back > 0.1 mm long, apex entire or minutely bifid, the cleft to 0.2 mm lone: awns 3.1-6.5 mm long, arising 0—0.5 mm below lemma apex, straight, antrorsely scabrous. Paleas 11-12 mm long, shorter than lemmas, keels glabrous or ciliate, cilia to 0.2 mm long. Stamens 3; anthers 1-2.6(-3.4) mm long, dark brown. 2n = 28. Distribution and Habitat.—Native. Bromus richardsonii is generally distributed west of the continental divide from the Yukon territory to central Mexico, with disjunct populations known from the Cypress Hills in southern ieee aman in Blac a in Soup Dakota and western Texas (Peterson et al. 2002; Pavlick & Anderton 2007) hout British Columbia east of the coastal mountain ranges (Fig. 12C). Its distribution is sympatric with that of B. ciliatus (Fig. 3B), but B. richardsonii generally occurs at higher elevations (Peterson et al. 2002). Habitats include meadows, open woods and gravelly areas. Elevation: 289-2133 m Common Name.—Richardson's brome. Notes.—Bromus richardsonii has been variously recognized as a distinct species (Shear 1900; Hitchcock 1913; Wagnon 1952; Mitchell & Wilton 1965; Mitchell 1967; Hultén 1968; Welsh 1974; Pavlick 1995; Herrera Arrieta 2001; Peterson et al. 2002; Saarela et al. 2005; Pavlick & Anderton 2007; Saarela and Peterson, in press), as a synonym of B. ciliatus (e.g., Allred 1993; Hitchcock 1951; Soderstrom & Beaman 1968; Wilken & Painter 1993; Douglas et al. 2001) or as a variety of B. ciliatus (e.g., Kartesz 1999). In a detailed study of the complex, Peterson et al. (2002) identified distinctive morphological, cytological and genetic differences that support recognition of these taxa as distinct species (see Taxonomic Key). Representative Specimens Examined. CANADA. British Columbia: Sicamous, 3 Jul 1889, Macoun 30013 (CAN); below Wapta Lake, Rocky Mts, 6 Aug 1904, Macoun 64804 (CAN); Field, Rocky Mountains, 19 Aug 1904, Macoun 64803 (CAN); Sicamous, 16 Jul 1914, M. Malte s.n. (CAN, DAO); near Bow River, Be 1-2 SE 1914, A. S. Hitchcock 11537 (US); Field, 2 Aug 1914, A.S. Hitchcock s.n. (DÃO): E of Field, 1-2 Aug 1914, A.S. Hitchcock 11,552 gs, 2 Aug 1915, M.O. Malte s.n. (CAN, DAO); Prince Rupert, 1 Aug 1916, M.O. Malte s.n. (CAN); W and NW stones of Mt Selwyn, 4000 ft, 26 Jal 1932, H.M. Raup & E.C. Abbe 4160 (CAN); Burns Lake, 5 Jul 1935, OJ. ne 2588 (US); Watching Creek, Tranquille, 8 Jul 1936, E.W. Tisdale 6902 (DAO, UBC); Telkwa, 30 Jul 1937, J.D. Menzies s.n. (UBC); C Lake, Riske Creek, Chilcotin, 14 Aug 1938, H.A. Senn 1941 (UBC); Fairmont Hot Springs, 16 Jul 1941, J.W. Eastham s.n. (UBC, V); Mt Sieben area, Yoho National Park, 1 Aug 1943, W.C. McCalla 7807 (UBC); valley of Kicking Horse River, 3 mi E o Field, 14 Sep 1943, G.H. Turner 3833 (DAO); Fort St. James, 3 Jul 1944, J.W. Eastham s.n. (UBC); Ootsa Lake P.O., 8 Jul 1944, J.W. East- ham s.n. (UBC); Burns Lake, 16 Jul 1944, J.W. Eastham s.n. (CAN, UBC, V); Canoe Creek, Clinton area, 8 Aug 1946, C. Rogers 35 (DAO); Pavilion Lake, 29 Aug 1949, V.C. Brink 49-470 (UBC); ca. 5 mi on Gibson Pass rd to Paddy Lake, Manning Park, 28 Jul 1960, A. McLean s.n. [60-203?] (DAO); Chilcotin, plot no. 90, 3650 ft, Jul 1968, C. Beil 68-7-17 (UBC); Chilcotin, plot no. 110, Aug 1968, C. Beil 68-8-19 (UBC); area of 10 mile Carbon Creek, 18 Aug 1971, n.c. (UBC); Emkay, Bleasdell Mt, 10 Aug 1972, n.c. (UBC); near Fraser Lake, NW of Drvwilliam Lake, 2450 ft, 10 Aug 1973, VJ. Krajina & R. Revel s.n. (UBC); ridge, 1.5 mi SE of Tyhee Lake, 2300 ft, 14 Jul 1974, VJ. Krajina, J. Pojar & C. Parsons s.n. (UBC); Ashnola Road, mile 24, 29 Jul 1975, R. Hainault 7630 (DAO); Driftwood Creek, ca. 7 mi ENE of Smithers, 2400 ft, 17 Jul 1974, VJ. Krajina, J. Pojar & C. Parsons s.n. (DAO, UBC); Driftwood Canyon and Smithers, 19 Jul 1974, J. Pojar J.92 (UBC); above Salix Creek, vicinity of mouth of Cow Parsnip Creek, Gladys Lake area, 4700 ft, 3 Aug 1975, J. Pojar s.n. (UBC), W entrance to Sanctuary Pass, Gladys Lake area, 5000 ft, 19 Aug 1975, J. Pojar s.n. (UBC); ca. 1 mi E of Relay Creek Cabin, 18 Jul 1977, C. Selby plot 35 4399 (UBC); White Horse Bluffs area, S of Wells Gray Park, 25 Jul 1979, T. Goward 81-375 (DAO); Bissett Creek, 10 km SW of Chetwynd, 610 m, 3 Aug 1979, L.E. Pavlick 79-779 (V); Stones Bay Road, 7 km from Post Office, Fort St. James, 13 Jul 1981, A. Kruckeberg 101 (UBC); Beece Creek area, 1510 m, 10 Aug 1981, L.E. Pavlick 81-113 (V); Chilcotin area, Big Creek — Farwell Canyon road, near Big Creek, 14 Sep 1993, J. Cayouette 7620 (DAO); Chilcotin Mts, Taseko Lake area, close to Beece Creek, 15 Sep 1993, J. Cayouette 352 | tani i f Texas 2( 7621 (DAO); Road 5A, 1 km N of B Lake, 16 Sep 1993, J.C tte 7622 (DAO): Pope Mountain, near Ft. St. James, 792 m, 19 Jul 1996, J.L. Penny & Botany BC 1996 (UBO): 13 mi SW of Cintendust past Kelly le 26 jul 2004, P.M. Peterson, J.M. Saarela & S. Smith 18740 (CAN, US); 1 mi S of Iskut on Hwy 37 towards Meziadin Ee along Coyote Creek, 20 Jul 2004, P.M. Peterson, J.M. Saarela & S, Smith 18675 (CAN, US); Skihist Provinical Park, Hwy 8, ca m from Lytton, 22 Jun 2006, J.M. Saarela, C. Sears & J. Maze 728 (CAN, UBC); 5.5 km along North Shore Road (Stella Road), N d SEN 16, just W of Nadleh RE Se Ge Nation Reserve bound- ary, N of Fraser Lake, 18 Jul 2007, J.M. Saarela & D.M. Percy 1068 (CAN, UBC); along J aser Lake, ca. 100 km W of Prince George, along dirt road 12 km W of Nadleh Whuten Band First Nati rea M locally as “Red Beach", 18 Jul 2007, J.M. Saarela & D.M. Percy 1083 (CAN, UBC); old gravel mining area, just $ of Hwy 16, N Be Bulkley River, just E of Ross Lake, 19 Jul 2007, J.M. Saarela & D.M. Percy 1089 (CAN, UBC 13. Bromus riparius Rehm., Notiz. Veg. Gest. Schwartz Meer 85. 1872. (Figs. 110-P, 13A). Zerna iin (Rehm.) Nevski in Acta Univ. Asiae Med. Ser. 8b, Bot. 17:17. 1934. B (Rehm.) Holub, Folia Geobot. Phytotax. 8:1 1973. B. biebersteinii auct. Am., non Roem. & Schult. Tree: RUSSIA: ae s.n. (ISOTYPE: LE). Perennial; pitose with short rhizomes; decaying basal leaves reticulate-fibrillose, plants 34-120 cm tall. Culms 2.8-6 mm wide at base, smooth; nodes (2—)3—4, brown, smooth or pubescent with hairs 0.2-0.5 mm long. Leaf sheaths closed for most of their length, glabrous to moderately pubescent with soft, wavy hairs to 0.5-1.6 mm long: auricles absent; ligules 1.1-3 mm long, membranous, glabrous, erose; blades 11.5—32 cm long, 2-6.5 mm wide, adaxial surfaces glabrous or sparsely to densely pilose with hairs 0.5-2.5 mm long, abaxial surfaces pubescent with short hairs 0.3-0.5 mm long or longer hairs to 2 mm long, margins with robust, stiff hairs to 2.5 mm long that are widest at the base. Panicles 5.5-23 cm long, 1.5-10 cm wide, open, branches erect to ascending, branches shorter or longer than spikelet, scabrous, 1-3 spikelets per branch, lowest inflorescence node with (1)2-3 branches. Spikelets 1.9-3.6 cm long (including awns); florets 4-10. Glumes subequal, lanceolate, glabrous or sometimes pubescent with hairs to 0.7 mm long, hairs sometimes restricted to nerves; lower glumes 6—9.7 mm long, 1-nerved; upper glumes 7.5-11 mm long, 3-nerved. Lemmas 11-13 mm long, 1.3-1.6 mm wide, glabrous or pubescent throughout or margin- ally, hairs to 0.5(-1) mm long, midribs scabrous distally; awns 3.5—7.1 mm long, arising 0-1 mm below lemma apex, straight, antrorsely scabrous. Paleas 9-9.6 mm long, shorter than lemmas; keels ciliate along upper half to 2/3 of their length, the cilia 0.1-0.2 mm long. Stamens 3; anthers (3-)4-5.7 mm long; dark brown to orange. 2n = 70 Distribution and Habitat.—Introduced. Grassy meadows, pastures. Bromus riparius is known only from a single collection along Highway 3 in southeastern British Columbia (Fig. 13A). It is promoted in western North America for forage, but its precise distribution is not clear (Pavlick & Anderton 2007). See additional notes below. Common Name.—English: meadow brome; French: brome des prés. Notes.—Meadow brome (Bromus riparius) is native to southeastern Europe, Turkey, the Caucasus and central Asia. It was introduced into North America in the last century as a potential forage crop. A cultivar (Regar selected from material collected in Turkey was registered in the United States in 1966 and in Can- ada in 1980 (de Araújo & Coulman 2004). Bromus riparius is now promoted and grown widely for forage across western Canada and in adjacent regions of the U.S.A. There is substantial interest in this grass in the agronomic community and several cultivars (Fleet, ‘Paddock’, ‘Cash’ and ‘Montana’) have recently been developed (Knowles 1990a, b; Cash et al. 2002; Jensen et al. 2004). From its first release, the name applied (incorrectly) to meadow brome in the United States was B. biebersteinii Roem. & Schult., a name that is still widely used in many government agronomy publications and in the seed trade. Bromus biebersteinii is a Mid ded SENG AC NA (EIS ae 1976) that is not reported for North America. lly similar and closely related (Saarela et al. 2007) to B. inermis (smooth iom. see EE ee introduced species widely used for forage and hay, which is seriously invasive in native plant communities (Otfinowski et al. 2007). Smooth bromegrass is widely grown because it is drought-tolerant and sod-forming, but it has slow regrowth characteristics and is considered better suited for use as hay. Meadow brome has better re-growth characteristics and is thus considered better suited for forage in pasture conditions (e.g., Thompson et al. 2003). Several reports indicate that B. riparius is less Saarela, Bromus in British Columbia, Canada 353 Bromus riparius romus Bromus secalinus squarrosus n FE in British Columbia. À Bromus rinariucs. B. Bromus secalinus. C. Circl LR Bromus squarrosus var. villosus. Scale bars = 200 km. aggressive than B. inermis and not invasive (Anonymous 2000; de Araújo 2001; Ogle et al. 2004). Through da PEN two i iin EE have been developed and selected by Agriculture and Agri-Food anada: g from a B. inermis and B. riparius (Knowles”, Coulman 2004), the other originating Doni a B. inermis x B. riparius hybrid backcrossed with B. inermis (Success”, Coulman 2006). These hybrids combine characteristics of each of their parents; they are.considered suitable for use as both hay and pasture (Knowles & Baron 1990; Knowles et al. 1993; Coulman 2004, 2006). Because of their agricultural importance, numerous aspects of morphology, forage quality and genetic diversity have been characterized in B. inermis, B. riparius, and their hybrids (e.g., Ferdinandez and Coulman 2000, 2001, 2002; Ferdinandez et al. 2001; de Araújo & Coulman 2004). There are no reports of B. riparius being adventive in Canada. Bromus riparius was treated in the Flora of North America (Pavlick & Anderton 2007). It was included there shortly before the volume went to press, but a detailed distribution map for the taxon was not available. In Canada, Bromus riparius is recognized as part of the flora in Saskatchewan (determined incorrectly as B. biebersteinii Roem. & Schult.; Harms 2006). Almost certainly it is present in Alberta, and a recent collec- tion from Manitoba is cited below. It has not yet been reported for British Columbia. There is no mention of B. riparius (or B. biebersteinii) in recent treatments of the British Columbia flora (Qian & Klinka 1998; Douglas et al. 2001), nor in Pavlick's (1995) treatment of Bromus in North America. In May 2006 I collected B. riparius in southeastern British Columbia in a fenced-in meadow at a rest stop along Hwy 3 [Saarela 417 (CAN, UBC)]; this collection apparently represents the first record of the species in the province. Indeed, the species has been grown in field trials in British Columbia (e.g., at Devick near Kamloops; May et al. 1999). and cultivars of B. riparius are EU S p in oa Columbia and used for amelioration S. J. Darbyshire, pers. comm., March 2 may have been deliberately planted here as De or it may la been introduced by road traffic. Sg of B. riparius is sometimes found as a contaminant in smooth brome seed commodities (Allison et al. 2001; SJ. Darbyshire, pers. comm. March 2007). Whatever the source of the introduction, Bromus riparius should be recognized as a component of the British Columbia flora, and additional occurrences for this species should be looked for. The morphological description is based on my field collections from British Columbia and Manitoba, as 354 tani i Texas 2( well as several specimens of cultivated plants (mostly greenhouse grown at the Agriculture and Agri-Food Canada Central Experimental Farm in Ottawa, Ontario, Canada) in DAO. Some of this cultivated material represents the cultivars ‘Fleet? ‘Paddock’ and ‘Regar’, differentiating morphologically among these and the other cultivars was beyond the scope of this study. Bromus riparius is one of three Bromus species in British Columbia (including B. inermis and B. pumpellianus) with rhizomes and long anthers (usually > 4 mm), and it therefore is relatively straight forward to identify. These three species are members of an Old World al- liance of taxa (B. pumpellianus is the only native species in North America that is part of this group) whose evolutionary relationships with other North American perennial Bromus species are not yet clear (Saarela et al. 2007). Because B. riparius is a relatively recent addition to the New World flora, there is little information in the North American botanical taxonomic literature for the species. There are several morphological characters that distinguish Bromus riparius. In an effort to identify characteristics to help seed analysts distinguish florets of B. inermis and B. riparius, which are sometimes mixed in seed commodities, Allison et al. (2001) conducted a careful study of the floret morphology of these taxa and found useful variation in the cilia (described by them as ‘siliceous prickle hairs’) on the palea keels. In B. riparius, cilia on the palea keels are present only distally, whereas in B. inermis cilia are present on the keels along the entire palea length. Figures 1-4 and 9 in Allison et al. (2001) illustrate this cl ter clearly. Allison et al. (2001) did not examine this character for B. pumpellianus. In B. pumpellianus cilia are present along the entire length of the palea keels, like in B. inermis, or only along the lower 34. Bromus riparius is further distinguished by its longer awns, which are (3-)4-7.1 mm long [vs. 0-3 mm long in B. inermis and 1-4 mm long in B. pumpellianus]. Several vegetative characters are also useful in distinguishing the three species. Leaf blade margins of B. riparius are characterized by the presence of long, stiff hairs (to 2.5 mm long) that are conspicuously wider at their base (Fig. 14); such robust hairs do not occur in B. inermis and B. pumpellianus. In his keys to Bromus in the (former) U.S.S.R., Tzvelev (1976) distinguished B. riparius from B. inermis and B. pumpellianus by the degree of shredding of basal leaf sheaths. Examination of this character in Canadian material confirms that it is very useful for distinguishing B. riparius from the other two taxa. The basal sheaths of decaying leaves in B. riparius are reticulate-fibrillose (i.e., they have a ‘thatched’ ap- pearance), the readily decaying basal sheaths in B. pumpellianus are split into straight fibers and the basal sheaths of B. inermis, which do not appear to decay as readily as those in B. pumpellianus or B. riparius, also split into straight fibers (Fig. 15). Pavlick & Anderton (2007) distinguished B. riparius from B. inermis and B. pumpellianus based on culm height and rhizome length; they noted that time constraints did not allow them to fully clarify its similarities and differences with these other taxa. Bromus riparius is weakly rhizomatous, B. pumpellianus is moderately so and B. inermis is strongly rhizomatous. Pavlick & Anderton (2007) also noted that it is not clear how B. riparius differs from B. pumpellianus subsp. dicksonii W. W. Mitch. & Wilton, a taxon known only from the Yukon River drainage in northwestern North America (Yukon and Alaska); characterization of differences between these two taxa was beyond the scope of this study. Representative Specimens Examined. CANADA. British Columbia: Rampart Rest Area, 10 km E of Cranbrook, meadow under Pinus ponderosa and Juniperus scopulorum, 49º33"2'N 115º8"38' W, 856 m, 23 May 2006, J.M. Saarela 417 (CAN, UBC). Alberta: Lethbridge, 10.15.1987, S. Smoliak s.n. (DAO-540342); cultivated land, fallow, NE-19-42-10-W4, 10 mi S of Hardisty, 3 Jun 1993, T. Yorchak V93W- FC268 (DAO-67859). Manitoba: Mary Jane Reservoir, ca. 10 km NW of Manitou, open, relatively undisturbed prairie habitat S of the reservoir, 14 Jul 2007, J.M. Saarela 1001 (CAN). Ontario (cultivated): Ottawa, Central Experimental Farm, greenhouse, cv. Fleet, Aug 1992, S.J. Darbyshire s.n. plant #28 (DAO-668498), plant #17 (DAO-668495), plant 425 (DAO- dedi ES 48 E EN plant #9 (DAO-668493), plant #27 (DAO- 668497), plant #31 (DAO-668500), plant 429 (DAO-6684 499) Farm, greenhouse, cv. Fleet, planted 26 Nov 1996, received as breeder seed C-86-2778, accession number 1 183, 22 Nov Pid M. Murray 4) s.n. (DAO-747661); Ottawa, Central Experimental Farm, collected from greenhouse, cv. Fleet, Knowles in late 1987 from Saskatoon, 15 Apr 1988, S.J. Darbyshire s.n. (DAO-575082); grown at manana See CV. leet, received in Ottawa as live plant 17 Sep 1987, R.P. Knowles # Fleet PH 12-5 (DAO-547672); g askatoon, cv. Fleet, received in Ottawa as live plant, 17 Sep 1987, R.P. Knowles # Fleet PH 12-32 (DAO-547670); Ottawa, Central ind Farm, green- house, cv. Paddock, 7 Aug 1992, S.J. Darbyshire s.n. plant # 19 (DAO-668488), plant #31 (DAO- a plant 430 Gen 668490) piam #10 (DAO-668487), plant 417 (DAO-668486), plant #27 (DAO-668489), plant 432 (DAO-66849] xd greenhouse grown, cv. Paddock, 23 Oc 1991, S.J. Darbyshire 4337 (DAO); Ottawa, Central el Farm, renhe cv. Geess planted 26 Nov 1996, received as breeder seed C-94-2023, 10 Nov 1997, M. Murray s.n. (DAO-74330); Ottawa, Central Experimental Saarela, Bromus in British Columbia, Canada 355 L A H IL H re | "1 L A H al H AE PA bs C PF emm Fic. 14. Leaf blades of B iparius [S | 417 (CAN)], showing the long, Farm, greenhouse, cv. Paddock, planted 26 November 1996, received as breeder seed C-86-2787, accession number 182, 26 Oct 1998, M. Murray s.n. (DAO-747653); Ottawa, Central Experimental Farm, cold frames, 23 Jun 1989, S.J. Darbyshire s.n. (DAO-594764, DAO- o UNITED STATES OF AMERICA. WASHINGTON: Cultivated at ARS-Pullman [from seeds (PI Identifier 31587) collected at: Rus Federation, oe Stavropol Territory, 16 Jun 1966, N. I. Vavilov, All-Russian Scientific Research 2], 14 Aug 1996, S.M. ps s.n. (DAO-5949 14. Bromus secalinus L. op: PL 1:76. 1753. (Figs. 16A—E, 13B). Bromus mollis var. secalinus (L.) Huds., Fl. Angl., ed. 2. 2:49. 1778. Avena secalina va cali Prodr. Stirp. Chap. Allerton 22. 1796. Serraful linus (L.) Bab., Man. Brit. Bot. 1:374. 1843. Forasaccus secalinus (L.) Bubani, Fl. Pyren. 4:388. 1901. Tyre: EUROPE: EE in A agris secalinis arenosis, Anon. (NEOTYPE: LINN-93.1, designated by PM. Smith, Notes Roy. Bot. Gard. Edinburgh 42:498. 1985) Bromus submuticus Steud., Syn. Pl. Glumac. 1:351. 1854. Tere: U.S.A. Missouri: St. Louis, 1838, N. Riehl s.n. (isorvee: US-865472, frag- ment) Annual; rhizomes absent. Culms 23-130 cm tall, 1—5(—7) mm wide at base, smooth; nodes 3-5, blackish- brown to yellow-brown, finely to densely retrorsely pubescent, hairs 0.1-0.4 mm long, lower nodes covered by sheaths. Leaf sheaths glabrous or sparsely pilose, hairs soft, 0.5-0.6 mm long; auricles absent; ligules 1.7-2.8 mm long, membranous, glabrous, apex erose; blades 8-28.5 cm long, 2-9 mm wide, abaxial sur- face pilose, hairs to 1.5(-2) mm long, soft and wavy, adaxial surface glabrous with occasional hairs to 0.5 mm long, margins smooth. Panicles 11-22 cm long, 2-9 cm wide, open, branches stiffly erect, sometimes secund, one or more lower branches usually longer than spikelet, smooth to scabrous, 1-3 spikelets per branch, lowest inflorescence node with 1-6 branches; 6-36 spikelets. Spikelets 1.3-2.5 cm long, ovate- be, 15. Pattern of decay in basal leaf sheaths. A. Bromus riparius [Saarela 417 (CAN, UBC)]. Basal sheaths decay readily and are A -fibrillose. B. Bromus inermis [Brink s.n. (U 95)]. Basal leaf sheaths do not dec eni idR a ey do de- cay, they split into a . C, Bromus pumpel- lianus [n.c. (UBC-V221751)]. Basal leaf sheaths read- ily decay into straight fibers. Scale bars = 1.5 mm. lanceolate, terete to slightly compressed, rachilla zigzagged and vis- ible at maturity, pote iz ux cn slightly subequal, smooth or scabrous, keels distally; lower glumes 3.6—4.9 mm long, 3-5(-7)-nerved; — es 4.2—6.1 mm long, 7-nerved, sometimes mucronate, mucros 0.2-0.6 mm long. Lemmas 6-7.7 mm long, 1.6-3 mm wide, 7-9-nerved, nerves not conspicuous, ovate-lanceolate, backs glabrous and Pac Been PS distally, margins dinstinctly inrolled in fruit; apex enti y bifid, the cleft 0.1-0.2 mm deep; awns 1-6.5 mm long, occasion- ally reduced to mucro or absent, arising 0.5—0.7 mm below lemma apex, usually straight, occasionally twisted once near base, widest at base, antrorsely scabrous. Paleas + equal in length to lemmas, apex usually visible at lemma summit, backs glabrous, keels ciliate, cilia to 0.3 mm long. Stamens 2; anthers 1-1.3 mm long, dark brown. Caryopses U- or V-shaped (i.e., margins strongly infolded) in cross section. 2n = 28. Distribution and Habitat.—Introduced. In North America, B. secalinus has been collected throughout the U.S.A. and southern Canada. It is apparently absent from the Canadian prairie provinces. In the U.S.A. itis most common in the midwestern states (Pavlick & Anderton 2007). B ollected only a few times in British Columbia, mostly in the southwestern part of idis province (Fig. 13B); the most recent collection is from 1991. Habitat includes waste places and fields. English: cheat, cheat chess, cheat grass, chess, chess brome, ryebrome, rye brome; French: brome des seigles, brome sécaline, coquiole noire, séglin, seigle bátard, brome faux seigle, Common Anima PÇ seiglin. Representative Specimens Examined. learns suas Sane: Hazelton, oe illegible], G. Gervais s.n (UBC); Alberni, 1900, [ district of Renfrew, 19 Jul 1902, C.O. gedet 787 (CAN); EH Hill, 22 Jul 1913, J. Macoun s.n. (V); within 5 mi of Lillooet, 7 Jul 1916, J.M. Macoun 93,964 (CAN, V) Swan Lake, Vancouver Island, 13 Jul 1938, J.W. Eastham s.n. (UBC); New Westminster, 12 Jul 1991, F. Lomer 91-149 (UBC). KEY TO THE VARIETIES OF BROMUS SQUARROSUS B. squarrosus var. squarrosus 15b. Glumes and lemmas densely pubescent, hairs to 0.8 mm lon B. squarrosus var. villosus 15a. Glumes and lemmas glabrous or scabrous 15a. Bromus squarrosus L. var. squarrosus, Sp. Pl. 1:76. 1753. te 13C, 16F-D. erui squarrosus (L.) Bab., Man. Brit. Bot. 375. bani, Fi. Pyren. 4:385. 1901. Bromus japonicus var. paniculatus (cuz oben Fl. Turkmen. 1:171. 1932. Tyre: SPAIN: Loefling N-93.8, designated by Smith, Notes Roy. Bot. Gard. Edinb. 79a (LECTOTYPE: L 42:500. 1985). Annual; rhizomes absent; plants 10-43 cm tall. Culms 1-2 mm wide at base, smooth; nodes 2-3, dark brown, pubescent, hairs to 0.6 mm ong. Leaf sheaths closed for most of their length, open for 4.2~9 mm ramen Saarela, Bromus in British Columbia, Canada SSS > AYN q 1 cm 357 Fic. 16. Bromus secalinus. A. Spikelet. B. Habit. C. Caryopsis. D. Lemmas. E. Inflorescence. Bromus squarrosus. F. Inflorescence. G. Spikelet. H. Glumes. L Lemma. [| [| £ ul Dos H in LI Lda nd ET nar VII 358 near apex, densely pilose with soft or sometimes stiff hairs 0.5-2 mm long; auricles absent; ligules 0.5-2 mm long, membranous, pubescent, apex erose; blades 2.5-10 cm long, 1.53 mm wide, abaxial surface densely pubescent, hairs 0.5-1.5 mm long, adaxial surface glabrous or minutely to densely pubescent, hairs stiff gins smooth or serrulate. Panicles (527—10.5 cm long, 2.7-7 mm wide, open, lax and somewhat sparse, racemose, often appearing one-sided (secund), sometimes reduced to a single spikelet, one or more lower branches usually longer than spikelet, branches smooth to scabrous, 12) spikelets per branch, lowest inflorescence node with 1-4(-5) branches; 3-17 spikelets. Spikelets 1.43.5 cm long, broadly oblong to ovate-lanceolate, terete to moderately compressed, rachillas not visible at maturity, florets 7-18. Glumes subequal, smooth, margins sometimes hyaline; lower glumes 4.5—/ mm long, 3—5(—7)-nerved; upper glumes 6.1-9.5 mm long, 7-nerved. Lemmas 8.2-11 mm long, 1.8-3.1 mm wide (when compressed), 7-9-nerved, nerves not conspicuous, broadly obovate, strongly angled near the middle, backs glabrous or scabridulous or (rarely) densely pubescent with hairs to 0.5 mm long, hyaline margins 0.5-1.2 mm wide at widest point, puberulent, apex minutely bifid, the cleft 0.1-0.3 mm deep; awns 8-11.7 mm long (awn on lowest lemma shorter), arising 1.7-3 mm below lemma apex, strongly divergent, usually twisted once near base, widest at base, scabrous. Paleas 6.1-9 mm long, backs glabrous, keels ciliate, cilia 0.3-0.5 mm long. Stamens 2; anthers (0.5-)0.9-1.6 mm long, dark brown. 2n = 14. Distribution and Habitat.—Introduced. Bromus squarrosus var. squarrosus is native to Eurasia (Izvelev 1976). In North America it occurs across the Great Plains in the northern U.S.A., around the Great Lakes area and in other scattered locations (Pavlick & Anderton 2007). In British Columbia it is most common in the interior and in the Columbia basin (Fig. 13C). Habitats include grasslands, hillsides, disturbed areas and roadsides. Elevation: 311-823 m Common Names.—corn brome, downy brome, field brome, nodding brome, one-way brome. and short or like abaxial surface than : Representative Specimens Examined. CANADA. British Columbia: near Kamloops, 2000 ft, 27 May 1934, G. V. Copley 7 (DAO); Richter Pass, Jul 1936, E.W. Tisdale 8850 (UBC); Kamloops, 22 Jun 1939, E.W. Tisdale 8759 (UBC); Lac du Bois, Kamloops, 4 Aug 1959, E.W. Tisdale 8760 (UBC); Kamloops, Sep 1939, E.W. Tisdale 8761 (UBC); Lac du Bois, Kamloops, 2700 ft, 10 Aug 1939, E.W. Tisdale 40-8 (DAO); Ashcroft, 6 Aug 1950, VJ. Krajina s.n. (UBC); Tranquille, 1100 ft, 5 Oct 1962, V.C. Brink s.n. (UBC); Bonaparte River Valley, 15 mi N of Cache Feu 1200 ft, CE n^i 5, Y Bici 65062007 (UBC); St. Eugene Mission, Cranbrook, 23 Jun 1968, F. Fodor 647 (UBC); Kamloops, ] , 23 Jun 1980, S.G. Aiken 1071 (DAO); Dallas Cliff (Point), 17 Jun 1982, R.K. Scagel 82-28 (UBC); N of Copper Creek, towards Tranquille Lake, 9 Jul 1991, F. Lomer 91-139 (UBC), Kootenay, Wasa, 20 Jun 1993, L.E. Pavlich 93-044 (V); Cariboo-Chilcotin, Dog Creek area, Harper's Creek Nach ih a Mine 3 mi Mun T. GE rd & C. sale pu ~ ru cias Mt, id pis Ape near LA Miis 1994. F. Lomer 94-152 ( g region, Hwy 1, 17 Aug 1995, F. fome 95 166 (UBC); 8 km E of Kaml , 26 May 1998, P. Williston PW98-134 (UBC); W of Osoyoos, L 3 km up Richter Mountain Road, 19 Jun 2006, H M. Saar C. Sears SJ. Maze 628 (CAN, UBC). 15b. Bromus squarrosus L. var. villosus Roth, Neue Beytr. Bot. 1:109. 1802. (Figs. 13C, 17). Similar to var. squarrosus, but glumes and lemmas densely pubescent with hairs to 0.8 mm long. Distribution and Habitat — In North America B. squarrosus var. villosus is only known from the three southern British Columbia collections cited below. Additional occurrences in North America should be searched for in the field and in herbaria. Tzvelev (1976:335) indicated that, in its native range, the variety is somewhat rarer in distribution than the typical variety. Notes.—Nomenclature used here follows Kerguélen (1999). Bromus squarrosus var. vi reported here for the first time for continental North America. Material examined from British Columbia closely matches material seen from Russia le.g., Skvortsov s.n. (CAN-578016) collected in Volgograd]. The description for B. squarrosus by Pavlick (1995) and Pavlick & Anderton (2007) does not mention the presence of dense pubescence in the taxon, which suggests that these authors did not observe pubescent forms of the species. The 1993 record from British Columbia was collected by Pavlick, but this was likely long after he completed work on his revision of Bromus for North America (Pavlick 1995). —À losus Roth is Specimens Examined. CANADA. British Columbia: Kootenay, Grand Forks, Gilpin Forest Service Road, E of Grand Forks, 23 Jun 1993, L.E. Pavlick 93-143 (V); roadside pullout on W side of Hwy 1 (Trans Canada) at Hells Gate, 22 Jun 2006, J.M. Saarela, C. Sears & Saarela, Bromus in British Columbia, Canada 359 e O MA "a RANA Pg H E Al A H | 1 far a Fic. 17.1 hol f B. villosus [Saarela, a LA TS D JAA “11 = 1 =] ] J. Maze 736 (CAN, UBC); Waneta, confluence of Columl d g pposi ing [ Hwy 224, 1400 ft, 17 Jun 2007, F.L. Lomer 6268 (CAN, UBC, V). 16. Bromus sterilis I . ; OP: Pl. 1:77. 1753. (Figs. ISAE, e E sterilis (L.) Fr., Bot. Not. 131. 1843. Genea sterilis (L.) Dumort., Bull. Soc. Roy. Bot. Belgique 7:67. 1868 is (L.) Nevski, Trudy Sredne-Aziatsk. Gosud. Univ., Ser. 8b, Bot. 17:20. 1934. Tyre: ENGLAND. Surrey: Tothill, near Headley, scattered plants in derelict hay-field on heavy soil overlying chalk, 15 Jun 1932, C. E. Hubbard 9045 (conserveD Tyee: E, proposed by Sales, Taxon 41:584. 1992; ISONEOTYPE: K). Annual; rhizomes absent. Culms 25-110 cm tall, 2-4 mm wide at base, smooth; nodes 3-5, light to dark brown, glabrous, lower nodes usually covered by sheaths. Leaf sheaths closed for most of their length, open for 8-15 mm in length, moderately to densely pubescent with short stiff hairs to 0.5 mm, auricles absent; ligules 2.3-4.6 mm long, membranous, glabrous or pubescent, erose-lacerate; blades 4.5-22 cm long, 2-6 mm wide, adaxial surface glabrous, minutely pubescent or densely pilose with soft, wavy hairs to 2.5 mm long, long hairs sometimes only along the margins, abaxial surface minutely pubescent with small appressed hairs, nia nc E 10-28 cm long, 3-18.5 cm wide, dense and ascending when ng, b d lax; usually longer than spikelets, shorter when panicle reduced to 1(2, 3 pus 133) spikelets per branch, aet inflorescence mode with = 6 branches. Spikelets 4.5—6.7 cm ), cuneate, subcylindric-compressed, rachill visible at maturity, florets 3-11. e. CH oo 1 cm Fic. 18. Bromus sterilis. A. Habit. B. Inflorescence. C. Glumes. D. Lemma. E. Spikelet. Bromus tectorum. F. Ligule. G. Spikelet. H. Lemma. I. Inflorescence. Bromus vulgaris. J. Spikelet. K. Lemma. L. Inflorescence. Saarela, Bromus in British Columbia, Canada 361 mus Bromus Bromus sterilis tectorum vulgaris Fic. 19. G hic distribution in British Columbia. A. Bromus sterilis. B. Bromus tectorum. C. Bromus vulgaris. Scale bars = 200 km. Ei P Glumes subequal, lanceolate, smooth, keel th or scabrous, margins hyaline; lower glumes 10—14 mm long, 1-3-nerved; upper glumes 7.5-21 mm long, 3—5-nerved. Lemmas 13-22 mm long, 1.1-1.7 mm wide, 7-nerved, nerves conspicuous, lanceolate, backs smooth or scabridulous, scabrules sometimes increasing in density distally, margins not distinctly inrolled in fruit, apex bifid, the cleft 1-3 mm deep; awns 15-30 mm long, arising about 1.5 mm below lemma apex, straight, antrorsely scabrous. Paleas 9.5-13.5 mm long, much shorter than lemma, backs glabrous, keels glabrous or ciliate, cilia often present only distally, cilia to 0.2 mm long. Stamens 3; anthers 1.3-1.7 mm long, dark brown. 2n - 14, 28. Distribution and Habitat.—Introduced. Bromus sterilis is native to Europe, northern Africa and south- western Asia (Sales 1993). In North America it occurs in western and eastern North America (Pavlick & Anderton 2007). In British Columbia B. sterilis occurs only in the extreme southwest of the province in the Vancouver area and on central to southern Vancouver Island (Fig. 19A). Habitats include roadsides, sandy beaches, meadows and waste places. Elevation: 0-425 m. Common Names.—barren brome, poverty brome. Notes.—Several intraspecific taxa that have been previously recognized are mentioned by Sales (1993:5-6), but she did not recognize these; none have been recognized in the North American flora. Representative Specimens Examined. CANADA. British penne pd of Victoria, E 1908, Macoun 76,833 (CAN); Victoria, Anderson's farm, 27 May 1893, Macoun 117 (CAN); Victoria, 13 Jun 1908, ^ 35 (CAN 1, Saanichton, 15 May 1913, Macoun 88,540 (CAN); Oak Bay, Victoria, 17 Jun 1913, Macoun 88,541 (CAN, V); V Victoria, 6 Jun 1908, Macoun 76,834 (CAN); Victoria, 11 May 1920, G.W. Copley 1-14 (DAO, UBC); Sidney, 30 Sep 1930, H. Groh s.n. (DAO); Shawnigan Lake, 27 May 1931, H. Groh s.n. (DAO); Crescent Beach, 31 May 1937, J.W. Eastham s.n. (CAN); Victoria, 2 Jun 1938, J.W. Eastham 431 (DAO, UBC); W Point Grey, Jun 1939, J.W. Eastham s.n. (DAO); W Point Gray, Vancouver, Jun 1939, J.W. Eastham 429 (UBC); Saanichton, 21 Jun 1939, W.R. Foster s.n. (DAO, UBC, V); James Island, off Saanichton, 5 Aug 1939, H. Groh 424 (DAO); No. 5 road, Lulu Island, 0.25 mi S of Twigg Island, 4 Jun 1944, J.W. Eastham 11426 (DAO, UBC); Whiffen Spit, Sooke, 19 May 1948, V.C. Brink 48-28 (UBC); Victoria, Finlayson Point, 20 May 1950, 10 m, V. Krajina & R.H. Spilsbury 3921 (DAO); Esquimalt Lagoon near Victoria, 7 May 1962, J. Hett 90 (DAO); Happy Valley near Victoria, 30 May 1962, J. Hett s.n. (DAO); Henderson Point, Brentwood, 16 Jun 1967, A. S. Harrison 20.1 (CAN); Victoria, Finlayson Point, 10 m, 20 May 1950, V. Krajina & R.H. Spilsbury 3921 (UBC), 2 mi E of Duncan, 19 May 1955, D.R. Lindsay & W. Woodbury 737 (DAO); John Dean Park, Saanichton, 14 Jun 1962, J. Hett 16 (DAO); Little Saanich Mtn, 350 ft, 17 May 1965, B.W. Davies 389 (DAO); Mt Finlayson, 425 m, 28 Jun 1978, L. Pavlick 78-231 (V); Mt Finlayson, 300 m, 16 May 1979, 300 m, L.E. Pavlick 79-6 (V); Saltspring Island, Mt Tuam, 3 Jul 1982, L.E. dne 82- a E Vancouver Se Mr O E of Goldstream Provincial Park, 1 Jul 1982, S.G. Aiken, S.J. Darbyshire & L.E. Pavl Park, Saanich Peninsula, 4 Jul 1988, S.J. Darbyshire 3792 (CAN, DAO); S end 362 t tani i Texas 2( of Tsawassen Drive S, Tsawassen Indian Reserve #0, 28 May 2006, J.M. Saarela & C. Sears 546 (CAN, UBC), Mayne Island, Bennett Bay, Gulf Islands National Park ed of po» Wilkes Road off Bennett Bay Road, 3 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 826 (CAN, UBC); Mayne Island, cliff int Park, ca. 1 km W of highest point where radar tower is, 3 Jun 2007, J.M. Saarela, D.M. Percy, Y. Chang & Q. C. E. Cronk 808 (CAN, UBC); Vancouver Island, es Lake De Park, sue N of Hwy 1, N of Langford, 4 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 861 (CAN, UBC); V. d, just S o g Hwy 1A, 6 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 909 (CAN, UBC); Vancouver eee , just off Hwy 19A along Crane Road, N of Guálienm Beach, at confluence of Nile Creek and the Straight of Georgia, 7 Jun 2007, J.M. Saarela, D.M. Percy €» Y. Chang 934b (CAN, UBC). 17. Bromus tectorum L., Sp. Pl. 77. nd (Figs. 18F-I, 19B). Zerna tectorum eio. Denkschr. Konigl. Akad. Wiss. Munchen 1813:297. 1814. Sched (L.) Fries, Bot. Not. 131. 184 (L.) Nevski, Áct. Univ. Asiae Med 8b., Bot. 17:20, 22. 1934. Genea tectorum (L.) Dumort., Bull. Soc. Bot. bs 7:67. 1868., in adnot. e: EUROPE: Anon. (LECIOTYPE: LINN 93/25, designated by Smith, Notes. Roy. Bot. Gard. Edinb. 42:500. 1985). Bromus tectorum var. nudus Klett & Richt., Fl. Leipzig 109. 1830. Bromus tectorum L. subsp. nudus (Klett & Richt.) Piper & Beattie, FI. 51. 1915. Bromus tectorum f. nudus (Klett & Richt.) St. John, Fl. S. E. Washington (Piper & Beattie) 36. 1937. Bromus a gras Ge John, Fl. S.E. Washington (Piper & Beattie), ed. 2. 536. 1956. Tyre: not known at. Sci. Phila. Proc. 1892:98. 1862. Tyre: U.S.A. Texas: Buckley s.n. (LECTOTYPE: US- 8654741. designated by Hitchcock, Man. Grass. U. S. 817. 1935). Annual; rhizomes absent; 5—80 cm tall. Culms 0.7-2.5 mm wide at base, smooth; nodes 2—4, blackish brown, ] glabrous or minutely pubescent. I 1 for most of their length, glabrous or var iously pubescent with dense, stiff hairs to 0.5 mm or a soft, wavy hairs to 1 mm long, longer hairs to 2 mm long sometimes present ds sheat] bsent; ligules (1223 mm long, membranous, glabrous, erose-lacerate; blades 1.3-14 cm long, E 5 mm wide, adaxial surface pubescent with sparse to dense short, stiff hairs to 0.3 mm long, abaxial surface puberulent with minute, appressed hairs ca. 0.1 mm long, hairs sometimes denser towards blade apex, margins smooth or with hairs like adaxial leaf surface. Panicles 4-22 cm long, 1-13.5 cm wide, condensed and ascending when young, becoming lax and nodding, sometimes reduced to a few spikelets, branches longer or shorter than spikelets, scabrous to densely pubescent, 1-14 spikelets per braiei; lowest SE node with 1-8 branches: Spikelets 2.5-3.5 cm long (including awns), cuneate, t maturity, rachil visible at maturity; florets 3—6(-8). Glumes subequal, lanceolate, apices + bifid, the clef 0.1-0.5 mm deep, glabrous or muy to densely pubescent with short, appressed hairs, M Gg Mio soft hairs to 1 mm long, tl ccurring in a line just inside margins, g ooth or scabrous; lower glumes 4-9 mm long, 13) need occasionally minutely awned, awn to d 1 mm long; upper glumes 7-13.5 mm long, 3(-5)-nerved. Lemmas 9-13.8 mm long, (0.6-)0.9-1.5 mm wide, 7-nerved, backs glabrous, scabrous, or pubescent with long, soft hairs to 1 mm long, sometimes denser along margins and sometimes only pubescent on upper 2/5; keels smooth or serrulate, apex bifid, the cleft 1-3 mm deep; awns 8-18 mm long, arising 1.1-2.7 mm below lemma apex, straight, antrorsely scabrous. Paleas 7-8.7 mm long, shorter than lemma, backs glabrous and translucent, keels ciliate, cilia to 0.6 mm long. Stamens 3; anthers 0.5-0.9 mm long, dark brown. 2n = 14. Distribution and Habitat.—Introduced. Bromus tectorum is widespread across the United States and southern Canada, with scattered collections from locations in Alaska, Yukon and the western Northwest Territories (Pavlick & Anderton 2007; Valliant et al. 2007). The species is most common in the southern half of the province and it has been collected sporadically in northwestern British Columbia [Fig. 19B; see Valliant et al. (2007) for another recent distribution map]. Habitats include roadsides, fields, meadows, arid slopes, woodlands, cliffs and talus slopes, hillsides, dry rangeland, pastures, sandy beaches, railroad tracks and other waste places. Elevation: 0—1800 m. Common Names.—English: cheatgrass, cheat grass, downy brome, drooping brome, early chess, military e e grass, thatch bium duds EE SE des toits, brome des murs. Notes. — Bromu ive weed. It was first recorded in North America from in 1790, and the first EE in western North America is from S 's Bridge, British Columbia Mac 1981; specimen cited below). À tudies | indicated that B. tectorum in North America is highly inbreeding and has very low levels A genetic variation across its North American range (e.g., Novack et al. 1 Saarela, Bromus in British Columbia, Canada 363 1991; Bartlett et al. 2002; Ramakrishnan et al. 2006). Genetic variation within populations, however, shows strong regional e anm with opos: in British Columbia having the Apia level of piai population h America (Bartlett et al. 2002). ! of B. tectorum into North P have been documented using genetic data (e.g., Mack 1981: Novack et al. 1991; Novack & Mack 1993, 2001; Bartlett et al. 2002). Valliant et al. (2007) provide an account of the introduction history of B. tectorum in British Columbia and Canada; based on allozyme data they identified a minimum of five independent introductions of B. tectorum into western Canada. On fT ARTS AA A R S E ined. CANADA. British Col bia: S 's Bridge, 4 Jun n 1889, M A 1900, Fletch 1 And 22132 (DAO): Ol mad Jul 1912, E. Wilson 1061 (UBC); Sidnei: 7 Jul 1913, M.O. Malte 27 (CAN); Spence's Bridge, 15 Jul 1913, H Davidson s.n. -(UBO); Twaal Valley, 21 Jun 1914, J. Davidson s.n. (UBC); near Moren's Ranch, Spence's Bridge, 21 Jun 1914, J. Davidson s.n. (UBC); Point Grey, 2 May 1915, J. Davidson s.n. (UBC); Pemberton Trail, 24 Jun 1916, J.M. Macoun 93962 (CAN); Lillooet, 2 Jul 1916, J.M. Macoun 93963 (CAN); Hazelton, Skeena River, 12 Jul 1917, J.M. Macoun 93975 (CAN); Okanagan Lake, 17 Jun 1918, A. H. Hutchinson s.n. (UBC); Courtenay Lake, 21 Jun 1918, Hutchinson s.n. (UBC); Powell River, jun 1923, J. B. Munro s.n. (UBC); White Rock, 25 May 1924, T.R. Ashlee s.n. (UBC); Eburne, 23 May 1931, H. Groh s.n. (DAO); Langley Prairie, 30 May 1931, H. Groh s.n. (DAO); Oliver, 3 Jun 1931, H. Groh s.n. (DAO); Kamloops, 4-7 Sep 1931, V. Kujala & A. Cajander s.n. (CAN); E Lytton, 500 ft, 11 Jul 1932, E.P. Hume s.n. (UBC); near Kamloops, 2000 ft, 27 May 1934, G. V. Copley 8 (DAO); near Cranbrook, 13 Se 1934, E.H. Moss 2942 (DAO); Spence's Bridge, Kamloops district, 24 May 1935, D.W. Thompson s.n. (DAO); Tranquille, Kamloops, Jun 1935, E.W. Tisdale 40-11 (DAO); Lac du Bois, Tranquille, 21 Jul 1935, E.W. Tisdale 40-11 (DAO); Squilax, 10 Aug 1935, P.P. Henson s.n. (DAO); Crescent Beach, 31 May 1937, J.W. Eastham s.n. (CAN); C. N. yards, Vancouver, 31 May 1937, W. Sandall s.n. (UBC); Nanaimo, 31 May 1939, J.W. Eastham s.n. (UBC); — 14 dus rs jin "dd 1945 (DAO); Kamloops, 21 Jun 1937, J.W. Eastham s.n. (DAO W Point Grey, Vancouver, 20 Jun 1937, JW. Mts, Hat Creek Valley, 3000 ft, 26 Jun 1938, J.W. Thompson GE. M. Thompson 227 (CAN, US); Lower See range, Kantons. 22 May 1939, E.W. mE s.n. (DAO); Prince Rupert-Terrace, Jul 1939, C. C. King s.n. (UBC); Terrace, 11 Aug 1939, H. Groh 533 (DAO); Kamloops, 24 Jun 1940, M.G. Dudley 246 (DAO); Locarno Park, Van- couver, 11 Jun 1944, J.W. Eastham s.n. (UBC); Gray Creek, 1 Jul 1944, W.C. McCalla "a (UBC); Fort Sede 12 Jul 1944, W.B. Johnstone 495 (UBC); Kimberley, F. Fodor 28, 14 Jul 1946 (UBC); Osoyoos, 3 Jul 1947, V.C. Brink s.n. (UBC); Lytton, 175-500 m, 18 Aug 1949, VJ. Krajina 1207 (UBC); Anarchist Mt, 19 Aug 1950, VJ. Krajina s.n. (UBC); Princeton, 19 Aug 1950, VJ. Krajina s.n. (UBC); Coyle, Merritt, Aug 1950, V. Krajina s.n. (UBC); Chase, Kamloops, 28 Aug 1950, V.J. Krajina s.n. (UBC); 1 mi S of Osoyoos on W side of lake, 19 May 1953 p Á. april & D.B. i gege 7899 RE e mi SE B ne along Similkameen River, 26 May 1953, J.A. Calder & D.B.O. Savile idge, 31 May 1953, J.A. Calder & D.B.O. Savile 8457 (DAO); about 4 mi E of NS on N side of 5 Thompson River, 2 Jun 1953, JA, Calder & D.B.O. Savile 8590 (DAO); about 4 mi S of Sicamous on E side of Mara Lake, 5 Jun 1953. J.A. Calder & D.B.O. Savile 8749 (DAO); Westbank, 8 Jun 1953, n.c. (DAO); 24 mi S of Kootenay Bay along high- way to Creston, 12 Jun 1953, J.A. Calder & D.B. a AE diii (DAO); oor River ne at Pads near Creston, 17 Jun 1953, J.A. Calder & D.B.O. Savile 9285 (DAO, US); 2.5 mi E , 20 Jun 1953, J. A. Calder & D.B.O. Savile 9428 (DAO, UBC); Christina Lake, 29 Jun 1953, G. H. Collin s.n. (DAO): Westbank, 8 Jul 1953, J.M. Smith s.n. (DAO); Invermere townsite, 14 Jul 1954, A. McLean s.n. (DAO); Parksville, Vancouver Island, May 1955, V.C. Brink s.n. (DAO, UBC); Osoyoos, 12 May 1955, D.R. Lindsay & W. Woodbury 595 (DAO); near Bowser, Qualicum Bay, Vancouver Island, 16 May 1955, D.R. Lindsay & W. Woodbury 637 (DAO); Port Alberni, 17 May 1955, D.R. Lindsay & W. Woodbury 660 (DAO); 20 mi N of Victoria, 19 May 1955, D.R. Lindsay & W. Wood- bury 725 (DAO); Patricia Bay Highway, one-half mi N of Royal Oak Corner, 20 May 1955, D.R. Lindasy & W. Woodbury 753 (DAO); S of Steveston, Lulu Island, 24 May 1955, D.R. Lindsay & W. Woodbury 770 (DAO); Abbotsford, 25 May 1955, D.R. Lindsay & W. Woodbury 795 (DAO); Boston Bar, 30 May 1955, D.R. Lindsay & W. Woodbury 861 (DAO); 4 mi S of Spence's Bridge, 31 May 1955, D.R. Lindsay and W. Woodbury 884 (DAO); 5 mi N of Kamloops, 1 Jun 1955, D.R. Lindsay & W. Woodbury 897 (DAO); 10 mi N of Merritt, Nicola Valley, 2 Jun 1955, D.R. Lindsay & W. Woodbury 906 (DAO); Williams Lake, 5 Jun 1955, D.R. Lindsay & W. Woodbury 971 (DAO); near Pavilion, 7 Jun 1955, D.R. Lindsay & W. Woodbury 995 (DAO); 2 mi N of Vernon, 10 Jun 1955, D.R. Lindsay & W. Woodbury 1023 (DAO); Lumby, 14 Jun 1955, D.R. Lindsay & W. Woodbury 1073 (DAO); 10 mi S E Kee Ge 1955, DR Miis & W. Niels n 1096 (DAO); 31 mi S of Williams Lake, 22 Jule iss G. E a Ln & W. Woodbury 1762 ( liver Rd, via Green Lake, 1650 ft, 11 May 1958, K. B 1 Lill p 1958, T.M.C. Taylor V. Krajina & F. M (UBC); Cranbrook, 12 Jul 1959, E.W. Sullivan 559430 (DAO): Kaslo, See 2200 ft, 21 Aug 1959, M. Bell s.n. (UBC); Indian Reserve E of Oliver, 4 May 1960, 1600 ft, K. Beamish €> E. Vrugtman 60102 (CAN); Indian Reserve E of Oliver, 4 May 1960, J.W. Eastham 60102 (US); between Win- dermere and Fairmont Hotsprings, ca. 3000 ft, 13 May 1960, K. Beamish & F. Vrugtman 60318 (UBC); Oliver campsite, 1100 ft, 17 May 1960, K. Beamish & P Vrugtman 60423 (UBC); Boy's Town, Hope-Princeton Hwy, 2300 ft, 11 Jul 1960, K.I. Beamish & F. Vrugtman 60648 (DAO); Mt Kobau lookout trail, S Okanagan, 3500 ft, 8 Jul 1961, J.W. Eastham 610412 (CAN, UBC); Esquimalt Lagoon, near Victoria, 7 May 1962, J. Hett s.n. (DAO); coast of Saanich wees near Victoria, 11 May 1962, J. Hett s.n. (DAO); Esquimalt Lagoon Near Victoria, 29 May 1962, J. Hett s.n. (DAO); ^ ark, about 30 mi no of Princeton and 90 mi E of Vancouver, 4000-5000 ft, 26 Jun 1964, H.J. Scoggan 15093 (CAN); Ol ] ticton, 26 Jun 1964, H J. Scoggan 15044 (CAN); Nano- ose Bay, 10 ft, 6 May 1965, B.W. Davies 256 (DAO): Little e Mtn, 400 a 17 May 1965, B.W. Davies 390 (DAO); Esquimalt Lagoon, 1 Jun 1965, B.W. Davies & R.A. Keller 700 (DA I and Trapp Lakes, 2300 ft, 23 Jun 1965, VJ. Krajina A 364 Jeun a (DAO); NW slopes in E end of Nicola pni 40 mi. S of Kamloops, 1900 ft, 23 Jun 1965, VJ. Krajina 650623139 (UBC); 34 mi N of Princeton, 23 Jun 1 Krajina 6506231 C); Beaver Pond, Manning Park, 14 Jul 1965, L L Bayly 65-129 (DAO); S of Kamloops, 2830 ft, April 1966, so a 65062303 (CAN, UBO); Island View Beach, Vancouver Island, 25 ft (7.6 m), 25 May 1966, A.S. Harrison 29.1 (CAN); 2 mi N of Princeton, 2300 ft, 31 May 1966, J. F. Alex 2258 ( DO aa Grove, on W shore of Kidd Lake, oro mid no. 5, 3000 ft, 22 Aug 1969, J.H. Soper, M.J Shchepaneh & A.F. Szczawinski 12,662 (C Kel , 14 Jun 1970, S. Kojima s.n. ( Park, along Hope-Princeton Hwy, Beaver Pond area, 1189 m (3900 ft), 22 Jun 1973, C.C. Chuang 118 (CAN); between Dee e Li and Telegraph Creek, 17 Jul 1973, VJ. Krajina, R.G. McMinn, K. o M. Annas s.n. (UBC); Botanie Valley road near ce 1200 ft, 18 May 1974, J. Pinder-Moss & J. Pojar 657 (UBC); Nanoose Hill, E side of Nanoose Bay, ca. 15 mi N of Nanaimo, Vancouver Island, < 500 ft, 27 May 1975, KI. Beamish, E. DON ee did (DAO); Ashnola Road, mile 23, 24 Jul 1975, R. Eon 7639 (DAO); Ca- thedral Park, 2 Jul 1976, R lt fO Hwy 97, 16 Jul 1976, M. Barkworth 1640 (DAO); Okanagan Game Farm, 1 mi NE of kaden 1800 ft, 2 May 1977, B. Wikeem & S. Gale s.n. (UBC); W of Prentice Lake trail and N of Relay Creek, 1800 m, 16 Jun 1977, C. Selby 20 (DAO, UBC); Hat Creek valley, 3300 ft, 23 Jun 1977, R.K. Scagel 77-0-83 (UBC); above trail to Prentice Lake, plot 134 #880, 1580 m, 8 Tk 1977, C. Sec s.n. (DAO); Nanoose Hill, N of Nanaimo, 2-4 May 1980, R. Turking- ton s.n. (UBC); Kamloops, ! . 23 Jun 1980, S.G. Aiken 1062 (DAO); Kamloops Road to Todd Mt, 2800 ft, 30 Jun 1980, S.G. Aiken 1118 (DAO): Opax Mountain, near Kamloops, 1 Jul 1980, S.G. Aiken 1130 (DAO), Prison Camp, 20 km N of Clearwater Village, 8 Sep 1980, T. Goward 81-815 (DAO); E of Princeton on Hwy 3 along Similkameen River, 31 Jul 1980, H.L. Dickson & M. Picher 3757 (DAO). Kent District, Harrison Lake, Harrison Hot Springs, 10 Jul 1982, 5.6. Aiken & S.J. Darbyshire 2237 (DAO); Kalamalka Lake Park, near Vernon, 23 May 1984, A. Ceska and R.T. SC See Gees Nee A Hot ee N of Pem- berton, Lillooet District, 650 m, 9 May 1987, A.A. Rose s.n. (UBC); below “The Thro N of Osoyoos Lake, 19 May 1987, WJ. Crins & S.G. Cannings 7380 (UBC), nea 30 Jun 1988, J. E. Whiting 36 (UBC): Gegen area, Haynes Lease idis 13 May 1990, T. Goward & T. Lea 90-593 (UBC), Christina Lake, 28 Aug 1990, DM Tiedje 5581 (DAO), Abbotsford Distr. Munic Mt, 60- El m, 25 May 1995, H. a F.R. pene M. Manna D. eege Goertzen 4849 (UBC); 1.9 km up K Road, S of Old Ri Pass Road, W of Osoyoos, 26 May 1995, H.I 9 (DAO); Hope, above Fraser River, ca. 2001 m., 1 Jun 1995, H. Beef, E.R. Ganders & L. Thacker 4892 (UBC); Texada ss off Harwood Point, at SE end of Dick Island, 2—5 m, 19 Jun 1997, H. Janszen, P. Janszen & M. Fontaine 2659B (UBC); 8 km E of oe 600 m, 11 de ee P. Williston s.n. (UBC); 12 km SW of Merritt, 27 Jul 1998, P. Williston PW98-400a (UBC); W 18), 14 May 1999, R. Bandringa 89 (UBC); Kootenay, Grand Forks, 30 Aug 1999, R. Hebda s.n. (UBC); edge of beach bo wrack line, below University of British Columbia, 25 May 2005, J.M. Saarela 316 (CAN, UBC); along Hwy 3, E of Hedly, 2.5 km W of Keremeos, gravel picnic area (rest stop), 18 May 2006, J.M. Saarela 336 (CAN, UBC); Rocky Mountain Forest District, along Hwy 3, 1.5 km E of JM. Saarela 413 (CAN, UBC); Rampart Rest Area, 10 km E of Cranbrook, 23 May 2006, J.M. Saarela 416 (CAN, UBC); ca. 2 km E of Er- == nb mu 24 May 2006, J.M. Saarel dii (CAN, UBC): 21.1 km W of Creston on Hwy 3A, 24 May 2006, J.M. Saarela 446 (CAN, t Area, K Society, 24 May 2006, J.M. Saarela 450, 451 (CAN, UBC); Shady Lake Campground, 4.7 a S of Osoyoos at U.S.A./ Canada border, 25 May 2006, J.M. Saarela 471 cs E Mie iu. Has Provincial Park, 25 May 2006, J.M. Saarela 485 (CAN, UBC); Richmond, London Farm, Gilbert Beach, just 28 May 2006, J.M. Saarela & C. Sears 525 (CAN, UBC); Ladner, S side of Canoe Passage, W terminus of River Road, 28 May 2006, J. M. p & C. Sears 539 (CAN, UBC); 10 km W of Princeton on Hwy 3, 19 Jun 2006, J.M. Saarela, C. Sears & J. Maze 587 o no Princeton, across from Princeton beyond TTO. A High School, 19 Jun 2006, J.M. Saarela, C. Sears & J. Maze 596 (CAN, 2nd fence, 19 Jun 2006, J.M. Saarela, C. Sears & J. Maze 603 (CAN, UBO): 1.9 km up S inder Mountain Road, 19 Jun 2006, J.M. Saarela, C. Sears & J. Maze 608 (CAN, UBC); Fairview Road, ca. 4 km NE from intersection with Lowe Road, 19 Jun 2006, J.M. Saarela, C. Sears & J. Maze 622 (CAN, UBC); University of British Columbia, beside Biological Sciences Building, 8 Sep 2006, J.M. Saarela 761 CAN) 18. Bromus vulgaris (Hook.) Shear, Bull. Div. Agrostol., U.S.D.A. 23:43, f. 24. 1900. (Figs. 18]-L, 19C). Bromus purgans var. vulgaris Hook., Fl. Bor.-Amer. 2:252. 1840. Bromus eximius subsp. umbraticus Piper, Contr. U. S. Natl. Herb. 11:144. 1906, nom. illeg. superfl. Bromopsis vulgaris (Hook.) Holub, Folia Geobot. Phytotax. 8:169. 1973. Zerna vulgaris (Hook.) Henrard, Blumea 4:498. 1941. Tree: U.S.A.: Columbia River, Scouler s.n. LECTOTYPE: K, designated by Wagnon, Brittonia 7:459. 1952; ISOTYPE: Masi LR ex. i 1840:252 ti hen he described B. purgans var. do id Eeer Dr d qd pu River. Dee » Ell E D Columbia. Dr Scoiller* and he cited “Fort Pos Dr pal g I y vas the first ] LI E +1 c 1 s PS | AJ al E mtl +1 + to recognize B. he included as SOME of B. "mu" p e., B. ae var. v. pauciflorus s dn in Beal and B debilis Nutt. ex Shear). Piper (1906) cre- B. purgans var. vulgari ated a new name dude eximius var ] : Hook., ft) l that Hoo! ined is indeed B ee in me sense of Shear (1900). o i on E Peu that de (1952) recognized B. add oni and M ANM Piper’ | y t “Goldie. Dr. Richardson." H 1 th from “Red River” is B. halmii Gray; Richardsons “Fort Norman” collection is B. inermis var. purparascens (=B. inermis Leyss X and D he sen (1900), who saw a SCH Scouler’s collection from “Columbia [River]” is B. vulgaris, i Saarela, Bromus in British Columbia, Canada 365 from the same E Re ee e B. Ea has ew taken up by all subsequent authors. Wagnon (1952) further noted that Richard ky Mountains, outside the = of B. in is o EE ted f. R + 7: 450 ass I PE + hs tl 1 5 11 FEL 1:1 1 fen pe] TIA 1 H D 211 RS " LT; 1 : Eus as te EIE ectorype e B. Ste var. vulgar e B. r 1 : ehe wur AE : : 4 CE: dal to distinguish t] ixed taxa da Hooker a Pavlick et bal (2003) "n katei that the correct name for B. nius Gage? Shear à is B. eximius SE Wee SE and ed not Inoue ES waon a ee B. purgans var. vulgaris, g VEDI Richardson as a single collection: Bromus vulgaris var. eximius Shear, Bull. Div. ue U.S.D.A. 23:44. 1900. Bromus eximius (Shear) Piper, Contr. U.S. Natl. Herb. 11:143. 1906. Tree: U.S.A. OREGON. \ , OI 4 mi above Wallowa I Lake, 23 Aug 1899, C.L. Shear 1791 (noLorvee: US-81575! Bromus vulgaris var. robustus Shear Bull. Div. Agrostol., U.S.D.A. 23:44. nd Bromus eximius var. iun (Shear) Piper, Contr. U.S. Natl. Herb. 1:143. 1906. Tyre: U.S.A. Orecon. Clatsop C aside, 11 Aug 1899, C.L. Shear & EL. EE Se E US- iid B S 1., Deutsche Bot. Monatsschr. 19:93. 1901. Tree: U.S.A. WASHINGTON. Skamania Co.: 22 Jul 1894, WN. Suksdorf 2335 — GH-234940, US-228545! with the additional locality information: “near the Ice Cave"). Perennial; rhizomes absent; plants 45-125 cm tall. Culms 2-5(-9) mm wide at base, smooth; nodes (324—6(—7), brown, glabrous or finely to densely retrorsely pubescent, hairs to 0.7(-1) mm long, occasion- ally with denser hairs above nodes; lower nodes covered by sheaths. Leaf sheaths closed for most of their length, glabrous or sparsely to densely pilose with hairs to 1.2 mm long; auricles absent; ligules (2-)3—6(7) mm long, membranous, glabrous, erose; blades 12-33 cm long, 3-16 mm wide, thin and papery when dry, adaxial surface sparsely to densely pilose, hairs to 1 mm long, soft and wavy, abaxial surface glabrous, margins smooth or serrulate. Panicles 8-25 cm long, 1—5 cm wide, narrow, branches erect to ascending or drooping, shorter or longer than spikelets, branches scabrous-pubescent, 1(-2) Spies pa branch, lowest inflorescence node with 1-5 branches. Spikelets 1.53 cm long, ovate-lanceolate, t ghtly compressed, rachilla sometimes visible at maturity; florets 3-9. Glumes subequal, lanceolate, bs or pubescent, hairs sometimes restricted to margins and keel; lower glumes 4-9 mm long, 1(-3)-nerved, lanceolate; upper glumes 5—11 mm long, 3-nerved, sometimes mucronate, mucros to 0.5 mm long. Lemmas 10-16 mm long, /-nerved, 1.2-2 mm wide, margins glabrous to sparsely or densely pubescent, backs glabrous to variously pubescent, hairs to 1 mm long, sometimes denser on lower 14; apex entire or minutely bifid, the cleft to 0.3 mm deep; awns (46-11 mm long, arising 0-0.5 mm below lemma apex, straight, antrorsely scabrous. Paleas 7.5—11 m long, shorter than lemmas, keels densely ciliate, cilia to 0.5 mm long. Stamens 3; anthers 2—3.5(-4) mm long, dark brown. 2n = 14. Distribution and Habitat. — Native. In North America, B. vulgaris o in British Columbia, soutl PONO and da to western Wyoming and south-central California (Pavlick & Anderton 2007). Bromus t the southern half of British Columbia (Fig. 19C). It was only recently reported from Wio erby Island (Queer Charlotte Islands; Lomer & Douglas 1999) A SE collection [Saarela & Percy 1169 (CAN, UBC)] represents the first record of B. vulgaris from tl ] Island. Habitats include moist woods, stream banks, damp thickets, along trails, meadows and rody banks. Elevation 0-2286 m. Common Names.—Columbia brome, narrow-flowered brome, common brome. Notes.—In herbarium material of B. vulgaris, leaf blades tend to appear conspicuously thin and papery compared to those in other species of B. sect. Bromopsis, though it is difficult to adequately describe this qualitative characteristic. This characteristic does not seem to be as evident in the field. At present, there are no leaf blade anatomical data for North American B. sect. Bromopsis species which might allow further evaluation of this character. Representative Specimens Examined. CANADA. British Columbia: Selkirk Range, 17 Aug 1885, Macoun 30,005 (CAN); Roger's Pass, Selkirk Mtns, 18 Aug 1885, J. Macoun 30005; Nanaimo, 7 Jun 1887, J. Macoun s.n. (US); Goldstream, Vancouver Island, 5 Jul 1887, Macoun 30009 (CAN); Goldstream, Vancouver Island, 5 Jul 1887, Macoun 30,010 (CAN, US); Deer Park, 1 A ' Lake, 18 Jun 1890, Macoun 30014 (CAN, US); Middle Selkirk Mts, Rogers Pass, 4500 ft, 1 Aug 1890, Macoun 11084 (CAN, US); Selkirk Mts, Rogers Pass, 4 Áug 1890, Macoun 30007 (CAN); Nanaimo, Vancouver Island, 14 Jul 1893, Macoun 135 (CAN); Cameron Lake, Vancouver Island, Jul-Aug 1899, 366 Jou 5 (DAO); Chilliwack River, 19 Jun 1901, J. M. A 26060 (CAN, US); Chilliwack Valley, 21 Jun 1901, 1800 ft, J.M. Macoun 26063 3 (US): Chilwack Lake, 2500 ft, 22 Jul 1901, J.M. Macoun 26062 (CAN, US); Chilliwack River, 2 Aug 1901, J.M. Macoun 26064 (CAN, US) River, 8 Aug 1901, 2000 ft, J.M. Macoun 26061 (CAN, US); Vancouver Island, 31 Jul 1902, C.O Rosendahl 856 (US); a 22 Jul 1902, C.O. Rosendahl 802 (US); Skagit Valley, 24 Jun 1905, J.M. Macoun 72949 (CAN, US); Glacier, Illecillewaet Valley, Loop Trail, 3500-4000 ft, 20 Jul 1906, S. Brown 656 (CAN); vicinity of Victoria, 18 Jun 1908, J. Macoun 78833 (US); head of Departure Bay, 23 Jun 1908, J. Macoun 76,848 (US); Departure Bay, near aces Station, 3 Jul 1908, Macoun 76846 (CAN, US); Mt Benson near Nanaimo, 13 Jul 1908, J. Macoun s.n. (US); vicinity of Victoria, 20 Jul 1908, J. Macoun 76845 (US); Colquitz River near Victoria, 20 Jul 1908, Macoun 76,850 (CAN, US); S shore of Cowichan Lake, 15 Jun 1909, C.O. Rosendahl 1819 (US); Bear Lake, 3 Jul 1911, W. Spreadbor- ough 87613 (US); Bear Lake, 5 Jul 1911, W. Spreadborough 87601 (US); Eburne, 3 Jun 1912, J. Davidson s.n.(UBC); Sidney, 7 Jul 1913, MO Malte 28 (CAN); vicinity of Sidney, 12 Jun 1913, J. Macoun s.n. (V); Mayne Island, 4 Jun 1914, J.M. Macoun 90083 (CAN); Oak Bay, near Victoria, 17 Jun 1913, J. Macoun 88,528 (CAN); Vancouver Island, Alberni, Jun 1916, W.R. Carter 906 (CAN); Cascade Range, near head of McGilliv Se Ge 5000 do 10 nas p J.M. Macoun 94,020 (CAN, US); Cascade Range, within n miles of o 22 Jul 1916, 3000 ft, IAM) R within five mi of Lillooet, 2500 ft, 22 Jul 1916, J 93,970 (CAN); Cascade Range, near head of McGillivray Cede 5000 ft, 1 Aug 1916, J.M. Macoun 93,976 (CAN); gen a Sen 20 Jun 1918, J.W. Kidman s.n. (UBC); Sidney, 4 Jul 1921, M.O. Malte 106,800 (CAN, DAO); Vancouver Island, Cowichan Lake, 25-26 Aug 1931, V. Kujala and A. Cajander s.n. (CAN); Vancouver Island, Victoria, Goldstream, 11-12 Sep 1931, V. Kujala & A. Cajander s.n. (CAN); Tabor Lake (Six-mile Lake), 6 mi E of Prince George, 14-18 Jul 1931, V. Kujala & A. Cajander s.n. (CAN); Mt between Chilliwack and Hope, 3000 ft, 9 Jul 1932, E. P. Hume s.n. (UBC); Athabasca Mill, Nelson, 4000 ft, 16 Jul 1937, J.W. Eastham 403 (UBC); Venus Mine, Nelson, 5000 ft, 18 Jul 1937, J.W. Eastham 404c (UBC); Hollyburn Trail, 1 Jul 1938, J.W. Eastham 406a (UBC); Beacon Hill Park, Victoria, 2 Jun 1938, J.W. Eastham 849 (UBC); Rossland Highway, Grand Forks, 18 Jul 1938, J.W. Eastham s.n. (DAO, UBC); Victoria, 11 Jun 1939, J.W. Eastham s.n. (DAO); Qualicum Falls Park, Vancouver Island, 31 Jun 1939, J.W. Eastham 401 (UBC); Mt Revelstoke, 3000 ft, 18 Jul 1939, H. Groh 186 (DAO); Goldstream Lake, Vancouver Island, 13 Jun 1940, I. Cowan ape A Mavis Lake, Vancouver Island, 13 Jun 1940, I. Cowan 8166 (UBC); Long Beach, Nelson, 17 Jun 1940, J.W. Eastham 7384 , 25 Jun 1940, J.W. Eastham 7540 (UBC); Lightning Peak Road, Monashee, 5000 ft, 14 Jul 1940, J.W. Eastham 7851 (UBC), Parkes ille, Vancouver Island, 10 Jun 1941, J.W. Eastham 8887 Ge Cultus Lake, Chilliwack, 11 Jun 1941, J.W. Eastham 8897 (UBC, US); Seymour Park, N shore, 21 Jun 1941, J.W. East} Alberni, Vancouver Island, 10 Aug 1941, J.H. Soper 8378 (DAO); Arrowsmith Trail, Cameron Lake, 14 Aug 1941, J.W. Eastham 9436 nn W Vancouver, 15th Street, 20 Jun 1943, JW. Eastham 10834a (UBC); Manning Park, 4000 ft, 25 Jul 1945, G.A. Hardy 14240 (UBO); Aleza Lake, 23 Jul 1946, J.W. Eastham 14809 (UBC); Vancouver, 120 ft, 19 Jul 1949, V. Krajina 74 (UBC); Botanic Valley, 22 Aug 1949, VJ. Krajina s.n. (UBC); Salmon River Valley 15 mi S of Sayward, 65 m, 3 Jun 1950, V. Krajina & R.H. Spilsbury 4302 (UBC), Qualicum Beach, 25 Jun 1950, 10 m, V. Krajina, R.H. Spilsbury & A. Szczawinski 4591 (DAO, UBC), Vancouver Island, Wolf Mountain, 300 m, 27 Jul 1950, V. rie i R.H. deus Lá & A. bei 5171 RE MacMillan Park, 185 m, 8 Aug 1950, V. Krajina, R.H. Spilsbury & A. Szczaw- inski 5331 (UBC); d Penticton, 5-7500 ft, 27 Jul 1950, D Martin s.n. (UBC); 0.5 mi W of 2nd Lake, 29 Jul 1950, V. Krajina, R.H. Ste GA, Seele 5195 (DAO); Vancouver Island, MacMillan Park, 185 m, 8 Aug 1950, V. Krajina, R.H. Spilsbury & A. Szczawinski 5329 (UBC); under Monashee Pass, 10 Jul 1952, VJ. Krajina s.n. (UBC); middle Nanaimo River Valley, 700 ft, 16 Jun 1955, D. Mueller-Dombois 22-8 (UBC); Ganges. as 1956, T.R. Ashlee 36 (UBC); Nanaimo River Valley, 700 ft, 11 Jul 1955, D. Mueller-Dombois 40-1 (UBC); Boundary Creek 10 mi N d, 30 Jul 1956, A. McLean s.n. (DAO); W Nanaimo River Valley, 1115 ft, 6 Aug 1957, D. Mueller-Dombois 18-1 (UBC); 9.5 mi on road Ges Beaver Lake to Likely, NW of Williams Lake, 28 Aug 1956, J.A. Calder, J.A. Parmelee & R.L. Taylor 20505 (DAO); Du Hamel Lakes, Nelson, Kootenay, 26 Jul 1958, M. Bell s.n (UBC); Sugar Lake, Cher- ryville, 28 Jun 1958, M. Bell s.n. (UBC); Big Qualicum River, Vancouver Island, 13 Jul 1959, L. Holm 524 ( (UBC); Nakusp, Kootenay, 2400 ft, 22 Jul 1959, M. Bell s.n. (UBC); E side of Lightning Lake, Manning Park, 4000 ft, 13 Jul 1960, K.I. Beamish & F. Vrugtman 60756 (DAO, UBC): Head of Finlayson Arm, N of Victoria, 16 Jul 1961, J.A. Calder & KI Mackay 31521 (DAO); Gail Falls on Kelsey Bay-Campbell River road, 12 Aug 1961, J.A. Calder & K.T. MacKay 32432 (DAO); Sarita River, 50 ft, 14 Jun 1963, A. Yound & W. Hubbard 395 (DAO); 1 mi E of Pilot Mountain Lookout, ca. 15 mi NE of Pri George, 1 Jul 1963, A. Auclair 15 (DAO); Kennedy River area, drainage of Ken- nedy River, 2 Jul 1963, A. Young & W. Hubbard 226 (DAO), Pilot Mountain Road, 1 mi E of the lookout, 2000 to 2400 ft, 2 Jul 1963, A. Auclair & S. Eis s.n. (DAO); Lawless Creed Road, 13.6 mi W of Tulameen, NW bari Princeton, 4500 ft, 16 Jul 1964, A. McLean s.n. (DAO); Mt Revelstoke National Park, 2040 ft, 22 Jun 1965, VJ. Krajina 65062233 (UB Mtn Road, 2.7 mi S of rock cut, S of Princeton, 5800 ft, 6 Aug 1964, A. McLean s.n. (DAO); E end of Durrance Lake, gege island. 134 m, 18 Jul 1967, A.S. Harrison 37.2 (CAN); McLeod Lake, 2250 ft, 19 Jul 1967, VJ. Krajina & Revel s.n. (UBC), Vancouver Island, Strathcona Provincial Park, 9 Sep 1967, K. Beder, V. Bedanr, S. Kojima & VJ. Krajina s.n. (UBC), Vancouver Island, Mill Hill, near Langford, 1 Jul 1969, J.H. Soper, M.J. Shchepanek & A.F. Szczawinshi 12243 (CAN); Vancouver Island, MacMillan Park, near Cameron Lake, 650 ft, 8 Jul 1969, J.H. Soper, M.H. Shchepanek & A.F. cal 12403 (CAN); Strathcona prenne ied 250 m, 8 Jul 1969, S. Kojima s.n. (UBC); Upper Sage Creek Valley, 5200 ft, 20 Jul 0, K.I. Beamish, M. Bell & N. Anderson 756 (1 Creek, 20 Jul 1970, K. Beamish, M. Bell & N. Anderson 756 (CAN); Mount E Robson Park, 3350 ft, 20 ec 1970, V. Krajinas n. . (UBC) National S d Trail no. 10, mile 7, Summit Road, 3,750 ft, 30 Jul 1970, J.H. Soper & M.J. Shchepanek 12740 ( ke Nat ios en (Silver Creek), along an no. 6, W Fork Woolsey Trail, 3 Aug 1970, J.H. Soper & MJ. Shchepanek 12809 (CAN o Gl National Park, Bald Mountain Trail to Grizzly Creek, at Beaver River crossing, 3600 ft, 27 Jul 1972, E. Haber & MJ. Shchepanek 1634 (CAN); Manning Park, Poland Lake area, 1768 m, : io 1973, i > cin 1056 Ge Point No Point, 23 Jul 1974, Hainault & Tanguay 7691 (DAO): ca. 2.25 mi NW of Prince George on “big bend”, 2000 ft, 30 Jul 1974, VJ. Krajina, J. Pojar & C. Parsons s.n. (UBO); Quesnel Saarela, Bromus in British Columbia, Canada 367 Region, 17 mi E of Quesnel on Barkerville Road, 17 Jul 1976, M. Barkworth 1634 (DAO); C.F.B. as Mt Lolo, 22 Jul 1976, M. Barkworth 1742 (DAO); Power River, W coast of V; Isl n 1979, C.E. Jones s.n. (DAO, UBC); W central Vancouver Island, 195 m, 20 Jun 1980, D. Gagnon & G. A. Spiers 80-60-11 (UBC); Mt EN 3 Jul 1980, B. Brookes 1171 (DAO); Vancouver Island, Shawnagan Lake Port Renfrew Road ca. 12 mi W of Shawnagan Lake, 2 Aug 1980, H.L. Dickson, M. Piche, D. Moennich, C. Barnard & R. Barnard 3790 (DAO); Lolo Mtn, 12 km E of Kamloops, 23 Aug 1980, B.M. Brooke s.n. (DAO); Vancouver Island, Goldstream Provincial Park, 30 Jun 1982, S.G. Aiken, S.J. Darbyshire & L.E. Pavlick 2104 (DAO); W coast Vancouver Island, Pachena Bay, SW of Bamfield, West Coast Trail, 7 Jul 1982, S.G. Aiken, S.J. Darbyshire & L.E. Pavlick 2214 (DAO, UBC); E of Kersley on Hwy 97, S of Quesnel, 2400 ft, 14 Jul 1982, S.G. Aiken, S.J. ei ixi & A. Roberts ae Wie Ge District, Timothy Mountain near Lac La na 5454 ft, 18 Jul 1982, S.G. Aiken La J (DAO); E si thew River, 930 m, 17 Aug 1982, L.E. Pavlich 82-346 (V Valley, Mount McGuire, NE ridge, 6 Sep 1984, Ceska 18066 M ied Goldstream Provincial Se E Sdn of Finlayson ir just outside park boundary, 30 Jun 1988, S.J. Darbyshire 3739 Island, S end, M LB 3 Jul 1988, S.J. Darbyshire & J. A. Darbyshire 3769 (CAN, DAO); Skye Meadows Ecological Reserve, SE of Chilliwack, 2 Jul 1989, G.B. Straley, R. Klinkenberg & B. Klinkenberg 5561 (UBC); Vancouver Island, Goldstream Park, 31 Aug 1993, J. Cayouette & S.J. Darbyshire C7503 PES Vancouver ae 2-3 Es E of Lizard Lake, 2 Sep 1993, J. Cayouette, S.J. Darbyshire & L.E. Pavlick C7516 (DAO); Vi Island, Port Alberni, River, 11 Sep DE J. Cayouette C7586, C7600 (DAO); oem Charlotte cape See Island, 6 km SSE of nae ee next to Blaine Ck., 3 m, 18 Jul 1997, F. Lomer & N. Grove 97403 (1 Lake, 12 mi NE Cranberry Junction, 22 Jul 2004, P.M. Peterson, J.M. Sg, & S. Smith 18694 (CAN, US); 9 mi NE of New EE? on road towards Cranberry Junction, 22 Jul 2004, P.M. Peterson, J.M. Saarela & S. Smith 18699 (CAN, US); Maroon Mountain Trail from end of road to ridge top, 23 jul 2004, P.M. Peterson, ad Saarela & S. pde 18726 (US, UBC); Mayne Island, Bennett Bay, Gulf Islands National Park ee dien Wilkes R y Road, 3 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 822 (CAN, UBC); Vancouver Island, [D k Road and Norris Road, S of Deep Cove, 4 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 838 (CAN, UBC); Vancouver landi Thetis Lake Regional Park, just N of Hwy 1, N of Langford, 4 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 853 (CAN, UBC); Vancouver Island, along Deering Road ca. 3 km N of Port Renfrew, 5 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chan ng 886 (CAN, UBC); Vancouver Island, 7.9 km along Horne Lake Road W of Hwy 19, 8 Jun 2007, J.M. Saarela, D.M. Percy & Y. Chang 975 (CAN, UBC); Queen Charlotte Islands, Graham Island, Towhill Ecological Reserve, along small ecozone road to beach, Naikoon Provincial Park, 22 Jul 2007, J.M. Saarela & D.M. Percy 1169 (CAN, UBC); Queen Charlotte Islands, Moresby Island, 21.5 km SW of Sandspit, E of Skidegate Lake, 0.3 km W of bridge at km marker 13, 2.2 km W of Y junction, 25 Jul 2007, J.M. Saarela & D.M. Percy 1294 (CAN, UBC); Queen Charlotte Islands, Moresby Island, 12.7 km S of Sandspit, 25 Jul 2007, J.M. Saarela & DM Percy 1296 (CAN, UBC); Queen Char- lotte Islands, Moresby Island, 11.2 km S of engin SE to e rdi 25 Jul 2007, J-M. Saarela & D.M. Percy 1300 (CAN, UBC); W of Smithers, along Hudson Bay Mountain Road, J-M. Saarela & D.M. Percy 1344 (CAN, UBC). Notes on Excluded Species There is a single, early collection of B. arvensis L. (field brome) from British Columbia [Invermere, 29 Jul 1915, M.O. Malte 108375 (CAN!, DAOU) In his distribution maps, Pavlick (1995) indicated that B. arvensis occurs in British Columbia, probably based on this collection. The annual B. arvensis, a species classified in B. sect. Bromus, is native to Europe; it occurs in scattered locations across North EE more commonly in the eastern U.S.A. (Pavlick & Anderton 2007). It is not known if B. o i ists in British Columbia; it therefore is not formally treated here. Should the species be collected SECH in Bh Columbia, it can be distinguished from other B. sect. Bromus species in the province as follows: anthers 2.5—5 mm [vs. anthers < 2.53) mm]; linear-lanceolate spikelets that narrow only slightly towards their apex [vs. ovate-lanceolate spikelets that narrow conspicuously towards their apex]; lower leaf sheaths with dense, softly appressed hairs 0.4—0.6 mm long [vs. lower leaf sheaths glabrous, sparsely to densely pilose with stiff hairs 0.5-0.6 mm long, or with soft and wavy hairs to 1.2 mm long]; and often purple-tinged spikelets (this colouration is not always present in dried specimens). Taylor and Macbride (1977) reported B. rubens L. [=B. madritensis subsp. rubens (L.) Husn.] from Brit- ish Columbia. Following this report, Salo (2005) indicated that red brome occurs in British Columbia, but she did not include any specimens on her distribution maps. There is a single collection of this species from British Columbia, collected over a century ago (Nanaimo, Vancouver Island, 13 Jul 1893, Macoun 118 CAN-39043!, photo: DAO-56101!). The species seems not to have persisted in British Columbia and it is therefore not treated here. Scoggan (1978:252) noted that B. orcuttianus Vasey had been previously reported from near Victoria, Vancouver Island. Bromus orcuttianus does not occur in British Columbia (see Saarela et al. 2005). 368 Jou ACKNOWLEDGEMENTS Field and herbarium work was supported by research grants from the Robert Bateman Arctic Research Fund (Smithsonian Institution), Genome Canada, the Lawrence R. Heckard Fund of the Jepson Herbarium and the Canadian Museum of Nature, and scholarships to JMS from the Natural Sciences and Engineer- ing Research Council of Canada and The University of British Columbia. Jack Maze provided funding for a three day collecting trip in and around the southern Okanagan Valley. I am grateful to Paul M. Peterson, Stephen Smith, Christopher J. Sears, Jack Maze, Diana M. Percy (and Dougal) and Ying Chang for help and discussion in the field; Linda Jennings, Cindy Sayre, Jennifer Doubt and Micheline Bouchard for help in the herbarium; and the curators of DAO, UBC, US and V for access to their specimens. Stephen J. Darbyshire kindly provided information on Bromus riparius and Frank Lomer provided information for several species. I thank S.J. Darbyshire, P.M. Peterson and Jacques Cayouette for their careful and insightful reviews of an earlier version of the manuscript. The illustrations are the work of Cynthia T. Roché, used with permission of the copyright holder, Utah State University. I thank Mary E. Barkworth for facilitating their use. REFERENCES Acto, C. and F. Llamas. 1999. The genus Bromus L. (Poaceae) in the Iberian Peninsula. Phanerog. Monogr. 22:1 -293. AiNoUcHE, M.L. and RJ. Bayer. 1997. On the origins of the tetraploid 8 species (section Bromus, Poaceae): insights from internal transcribed spacer sequences of nuclear ribosomal DNA. 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Remesh Forest ii^ ie o Conservation Division Forest Research Institute E e 653, Thrissur, Kerala, INDIA muktesh@kfri.o ABSTRACT M hloa (P Bam! ideae) g j tedand d ibed here | ] bamboo, which formerly was described under the genus Oxytenanthera Munro. A new combination, M hl itchiei (M ) M. Kumar & R | nov., is proposed RESUMEN A. 1.1 fT) D 1 J A H J 1 1 ] 1 1 1 1 lg Oxyt 1 A Cc E 1 RA. 1.1 4 ih "a F CT h, comb nov INIRODULG LION While working on revisionary studies e the BETEN Oxytenanthera Munro, Pseudoxytenanthera Soderstr. & Ellis, it ] | that Pseud iei (M ) Naithani, possessed EES not included in the genus P. 1 yt th Detalle ] I hol gl l studi led that tl number of 1 4 ] 1 li ti +] ` ] " af which are very distinct from the g ME E and Pseudoxytenanthera. Therefore. we are erecting a new genus to accommodate the species. It was Colonel Munro (1868) who first described the new species in his monograph on Bambusa. Bambusa ritcheyi, was based on a herbarium specimen collected by J.C. Ritchie from Kala Nuddi, Bombay, India. In the addenda of the same publication, Munro corrected the spelling of the ritcheyi to 'ritchiei' to commemorate Ritchie's name. After a lapse of five years Beddome (1873) collected this bamboo from Anamalai and included it under the genus Oxytenanthera Munro. Due to the presence of a monostigmatic ovary, Beddome named this species as Oxytenanthera monostigma. This species was subsequently reported by various workers from Western Ghats, hills of south west India from Ra NE to Anamalai up to Palakkad gap such as Brandis from Sattara Ghats in 1870; R.S. Fagan at Mahabalesl in 1892; WA. Talbot from North Canara in 1884 and 1889; R.C. Wroghton, from Pune district and A.D. Wilkins from Ahamed Nagar in 1892 (Gamble 1896). Gamble stated that Munro himself had also agreed with the new treatment as Oxytenanthera monostigma. Later, Brandis (1906) and Bourdillon (1908) followed this treatment. Gamble, noted that the species has very well-marked velvety culms, a narrow culm sheath, long narrow spikelets with only one flower, and a glabrous ovary and style. While working on this species at a later day, Gamble commented, "I am rather in doubt whether Munro's specific name should not und pee. one Es doubt was clearly noted by Blatter and McCann (1929) and y (Munro) Blatt. & McCann. Without considering the ee species name as "itchier. several workers erroneously spelled the epithet as ritcheyi in their publications. Nayar and Ansari (1982) also followed the species name spelling it ritcheyi. While preparing an account of the enumeration of Indian monocotyledons, Majumdar (1989) came to the conclusion that the Indian species of Oxytenanthera do not fit the generic description of Oxytenanthera. He erected a new genus, Pseudotenanthera, to include species having sub-scandent to scandent branches in tufts, no resting central bud, a thin solid style, a pericarp thin and separable from seed, and made a new combination Pseudotenanthera ritcheyi (Munro) R. Majumdar. Subsequently, Naithani (1990) treated ritcheyi in the genus Pseudoxytenantl f Soderstrom and Ellis T | Rot Dee Inst Tayac 2(1 Y 371. 378. 2008 374 I E Cel Dad H In LI bidit f Texas 2(1) (1988) to accommodate Oxytenanthera monadelpha (Thwaites) Alston which has a vine like culm, numerous basal branches, capitate cluster of spikelets, and six stamens. Naithani (1990) maintained that Majumdar's new genus, Pseudotenanthera, coincidentally possesses similar characters of Pseudoxytenanthera Soderstr. & Ellis. Therefore, Naithani considered the genus a superfluous name and transferred all the species known of Oxytenanthera from India to the genus Pseudoxytenanthera Soderstr. & Ellis. During revisionary studies on Indian Bamboos the authors conducted a detailed comparative account on generic characters of Oxytenanthera, Pseudoxytenanthera and noted that the Pseudoxytenanthera ritchiei (Munro) Naithani, possesses a number of distinct characters such as, completely solid culms, prophyllum buds without keels, inflorescence with strongly capitate heads, with leaf sheaths, palea without keels, and a EEN ovary. The other species Hodie under the genus Oxytenanthera possesses hollow culms, keeled p y of buds, Is without leaf sheaths, palea two keeled, and three stigmas. The L distinsvishimg characters of the species also were compared with other genera of the sub tribe Bambusinae such as Dendrocalamus, Bambusa and Gigantochloa It has been noted that the diagnostic characters like solid culms clothed with silky white tomentum; prophyllum buds and palea without keel; monostigmatic ovary, and united filaments, together make this species distinct and therefore, belongs under a new genus Munrochloa. Inter generic affinities The genus Munrochloa shows some affinities with Dendrocalamus, Oxytenanthera, Pseudoxytenanthera and Gigantochloa. (Table 1). The habit and habitats of this taxon resembles Denrocalamus stocksii and Dendro- calamus strictus both having thick walled culms for those clumps especially growing in dry areas. Though Munrochloa ritchiei grows even in fully eee areas a tudy solid culms. = nature of inflorescence is Ld; ——— M To almost similar to Dendrocalamus, Oxyt thera but in A are oe EEN wiii lear eai: The ide are ingle flowered. The palea of all the other members int keeled. In} lea is without any keels. The stigma of the species of Oxyt thera and Pseudoxyt thera is divided in to ye and Munrochloa has a monostigmatic ovary similar to Dendrocalamus and egen oa. The inter generic affinity of the Munrochloa shows that there is an interlink between Dend d Oxytenanthera and also having distinct generic status with few key characters like, imperfectly keeled prophyllum buds, fully solid culms, palea without any keels. = KEY TO MUNROCHLOA AND ALLIED GENERA 1. B apnd floret Md undivided stigm 2. Culms fully midculm propyl ana pateat Don D keeleo Munrochloa 2. me hollow, midcul 3. Inflorescence acne packed in semiverticellate clusters of spikelets, filaments of the anther free Dendrocalamus 3. Inflorescence loosely packed in semiverticellate clusters of spikelets, filaments of the anther e Gigantochloa 1. Bamboo floret with stigma divided into three. 4. Bamboos erect Oxytenanthera 4. Bamboos scandent or climbing Pseudoxytenanthera Munrochloa M. Kumar & Remesh, gen. nov. T Munrochloa ritchiei (Munro) M. K & Remesh. Oxytenanth imil tu fl t do, culmis us tomento albo-sericeo tectis, prophylli gemmis in culmo ti p ecarinata I , H DO An erect medium sized gregarious, bamboo forming loose clumps. Rhizome is Ln ien Culms are erect and solid. m Bp keeled. ws sheaths narrow, conical in shape. B Imost equal size | ai fl ompound paa terminal panicle, arises on the nodes of branches as d e 1 iated witl lonas Spikel 1] fl d, sometimes D 2, lanceolate, better? linear-lanceolate, glabrous. Palea membranous, E not keki Stamens 6, 375 Munrochloa F sisdoÁJe» prosdl||3 3[Buls Aey pue Duo? pau juesqy P9[99% ION SNOIGE|D P3/9MO|Y /BUIS JUsUILUOId ale syjeays Jea| pue ajeyides ABUS P3/32% JON 38195 [L10 INOUUM Aje\uawipnd 10 juasay 3SOJU3UJO] pue ubnoy sisdoKJe» prosdi[[3 SEIN Ailey pue Duc? Paun juəsqy Ppajsax c Aey Ajasseds POJIMON-Z JU3sqe yleaus Jeo| pue ajeydes ABuons Poe c 30135 [JO 1M pedo[o^ep |I9M snoJge|b pue u1oouJg sisdoÁ1e» posd1]13 əy Kley pue Duc? Dour 1U9SQY Pal99x c Aney Ájosieds PaJaMmo|-£—| juasa!d ÁjaJeJ u1eaus jes] pue ajeude» A[Buong p3[9o» c 38195 |210 JNOUZIM pedo[e^ap ||2AA snoJqe|b pue uoouisg sisdoÁ1e» Dose JO 2upuu4"? det Aney pue Duc paun Sa/D3ds Maj ui JUSSaJd inq 1ussqe ÁAnsow pjs) c ÁJey Ajasieds JO snouqes peJaMo|-p-c Juəsqe yleaus je] pue peau e ul may 'ajeude» Ajasoo7 P2|99> c 38]95 |BJO 1nOU1IM JO ULM pado|3A9p ||9AA snolqe¡6 pue yl00uus sisdoKue» asoqo|5 3|buis Aey pue Duc? SEI juasay p3[99» c Aey Ájesieds peISMOJ-£-c juasge yleays Jes| pue 31e3ide» Ajbuons Pelee c 3025 |BJO INOYAM JO YUM pedo|s^ap ||3AA snoJ4ge[6 pue u1oouug Tu eublis SI SIS syuəwejy ayuy sa|A5IPO] 19Jexids Uieaus Jeo| JO aduasald pue Bulysueig e»ua2saJonu| si&udoud wn pr opune ueaus wind 32epns WIND Dos Aus MOIOH MO||OH MO||OH PI]OS/MO]|OH un pex»eu Huo] Jo pe»x»ou JJOYS pay)au pay>3u pex»eu Bug] JeipoduwAs pex»eu 110us |eipoduuAg JO Duo jeripoduuÁgs Lous [eipoduJÁAgS HOYS ¡erpodwÁs SQWOZIUY 313 ` Pulquuipiuues pue juapueos 19913 poy 19913 Op TO ET TO DIS3Y3UDUIJÁXOPN3SG DISYIUDUIJÁXO DOo[u20]ubDic) snuiD[pooupuaq SIS)RIe y) ‘elouab Dote YUM eOo|YdOJUNW Jo uosueduioo v *| av] 376 t tani ti Texas 2( monadelphous, strongly apiculate. Ovary glabrous, ovate; stigma single, curved plumose. Fruit a caryopsis, linear-oblong, faintly grooved with a small beak. Similar to Oxytenanthera Munro in general ap] solid culms clothed with silky white tomentum on the surface, imperfectly keeled mid culm prophyll buds, palea without keels and monostigmatic ovary. Etymology.—The generic name is to commemorate the name of Colonel William Munro (1818-1880), who had contributed the most valuable monograph on Bamboos of the world. This genus is represented only with a single species and endemic to Western Ghats of India. 1:4f type but differs in presence of Munrochloa ritchiei (Munro) a Kumar & igi diui nov. (Fig. 1). Bambusa ritchiei Munro, Trans. Linn. Soc. London 26:157. 1868. Oxyt J. Bombay Nat. Hist. Soc. 33:773. 1929, *ritcheyi"; VJ. Nair & R. Ansari, J. Econ. Tax. "Bet 3:616. 1982. Pseudotenanihera ri ritchiei a Gu R.B. Majumdar in Karthikeyan et al., Fl. In d. ser. 4, 2(Enum. Monocotyl.): 280. 1989. Pseud Naithani y Bombav Nat. Hist. Soc. 87:440. 1990; D.N. Tewari, Monogr. Bamboo 127. 1992; KK. Seethal. & M. Kürdar. Bamboos India 225. 1998. Pseudoxytenanthera ritchiei (Munro) Ohmb., Bamboos World 313. 1999, an isonym. Tere: INDIA: Bombay, Kala Eege C. SE 820 (LecrorvrE, here designated: K). O ld., Fl. Sylv. S. India 3:233. 1873. Gamble, Ann. Roy. Bot. Gard. Calcutta 7:74, t. 65. 1896 & in Hook. f., EI Brit. India 7: 402. 1897. ends Indian Trees 674. 1906. Bourd., Forest Trees Travancore 400. 1908. C.E.C. Fisch. in Gamble, Fl. Madras 3:1861. 1934. Tree: INDIA: Anamalai, R.H. Beddome s.n. (HOLOTYPE: K)]. Vernacular names.—Choomaree, chiwa, chiwan, Huda, manga, udhe, thandali (Marathi) Erankol and Korna (Malayalam) Specimens examined: INDIA. KARNATAKA: Uttara Kannda Dist. s. loc., 1884, WA Talbot 583 (BSD); Feb 1889 WA Talbot 906 (CAL); Dandeli, 21 Jan 1924, RN Sarkar 2142 (DD). KERALA: Kannur Dist. Panathur, 28 Jan 1979, VJ Nair 59948 (MH); 28 Jan 1979, Panathady, 28 Jan 1979, VS run 59291 EE Malappuram Dist. Nilambur, 11 EN HG aech 1135 (DD); Manikkamudy, 6 Feb 2001, M Remesh & NU FRI); Vazhikkadavau, 7 Aug 1983, P! 63 (CALI). Palakkad Dist. Manthanpotti, 20 Nov 1999, M. Kumar & Stephen Sequiera 20635 (KFRI) Distribution and ecology.—This species is endemic to Western Ghats. It is distributed in northern Kerala and Karnataka. It was also reported from Maharashtra. It is found growing from an altitude of 200-1100 m. It is a component of moist deciduous forests and also found as pure patches. Sporadic flowering is common during summer months. Gregarious flowering was observed in the year 2001 at Nilambur forests, Malappuram district, Kerala. Conservation status.—This potential bamboo of south India is extracted for various uses. The recent study by the authors revealed that the species is Conservation Dependent as per the IUCN standards and needs appropriate tion and management strategies for a sustainable utilization of this species. es e solid culms of this bamboo are used for making furniture, lathi, etc. It is also used as a support for betal plants, for making baskets, umbrella handles and walking sticks. ACKNOWLEDGMENTS The authors wish to acknowledge J.K. Sharma, Director, Kerala Forest Research Institute, Peechi for his keen interest and encouragement and for the facilities. The help rendered by S.A. Renvoize, Herbarium, ie Botanic garden, Kew for ae the cibachrome sheets of the type specimen deposited in K and the authorities of the Indian Herbari as CAL, CALI, DD, BSI, MH, and BLAT for the permission to consult the specimens deposited in the respective herbaria are gratefully acknowledged. The authors express their gratitude to J.F Veldkamp, Nationaal Herbarium, Nederland, Leiden The Netherlands, for providing the Latin diagnoses. The authors are also thankful to the Ministry of Environment and forest, Govt. of India, New Delhi, for the financial support under AICOPTAX Scheme for undertaking this study. We thank Kanchi Gandhi and two anonymous reviewers for their help. 377 : Munrochioa, : D Inflorescence; E Spikelet; d H. Floret; I. Style; J. Fruit. ; G. Palea; F. Lemma; 378 REFERENCES BeDDome, R.H. 1873. The flora Sylvatica for southern India. Madras. 3:232-233. Damp E. and McCann. 1929. Gramineae. In: Blatter E. Revision of the flora of the Bombay Presidency Part X. J. Bombay Nat. Hist. Soc. 33:753-775. BOURDILLON, J.F. 1908. The forest trees of Travancore. The Travancore Govt. Press, Trivandrum. Branois, D. 1906. Indian trees. Bishen Singh Mahendra Pal Singh, Dehra Dun. GAMBLE, J.S. 1896. The Bambuseae of British India. Ann. Royal Bot. Gard., Calcutta 7: 75-76. Munro, W. 1868. A monograph of Bambuseae. Trans. Linn. Soc. London 26:126- p MaAJumDAR, RB. 1989. In: S. Karthikeyan, et al. Flora Indicae, ser. 4, vol. 1. Enumera tio tyled Botanical Survey of India, Howrah, Calcutta. Pp. 274—283. NarrHANI, H.B. 1990. Nomenclature of Indian species of Oxytenanthera Munro. J. Bombay Nat. Hist. Soc. 87:439-440 Nair VJ. and R. Ansari. 1982. Correct name of Oxytenanthera monostigma Bedd. (Bambusaceae). J. Econ. Taxon. Bot. 3:616. SopersTROM T.R. and R.P. Eus. 1988. The woody bamboos (Poaceae: Bambuseae) of Sri Lanka: A morphological- anatomical study. Smithsonian Institution Press, Washington. TWO NEW SPECIES OF CYPERACEAE FROM PENINSULAR INDIA M.A. Wadood Khan Department of Botany Majalgaon Arts, Science & Commerce College Majalgaon-431131, District: Beed Maharashtra, INDIA P Lakshminarasimhan Central National Herbarium Botanical Survey of India PO. Botanic Garden, Howrah-71 1103 West Bengal, INDIA ABSTRACT Two new species, Cyperus karthikeyanii (from Karnataka) and Fimbristylis naikii (from Maharashtra and Ándhra Pradesh), are A .I A = 27] + aux]. pz t jus ralh l . E së TE: Es asas tis A: +t T 11: J : 1 SE BEER | r + E RESUMEN I r = 7 LO E st se ei An] 1 A 11 nh ] Ly ADIA 1 t Tos 1 $2. kac F 5 VE J LOUER Bot ¥ELO. INL RODUCTION Vary 1 ] ^ A EPA a The genera Cyperus L. and Fimbristylis Vahl consist of ca. 300 and 250 in tropical and warm regions of the world (Mabberley 1997). Karthikeyan et al (1989) in his enumeration of Monocotyledons, listed 86 taxa (71 species) of Cyperus and 102 taxa (90 species) of Fimbristylis from India. During the course of botanical explorations of the tropical evergreen forests in the Western Ghats, two interesting specimens belonging to Cyperaceae were collected viz., Cyperus (Section Mariscus) and Fimbri- stylis (Section Fimbristylis, Series Dichotomae). Based on a perusal of the literature, comparison with the herbarium collections at Herbarium of Cyperaceae, Majalgaon College, Majalgaon (HCMCM), BSI & CAL, and our own field observations, the specimens were found to be very distinct and both are described here as new species viz., Cyperus karthikeyanii and Fimbristylis naikii. 1. Cyperus karthikeyanii Wad. Khan & Lakshmin. Pa nov. (Fig. 1). Tyee: INDIA. Karnataka STATE: Udupi, 20 Sep 1999, Sardesai s.n. (HOLOTYPE: CAL; ISOTYPES: HCMCM, Aahterte Cyperus clarkei T. Cooke et C. aed es Boeck. habitu stolonifero uci ad d cormo ied pub pide: gluma , sed habitu tenuissimo, st I | fructifera involuta dorsalite y dado E m 3 1 1 1 x 14 Az, " ALL ALLA LK, q 31: spicis brevioribus (4—6 mm), | longioribus (ca. 2.5 if tol tol lender, wiry, 1-2 cm long, 1.25-1.5 mm thick, Perennials, 15-35 cm tall with tul covered with purple-brown, strongly nerved, ovate or ovate-lanceolate scales (2.5-4 x 1-2.5 mm) which soon become fibrous; tubers ellipsoid, purple-brownish, 4-5 mm long and broad, covered with fibrous remains; stem trigonous, filiform to rather slender, 0.5-0.7 mm thick, glabrous, with often corm like or enlarged base. Leaves basal, 0.5-1 mm wide, flexuous, shorter to, as long as or rarely overtopping the stems, scaberulous on margins in upper half, caudate-tipped; sheath pale to dark purple-brownish, fibrous with age. Inflorescence of single, terminal capitate head of 2—5 sessile clustered spikes under 1 cm long and broad; involucral bracts 3—5, setiform, the others foliaceous, the longest one 6—20 cm long, spreading or sub- erect, glabrous, subcylindricaly oblong, 4-5 mm long, greenish with up to 15 spikelets. Spikelets trigonous, ellipsoid, deciduors, 3.8—4 x 0.5-0.7 mm, narrowed towards both the ends, curved, whitish, tinged with purple near base or purple dotted, one flowered. Glumes 4, empty glumes 2, persistent on rachilla, broadly hyaline margined, unequal, the larger one broadly to suborbicular-ovate, ca. 1 mm long, several nerved, the second smaller one lanceoiate; fruiting glumes 2, subequal, the basal one 3.8—4 x 0.7 mm, enrolled, tightly wrapping the nut, multinerved, herbaceous, acute, rounded on the back, the upper one narrowly | Ent Ras Inst Tawas 2(1): 170 _ 384. 2008 380 a ett ae ^ E et? + HP "Wad Khan & Lakshmin. 1a—b. Habit; 2. S Fic. 1.6 d wai 4, Nut; 5. Style with stigmas; 6. Stolon scale; 7-8. Empty glumes. Li Id El - E [| , Al H E e E [| y: 381 lanceolate, hyaline, uninerved, acuminate (with awn-like tip). Stamens 3. Styles 3—fid, filiform, 0.8-1 mm long, glabrous; stigmas shorter than the styles. Nuts acutely trigonous, ellipsoid- EE 2 qe SE é 4-0.6 | mm, curved, narrowed towards both the ends, pale orange-yellow, puncticulate with cells; gynophore obscure. In its stoloniferous habit with usually corm-like base of stems, the one fruited spikelets and involute fruiting glume with rounded back, it shows affinities with Cyperus clarkei 1. Cooke and C. paniceus (Rottb.) Boeck. but differs essentially by the extreme slender habit, the tuberiferous slender stolons covered with fibrous remains, the smaller (4-6 mm long) spikes, the some what longer nuts (ca. 2.5 mm) and the 3 sta- mens. Habitat and ecology.—Occasional under shade of thick semi-evergreen forests (1000 m above m.s.1.). Flowering and fruiting.—September-November. Etymology.—This species has been named after Dr. S. Karthikeyan (ex Deputy Director, Botanical Survey of India) in appreciation of his contribution to the field of systematic botany. PARATYPES: INDIA. KARNATAKA STATE: Londha Forest, 27 Nov 2000, Khan 4759 (HCMCM); Ramnagar Forest, 22 Nov 2007, Khan 1205 CMCM KEY TO NEW AND ALLIED SPECIES . Stolons tuberiferous, 1—2 g; st to rather slender, 0.5-0.7 mm thick; spikes 4-5(-6) m long; nuts ca. 2.5 mm long C. karthikeyanii . Stolons etuberiferous, 5-10 cm long; stems and leaves not filiform or if stems sometimes slender (0.5-1 mm thick in C. paniceus var. paniceus) the leaves (1-)2-6 mm wide; spikes 8-15 mm long; nuts 1.5-2.2 mm long. n — —À 2. Stems distinctly quasi bulbosely thickened at base; leaf-sheaths straw colored; rays of inflorescence if present up to 4 mm long; nuts 1.5-1.7 mm lon C. clarkei 2. on not or slightly bulbose at base; E often purplish; rays if present up to 4 cm long; nuts 1.8-2.2 mm long C. paniceus 2. Fimbristylis naikii Wad. Khan - Lakshmin., sp. nov. (Fig. 2). Tyre: INDIA. MaarasHT Ra: Kolhapur-Amboli Road, 18 Oct 1995, Khan 4341 (HoLorrre: C I Fimbristylis t Vahl valde similis sed gynopl i pl | f EEN 0.1-0.2(-0.3) mm longo latoque, nuce late bovoid ficie ordinati btrabecul I l linati bus 15-18 verticalibus E Cp UAM constante, nuce ad apicem Seege mutica Ke E e apiculata, EE 3-9 in inf] dichotomam (L.) Vahl etiam ut glumaru il joribus (1.2-1.6 x 08-12 mm) atque series plus quam 15 vertical Jul jetil ibus i is epid lium fi ibus differt Annuals, 10-35 cm tall; stems slender, trigonous to 1 trigonous, 0.3-1 mm thick, ribbed, glabrous, rather rigid. Leaves ligulate, subbasal, 1-2 mm wide, Y 2 to 3/4 or more as long as the stems, flat with often thick rib-like margins, hairy with long white cilia or glabrescent, acute; sheaths densely hairy, glabrous or glabrescent, pale ferrugineus or dark brownish. Inflorescence small, simple to subcompound, 1.5-3.5 cm long and broad with usually 3-9 spikelets; rays if present 1—3, each one bearing usually paired (rarely 3) spikelets; involucral bracts up to 6, 1-2 foliaceous, the other ones setiform, the longest one as long as to much overtopping the inflorescence up to 6 cm long, long ciliate at least on margins and base. Spikelets all solitary, ovoid or ovoid-oblong, 3-7 x 23.5 mm, acute, brown-white variegated, many flowered; peduncles 2—5 mm long; rachilla EEN Glumes spiral, some what loosely imbricated, ovate to broadly ovate, 2-2.8 x longer than broad, multistriate on sides, obtuse or acute, distinctly + T; A | Pd apiculate, often enrolled in lina, seddish: brown with whitish keel portion on the back, broadly hyaline margined; keel 3-5-nerved with strong mid nerve. Stamens 1 or 2; anthers elliptic-oblong, 0.4-0.5 mm long, apiculate. Styles flat, shorter than the nuts, 1-1.2 mm long, conspicuously broader than the staminal filaments, fimbriate throughout, not hyaline margined, pyramidally dilated at base or somewhat gradually narrowed upwards; stigmas shorter than the styles, shortly hairy. Nuts biconvex, broadly obovoid, 1.5—1.6 x 0.8-1.2 mm, turgid in the centre, subtrabeculate with 15-18 vertical striations in between transversely oblong thin walled epidermal cells, distinct in the centre and on the shoulder, nearly quadrate near base 1.8-2.2 mm, almost as] 382 Ae v TC schéi Ca A. y » H e MA e IER ti ios PEN INTR 1 Ba der IA wit S. ARI SUNT MP "i f e oe deg lk dir inu M PANE nut ve ons a pė =, we ra A id A a ae E r m a re a A od e & p a | G 4 EH are ` ek: a A A ae Ka 7 Style: 8 St C 3-4. Glumes; 5—6. Nut 9a—c. Fimbristylis tomentosa Vahl — nut, gynophore and epidermal cells (for comparison). * F Fic. 2. Fimbristylis naikii Wad. Khan & Lakshmin. 1. Habit; 2. Spikelets Stamen; , WE FI 1.1 q » [| ; RE D Er. £ [| ye 383 and apex, and on shoulder, apex rounded to subtruncate, muticous; gynophore discoid or cupoid, small, 0.1—0.2(-0.3) mm long and broad, not or obscurely lobed, often reddish-brown (very rarely dull whitish), its thin rim often appressed to the nut base. It is very similar to F. tomentosa Vahl but differs from the latter by the smaller, usually reddish-brown, 0.1-0.2(-0.3) mm long and broad iad abe the ee? E nut with subtrabeculate, tuberculate surface pattern with 15-18 vertical f cells bt muticous or obscurely apiculate apex together with 3-9 spikelets in smaller misen Apparently it SES approaches F. dichotoma (L.) Vahl and differ in having many nerved sides of glumes, the larger (1.5-1.6 x 0.8-1.2 mm) nuts with more than 15 vertical rows of thin walled epidermal cells. Habitat and ecology.—-Occasional in wet open grassland and eutrophicated marshes. Flowering and fruiting — October-November. Etymology.—The species has been named after Dr. V.N. Naik (teacher of senior author) in appreciation of his outstanding contribution in the field of taxonomy. PARATYPES: INDIA. MAHARASHTRA. Kolhapur District: Shivaji University Campus, 18 Oct 1995, Khan 43374 (HCMCM); way to Gadh- inglaj to Ajra, 11 Oct 2004, Chavan 5314, 5317 (HCMCM). Anpura Prapesu. Medak District: Patancheru, 30 Oct 2005, Solanke 824 (HCMCM) Notes.—This and other 3 taxa viz., F. tomentosa Vahl (= F. podocarpa sensu C.B. Clarke, 1893; = F. dichotoma (L.) Vahl subsp. podocarpa sensu T. Koyama, 1985; = F. dichotoma var. pluristriata sensu Napper 1971), F. alboviridis C.B. Clarke, and the African F. striolata Napper are very closely related to each other. Authors such as Clarke (1893), Kern (1974), Kral (1971), and a few others treated E tomentosa distinct specifically from F. dichotoma. Va With its smoothly t surface pattern having more than 15 fine close vertical striations or rows of thin walled cells and conspicuous tuberculations, F. naikii is very unlike to F. dichotoma or F. merrillii J. Kern, but somewhat approaches to F. alboviridis C.B. Clarke. The latter is distinguished by the smaller nuts (0.8—1 x 0.5-0.7 mm) and white-brown variegated nerveless glumes (ca. 2 mm long). The new species is undoubtedly more close to F. tomentosa and can be thus distinguished from the latter and other allies as under. KEY TO NEW AND ALLIED SPECIES 1. Nuts with 15-24 fine close p SE or rows of thin walled EDO NON E smoothly reticulate or subtrabeculate, usually large, pA TOPE large, ae 0.7 mm lo cu robconical, omen produced below, 3-lobed, whitish with thick t Dase; Nuts broadly ellipsoid or suborbicular, 1.9-2.2 mm long, acutish at apex, etuberculate (rarely with 1-3 faint tubercles), ie from isodiametric hexagonal or rounded cells, arranged in 20-24 vertical rows or striations F. tomentosa 2. Gynophore small 0.2-0.3 mm long, cupoid, not lobed, always brownish, with thin rim appressed to nut base (appearing continuation of nut-body); nuts broadly obovoid, 1.5-1.6 mm long, rounded to subtruncate at apex, densely tuberculate, subtrabeculate from some what transversely oblong or elliptic cells arranged in 15-18 vertical rows or striations F. naikii . Nuts with 5-10(-12) vertical ridges or rows of thick walled transversely oblong (bar-like) epidermal cells, i.e. coarsely trabeculate, 0.8-1.2 mm long; glumes be on GES more or less hairy throughout the leng |... F. dichotoma 3. Styles e or almost glabrous with sometimes few cilia at top, not hyaline anna nuts Deeg tuberculat | F. merrillii ACKNOWLEDGMENTS 1 L] 4 1: The authors are grateful to David Simpson, Royal Botanic Gardens, Kew for his g g the species. Help rendered by Wilmot-Dear (Melanie Thomas) and Katherine Challis, Royal Botanic Gardens, dens, Kew for the Spanish Kew for Latin En MATS 1 Lourdes Rico and Patricia Malcom, Royal Bot T J 1. The senior author is thankful to University Grants Commission for financial assistance nde Minor and Major Research Projects and M.B. Gadhave, Principal, Majalgaon Arts, abstract is also e 384 Journal of tl tanical h Insti Texas 2( Science & Commerce College, Beed for encouragement. One of us (PL) is thankful to the Director, Botani- cal Survey of India, Kolkata for providing facilities and to M.S. Mondal, Joint Director, Central National Herbarium, Botanical Survey of India, Howrah for constant encouragement. REFERENCES Case, C.B. 1893. Cyperaceae. In: J.D. Hooker, ed. The flora of British India 6:585-748. L. Reeve & Co., London (vol. published in 1894). KARTHIKEYAN, S., S.K. JAIN, M.P. Nayar, and M. SANJAPPA. 1989, Florae Indica ti tyled Botanical Survey of India, Calcutta. Kern, J.H. 1974. Cyperaceae. In: C.G.G.J. van Steenis, ed. Flora Malesiana 1, 7(3):435-753. Noordhoff International Publishing, Leyden, The Netherlands. Koyama, T. 1985. Cyperaceae. In: M.D. Dassana yake & FR SEI eds. Arevised handbook to the flora of Ceylon 5:125-405. Published for Smithsonian Institution and tl | | Science Foundation, Washington, D.C., by Amerind Publishing Co. Pvt. Ltd., New Delhi. KraL, R. 1971. A treatment of Abildgaardia, Bulbostylis and Fimbristylis (Cyperaceae) for North America. Sida 4:57-227. Mapp pe DJ. 1997. The plant book. ed. 2. (Reprinted with corrections, 1998, 2000). Cambridge University Press. Napper, DM. 1971. Fimbristylis, Scleria and Diplacrum (Cyperaceae) in tropical West Africa. Kew Bull. 25:435-446 LYONOTHAMNEAE, A NEW TRIBE IN THE ROSACEAE (ROSALES) Luc Brouillet Herbier Marie-Victorin Institut de SE en Se végétale rsité de Montréal 4 " I rue ES Québec, CANADA, H1X 2B2 luc.brouilletgumontreal.ca ABSTRACT Recent molecular yir aim of Rosidae and Rosaceae, notably the Potter et al. (2007) paper, have led to changes in the circumscription and classification of Rosaceae. In the Potter et al. Pape Ke was ge outside SC STEE di It had been traditionally classified in pande Quillaja is now a member o is erected to house Lyonothamnus. ABSTRACI Recientes análisis ee a E Rosidae y aes o el Ge de Potter et e i ge EE cambios en la ci E y et al., L j ] tribu reconocida Xa sx (el ] fe? sl t qos 1 Q :11 j : Q ;1] ij 1 Ja] H d | Q 211 j (F 1 les). Se ] tril gené ica, I y tl , para albergar a Lyonothamnus Molecular phylogeneti l f the angi d, m ifically, of Rosidae Tahkht. (summarized in eege Phylogeny oon 2003) eg Rosaceae ee. (e.g., Morea et al. 1994) have substantially changed our perception of the relationships of Rosaceae, and have resulted in slight changes to its circumscription. Traditional Rosales Bercht. & J. Presl (e.g., Cronquist 1981) included families that are now dispersed in many orders and subclasses (Angiosperm Phylogeny Group 2003). Rosales now has a circumscription that includes members of the former Rhamnales Linnk and Urticales Juss. ex Bercht. & J. Presl (Angiosperm Phylogeny Group 2003). Similarly, phylogenetic analyses have led to a new classification of the Rosaceae deeg et al. id IS a reorganization of subfamilies and, to a lesser extent, tribes. Hypotheses of rela- + LI 1:64 Se ke ALIAS MMS AA LL AS d subfamilies w as a result of this analysis. One may pa for instance, the traditional classification used by Kalkman (2004) to that of Potter et al. (2007): the classic subfamilies Spiraeoideae Arn., Rosoideae Arn., Prunoideae Horan., and Maloideae Weber, rec- ognized in the former, were reduced to three in the latter, Rosoideae, Dryadoideae Sweet, and Spiraeoideae. Ihe last subfamily now incorporates genera formerly included in three subfamilies, including Prunoideae and Maloideae. In the phylogenetic analysis of Potter et al. (2007), Lyonothamnus A. Gray appeared at the base of Spiraeoideae, sister to all other members of the clade. In their classification, Lyonothamnus was not assigned to a tribe. Historically, Lyonotl has been considered part of tribe Quillajeae Endl., along with Quillaja Molina. Quillaja is now treated as the only genus in o ae D. Don of the Fabales Bromhead (Angiosperm Phylogeny Group 2003). The subfamilial and tribal classification of Potter et al. (2007) is be- ing used as the basis for EE Seege in the Flora of North America project (volume 9, Rosales in part). Currently, no tribal f (Pankhurst 2007; Reveal 2006). Therefore, I am proposing the creation of a tribe, irons made to accomodate Lyonothamnus. Tribus Lyonothamneae Brouillet, tribus nov. Te: Lyonothamnus A. Gray, Proc. Amer. Acad. Arts 20:291. 1885. Arbores inermes Folia opposita, i plici 1 t | IH t ta El : icalyx (1-)2—3-brac teolatus; o S torus nullus: carpella 2), distincta, libera, styli terminales, distin e 2- 4 vel plura, fasciculat [elo e] Trees unarmed. Leaves opposite, DST or compound, stipules caducous, free; venation pinnate. Flowers J. Bot. Res. Inst. Texas 2(1): 385 — 386. 2008 386 J the Botanical h Insti Texas 2(1) perigynous; epicalyx of (1-)2-3 bractlets; hypanthium campanulate; torus none; carpels 2(—3), distinct, free, styles terminal, distinct; ovules 2-4 or more, clustered. Fruits aggregated follicles (follicetum), styles tardily falling. x = 9. ACKNOWLEDGMENTS The author wishes to thank David Morgan, Jim Reveal, and John Strother for their helpful comments on the first draft of this paper. REFERENCES ANGIOSPERM PHYLOGENY GROUP. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG ll. Bot. J. Linn. Soc. 141:399-436. Cronauist, A.J, 1981. An integrated system of classification of flowering plants. Columbia University Press, New York. KALKMAN, C. 2004. Rosaceae, In: K. Kubitzki, ed. Flowering plants. Dicotyledons: Celastrales, Oxalidales, Rosales, Cornales, Ericales. Vol. VI in K. Kubitzki, series ed., The families and genera of vascular plants. Berlin. Pp. 343-386. Morgan D.R, D.E. Soutis, and KR. RoBERTSON. 1994. Systematic and evolutionary implications of rbcL sequence variation in Rosaceae. Amer. J. Bot. 81:890—903. PANKHURST, R. 2007. Lyonothamnus. In: Rosaceae database. Provisional global plant checklist. International Orga- nization for Plant Information (IOPI). [http//bgbm3.bgbm fu-berlin.de/iopi/gpc/P TaxonDetail.asp?NameCa che=Lyonothamnus&PTRefFk=18 (accessed Oct. 2, 2007)]. Porter, D., T. Eriksson, R.C. Evans, S.-H. Oh, LEE. Smepmark, D.R. Morgan, M. Kerr, KR. ROBERTSON, M ARSENAULT, T.A. Dick- INSON, and C.S. CampeELL. 2007. Phylogeny and classification of Rosaceae. Pl. Syst. Evol. 266:5-43. ReveaL, J.L. 2007. Rosaceae. In: Indi i prag i plant vascularium. [http://www.plant- ystemati : l/pbio/fam/famRS.html (accessed Dec. 11, 2007)]. THE TAXONOMY OF NORTH AMERICAN LOTI (FABACEAE: LOTEAB): NEW NAMES IN ACMISPON AND HOSACKIA Luc Brouillet Herbier Marie-Victorin Institut de recherche en biologie végétale jal 4101 Sherbrooke St. E Montreal, Quebec, CANADA H1X 282 luc.brouilletgumontreal.ca ABSTRACT ke ëch +l 4 4 £ ¿1 H £ +1 J EZ fend T Aë 1 J Cal 4 4 fel f 1 gi ENT aL e I America, the author "eo two genera within North EN Loteae: Hosackia and Acmispon (including Ottley d Syrmatium) New names are provided for varieties within Hosachia, and I I ly included within Acmispon RESUMEN E 1 Ease 3 ral 1 Ian Ant D 1.1 — Es 1, El a PP A pret g The Jepson Manual” y grupo[ , l aut dos gé las Lot | j Hosackia y Acmispon (que incluye Ottleya y Syrmatium). Se aportan nuevos nombres para variedades de Hosackia, y para especies y variedad incluidas previ Acmispon The taxonomy of Lotus s.l. (Fabaceae: Loteae) has been controversial, particularly with respect to the issue of including or excluding the North American species from the genus; Cullen (1959) and Sokoloff (1999, 2000) together provide a summary of the taxonomic history involved. Following Greene (1890), it became usual (though not unanimous) to treat North American species within Lotus s.l., as was done, among others, by Ottley (1923, 1944, cl (1981, 1993), Barneby (1989), Arambarri (2000), and Stenglein et al. 2003). Traditionally, three subgenera have been recognized in North America: Hosackia, Syrmatium, and Acmispon, the latter subdivided into sections Microlotus and Simpeteria. In the last 10 years, however, morphological studies have shown that North American taxa were distinct from Eurasian Lotus s.s. Therefore, Sokoloff (1999, 2000a, b; Sokoloff & Lock 2005) proposed the recognition of four genera of North American Loteae: Hosackia, Acmispon, Syrmatium, and Ottleya (section Simpeteria segregated), corresponding with the four traditionally used subgeneric groups mentioned above. This taxonomy was accepted by Arambarri et al. (2005) in their numerical taxonomic study of the American species. The molecular phylogenetic analyses of Allan and Porter (2000) and Allan et al. (2003), based on the nrDNA internal transcribed spacer (ITS), confirmed that the North American taxa were distinct from Lotus s.s. and resulted in the recognition of two clades among American species: pas EE clade and the clade comprising the subgenera icto id d. arlade Syrmatium, Sé section S pet ] Ifrerealled the A cmispo etic on both morphological (e.g., aporte pollen, leafy stipules) and molecular bind except for he unexplained position of L. aboriginus in Allan and Porter (2000). Nomenclatural transfer of some varieties recognized in Hosackia species has yet to be done, however. The taxonomic disposition of species within the Acmispon clade (Allan & Porter 2000; Allan et al. 2003) (with 4-aperturate pollen and gland-like stipules, among others) appears less clear-cut. While there was some resolution within the clade in the ITS-based phylogenetic study of Alan and Porter (2000), Allan et al. (2003) showed a well-supported but unresolved Acmispon clade. In the Allan and Porter analysis, sub- genus Syrmatium appears monophyletic, with L. salsuginosus (Acmispon maritimus, sect. Acmispon, Sokoloff 2000) — L. aboriginus (Hosackia) as sister to the subclade. A second subclade includes members of sections Simpeteria (Ottleya, Sokoloff 1999) and Microlotus (L. wrangelianus; Acmispon sect. Anisolotus, Sokoloff 2000). It is worth noting that Ottley (1944) included her new section Simpeteria within Lotus subgenus Acmispon. J. Bot. Res. Inst. Texas 2(1): 387 — 394, 2008 388 D [| Cal D ad H ip LI nr f Texas 2(1) Thus Acmispon, as defined by Sokoloff (2000) and as traditionally understood, appears polyphyletic in the ITS phylogeny, each section belonging to a separate clade. Only two Va dr ein dde were included in the study, which prevents further conclusion. i contrary to what y suggest, it does not seem possible to clearly delimit tl ps within the clade. Therefore, I am suggesting that two genera should be recognized among d mcum) Loti at the present time, corresponding to the two clades recognized in molecular analyses: Hosackia and Acmispon (the oldest generic name available for the Acmispon clade). Given the ambiguity concerning the monophyly of lineages within Acmispon, I prefer not to recognize subgeneric taxa at the present time, even though the Syrmatium group appears monophyletic. The delimitation of such taxa will have to be evaluated using more complete molecular and morphological data sets. The purpose of this paper is to provide the nomenclatural combinations that are needed in Hosackia and Acmispon, particularly in view of the upcoming treatments of these genera for the second edition of The epson Manual and for the Flora of North America project. I am mainly following the taxonomy of Isely (1981, 1993) for the North American species. A list of already accepted names in Hosackia and Acmispon for North America north of Mexico is provided. NEW COMBINATIONS IN HOSACKIA Hosackia crassifolia Benth. var. otayensis (M ex Isely) Brouillet, comb. nov. B Lot Greene var. otayensis Moran ex Isely, Brittonia 30: 466-467. 1978. Hosackia oblongifolia Benth. var. cuprea (Greene) Brouillet, comb. nov. Bastonym: Lotus cupreus Greene, Leafl. Bot. Obs. Crit. dd 74, 1904. Hosackia cuprea (Greene) Smiley, Univ. Calif. Publ. Bot. 9:257. 1921. Lotus oblongifolius (Benth.) Greene ) Ottley, Univ. Calif. Publ. Bot. 10(3):206. 1 Hosackia stipularis Benth. var. ottleyi (Isely) Beni comb. nov. B Lotus stipularis (Benth.) Greene var. ottleyi Isely, Brittonia 30:468—469. 1978 NEW COMBINATIONS IN ACMISPON Acmispon americanus (Nutt.) Ry db. Var. Melleri a aa comb. nov. BASIONYM: tots halen Britton, Bull. Torrey Bot. Club 17:312-323. 1890. F Jahrb. Syst. 25:228. 18 & Oosting, Spring Fl. enn 79. 1940. Acrispon helleri Min A. Heller, Cat. N. Amer. Pl. 205. 1914. Acmispon helleri (Brit- on) Small, Man. S.E. Fl. 687. 1933. Lot Clem. & E.G. Clem. var. helleri (Britton) Isely, Brittonia 30:468. 1978. Lotus IU Benth. var hell 30 itton) Kartesz & Gandhi, Phytologia 71:96. 1991 Acmispon argophyllus (A. Gray) Brouillet, comb. nov. Basion: Hosackia argophylla A. Gray, Pl. Nov. Thurb. 316. 1854. Syrmatium argophyllum (A. Gray) Greene, Bull. Calif. Acad. Sci. 2:147. 1886. Lotus argophyllus (A. Gray) Greene, Pittonia 2:149. 1890. Acmispon argophyllus (A. Gray) Brouillet var. argophyllus. Hosackia argophylla A. Gray var. argophylla. Hosachia argentea Kellogg, Proc. Calif. Acad. Sci. 3:38. 1863. Hosachia M n Gray var. decora I.M. Johnst., Bull. S. Calif. Acad. Sci. 17(2):63-64. 1918. Lot llus (A. Gray) Greene var. decorus (I.M. Johnst.) Ottley, Univ. Calif. Publ. Bot. 10(3):237-238. 1923. Lotus argophyllus (A. n Greens subsp. decorus (I.M. eg Munz, Fl. S. Calif. 446. 1974. o argophyllus (A. Gray) Brouillet var. adsurgens (Dunkle) Brouillet, comb. nov. Basionvm: Lotus argo- us (A. Gray) Greene var. adsurgens Dunkle, Bull. S. Calif. Acad. Sci. 39:197. 1941. Lotus argophyllus (A. Gray) Greene subsp. inia (Dunkle) PH. Raven, Aliso 5:326. 1963 Acmispon argophyllus (A. Gray) Brouillet var. argenteus (Dunkle) Brouillet, comb. nov. B Lotus argophyll (A. Gray) Greene var. argenteus Dunkle, Bull. S. Calif. Acad. Sci. 39:176. 1940 Hosacki ithopus G , Bull. Calif. Acad. Sci. 1:185. 1885. $ ti ithopus (G )G , Bull. Calif. Acad. Sci. 2:148. 1886 Lot (G , Pittonia 2:149. 1890. Lot gophyllus (A. Gray) G Publ. Bot. 10: 238. 1923. Lot g (G ) PH Raven, BE 5:326 1963. Lotus argophyllus (A. Gray) Greene var. hausse Durkis Bull s.d Calif hes Sci. 39:175. 1940. Hosackia venusta Eastw., Proc. Calif. Acad. Sci., Ser. 3, 1:103, t. 8. 1898. Hosackia ornithopus Greene subsp. venusta (Eastw.) Abrams, Ill. Fl. Pac. States 2:550. 1944. Lotus venustus (Eastw.) A. Heller, Cat. N. Amer. Pl. 7. 1900. Syrmatium venustum (Eastw.) Davidson & Moxley, Fl. S. Calif. 199. 1923. Brouillet, New n in Acmi JH ki 389 Acmispon argophyllus (A. mel epo var. fremontii (A. Gray) Brouillet, comb. nov. Basionym: Hosackia argophylla A. Gray var. fremontii A. Gray, Proc. Acad. Nat. Sci. Philadelphia 15:347. See Hosackia lala (A. Gray) Abrams, Ill. Fl. Pacific States 2:550. 1944. Lotus pu (A Gray) A Heller, Cat. N. Amer. Pl. 7. 1900. S ti (A Gray) A Heller, Muhlenbergia :67. 1913. Lotus argophyllus (A. Gray) Greene var. fremontii (A. ES Ginter Univ. Calif. Publ. Bot. 10(3):237—238. 1923. + IY 1 E C Watco YG Pittonia 2:148 Syrmatium niveum Greene, Bull. Calif. Acad. Sci. 2:148. 1863, 1890. Lotus argophyilus (A. Gray) Greene var. ni (G ) Ottley, Univ. Calif. Publ. Bot. 10(3): 237-238. 1923. otis ns (A. Gray) Greene subsp. niveus (Greene) Munz, Fl. s. Calif. 446. 1974. Acmispon Sr An (Greene) Brouillet, comb. nov. Basionya: Hosackia argyraea Greene, Bull. Calif. Acad. Sci. 1(44):184. 1886 11885]. Lotus a (Greene) Greene, Pittonia 2:144. 1890. Anisolotus argyraeus (Greene) A. Heller, Muhlenbergia 8:47. 1912. SC argyraea (Greene) D.D. Sokoloff, Feddes Repert, 110 (1-2):94. Acmispon argyraeus (Greene) Brouillet var. multicaulis EE Brouillet, comb. nov. Basiowvw: Lotus wrightii (A. Gray) Greene var. multicaulis Ottley, Univ. Calif. Publ. Bot. 10:211, pl. 923. Hosackia wrightii A. Gray subsp. multicaulis (A. M. Ottley) Abrams, Ill. Fl. Pac. States 2:543. 1944. Lotus argyraeus (Greene) Ee SE multicaulis (Ottley) Munz, Aliso 4:93. 1958. Lotus argyraeus (Greene) Greene var. multicaulis (Ottley) Isely, Brittonia 30:466 Acmispon argyraeus (Greene) Brouillet var. notitius (Isely) Brouillet, comb. nov. Basen: Lotus argyraeus (Greene) Greene var. notitius Isely, Brittonia 30:466. 1978 Acmispon cytisoides (Benth.) Brouillet, comb. nov. Basionwm: Hosackia cytisoides Benth., Trans. Lin. Soc. London 17:366. 1837, non Lotus cytisoides L. 1753. Syrmatium cytisoides (Benth.) Greene, Bull. Calif. Acad. Sci. 2:147. 1886 Hosachia s Benth. var. rubescens Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1(2):324. 1838. Hosackia rul (Torr. & A. Gray) Anon., ., Proc. Biol. Soc. Wash. 13:203. 1900. Depranolobus rubescens Nutt. ex Torr. & A. Gray, Fl. N. ner 1(2):324. 1838. 90 Lotus ede , Pittonia 2(9):148. 18 Acmispon dendroideus (Greene) Brouillet, comb. nov. Basionym: Syrmatium dendroideum Greene, Bull. Calif. Acad. Sci. 2(6B):146. 1886. Lotus dendroideus (Greene) Greene, Pittonia 2:148. 1890. Hosackia dendroidea (Greene) Abrams, Ill. Fl. Pacific States 2:552. 1944 Acmispon dendroideus (Greene) Brouillet var. traskiae (Noddin) Brouillet, comb. nov. Basionm: Syrmatium traskiae Eastw. ex Noddin Fl. Los Ang. 201. 1917. Lotus scoparius (Nutt. ex Torr. & A. Gray) Ottley var. traskiae (Eastw. ex Noddin) Ottley, Univ. Calif. Publ. Bot. 10(3):229. 1923. Lotus scoparius (Nutt. ex Torr. & A. Gray) Ottlev subsp. traskiae (Noddin) PH. Raven, Aliso 5:326. 1963. Lotus dendroideus (Greene) Greene var. traskiae (Noddin) MM Brittonia 30:467. 1978. inan um dendroideus (Greene) Brouillet var. veatchii (Greene) Brouillet, comb. nov. Basonvm: Hosackia veatchil Bull. Calif. Acad. Sci. 1:83. 1885. Syrmatium veatchii ipis Greene, Bull. Calif. Acad. Sci. 2:148. 1886. Lotus veatchii aep Greene, Pittonia 2:148. SE Lotus ne (Nutt. ex Torr. & A. Gray) Ottley var. veatchii (Greene) Ottley, Univ. Calif. Publ. Bot. 10(3):228. 1923. Lot ) Greene var. veatchii (Greene) Isely, Brittonia 30:467. 19 Syrmatium patens Greene, Bull. Calif. Acad. Sci. 2:147. 1886. Acmispon distichum (Greene) Brouillet, comb. nov. Basionym: Hosackia disticha Greene, Bull. Calif. Acad. Sci. 1(4A):186. 1885. Syrmatium distichum (Greene) Greene, Bull. Calif. Acad. Sci. 2(6B):148. 1886. Lotus distichus (Greene) Greene, Pittonia 2(9):149. 1890. Acmispon flexuosus (Greene) Brouillet, comb. nov. Basionym: Hosackia flexuosa Greene, Bull. Calif. Acad. Sci. 1:82. 1885, non Lotus flexuosus Desr. 1792. Ottleya flexuosa (Greene) D.D. Sokoloff, Feddes Repert. 110(1-2):94. 1999 Lotus cedrosensis G , Pittonia 2(9):144. 1890. iid ie oss (Vogel) Brouillet, comb. nov. Basownt: Syrmatium glabrum Vogel, Linnaea 10:591. 1835-1836. Lotus glaber Vogel) Greene, Pittonia 2:148. 1890, nom. illeg., non Lotus glaber Mill. 1768. Hosackia glabra (Vogel) Torr., U. S. Expl. Exped., uid 2:274. 1874. Hosackia . Nutt. ex Torr. & A. Gray subsp. glabra. Anisolotus glaber (Vogel) M. Armstr. & Thornber, Field Book W. Wild Fl. 244. 1 Hosackia scoparia Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1:325. 1838. Lotus scoparius (Nutt. ex Torr. & A. Gray) Ottley, Univ. Calif. Publ. Bot. 10 (3:229, pl. 77 1923. Hosachia scoparia Nutt. ex Torr. & A. Gray var. diffusa A. Gray, Proc. Acad. Nat. Sci. Philadelphia 1863: 346. 1864. Hosackia glabra (Vogel) Torr. var. diffusus A Gray, Proc. Acad. Nat. Sci. Philadelphia 1863: 346. 1864. Lotus diffusus (A. Gray) A. Heller, Cat. N. Amer. Pl 7.1900 390 ¡ m noun glabrus (Vogel) Brouillet var. brevialatus (Ottley) Brouillet, comb. nov. Bastonyn: Lotus scoparius (Nutt. rr. & A. Gray) Ottley var. dcm Ottley, Univ. Calif. Publ. Bot. 10(3):227, pl. 77, fig. 10-16. 1923. Hosackia glabra (Vogel) Torr. BS brevialata (A. M. Ottley) Abrams, Ill. Fl. Pacific States 2:552. 1944. Lotus scoparius (Nutt. ex Torr. & A. Gray) Ottley subsp. brevialatus (Ottley) Ee E S. Calif. 450. 1974. A cmi lifl (Benth.) Brouillet, comb. nov. B Hosackia grandiflora Benth., Trans. Linn. Soc. London 11:566. 1837. p us grange Dean Greene, Pittonia 2:145. Anisol lifl (Benth.) A. Heller, Mulhenbergia 8(1-2):20. 1912 D. Sokoloff, Feddes Repert.110(1-2): 94. 1999. Hosackia ochroleuca Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1(2):323. 1838. Hosachia grana on Stee var. anthylloides A. Gray, Proc. Acad. Nat. Sci. Philadelphia 1863: Millsp., Publ. Field Columbian Mus., Bot. Ser. 5:150. 1923. Hosackia occulta Greene, Bull. Calif. Acad. Sci. 2:394. 1897. 350. 1864. Hosackia anthylloides (A. Gray) Lotus M LU Greene, Pittonia 2:144. 1890. ee nre Mn aon o Bot. o e 25:228. 1898. , Pittonia 2:145. 1890. A (Greene) Abrams, Ill. Fl. Pacific States 2:542. 1944. Lotus confinis Greene, wn 1(12):258. 1893. 8. 1912. Hosackia leucophaea Lotus grandiflorus (Benth.) Greene var. mutabilis Ottley, Univ. Calif. Publ. Bot. 10(3):208, pl. 68, fig. 2-6, pl. 69, fig. 1. 1923, Acmispon grandiflorus (Benth.) Brouillet var. macranthus (Greene) Brouillet, comb. nov. Basionym: Lotus grandiflorus (Benth.) MM Var. Ne Geen SE GEN 30:407. Sci. 1:81. 1885. I 1978. Hosackia da Cai Greene, Bull. Calif. Acad. 8(4):48. 1912. A. Heller, Mulhenbergia Acmispon greenei (Wooton & Standl.) Brouillet, comb. nov. Basin: Anisolotus greenei Wooton & Standl., Contr. U.S. Natl. Herb. 1(44):185. 1886 [1885]. Hosackia greenei (Wooton & Standl.) Wiggins, Contr. Dudley Herb. 4(2):19. 1950. Ottleya greenei (Wooton & Standl.) D.D. Sokoloff, Feddes Repert. 110 (1-2):94. 1999. Hosackia mollis Greene, Bull. Calif. Acad. Sci. 1:185. 1885, nom. illeg., non Nutt. 1838. Anisolotus mollis (Greene) A. Heller, Muhlen- bergia 8:60. 1912. Lotus mollis (Greene) Greene, Pittonia 2:143. 1890, non Balf. f. 1882. Lotus greenei Ottley, J. Wash. Acad. Sci. 29(11):483. 1939, nom ?Lotus neomexicanus E , Pittonia 2(9):143. 1890. Hosackia neomexicana nid us Bot. Ru ee 25:229. 1898. Anisolotus ) A Heller, M 1.1 O EVEN 1019 A , Contr. U.S. Natl. Herb. 16(4): 135. 1913. Acmispon haydonii (Orcutt) Brouillet, comb. nov. Basowvw: Hosackia haydonii W. Amer. Sci. 6:63. 1889. Lotus haydonii (Orcutt) Greene, Pittonia 2(9):149. 1890. Syrmatium haydonii (Orcutt) Brand, Bot. Jahrb. Syst. 25:228. 1898. Syrmatium haydonii (Orcutt) A. Heller, Muhlenbergia 9(5):67. 1913. Acmispon a (Greene) Brouillet, comb. nov. Basionym: Hosackia heermannii Durand & Hilg., Pl. Heermann. 39. 1854. Syrmatium heermannii (Durand & Hilg.) Greene, Bull. Calif. Acad. Sci.2(6B):148. 1886. Lotus heermannii (Durand & Hilg.) Greene, Pittonia 2(9):150. 1890. Lotus eriophorus Greene var. heermannii (Durand & Hilg.) Ottley, Univ. Calif. Publ. Bot. 10(3):232-234. 1923 Hosackia decumbens Benth. var. glabriuscula Hook. & Arn., Bot. Beechey Voy. 137. 1832. Hosackia t tosa Hool uscula (Hook. & Arn.) Abrams, 111. Fl. Pacific States 2:546. 1944 & Am subsp glabri- Acmispon heermannii (Greene) Brouillet var. orbicularis (A. Gray) Brouillet, GE nov. Bason: Hosackia heermannii Durand & Hilg. var. orbicularis A. Gray, Proc. Acad. Nat. Sci. Philadelphia 15:348. nii (Durand & Hilg.) Greene var. orbicularis (A. Gray) Isely, Brittonia 30:467. 1 Lotus icm Greene, n iei 1(10): Pak Bd M" REIS uae & 2. Greene var. eriophorus (Greene) Ottley, Brittonia Hosackia tomentosa Hook. a A. Bot. desen n 137. 1832. 2 osadía tomentosa m & Arn. subsp. tomentosa. Lotus tomentosus (Hook. & Arn.) Greene, Pittonia 2:150. 1890, nom. illeg., non Desr. 1792. Syrmatium tomentosum (Hook. & Arn.) Vogel, Linnaea 10:59]. 1835-36. Drepanolobus lanatus Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1(2):324. 1838. Acmispon intricatus (Eastw.) Brouillet, comb. nov. Basiowvw: Lotus intricatus Eastw., Leafl. W. Bot. 3(7):159. 1942. Ottleya intricata (Eastw.) D.D. Sokoloff, Feddes Repert. 110(1-2):95. 1999. Acmi d Hosacki 391 Brouillet New nam H Benth., Trans. Linn. Soc. London 17:366. 1837. Syr- Acmispon junceus (Benth.) Brouillet, comb. n matium junceum (Benth.) Greene, Bull. Calif. Acad. Sci. 2(6B):147. 1886. Dons junceus (Benth.) Greene, Pittonia 2(9):148. 1890. AA PUDO en ittonia Acmispon junceus (Benth.) Brouillet var. biolettii (G ) Brouillet, comb. nov. B Lot 2(114):222. 1892. H , West Coast Botany 34 1808 S ti ttii (G ) A. Heller, Malileribérgia 9(5):67. 1913. Lotus j (Benth.) (cene var. biolettii (G ) Ottley, Univ. Calif. Publ. Bot. 10): 231-232. 1923. To eng 1 H rn 1 H “11 Acmispon maritimus var. brevivexillus Aur Broidit oe nov. B g Ottley, Univ. Calif. Publ. Bot. 10(3):217, t. 73. 1923. Lot | illus (Ottley) Munz, FI. S. Calif. 449. 1974. Lotus humilis Greene, Pittonia 2(9):140. 1890. Hosackia humilis (G ) Abrams, Ill. Fl. Pacific St 2:545. 1944. aces! sean: mearnsii (Britt. ) e erter nov. arid es mearnsii Britton, Trans. New York Acad. Sci. 8(3—4):65. , Pittonia 2(9):144. 1890. Otel ii (Bri 3D D ne Feddes E Repert. 110 Se 1999. pd Acmispon mearnsii (Britt.) Brouillet var. equisolensis (J.L. Anderson) Brouillet, comb. nov. Basionym: Lotus mearnsii (Britt.) Greene var. equisolensis J.L. Anderson, Madroño 43:261—262. 1996. Acmispon micranthus (Nutt. ex Torr. & A. Gray) Brouillet, comb. nov. Basin: Hosackia micrantha Nutt. ex Torr. € A. Gray, Fl. N. Amer. 1(2):324. 1838, non Lotus micranthus Benth. 1837. Syrmatium micranthum (Nutt.) Greene, Bull. Calif. Acad. Sci. 2(6B):147. 1886. Lotus hamatus G , Pittonia 2(9):150. 1890. nom. nov., non L. micranthus Benth. 1836. Acmispon nevadensis (S. Watson) Brouillet, comb. nov. Basionym: Hosackia decumbens Benth. var. nevadensis S. Watson, Bot. California 1:138. 1876. Lotus nevadensis (S. Watson) Greene, Pittonia 2(9):149. 1890. Lotus douglasii Greene var. nevadensis (S. Watson) Ottley, Univ. Calif. Publ. Bot. 10(3):234-236. 1923. Syrmatium nevadense (S. Watson) Greene, Bull. Calif. Acad. Sci. 2(6B):148. 1886. Hosackia nevadensis (S. Watson) Parish, Pl. World 20(7):220. 1917. Hosachia decumbens Benth., Trans. Linn. Soc. London 17:366. 1837, non Lotus decumbens Poir. 1814. Syrmatium decumbens (Benth.) Greene, Bull. Calif. Acad. Sci. 2(6B):148. 1886. Anisolotus decumbens (Benth.) Thornberg, Field Book West. Wild Fl. 244. 1915. Lotus douglasii Greene, Pittonia 2(9):149. 1890, nom. nov., non L. decumbens Poir. 1814. Lotus nevadensis (S.S. Watson) Greene var. Mio Ge Ottley, Brittonia 5:81. 1944. Univ. Calif. Publ. Bot. 10(3):235-236. 1923. Lotus nevadensis (S. Watson) Greene var. congestus (Ottley) Ottley, Brittonia 5: 81. 194 4. Lotus leonis Eastw., Leaflets W Bot. 2(1):7-8. 1937. Acmispon nevadensis (S. Watson) Brouillet var. davidsonii (Greene) Brouillet, comb. nov. Bastonym: Lotus davidsonii Greene, Erythea 1(10):207. 1893 nom. nov. [based on L. sulphureus Greene]. Lotus sulphureus Greene, Pittonia 2:293. 1892, nom. illeg., non Boissier 1843. Syrmatium sulphureum Brand, Bot. Jahrb. Syst. 25:231. 1898, nom. superfl. & illegit. for Lotus davidsonii Greene 1893. Hosackia sulphurea (Greene) Abrams, Ill. Fl. Pacific States 2:547. 1944. Syrmatium davidsonii (Greene) A. Heller, Muhlenbergia 9(5):67. 1913. Lotus argophyllus (A. Gray) Greene var. davidsonii (Greene) Jepson, Fl. Calif. 2(3):326. 1936. Lotus nevadensis var. davidsonii (Greene) Isely, Brittonia 30:467—468. 1978. Acmispon niveus (S. Watson) Brouillet, comb. nov. Basiown: Hosackia nivea S. Watson, Proc. Amer. Acad. Arts Sci. 22(2):470. 887. Ottleva nivea (S. Watson) D.D. Sokoloff, Feddes Repert. 110(1—2):95. 1999 Hosackia bryantii Brandegee, Proc. Calif. Acad. Sci., Ser. 2, 2:144. 1889. Lotus bryantii (Brandegee) Ottley, Brittonia 5:98. 1942. Acmispon nudatus (Watson) o comb. nov. B ] Pittonia 1:173. 1888. Lotus nudatus (Greene) Greene, Pittonia 2(9):148.1890. Hosacki (G T MM MN Contr. U.S. Natl. Herb. 1(1):14. 1890. Acmispon oroboides (Kunth) Brouillet, comb. nov. Basiony: lephrosia oroboides Kunth in A. von Humbolt et al., Nov. Gen Sp. 6(fol.): 362. 6(qto.): 462, pl. 578, 579. 1824. Lotus oroboides (Kunth) Ottley in Kearney & Peebles J. Washington Acad. Sci. 29(11):483. 1939. Cracca oroboides (Kunth) Kuntze, Revis. Gen. Pl. 1:175. 1891. Ottleya oroboides (Kunth in A. von Humboldt et al.) D.D. Sokoloff, Feddes Repert. 110(1-2):95. 1999. Hosackia angustifolia G. Don, Gen. Syst. 2:200. 1832. Hosackia puberula Benth., Pl. Hartw. 305. 1849. Anisolotus puberulus (Benth.) Wooton & Standley, Contr. US Nat. Herb.16(4):135. 1913 Lotus puberulus (Benth.) Greene, Pittonia 2:142. 1890. Hosackia no SE Proc. E wa 2:144. 1889. Lotus geg (Kunth) var. plebeius (Brandegee) Ottley, Brittonia 5:107. 1944 g 1 Basin Naturalist 46:2. 392 Hosackia rigida Benth. var. nummularia M.E. Jones, Proc. Calif. e Sci. 2:633. 1895. Anisolotus nummularius (M.E. Jones) Wooton & Standl., Contr. U.S. Natl. Herb. 16:135 M.E. Jones) Tidestrom, Contr. U.S. Natl. Herb.25:303. 1925. Lotus KMS Dayton, Proc. Biol. Soc. Washington 40:119. 1927. M GE (Kunth in A. von Humboldt et al.) var. nummularius (M.E. Jones) Isely, Brittonia 30:468. 1978. Lotus ramulosus M.E. Jones, Contr. W. Bot. 15:138. 1929. Lotus oroboides (Kunth) var. ramulosus (M.E. Jones) Ottley, Brittonia 5:107. Lotus em (Kunth) Ottley ex Kearny & Peebles var. nanus (A. Gray) Isely, Brittonia 30:468. 1978. a prostratus (Nutt. ex Torr. & A. Gray) Brouillet, comb. nov. Basionwm: Hosackia prostrata Nutt. ex Torr. & Fl. N. Amer. 1(2):325. 1838., non Lotus prostratus L. 1758. Syrmatium prostratum (Nutt. ex Torr. & A. Gray) Greene, Bull. Calif er Sci. 2:147. 1886 Lotus nuttalianus Greene, Pittonia 2:150. 1890. Acmispon procumbens (Greene) Brouillet, comb. nov. Basiowvw: Hosackia procumbens Greene, Bull. Calif. Acad. 1(3):82. 1885. Lotus procumbens (Greene) Greene, Pittonia 2(9):149. 1890. Syrmatium procumbens (Greene) Greene, Bull. Calif. Acad. Sci. 2(6B):148. 1886. edad sericea Benth., Trans. Linn. Soc. Contes 17:367. M non Lotus sericeus Moench 1802, non DC. 1813, non Pursh 1814. nth.) Greene, Bull. Calif. Acad. Sci. 2(6B):147 Mod Lotus SEN phyll Brand, Bot. Ja subsp sericed. $ lus Greene, Pittonia 2(9):149. 1890, nom. nov., non L. sericeus Moench 180 Syst. 25:229. Acmispon procumbens var. jepsonii (Ottley) Brouillet, comb. nov. Basin: Lotus leucophyllus Greene var. = in Univ. Calif. bn Bot. 10(3):227. 1923. Lotus procumbens (Greene) Greene var. jepsonii (Ottley) Ottley, Brittonia 5:81. 1 subsp ien«snnii (Ottley) Abrams, Hi Fl Pacific States 2:552 194 + L H ida Benth., PL Hartw. 305. 1849. Lotus rigidus (Benth.) Acmispon rigidus (Benth.) Brouillet, comb. nov. Greene, Pittonia 2(9):142. 1890. Anisolotus rigidus (Benth.) Rydb., Bull. E Bot. Club 33:144. 1906. A. rigidus (Benth. ) A Heller, Muhlenbergia 8:20. 1912. Otileya rigida (Benth.) D.D. Sokoloff, Feddes Repert. 110(1—2):95. 1999. Hosackia puberula A. Gray, Smithsonian Contr. o Se 42. nid nom. imi non esr 1849. Lotus argensis Coville, Contr. U.S. Natl. Herb. 4:83. 1 g ller, Muhlenbergia 9(5):67. 1913. — strigosus oh ex o o A. Gray) Brouillet, comb. nov. Baan: Hosackia strigosa Nutt. ex Torr. & A Fl. N. Amer. 1:326. 1 g (Nutt. ex Torr. & A. Gray) Greene. Lotus strigosus (Nutt. ex Torr. & A. Gray) Greene Var. E Anisolotus strigosus Mr ex Torr. & A. Gray) A. Heller, Muhlenbergia 3(6): 101. 1907. Ottleya strigosa (Nutt. ex Torr. & A. Gray) D.D. Sokoloff, Feddes Repert. 110(1—2):95. 1999 Lotus hirtellus Greene, Pittonia 2:142. 1890. Hosackia hirtella (Greene) Brand, Bot. Jahrb. Syst. 25:228. 1890. Lotus strigosus var. hirtel- lus (Greene) Ottley. Hosackia strigosa var. hirtella (Greene) H.M. Hall, Uiniv. Calif. Publ. Bot. 4(12):199. 1912. Anisolotus hirtellus (Greene) A. Heller, Muhlenbergia 8:60. 1912. I (Nutt. ex Torr. & A. Gray) Greene var. hirtellus (Greene) Ottley, Unix. Calif. Publ. Bot. 10(3):214, t. 71. 1923. Hosackia maritima Torr. in WH. Emory, Rep. U.S. Mex. Bound. 2(1):55. Hone Hosackia nudiflora Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1:326. 1838. 1 1890. Lotus strigosus (Nutt. ex Torr. & A. Gray) Greene var. nudiflorus (Nutt. ex Torr. & A. Gray) Jepson, Fl. W Calif. 302. 1901. Anisolotus nudiflorus (Nutt. ex Torr. & A. Gray) A. Heller, Muhlenbergia 7:139. 1911 (1912). Hosachia rubella Nutt. ex Torr. & A. Gray, Fl. N. Amer. 1:326. 1838. Lotus rubellus (Nutt. ex Torr. & A. Gray) Greene, Pittonia 2:141. 1890. Anisolotus rubellus (Nutt. ex Torr. & A. Gray) A. Heller, Muhlenbergia 7:139. 1911 (19 _ Pittonia 2:140. 1890. Hosackia tomentella (Greene) Brand, Bot. Jahrb. So 25:231. 1898. Hosackia tomentella 944. (Nutt. ex Torr. & A. Gray) Greene, Pittonia 2:141. Lotus tomentellus G + ne) Abrams, Ill El] Pamfia Crar Acmispon utahensis (Ottley) Brouillet, comb. nov. Bason: Lotus utahensis Ottley, Brittonia 5:108-109. 1944. Ottleya utahensis (Ottley) D.D. Sokoloff, Feddes Repert.110(1—2):96. 1999. Acmispon wrightii (A. Gray) Brouillet, comb. nov. Bastonym: Hosackia wrightii A. Gray, Smithsonian Contr. Knowl. 5(6):304. 1853. Lotus wrightii (A. Gray) Greene, Pittonia 2(9):143. 1890. Anisolotus wrightii (A. Gray) Rydb., Bull. Torrey Bot. Club 33:144. 1906. Ottleya wrightii (A. Gray) D.D. Sokoloff, Feddes Repert. 110(1—2):96. 1999 Brouillet, N à in Acmis| IH ki 393 LIST OF ACCEPTED NAMES IN HOSACKIA FOR NORTH AMERICA NORTH OF MEXICO Hosackia alamosana Rose, Contr. U.S. Natl. Herb. 1(4):96. 1891. Hosackia crassifolia Benth., Trans. Linnean Soc. London 17:365. 1837. Hosackia crassifolia Benth. var. crassifolia Hosackia gracilis Benth., Trans. Linnean Soc. London 17:365. 1837. Hosackia incana Torr., Pacific Railr. Rep. 4:79, pl. 4. 1857 Hosackia oblongifolia Benth., Pl. Hartw. 305. 1849. Hosackia oblongifolia Benth. var. oblongifolia Hosackia pinnata (Hook.) Abrams, Ill. Fl. Pac. States 2:541. 1944. Hosackia rosea Eastw., Proc. Calif. Acad. Ser. 2, 6:424, pl. 55. 1896. Hosackia stipularis Benth., Trans. Linnean Soc. London 17:365. 1837. Hosackia stipularis Benth. var. stipularis Hosackia yollaboliensis (Munz) D.D. Sokoloff, Kew Bull. 55:1009. 2000. LIST OF ACCEPTED NAMES IN ACMISPON FOR NORTH AMERICA NORTH OF MEXICO Acmispon americanus (Nutt.) Rydberg, Ann. Bot. Fennici 37:129. 2000. Acmispon americanus (Nutt.) Rydb. var. americanus Acmispon brachycarpus (Benth.) D.D. Sokoloff, Ann. Bot. Fennici 37:130. 2000. Acmispon denticulatus (Drew) D.D. Sokoloff, Ann. Bot. Fennici 37:130. 2000. Acmispon maritimus (Nutt.) D.D. Sokoloff, Ann. Bot. Fennici 37:129. 2000. Acmispon parviflorus (Benth.) D.D. Sokoloff, Ann. Bot. Fennici 37:129. 2000. Acmispon rubriflorus (Sharsm.) D.D. Sokoloff, Ann. Bot. Fennici 37:130. 2000. Acmispon wrangelianus (Fish. & C.A. Mey.) D.D. Sokoloff, Ann. Bot. Fennici 37:129. 2000. ACKNOWLEDGMENTS The author thanks Kanchi Gandhi, Harvard University Herbaria, R. Kiger, Hunt Institute for Botanical Documentation, and James Zarucchi, Missouri Botanical Garden, for their help with nomenclatural and bibliographic matters, as well as Martin Wojciechowski, Arizona State University, for his comments on the manuscript. REFERENCES BUM Gë and d iua o Mood SEU ane DOR GEHEN relationships of Lotea and Coronilleae lear ri s. Amer. J Bot. 87:1871- 1881. ALLAN, GJ, E.A. READER im Weg ane D.D. Soxotorr. 2003. Molecular Pu HE e of tribe Loteae (Le- phy. In: B.B. Kli | A. Bruneau, eds. Advances in legume systematics, part 10, Higher Level mena Royal Botanic eugene Kew. Pp. 371-393. ARAMBARRI, A.M, 2000. A cladistic analysis of the New World species of Lotus L. (Fabaceae, Loteae). Cladistics 16:283-297. ARAMBARRI, A.M., S.A, STENGLEIN, M.N. Colares, and M.C. Novoa. 2005. Taxonomy of the New World species of Lotus (Leguminosae: Loteae). Australian J. Bot. 53:797-812. BARNEBY, R.C. 1989. Intermountain flora, vol. 3(B): Fabales. New York Botanical Garden, Bronx, NY. CULLEN, E.O. 1959. Studies in the genus Lotus (Leguminosae) |. Limits and subdivisions of the genus. Canad. J. Bot. 37:157-165. Greene, E.L. 1890, Enumeration of the North American Loti. Pittonia 2:133-150. IseLy, D. 1981. Leguminosae of the United States. Ill. Subfamily Papilionoideae: Tribes Sophoreae, Podalyrieae, Loteae. Mem. N.Y. Bot. Gard. 25(3). 394 | ti tanical Texas 2( Isety, D. 1993. Lotus. In: J.C. Hickman, ed, The Jepson manual: higher plants of California. University of California Press, Berkeley. Pp. 616-622. Omey, AM. 1923. A revision of Californian species of Lotus. Univ. Calif. Publ. Bot. 10:189-305. Orey, AM. 1944, The American Loti with special consideration of a proposed new section. Brittonia 5:81-1 23 SokoLorr, D.D. 1999. Ottleya, a new genus of Papilionaceae-Loteae from North America. Feddes Repert. 110:89-97. Soe opt. D.D. 2000a. New combinations in Hosackia Douglas ex Benth. (Leguminosae: Loteae). Kew Bull. 55:1009-1010. SokoLorr, D.D. 2000b. New combinations in Acmispon (Leguminosae, Lotae). Ann. Bot. Fennici 37:125-131. SoxoLorr, D.D. and J.M. Lock. 2005. Loteae. In: G. Lewis, B. Schrire, B Mackinder, and M. Lock. Legumes of the world. Royal Botanic Gardens, Kew. Pp. 455—465. STENGLEIN, S.A., A.M. ARAMBARRI, M.N. CoLares, M.C. Novoa, and C.E. Vizcaino. 2003. | eaf epidermal characteristics of Lotus: subgenus Acmispon (Fabaceae: Loteae) and a numerical taxonomic evaluation. Canad. J. Bot. 81:933-944. MICRANTHES NELSONIANA VAR. PORSILDIANA (SAXIFRAGACEAB), THE PROPER NAME AT THE VARIETAL LEVEL Luc Brouillet Herbier Marie-Victorin Institut de recherche en biologie végétale niversité de Montréal 4101 rue SEN Montréal, Quebec, CANADA H 1X 2B2 luc.brouilletgumontreal.ca ABSTRACT TT E + ei dos Mi d J i p ildi l lori . pacifica. A new combination is therefore presented a th bination M peti laris (Raf.) Bush i id 1 valid. RESUMEN Cuando se 1 las d dades, el bre M I r. porsildiana tiene prioridad sobre var. pacifica. Por ello se t bi id Además, | bi idn M petiolaris (Raf) Bush se considera valida. EU Micranthes nelsoniana Recently, Brouillet and Gornall (2007) presented new combinations in Micranthes for the North American flora. In the treatment of the genus for the Flora of North America, Elvander and Brouillet (ined.) are merg- ing two iras did taxa e Micranthes cn (D. Don) Small in had traditionally been considered distinct: let ifi I ildi Sul E ifi 1 varietal names exist for both entities in Saxifraga. When merging ther taxa at tl bspecific level, t] pacij At the varietal ievel, however, porsildiana has priority over pacifica ( y below). Dicam the varietal level is being used for the Flora of North America EECH M publishing new names, however, Brouillet and Gornall (loc. cit.) made a combination for var. pacifica but not for var. porsildiana. A new combination in Micranthes for the latter is therefore required and is presented here. Since Gornall and Ohba (ined.) had prepared combinations for all varieties in Micranthes nelsoniana, I am attributing the new combination to these authors. Micranthes d (D e See var. ra (Calder & Savile) Gornall & H. Ohba, comb. nov. Basic subsp. porsildiana Calder & Savile, Canad. J. Bot. 38:429. 1960. iride EN var. ei vd ile) B Boi n, Naturaliste Canad. 93:646. 1966. Saxifi 1 D. Don subsp Not. Bot. 126: 494. 1973. Saxifraga nelsoniana var. porsildiana (Calder & Savile) H. Ohba, J. jap: Bot. 68: 336. 1993. Micranthes nelsoniana (D. Don) Small var. pacifica (Hultén) Gornall & H. Mie Je Bot. Res. Inst. Texas 1: ud i M scien MM L. subsp. pacifica Hultén, Acta Univ. Lund. ns 41:928. 1945. Saxif D. Don subsp. paci nelsoniana var. pacifica (Hultén) H. Ohba, J. Jap. Bot. 68:336. 1993. ` Micranthes petiolaris Brouillet and Gornall (2007) made a new combination for Micranthes petiolaris. The combination by Bush (1928), however, is currently considered valid and has priority. Therefore, Micranthes petiolaris (Raf.) Brouillet & Gornall must be considered an isonym. Micrantl iolaris (Raf.) Bush, Amer. Midl. Naturalist 11:225. 1928. Bastonym: Hexaphoma petiolaris Raf., Fl. Tellur. 2:67. 1837. Saxi fraga iaa Britton ex Small & Vail, Mem. Torrey Bot. Club 4:118. 1894. Micranthes petiolaris (Raf.) Brouillet & Gornall, J. Bot. Res. Inst. Texas 1:1021. 2007 J. Bot. Res. Inst, Texas 2(1): 395 — 396. 2008 ACKNOWLEDGMENTS The author thanks John Strother, Herbarium, University of California-Berkeley, Kanchi Gandhi, Harvard University Herbaria, and James Zarucchi, Missouri Botanical Garden, for their help with nomenclatural matters, and Richard Gornall for having made his work available. James Zarucchi and Elizabeth Fortson Wells kindly reviewed the paper. REFERENCES BrouiLLET, L. and R. Gornal. 2007. New combinations in Micranthes (a segregate of Saxifraga, Saxifragaceae) in North America. J. Bot. Res. Inst. Texas 1:1019-1022. Bush, B.F 1928. Some species of Saxifraga. Amer. Midl. Naturalist 11:213-235. NOMENCLATURAL CHANGES IN NEMACLADUS (CAMPANULACEAE) Nancy R. Morin! Biology Department Northern Arizona University Flagstaff Arizona 8601 1, U.S.A. ABSTRACT Based ical and DNA sequence evidence, Parishella calif | f | to Nemacladus, and Nemacladus glandulif var. orientalis, Nemaclddis glandulferus v var. basins and Newiarladus rubescens var. tenuis are recognized as species. Two new varieties and one new s] tenuis var. aliformis, Nemacladus secundiflorus var. robbinsii, and Nemacladus calcaratus. RESUMEN n pa 1 L1. : TYXNTA D - AH LE H E AT nrientatic LA, E h A EN nd EE glanduliferus var. NEE y Nenaad rubescens var. t y una nueva especie: Nemacladus tenuis var. aliformis, Nemacladus secundiflorus var. robbisi. y Nemacladus calcaratus. Nemacladus is an isolated genus in Campanulaceae p a B H.G. Gustafsson, Lammers 2007a, 2007b, or Nemacladaceae Nuttall, Takhtajan 1997, which include ladus McVaugh and Parishella A. Gray) basal to the Campanuloideae and Lobelioideae (Cosner et al. 1994) or sister to a clade comprising Cyphiaceae, Cyphocarpaceae, and Lobeliaceae (Gustafsson & Bremer 1995). Distributed in southwestern U.S. and northern Mexico, extending south to Sonora and Baja California and north into southeastern Oregon and western Wyoming, it is most diverse in California. Filaments connate for part of their length distinguish it from Campanuloideae, free anthers distinguish it from Lobelioideae, and absence of a fluid filled cavity in the stigma separates it from Cyphioideae. A detailed study of Nemacladus has been in progress since the author prepared a treatment of it for the Jepson Manual (Morin & Milburn 1993). Extensive field work, study of herbarium specimens, and, in collaboration with TJ. Ayers at Northern Arizona University, analysis of DNA and study of floral and seed morphology using SEM, have been done for a monograph that is in preparation. In order to reflect the results of this work in the revision of the Jepson Manualin preparation, it is necessary to make some nomenclatural changes in advance of the full monograph. The monospecific Parishella A. Gray, found in the Mojave Desert and disjunct in mountains in Santa Barbara County, California, has long been thought to be the genus most closely related to Nemacladus and is here transferred into Nemacladus. In ITS (Haberle 1998), and atpB (Ayers in prep.) analysis Parishella californica is nested within Nemacladus, with closest alliance to N. rigidus Curran. Ás in other Nemacladus, large, clear cells are attached to the filaments; the non-resupinate, nearly actinomorphic flowers with white, cup-shaped corolla are similar to those of N. pinnatifidus Greene. Nemacladus californicus (A. Gray) Morin, comb. no ishella californicus A. Gray, Bot. Gaz. 7:94. 1882. Tyre: CALIFORNIA. SAN BERNARDINO Co.: Rabbit Springs, Mohave Desert, d 1328, May 1882 (HOLOTYPE: GH! isotypes: MOI, NY!, UCD. McVaugh (1939) recognized three varieties in N. glanduliferus based on their habit and the aspect of the pedicels, considering the flowers to be similar in all three. The flowers differ significantly, however, and these taxa are here recognized as species. Nemacladus glanduliferus and N. orientalis are sister taxa in the atpB and ITS analysis (Ayers, in prep.). Nemacladus glanduliferus Jepson has nearly OS pps E flowers that are mostly resu- pinate, petals white, all similar in size and shape, connate at | g a cuplike tube, free lobes reflexed; ‘Contact address, Nancy R. Morin, RO, Box 716 Point Arena, California 95468, U.S.A. J. Bot, Res. Inst. Texas 2(1): 397 — 400. 2008 I lafal Dos H In LI PERA £Tauee 311) 398 filament tube and style erect, arching over flower; capsule hemispheric with base and apex rounded, 2-2.3 mm diameter. It occurs in desert areas of California, Arizona, and northern Baja California and Sonora, Mexico. Nemacladus Heu di sad Ta outward-facing flowers that are not resupinate, petals white with markings, free nearly to base, lower two petals longer and narrower than upper three petals; filament tube and style deflexed between lower two petals; capsules hemispheric with base rounded and apex acute, 1.5-2 mm diameter. It is widely distributed in desert areas of California, Nevada, Arizona, Utah, north to Wyoming and south to Baja California and Sonora, Mexico. Nemacladus australis has highly zygomorphic, outward-facing flowers that are not resupinate, petals white with maroon markings, free nearly to base, lower two petals longer and narrower than upper three petals, filament tube and style deflexed between lower two petals; capsules spherical, 3.5-4 mm diameter. It is known from Rosario, Bajia de los Angelos, Santa Catarina, El Terminal, and Sierra Calvario, in Baja California. Nemacladus orientalis (McVaugh) Morin, comb. et stat. nov. Nemacladus glanduliferus var. orientalis McVaugh, Amer. Midl. Naturalist 22: 540. 1939. Tre: NEVADA. Cark Co.: shore of Lake Mead, near Boulder Dam, 2 May 1938, Percy Train 1566 (HOLOTYPE: NA-no. 52430; IsoryPE: UNIV). Nemacladus australis (Munz) Morin, comb. et stat. nov. Nemacladus rigidus var. australis Munz, Amer. J. Bot. 11:242, tab. 10. 192. Tyre: BAJA CALIFORNIA. Rosario, 1 May 1886, C.R. Orcutt 1348 (novorvee: GHI; rsorvees: MOI, UC!). Nemacladus glanduliferus var. australis (Munz) McVaugh, Amer. Midl. Naturalist 22:540. 1939 N ladus rul Greene is one of the most striki ies in the genus, with highly zygomorphic flow- ers -— conspicuous markings on the petals. McVaugh. game var. tenuis based on shape, size, and margins of the basal leaves, size of anthers, and nature of the clear cells attached to the filaments. Except for these characters, he believed the flowers were basically the same as typical rubescens. Field studies show that the flowers of tenuis are very different from those of rubescens and in ways that almost certainly Pee gene transfer between the two, and that tl re two different floral mophologies within tenuis. N tenuis is SES Geess asa el a two varieties. th have silvery grey s or reddish st with a silvery sheen, and yellowish green leaves. Nemacladus rubescens has oblanceolate to elliptic, mostly obtuse leaves with entire margins; flowers face outward, are highly zygomorphic, not resupinate, petals divided nearly to base, yellowish with maroon markings, elliptic to lanceolate and straight; filament a and style iain between two lower petals. Nemacladus tenuis has leaves oblanceolate, deeply pinnatifid; flov or outward, are resupinate, petals yellowish with maroon and yellow markings or white with pinkish or yellowish markings, lower three petals connate at base or divided nearly to base, together bowl-shaped; filament tube and style erect and arching over the flower. pata tenuis (McVaugh) Morin, uu et stat. nov. ? | tenuis McVaugh, Amer. Midl. Naturalist 939. Tire: CALIFORNIA. R Mt., Colorado Desert, 17 Apr 1905, H.M. Hall 5819 (Hororve: US 6149291; isotypes: ARIZ!, DS!, GH!, MOI, NY!, OSU!, P!, POMI, UC). Nemacladus tenuis var. tenuis Sepals 0.7-1 mm long, linear; corolla divided 1/2 length, lobes nearly alike; all lobes white with deep pink or yellowish tips, deltate, 0.9-2 mm long; filaments 1-2 mm long, arched, anthers 0.4—0.5 mm long. Mojave and Sonoran Deserts. Nemacladus tenuis var. aliformis Morin, var. nov. Tee: CALIFORNIA. San BERNARDINO Co.: 1 mi E of junction with Hwy. 127 on Old Spanish Gentry Road, 26 Apr 2003, Morin 589b (oo org: ASC). AN T T UNE ERE Y E ; ] te deltatus 2 li t , angustatus, arcuatus , filis longius, L as] f 1 et antherae 05-061 mm Geste differt. Morin, Nomendatural changes in Nemacladus 399 Sepals 172.5 mm long, narrowly deltate; corolla divided nearly to base, 2 upper lobes maroon or brownish, linear, arched, 0.9-2.5 x 0.3-0.5 mm long, 3 lower lobes white with yellow and brownish marks, 0.9-2.2 mm long, ciliate; filaments 2-3.5 mm long, anthers 0.5-0.6 mm long. Mohave Desert. Etymology.—The epithet “aliformis” (from ala, wing, and -formis, with the form of) refers to the winged appearance of the flowers, which resemble small flying insects. Robbins (1958) helped clarify the circumscription of Nemacladus species that have resupinate flowers with corollas that are mostly white or white with a lilac or pink tinge. One of these is Nemacladus secundiflo- rus G.T. Robbins, known from the South Coast Ranges and Greenhorn Mountains in California. He noted an unusual population (Raven 9138 and Bacigalupi 5649) from San Benito County but concluded it was not different enough to warrant recognition. Study of fresh Ra ana additional populations indicates that the flowers of these plants consistently differ from typical i in being much smaller and in having a narrower corolla tube. Nemacladus secundiflorus var. secundiflorus has flowers with a white corolla, sometimes with pinkish veins, tube 2.5-3.5 mm long, broadly cylindrical, upper 2 lobes widely spreading, 2-2.5 mm long, glabrous to hairy, lower lobes reflexed, 2.5-3 mm long; filaments + 2-2.5 mm long, anthers 0.5-0.7 mm long, with many long, ca. 0.7 mm hairs. South Coast Ranges in Monterey and San Luis Obispo counties, and the Greenhorn Mountains in Tulare and Kern counties. Nemacladus secundiflorus var. robbinsii Morin, var. nov. Tee: CALIFORNIA. San Benito Co.: just W of the junction to Pinnacles on the road from Hollister to Bitterwater, 5 May 1956, Raven, Stebbins, et al., 9138 (HOLOTYPE: JEPS 14554! isotypes: DS!, JEPS 78490!, NY!, RSA!, SBBG!, SLO!, US A Nemacladus secundiflorus Robbins var lif! fl parvioribus , llae tubus 0.5-0.8 mm longus, lobis 0.3-0.5 mm longus, antherae 0.1 mm longus differt. Corolla white or pale lavender; tube 0.5-0.8 mm long, narrowly cylindrical, upper 2 lobes spreading, 0.3-0.5 mm long, glabrous, lower lobes reflexed, 0.3-0.5 mm long; filaments + 0.8 mm long, anthers 0.1 mm long, hairs absent or few, ca. 0.5 mm long. South Coast Ranges in San Benito, San Luis Obispo, and Ventura counties with one population in the Greenhorn Mountains in Tulare Co. Em 08y. moo i is named in honor of Guy Thomas Robbins, 1916-1960, in recognition of his ] dies of Nemacladus. He received a B.S. from the University of California, Berkeley and an M.S. at University of Colorado, Boulder, after which he was on the faculty at University of Arizona and at Oklahoma State College, ine He then entered the Ph.D program at University of California, Berkeley and in 1952 took a full-time position as a Research Botanist in the Jepson Herbarium. In addition to his work on Nemacladus, he published a revision of North American Androsace (Robbins 1944). Additional information about his life can be found in the memoirs of Lavinia Pearl Butler Robbins (1882-1985), his mother, who donated her memoirs in 1972 to the Bancroft Library at University of California, Berkeley, in his memory. Les N lad I tus Morin, sp. nov. Tee: CALIFORN Co.: Sof CI C , ca. 9.3 air mi W of Hwy 395, sandy flats bet itic boulders, near 35.83N, 118.04W 1745 m, 4 Jun 2008, Morin 641 Gown ASC!; sory: UCI). e € : 11 ] * “1 Hs r D Plants 2-4 cm tall, branched from base or 0.5-1 cm from base, axis somewhat zig-zagged, stems widely spreading at a 60—75? angle to axis, sparsely hairy; internodes 0.5—1 cm. Basal leaves green with reddish margins, narrowly lanceolate to spatulate, 2-5 x 0.2-1.5 mm, remotely toothed, densely hairy, apex acute. Bracts 1-1.2(-2) mm long, ovate-lanceolate, entire or remotely and minutely toothed, sparsely hairy, + clasping pedicel at base, abruptly arched away from pedicel 1/3 to 1/2 from base, apex minutely apiculate; pedicels straight or arched, 1-1.2 cm long, + densely hairy at Mond more sparsely hairy distally. Flowers bilaterally symmetrical, not resupinate; hypanthium tube | to broadly obconic, ca. 0.5 mm long; calyx lobes lanceolate, glabrous, lowest calyx lobe reflexed and ese. at base to spur, 1.2-1.5 x 0.5 mm, 2 flanking lobes 1.2-1.5 mm long, upper 2 lobes 0.8-1 mm long; petals radiating on adaxial (upper) side of corolla, white with longitudinal red or maroon line on proximal half, ovate, apex acute, bearing slender, 400 Journal of tl 0.4 mm, downward pointing hairs abaxially and adaxially, lowest 2 petals connate basally 1 mm forming a spur, free portion 2.5 x 1.5 mm, 2 flanking and uppermost petals narrower, ca. 2.5 x 1 mm; filaments con- nate in distal 2/3, 1.8-2 mm, bearing 0.5—1 mm long hairs at apex, 2 narrow, yellow, stipe-like appendages extending from near base of fil ts terminated by semicircular pad from which 8-12 clear, narrow, acute cells ca. 0.3 mm long radiate, anthers 0.3-0.4 mm; ovary subinferior, broadly hemispheric, base obtuse, apex nearly flat, 3 glands alternating with filaments low, oblong, densely papillate; capsule ca. 3 mm long, calyx lobes splayed outward. Nemacladus calcaratus is known only from the Chimney Creek area, Tulare County, California at the southern end of the Pacific Crest. Etymology.—Calcaratus is in reference to the nectar spur. Ertter et al. (6331) noted the red stripes on the corolla, and Boyd and Bramlet (1931) pointed out the small spur. It is the first species of Nemacladus known to have a definite nectar spur, although the corolla of Nemacladus ramosissimus Nutt. 1s gibbous. The flat ovary apex and relatively large corolla with all petals n ad: on the upper part are also distinctive. CH ex Ar TGCADWIT A Tal fe 234 3 ¡ ENE Y 111 nm] A ,at W end of ridge, elev. 6200 ft, 9 Jun 1986, Ertter, Holland, and Dains 6331 (RSA, UC); Southern Sierra Nevada, Chimney TN Cups on, north Slope Peak 7155, south Chimney Meadow, elev. 6800 ft, 9 Jun 1986, Boyd & Bramlet 1931 (CAS, RSA). ACKNOWLEDGMENTS I thank Tom Lammers and Tina Ayers for their helpful comments on the manuscript. REFERENCES Cosner, M.E. 1993. Phylogenetic relationships in the Campanulales based on rbcL sequences. Pl. Syst. Evol. 190:79-95 Gustarsson, M.H.G. and K. Bremer. 1995, Morphology and phylogenetic interrelationships of the Asteraceae, Ca- lyceraceae, Campanulaceae, Ge ane related ES Ea Amer. J. Bot. 82:250-265. HABERLE, R.C. 1998. Phylogenetic syst e North America Cyphioids (Campanu- laceae, sensu lato). M.S. Thesis, Northern SE University, Flagstaff LAMMERS, T.G. 2007a. World checklist and bibliography of Campanulaceae. Royal Botanic Gardens, Kew. Lammers, T.G. 2007b. Campanulaceae. In: K. Kubitzki ed, The families and genera of Vascular Plants, Vol.8, Asteridae, ed. J.W. Kaereit and C. Jeffrey. Springer-Verlag, Germany. Pp. 26-56. McVauaH, R. 1939. Some realignments in the genus Nemacladus. Amer. Midland Naturalist 22:521-550. Morin, N.R. and J. MuBurn. 1993. Nemacladus. In: James C. Hickman, ed, The Jepson manual: higher Plants of California. University of California Press, Berkeley. Pp. 465—468. Roseins, G.T. 1944, North American Androsace. Amer. Midl. Naturalist 32:137-163. Rosens, G.T. 1958. Notes on the genus Nemaciadus. Aliso 4:139-147 TAkHTAJAN, A. 1997. Diversity and classification of flowering plants. Columbia University Press. New York. PHYLOGENETIC ANALYSIS OF NORTH AMERICAN PLUMS (PRUNUS SECT. PRUNOCERASUS: ROSACEAE) BASED ON NUCLEAR LEAFY AND S6PDH SEQUENCES Joseph R. Rohrer,' Megan A. O'Brien, and Julie A. Anderson Eas of Biology University of Wisconsin-Eau Claire Equ e Wisconsin 54702-4004, U.S.A. ABSTRACT 1 FT TZATW g e A 1 3 : : Vas i Dee L i | f North A ica (P t Prunocerasus). I The clad lting f i id Bayes imferenas are pas resolved and EE with clades based on rolas DNA sequences Evidence Gef both es genes oa Lei of P. texana in sect. Prunocerasus. Sequere ee P. americana and P. mexicana form a monophyletic group on dnd kor form a clade sister to P. tl 2 , but most P g cla The EE e of a 250-bp insertion within LEAFY intron 2 supports a cl I P. hortulana and P. murrayana. P g is, P. gracilis, P. maritima, and phyleti the LEAFY cladogram, basal to lad d of P. aneusti ifoli a and tifolia-lil { P. rivularis en P. munsoniana Data from LEAFY and s6pdh suggest o hybrid origins for P. rivularis and P. munsoniana, with P. pao as one progenitor. RESUMEN * [oes un e LA = secuencias de DNA de dos ni nins (LEAFY y s6pdh) para investigar | ] t. Prunocerasus). 1 g jue resultan de la máxima parsi f 1 E 5 1 ^ 1 J 1 A s ds TARTA sdlestiul T lend A + I + Eu bos g ] fi la colocación de P. texana en la sect. Prunocerasus. I ias de P. americana y P. mexicana forman filéti l clad de s6pdh, pero la mayot de P i ti d i r o div ten que juntas Ge un dado Bette de P. subcordata. La posesión compartida d i ión de 250-bp en el intrón 2 P. hortulana P. murrayana. P alleghaniensis, P. gracilis, P. maritima, y P. me ae filéti l clad P. angustifolia y , I E PEN y P munsoniana. Los dos de LEAFY y s6pdh sugi igen híbrido para P. rivularis y P. munsoniana, con P. angustifolia como uno de los progenitores. INTRODUCTION Prunus is one of the larger genera of the Rosaceae with about 200 species, which are most abundant in the North Temperate zone but also range into the tropics and Southern Hemisphere (Rehder 1940; Robertson 1974). It is important commercially as the source of cherries, almonds, peaches, nectarines, apricots, and plums; numerous species are grown ornamentally for their beautiful early-spring flowers. The classifica- tion of Prunus has been debated from Linnaeus to the present day, with some botanists arguing for a broad concept of the genus that includes all of the fruits mentioned above (e.g., McVaugh 1951; Robertson 1974) and others dividing Prunus into a variety of segregate genera including Amygdalus, Armeniaca, Cerasus, Laurocerasus, Padus, and Persica (e.g., Iwatsuki et al. 2001; Flora of China Editorial Committee 2003). The most widely used classification has been that of Rehder (1940) who recognized Prunus in its broad sense with five subgenera. At the species level there has been considerable taxonomic uncertainty among the North American plums (Prunus sect. Prunocerasus) In 1892 Bailey called Prunocerasus the "hardest puzzle in American pomology,” a memorable phrase quoted by Shaw and Small (2004) more than a century later in reporting the use of multiple cpDNA sequences to bear on this enduring puzzle. Shaw and Small (2004) provided an ‘Author for correspondence; email: jrohrer@uwec.edu J. Bot. Res. Inst. Texas 2(1): 401 — 414, 2008 402 t tani titute of Texas 2( extensive review and analysis of the taxonomic history of P and they noted that the unresolved classification of North American plums is the result of inters] imilarities, intraspecific variation, and probably interspecific hybridization. The earliest studies using DNA sequences to examine relationships in Prunus focused on a limited number of cultivated species (Badenes & Parfitt 1995; Uematsu et al. 1991). In 2001 two research groups sought to clarify phylogenetic relationships among the subgenera and numerous sections of Prunus by ex- amining species on a worldwide scale (Bortiri et al. 2001; Lee & Wen 2001). Unfortunately relationships among the species of section Prunocerasus remained unresolved. Both groups used ITS sequences and found slight variation among the native plums included in their studies. Analyses by Bortiri et al. (2001) using chloroplast trnL-trnF sequences ; also dia nd slight variation among six North American species. Rohrer et al. (2004) used alysis in an attempt to find molecular markers more variable among species of Prunocerasus than ITS or trn seque However, all pairs of species in the study shared fewer than half of their alleles, indicating greater geneti g mong the North American plums than expected given their similar ITS and trn sequences. Accessions of two or three individuals of the same spe- cies often did not cluster together, suggesting that microsatellites La ad too Ms to be informative in resolving species relationships in Prunocerasus. Seeking to test the] monophyly for Prunocerasus and to resolve relationships among the species, Shaw and Small (2004, 2005) turned to seven noncoding regions in the chloroplast genome shown to be highly variable across a wide array of taxa (Shaw et al. 2005). Given their taxonomically broad sampling of species both within the section and across all of Prunus, Shaw and Small (2004, 2005) provided unequivocal support for the monophyly of sect. Prunocerasus. Within the section the analyzed sequences varied little among the species. They demonstrated strong support for P. subcordata as sister to the rest of Prunocerasus. An unexpected result was the inclusion of the hairy-fruited P. texana within Prunocerasus positioned between P. subcordata and the other species, which formed three primary clades with little resolution within each clade. Many of the species have exemplars on more than one clade, and the chloroplast haplotypes show greater correlation with geography than species limits. Bortiri et al. (2002) demonstrated that the nuclear gene s6pdh was more variable among Prunus species than either ITS or trn and showed greater promise for resolving relationships in the genus. However, only three species of Prunocerasus were included in their analysis. In addition, Oh and Potter (2003) reported sequences from the second intron of the nuclear gene LEAFY were more phylogenetically informative than either ITS or cpDNA in the rosaceous genera Mine ane Stephanandra. In this study we explore the use of these low-copy nuclear genes for resolving relati among the species of Prunocerasus. Specifically we expand the use of s6pdh to all commonly recognized nE in KR section and include seq from the second intron of LEAFY. MATERIALS AND METHODS Plant Material —We were able to obtain at least one accession from each of the 15 (or so) species of North American plums. The exact number of pes is subject to the differences in opinion concerning species circumscriptions that ani ebate. Apricot (P. armeniaca), cherry plum (P. cerasifera), and two varieties of sand cherry (P. pumila var. pumila and P. pumila var. susquehanae) were used for outgroup com- parison. In all, we analyzed 37 accessions of North American plums plus the four outgroup taxa (Table 1). Some specimens were collected from wild plant populations; others were collected from cultivated plants of the University of California, Davis campus, the USDA National Clonal Germplasm Repository in Davis, and the USDA Southeastern Fruit and Nut Research Station in Byron, GA. All sources of DNA are documented by ] j ted in the University of Wisconsin-Eau Claire herbarium (UWEC). We verified TTT against published descriptions and authentic herbarium material. DNA Extraction, Amplification, and Sequencing.—Total DNA was isolated by grinding approximately 100 mg (wet weight) of fresh or frozen leaves in liquid nitrogen and using the DNeasy Plant Mini Kit (Qia- gen) according to the standard protocol. 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Intron 2 of the nuclear gene LEAFY was amplified using primers modified from Oh and Potter (2003): LFY5: 5'-CAGAACATTGCCAAGGAGC-3', LFY4: 5'-GGCTTGTTGATGTAGCT- TGC-3'; and nuclear gene s6pdh from Exon 2 to Exon 6 was amplified using primers modified from Bortiri et al. (2002) plus a new primer (0): sópdh-k: 5'-CAAGAGTGAAGCAGACGTTGG-3', s6pdh-h: 5-AGACCA- ATGCTACGAACTAGGCCG-3', s6pdh- o: 5'-AGAATAAGGTGTTGGACATAGACG-3', s6pdh- p: 3'-AGAGTG- GTCCTGGATTTCTTATCTA-3’. Thirty-five cycles of three-step PCR, preceded by an initial melti p of 4 min at 94°C and concluded by a final extension of 7 min at 72°C, were carried out as follows: den at 94°C for 30 sec, primer annealing at 52°C (LEAFY) or 54°C (s6pdh) for 1 min, and extension at 72°C for 2 min. Following electrophoresis we excised PCR products from 1% agarose gels and purified them using a QlAquick Gel Extraction Kit (Qiagen). We sent the fragments for direct sequencing using the PCR primers either to Davis Sequencing (Davis, CA) or the DNA Sequence Laboratory of the University of Wisconsin Biotechnology Center (Madison, WT). PCR products with unreadable sequences or a significant number of ambiguous base calls were cloned using the TOPO TA Cloning Kit for Sequencing (Invitrogen) following the manufacturer's instructions. Sequencing of all clones using T3 and T7 primers was done by the DNA Sequence ESO of the UW Biotechnology SEDE E wae one oM with posse PER aar cat La ege 1 removed from furtl each other I artifacts and to the original chromatogram obtained through e SE - When two or more clones yielded identical E SEH one was used in tog analyses. 1 for] ] ing Chromas 2.22 (Tech- nelysium Pty Ltd), and the Ses ess were aligned by eye in GeneDoc 2.6.002 (Nicholas & Nicholas 1997). The close relationship among P jes made manual alignment of the sequences straightforward. For data analysis, parsimony informative indels were coded as additional binary characters. Seven indels were included for LEAFY and eight for s6pdh. LEAFY sequences were trimmed to just Intron 2; s6pdh sequences were truncated to begin 18 bp downstream from the end of primer k, located in Exon 2, and end 14 bp upstream from the start of primer p in Exon 6. Sequences were submitted to GenBank (accession numbers listed in Table 1). Phylogenetic Analyses.—Both data sets were analyzed using maximum parsimony (MP) with PAUP* 4.0b10 (Swofford 2002) and Bayesian inference (BI) with MrBayes 3.1.2 (Huelsenbeck & Ronquist 2001; Ronquist & Huelsebeck 2003). Two heuristic MP searches with TBR branch swapping and the Multrees option were performed: the first with MaxTrees set to one million; in the second, the “nchuck” and “chuck- score" options were employed so that 10,000 trees were saved and swapped for each of 30 random addition replicates. The results of each search were summarized as strict consensus cladograms. Relative support for each branch was assessed by heuristic searches of 1000 bootstrap replications with 10 random addition replicates each and TBR branch swapping. For each random addition replicate 100 trees were saved. For BI analyses each data set was partitioned into nucleotide characters and indel characters. ModelTest 3.7 (Posada & Crandall 1998) was used to select the substitution model that best fit the nucleotide data using the Akaike information criterion: TVMef for LEAFY and HKY+I+G for s6pdh. Each of two BI runs for each gene consisted of three heated and one cold chain with samples drawn every 100 generations. We ran the LEAFY data for 1,000,000 generations discarding the first 500,000 generations, whereas for s6pdh we ran 1,500,000 generations discarding the first 750,000 generations. The runs for each data set were combined and a majority rule consensus tree constructed with a 6796 threshold. Determination of chloroplast haplotypes.—In order to compare our phylogenetic results from nuclear LEAFY and s6pdh with those of Shaw and Small (2004, 2005), we sequenced the trnL-trnF intergenic spacer from the chloroplast genome for our accessions. We amplified the spacer using the e-f primer pair of Taberlet et al. (1991) and used primer e, and sometimes primer f, for sequencing (GenBank accessions EU606152- EU606192). We aligned the 18 Prunocerasus exemplars used in Shaw and Small L 406 | ti tanical Insti lexas 2( (2004). The maximum parsimony tree from the combined dataset was used to place each of our native plum accessions into one of the primary chloroplast haplotypes (A, B, C, S, or T) of Shaw and Small (2005). RESULTS For clarity in the results and discussion sections we refer to specific sequences by their names on the phy- lograms, omitting the genus name and differentiating among clones from the same collection by decimels following the last three digits of the collection number LEAFY Alignment and Data Analysis.—PCR with primers LFY5 and LFY4 yielded fragments approxi- mately 750 or 1000 nucleotides in length, the difference being a 250-bp insertion. Only 1000-bp fragments were PCR amplified for Prunus hortulana and P. murrayana. All three individuals of P. rivularis and two of the four collections of P. munsoniana (513, 676) had both 750 and 1000-bp fragments (Fig 1). The other collections of P. munsoniana and the remainder of the species yielded only 750-bp fragments. Some of the cloned P. rivularis sequences appeared to be recombinations of the 750 and 1000-bp sequences, similar to those of P. angustifolia and P. hortulana/P. murrayana respectively (Fig 2). Putative recombinant sequences were omitted from the phylogenetic analyses. Trimmed to LEAFY Intron 2, the ingroup sequences varied in length from 658 bp in P. texana to 925 bp in one clone of P. rivularis (528.22). The final alignment used in the LEAFY data analysis included 930 nucleotide characters, of which 856 were invariant, 35 varied but were parsimony-uninformative, and 39 (4.296) were parsimony informative for the ingroup taxa. The strict consensus MP cladograms from the two parsimony analyses are identical (not shown), each of length 194 with consistency index (CI) of 0.634 and retention index (RI) of 0.796. Their topologies are similar to the Bayesian analysis phylogram (Fig. 3), but several nodes (indicated by the solid circles) are collapsed and there are minor rearrangements within Clades 3 and 4 (topology indicated by dotted lines). Species of Prunus sect. Prunocerasus form a monophyletic group split into three lineages: P. subcordata plus Clades 1 and 2. Clade 3 is composed of P. angustifolia and angustifolia-like sequences from P. munsoniana and P. rivularis. The latter two species (names in bold on Fig. 3) also have cloned sequences from the same individuals on Clade 2. Relationships among P. americana, P. mexicana, and P. nigra are unresolved; they diverge from a basal polytomy on Clade 2. Sequences of Clade 4, including those of P. hortulana, P. mur- rayana (except 5255), P. munsoniana, and P. rivularis, have a 250-bp insertion not found among the other sequences. s6pdh Alignment and Data Analysis.—The final ali t for the s6pdh analysis includes 1272 nucleotide characters, of which 1197 were invariant, 46 varied but parsi inf tive, and 29 (2.3%) were parsimony informative for the Prunus sect. Prunocerasus species. The Soest gthe ingroup were those of P. hortulana (628), P. maritima, and P. subcordata at 1204 Alice des including 7 or 8 IC repeats in Intron 3, whereas one P. americana clone (505.2) had the longest sequence at 1256 nucleotides with 31 TC repeats. Just as in the LEAFY analyses, the strict consensus MP cladograms from both parsimony analyses of s6pdh are identical (not shown) and have lengths of 153, consistency indices (CI) of 0.699, and retention indices (RI) of 0.854. The MP cladograms are nearly identical to the BI phylogram (Fig. 4). On the MP cla- dograms, Clades 5 and 6 are sister groups such that Prunus sect. Prunocerasus forms a monophyletic group, albeit with bootstrap support less than 5096, the branch indicated by the black circle is collapsed, and americana.684.1 is sister to americana.495.3/americana.505.23 on Clade 5 (indicated by dotted lines). Clade 5 includes P. subcordata plus subcordata-like sequences from five individuals of P. americana and one of P. munsoniana. These accessions (names in bold on Fig. 4) also have sequences on Clade 6, which includes all the other native plum taxa. There is poor resolution among these taxa. Prunus angustifolia, P. texana, and angustifolia-like seq from P. munsoniana and P. rivularis compose Clade 7. Clade 8 consists solely of P. americana and P. mexicana, plus a similar sequence from one individual of P. umbellata (520.4), which also has a sequence among the taxa of the Clade 6 basal polytomy (umbellata.520.3). Rohrer et al., Phyl ti lysis of North American P t. Prunocerasus 407 Li d - angu muns rivu hort 100 bp 10519 10513 10514 10598 Ladder —— = 1000 bp — —— — — 750 bp rivularis. 514.24 | ! — rivularis.527.24 | C GA * TECGGA 6 | rivularis.528.2 em rivularis.514.1 | CGA | CTTCAC A | rivularis.514.2 S | rivularis.527.22 | | ATG CTTCACA | rivularis.527 2 | j — | rivularis.528.1 | H gel A TG | *ICCGGA G | rivularis.514.21 rivularis. 527.3 | rivularis. 527.25 | | d * A TG C TTCACA | rivularis.528.3 Fic. 2. Graphi tati f cloned LEAFY from P. ivulari ions. Each bar re; ts DNA seq Dark gray seg kr See D Hit iflada 9 Fig 3) PL A l g if g a 5 fara sertio (alignment gap), t i p and 9-bp d lett tide diff type and angustifolia-ty -type sequences. The I laries bet tyr | to within about 75 bp DISCUSSION The nuclear genes LEAFY and s6pdh provide weak resolution of relationships among species of Prunus sect. Prunocerasus. The majority of taxa diverge from polytomies on the MP consensus and BI trees. The topology of these trees is different from the cpDNA cladogram of Shaw and Small (2005) where P. subcordata and then P. texana are sister groups to the remaining species, which segregate into three primary clades: American 408 7/4) hortulana.628 A munsoniana 513.2 C munsoniana.876.3 C rivularis.527.24 C murrayana.525S A Clade 4 93 100 murrayana.524 A Lm [m L——— rivularis.528.2 C 97 hortulana.598.4 C 0 — americana.495.4 C 64 | americana.587.3 C e americana.607 € 65! munsoniana.592.1 C o 95 | mexicana.511 A 93 mexicana.591 A C 3T americana.495.2 —— americana. 505 A | amerícana.508 C Clade 2 EM americana.587.2 C americana.684 mexicana.517 A es. | mexicana.518 mexicana.521 A | mexicana, 545 A nigra.606 A -—— ——— nigra633 A 90 geniculata.609 B y EA texana.685 munsoniana.513S C munsoniana.5924 C En munsoniana.676.1 C Prunussect. |. || | | || | JL. rivularis.527.25 C Ince Clade 3 angustifolia. 588.2 C 400 munsoniana.626.1 C munsoniana.628 | 8 rivularis.514.21 C rivularis.528.3 C a a rivularis.514S angustifolia.519 C 100 ic angustifolia.588.1 C 199 wade” alleghaniensis. 467 C 100 'umbellata. 520 C 66 umbellata.620 C poe emgoen 468 C | - gracilis.516 C ma.590 B subcordata. a S cerasifera 100 _; pumila var. pumila uod 100 ! pumila var. susquehanae.445 _ — 1 change it | hy! f B | | g t | d; g i t Support (%)f h clade; below brand AP I p val 5096 MP strict consensus li re labeled de 3. EFArt £T CL i H 1 al L Taxaa F D m [| E am J* ey f D d LI LI " Ll J £ e ll f^rhnrt X f [| H ER Al A . nm Rohrer et al., Phvi t. Prunocerasus 400 americana.495.2 C americana.505.2 americana.508.4 C 7; C americana.58 americana.684.2 C 100 mexicana.511 A 76 mexicana.517 Clade 8 — mexicana.518 A 92 mexicana.521 A —— umbellata. 520.4 € americana.607.3 angustifolia.519 C — munsoniana.5132 C munsoniana.592.24 C i rivularis.527.3 rivularis.528.4 C texana.685 T angustifolia.588 C 100 munsoniana.592.21 C Clade 6 ef 87 | rivutaris.527.2 C Na 92! munsoniana.513.1 C X——- 34 ¡nigra.606 A ! nigra. 633 A | alleghaniensis.467 C — alleghaniensis.468 C — geniculata.609 B gracilis.516 CG 3 E 3 D o ho » cara | murrayana.526 rivularis.514.1 C H rivularis.528.2 C — umbellata.520.3 C t umbellata.o20 C X. 00 [——— americana.495.3 C See ¡ 88 Lo americana.505.23 A L— ameriçana.684.1 C americana.607.2 C Clade 5 100 MM 61 " 98 66! munsoniana.592.23 C © 59 | americana.508.3 C subcordata.5 S 100 j pumila var. pumila.611 100 L— pumila var. susquehanae.445 cerasifera.541 —— 1 change Fic. 4. sópdh 67% majority rul phylogram from Bayesian analysis. Ingroup t italicized; outgroup t i ype. Support | L zt Bly A H | L O E (96) fi 1 | J d | L L aanl FEN | | Fans MP strict consensus Taxa are labeled Lea Jta E I | a L f J al L ^ | LA E sl) H aL A J4 MAJA L Lat 4 r pott 32M MK IK J» 410 | lof tl tanical h Instit Texas 2( (A), Beach (B), and Chickasaw (C). Clade 1 on the LEAFY tree and s6pdh Clades 5 and 7 consist largely of accessions with cpDNA haplotype C, otherwise taxa on the nuclear gene trees do not resolve into the A, B, and C clades of Shaw and Small (2005). fica qas all chloroplast and nuclear phylogenies is ge sorting, or long-branch attraction — common and typically attributed to such factors as | eo et » m Doyle 1997; Macdison 1997; Sang & Zhong 2000). e geographic distribution of e chloroplas d with chloroplast capture given the ease with which North American ds are ee to os endis (Medel 1911; Shaw & Small 2004). Although Prunus texana, known as peachbush or Texas wild peach, has historically been placed in sect. Amygdalus due to its hairy-skinned fruit, Shaw and Small d dée that it has a plum chloroplast genome. Evidence from the nuclear genes LEAFY and s6pdh tin sect. Prunocerasus. On the cpDNA cladogram P. texana was sister to the other pums of eastern North America, a position we could not confirm. It is sister to P. geniculata, the rare endemic of the Lake Wales Ridge in central Florida, on our LEAFY phylogram. However, this relationship is based on one character change and has weak bootstrap support. On the s6pdh phylogram P. texana is part of Clade 7 with P. angustifolia and the angustifolia-like sequences of P. munsoniana and P. rivularis. Prunus alleghaniensis, P. gracilis, P. maritima, and P. umbellata form a paraphyletic group on LEAFY Clade 1, basal to Clade 3. These four species clustered on the UPGMA dendrogram based on similarity of microsatellite markers in the study of Rohrer et al. (2004). These species closely resemble each other morphologically and were allied by early students of American plums such as Wight (1915). On the sópdh EE EE are part - the a Clade 6 basal bis Prunus 1 be difficult to sep hologically, especially when working with herbarium specimens. pue their LEAFY and s6pdh sequences were mia and without resolution into monophyletic P. americana and P. mexicana groups. On the LEAFY phylogram the P. americana and P. mexicana sequences diverge from a basal polytomy along with P. nigra, munsoniana.592.1, and Clade 4. LEAFY sequences americana.508 (WD, americana.684 (PA), mexicana.518 (TX), and mexicana.545 (cult.) differ trivially by the length of a poly-A run and some nucleotide characters heterozygous for one or two sequences but with only one of the two bases on the other sequences. Otherwise they are identical. Although our data do not separate Prunus americana from P. mexicana sequences on either LEAFY Clade 2 or s6pdh Clade 8, Boonprakob et al. (2001) calculated a UPGMA dendrogram, based on similarities of RAPD markers, on which all 13 of their accessions of P. americana clustered together and distinct from a cluster of all 11 accessions of P. mexicana. Clearly more collections with a wide geographic distribution and more genes will need to be examined and correlated with the morphological characteristics of those accessions before the taxonomic structure of the P. americana-mexicana complex is understood. In an earlier microsatellite study, Rohrer et al. (2004) included two collections from east Texas near Marshall Ge e P (americana x mexicana?) 10517 and 10518 in that publication) that are morpho- logically int typical Prunus americana and P. mexicana. The lack of differentiation between P. americana and P. mexicana LEAFY and s6pdh ee leaves their determination ambiguous. However, because neither 517 nor 518 have a subcordata-like s6p e and | t modern fl exclude P. americana from Texas, we have provisionally deeem ed these accessions to be P. mexicana. Two distinct sequences were isolated from five of the six accessions of Prunus americana by cloning their s6pdh PCR products. One sequence from each pair is on Clade 8 along with P. mexicana and the single sequence of P. americana oe the omen is on Clade 5 with P. subcordata. Given that Së P. americana se- quences form a ph; group sister to P. subcordata and that P. subcordat y in California and Oregon, far west of P. ge in eastern North America, it seems alle that che five P. americana collections with two s6pdh sequences are recent hybrids independently involving P. subcordata. If sópdh is a single copy gene in all diploid Prunus species as concluded by Bortiri et al. (2002) and if P. americana is diploid as it is reported to be (2n = 16, Love & Love 1982), then a high degree of heterozygosity is being maintained at a single locus. However, we cannot rule out the possibility that there are two heterologous Rohrer et al., Phyl g ti lysis of North A j p t. Prunocerasus 411 ancestral s6pdh loci across Prunus with only one or the other copy observed in most North American plums, or alternatively gene duplication in P. americana. Bortiri et al. (2002) found divergent s6pdh sequences in Prunus caroliniana and P. emarginata. They | thesized that divergent paralogs in P. caroliniana, a tetraploid species, may have arisen through EHE np lesion associated with polyploidization. In P. caroliniana the inferred polypeptides of the diver ger ent clones es at P» of 251 amino acids (Bortiri et al. 2002), whereas for each of our five accessions of Prunus americana with divergen t s6 , the inferred polypeptides of the two clones differ by only 1—4 out of 223 amino ads To Boer pan the existence of divergent sequences in P. americana, it would be useful to have chromosome counts from each of those accessions to verify that they are indeed diploid. Also because Southern blotting with a Malus domestica cDNA s6pdh probe by Bortiri et al. (2002) detected more than one region of sequence similarity in several genera of Rosaceae, the possibility of paralogous genes might be further investigated. Alleles closely related to those of Prunus americana, and possibly P. mexicana, were found in other plum species suggesting introgression. P iana 592 has both s6pdha EAFY sequences monophyletic with sequences from P. americana 607. On the sópdh tree munsoniana 592.23 is i with the five subcordata- like P. americana clones and has the same amino acid sequence as americana.495.3 and americana.505.23. On the LEAFY tree munsoniana.592.1 is sister to americana.607. Also P. umbellata 520 has an americana! mexicana-type s6pdh sequence (umbellata.520.4) in addition to a sequence similar to that of the P. umbellata 620 collection. Sequences from both LEAFY and s6pdh suggest that Prunus rivularis may be an allopolyploid with P. angustifolia as one parent. The LEAFY sequences further suggest that the other parent is most likely P. hor- tulana or P. murrayana and that recombination between parental sequences has taken place. Three or four distinct sequences were cloned from each of the P. rivularis accessions. LEAFY is believed to be a single-copy nuclear gene (Frohlich & Parker 2000; Oh & Potter 2003) and would be expected to have one or two alleles in a diploid species. At least one sequence from each P. rivularis accession is monophyletic with sequences of P. angustifolia (Clade 3 on the LEAFY phylogram and Clade 7 on the sópdh phylogram). The other se- quence is on Clade 4 of LEAFY along with sequences of P. hortulana and P. murrayana. In addition, some LEAFY recombinant sequences were cloned where part of the sequence matches P. angustifolia and the rest matches P. hortulana/murrayana (graphically illustrated on Fig. 2). Although it is possible for recombinant sequences to arise as PCR artifacts (Cronn et al. 2002; Posada et al. 2002), in our study they were isolated only from P. rivularis and they were found among the LEAFY sequences of all three P. rivularis accessions. This would seem to support the hypothesis that the recombinant sequences arose through meiotic recom- bination following hybridization between P. angustifolia and either P. hortulana or P. murrayana. Given the few phylogenetically-informative characters in the s6pdh data, if there were recombinant s6pdh sequences, we were unable to identify them. Plants in our study determined as Prunus munsoniana appear to be a mixture of hybrids having P. an- gustifolia as one of the parents. All four P. munsoniana collections have a LEAFY sequence on Clade 3 with P. angustifolia and P. rivularis, and three collections have s6pdh sequences on Clade 7, again with P. angustifolia and P. rivularis (we are lacking a s6pdh sequence for P. munsoniana 676). Two P. munsoniana collections (513 and 676) have a second LEAFY sequence with the 250-bp insertion found also among sequences B P. hortu- lana, P. murrayana, and P. rivularis on Clade 4. On the BI cladogram, munsoniana.513.2 676.3 form a trichotomy with rivularis.527.24, but on the MP cladogram they are sister to hortulana.628. Thus, these two P. munsoniana collections have LEAFY sequences closely related to those of P. rivularis, which lends support to the suggestion by Diggs et al. (1999) that P. munsoniana is simply a larger version of P. rivularis. However, the other two specimens of P. munsoniana (592 and 626) lack hortulana/murrayana-like LEAFY sequences, yet are very similar morphologically to 513 and 676. For P. munsoniana 626 we were only able to isolate angustifolia-like LEAFY sequences. Likewise we isolated only a single s6pdh sequence, which lies on Clade 7 with P. angustifolia and P. rivularis. P. munsoniana 513 and 592 each yielded three distinct s6pdh may be allopolyploids. P. munsoniana 592 keen sequences, suggesting that at least some P. munsoniana L 412 Jou has a sequence on s6pdh Clade 7 (592.24) with P. angustifolia, and on Clade 5 (592.23) sister to a clone of P. americana (607.2), just as on the LEAFY phylogram. The third sópdh sequence (592.21) is sister to rivu- laris.527.2 and these two are sister to munsoniana.513.1. To our knowledge we are the first to suggest a close phylogenetic relationship between Prunus mur- rayana and P. hortulana (Clade 4 on the LEAFY phylogram). P. murrayana.525L and murrayana.526 form a clade with hortulana.598.1 having 93% MP bootstrap support and 100% BI posterior probability. The s6pdh sequences do not confirm or refute the close relationship between P. murrayana and P. hortulana, because their sequences along with those of P. alleghaniensis, P. geniculata, P. gracilis, and several other species all are unresolved on the Clade 6 basal polytomy. Until a decade ago, P. murrayana was a poorly known species endemic to Texas west of the Pecos River and represented in herbaria by only a handful of specimens. When originally described, Palmer (1929) wrote that it might be most closely related to P. rivularis. Enquist (1997) formalized this relationship by designating P. murrayana as a synonym when he described P. rivularis var. pubescens Enquist. He discovered numerous plums on the western don e the Edwards Plateau SH pubes- cent first-year branchlets, pubescent pedicels, and pul ,an led that these plants were the same taxon as P. murrayana — from her west Many di Enquist's collections are morphologically well matched with authentic P. murrayana specimens. He explained how most of the apparent differences between P. murrayana and P. rivularis did not hold in light of the new collections and that the only distinguishing character was pubescence; the branchlets, pedicels, and petioles of P. rivularis are glabrous or Pd so (Enquist 1997). Clade 4 of ER LEAFY phylogram includes not only P. hortulana and P. from all three accessions of P. rivularis and a couple from P. munsoniana. Our P. Wu IA eden: are from Coke and Tom Greene Counties east of the Pecos River at localities where Enquist (1997) collected P. rivularis var. pubescens [= P. murrayana]. These collections have only Clade 4 LEAFY sequences, whereas our three collections of P. rivularis have both Clade 4 and Clade 3 sequences. SE P. Mii i accessions have s6pdh sequences only from the Clade 6 polytomy, whereas all three s have a cloned sequence on Clade 7 with P. angustifolia and from the basal polytomy of Clade 6. We believe that P. murrayana is a diploid species distinct from P. rivularis, which we believe is an allopolyploid derived from P. angustifolia and either P. murrayana or P. hortulana. ACKNOWLEDGMENTS Financial support for this research was provided by several Faculty/Student Research Collaboration Grants and a University Research and Creative Activity Grant from the UWEC Office of University Research and Sponsored Programs. The Faculty Sabbatical Leave Program of the University of Wisconsin allowed J. Rohrer to spend five months in the Department of Pomology at the University of California, Davis, where Dan Potter, Esteban Bortiri, and Sang-Hun Oh graciously shared their knowledge of molecular techniques and extended hospitality. We thank Lloyd Turtinen and Sasha Showsh of the UWEC Biology Department for use of equipment in their labs. We also wish to acknowledge the contributions of UWEC undergraduates Kelly Leinberger and Sarah Ivory to this project. Dick Okie, Carl Weekley, Bob O'Kennon, and Jim Everitt kindly provided hard-to-obtain plant material for sequencing. Three anonymous reviewers provided many helpful comments for improving the paper. REFERENCES ALBALADEJO, R.G, J.F. AGUILAR, A. Aparicio, and G.N. FeLiner 2005. Contrasting nuclear-plastid tic patterns in the recently diverged Iberian Phlomis crinita and P. lychnitis lineages (Lamiaceae). Taxon 54. 987-998. BADENES, M.L. and DE Parent. 1995. 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Phylogenetic relationships in the stone fruit group of Prunus as revealed by restriction fragment analysis of chloroplast DNA. Jap. J. Genet. 66:59-69. WicHt, WE 1915, Native American species of Prunus. Bull. U.S. Dept. Agric. 179:1—75. — d — l RECOGNITION OF THREE TAXA OF EASTERN NORTH AMERICAN “WALDSTEINIA” AND THEIR APPROPRIATE NAMES WHEN INCORPORATED INTO GEUM (COLURIEAE: ROSACEAE) Alan S. Weakley Kanchi N. Gandhi University of North Carolina Herbarium (NCU) Harvard University Herbaria (GH) North Carolina Botanical Garden, ^ Chapel Hill 22 Divinity Aven ampus Box 328 Cambridge, Massachusetts o | E -2094, U.S.A. Chapel Hill, North Carolina 99-3280, U.S.A, gandhi@oeb.harvard.edu eakley@unc.edu ABSTRACT We review tl ] I and taxonomic history of eastern North American st litionally treated in th E Waldsteinia Willd. (Rosaceae), and ] t g t (as has been don ith rar pti ) pues these in the genus Geum L. pas +l E 1 1 1 à M EOS ssl 4 ru 1 1 T. dad. us +l y WIEL 1 SE taxa, d most conservative treatment is to Ze them at the same (species) rank. A new combination is made in Geum CH the ted as W. M Cn im W. pagando Gg be var, Lini flora (Small) Fernald, W. doniana Tratt., or W. PONES ssp. ERC (Tratt.) T (Tratt.) Weakley & Gandhi, comb. nov. RESUMEN ict d A 1 lat lads] i «ber qp ui : i "E A 1:5; 1 1 P, Se revisa la | y E g Walds- teinia Willd. e y luye q leseabl (com ha hech pciones) y colocarlos en el género Geum | t fológ ] yl de d lecul los que apoy EE ee Le taxa elt ient 5 d ] ] limi y C41 n dedlendese de ine ted Undead rara W. Para Small, W. fragarioides (Michx.) Tratt. var. prior (Small) Fernald, W. doniana Tratt., o W. ia ssp. doniana (Tratt.) Teppner (este último no publicado efectivamente): Geum donianum (Tratt.) Weakley & Gandhi, comb. nov. Recent investigations in the taxonomy of tribe Colurieae Rydberg of the Rosaceae have led to the conclu- sion that Waldsteinia Tratt. is abu ou - embedded within Geum L. and Pedes be combined with it (Smedmark 2006). Smed 2006), Smedmark et al. (2003), and Smed ] Eriksson (2002) presented 11 1 a strong evidence that Waldsieiniái is involved in ancient from which parts of Geum have arisen, and they argue that the most reasonable GER of the genus is to include all the perennial herbs of Geinae involved in the reticulate evolution. Morphologically, Waldsteinia is comfortably congruent within a broadly defined Geum, differing only in the style dehiscent at its base; and Geum (in a narrower sense) already includes a wide diversity of style morphologies, apparently adaptations to a variety of pol- lination and dispersal Denon due Most North American ted three taxa of Waldsteinia for eastern North America: Taxon A (" fragarioides," sensu stricto), a northern EE entity with trifoliolate leaves; Taxon B “parviflora” r "doniana"), a southern smaller-petaled entity with trifoliolate leaves; and Taxon C (“lobata”), a Southern Aun endemic with small petals and lobed, rather than trifoliolate, leaves (Table 1). A notable excep- tion to the acceptance of Taxon A and Taxon B as distinct at some formal taxonomic level is the treatment of Radford et al. (1968), which lumps Taxa A and B and does not mention W. lobata, reported for the Carolinas by earlier authors. Smedmark (2006) made transfers into Geum of Waldsteinia species, but regarded Taxon B asa synonym of Taxon A (without explanation, but perhaps because of the nomenclatural and taxonomic confusion discussed below), leaving it without a name in Geum. Although their acceptance as taxa has been nearly universal, the appropriate taxonomic level at which to recognize these entities (species, subspecies, or variety) has remained controversial and variable (Table 1). J. Bot. Res. Inst. Texas 2(1): 415 — 418. 2008 fall Dos H In LI dede £T TAE! dl 416 | Taste 1. A comparison of recent taxonomic treatments of the "Waldsteinia" taxa of eastern North America. Treatment Taxon A: fragarioides Taxon B: doniana/parviflora Taxon C; lobata Small (1898) Waldsteinia fragarioides Waldsteinia parviflora Waldsteinia lobata Rydberg (1913) Waldsteinia fragarioides Waldsteinia doniana Waldsteinia lobata Small (1933) Waldsteinia fragarioides Waldsteinia doniana Waldsteinia lobata Fernald (1950) Waldsteinia fragarioides var. Waldsteinia fragarioides var. fragarioides parviflora [out of area treated] Gleason (1952) Waldsteinia fragarioides Waldsteinia parviflora fout of area treated] Teppner (1968) Waldsteinia fragarioides ssp. Waldsteinia fragarioides "ssp. fragarioides doniana" Waldsteinia lobata Radford, Ahles, & Bell (1968) Waldsteinia fragarioides [not treated) Gleason €: Cronquist (1991) Waldsteinia fragarioides var. Waldsteinia fragarioides var. fragarioides parviflora {out of area treated} Kartesz 1999 Waldsteinia fragarioides ssp. Waldsteinia tragarioides “ssp. fragarioides doniana” Waldsteinia lobata Smedmark (2006) Geum fragarioides Geum lobatum Weakley & Gandhi (this paper) Geum fragarioides Geum donianum Geum lobatum Phipps in FNA (in prep.) Waldsteinia fragarioides Waldsteinia doniana Waldsteinia lobata A review of literature and specimens reveals that tl taxa should | ized in eastern North America ii ge SE the three taxa eid from one another in details of leal lobing/division and petal size, 1 ] rrelated with coherent geographic distributions. Though imperfect herbarium specimens may be difficult to interpret, there seem to be no true iU ANN nor is their clinal variation in petal size. Moreover, the “Waldsteinia group" of i lly conservative, with rela- g taxa on different BN such) as the North Ámerican tively minor morphological characters set G. fragarioides and the Eurasian G. ternatum (Stephan) Smedmark. Indeed, the morphological relationships of the three taxa make the closer affinity of Taxon B problematic (if leaf lobing is the more phylogenetically revealing character, then Taxon B may be more closely related to Taxon A, but lacking molecular data it is plausible that the floral characters are more phylogenetically revealing, and taxa B and C are more closely related, forming a southern, small-petaled clade). For these reasons, we agree with Phipps (in prep.) that the best current (and conservative) taxonomic treatment of the “Waldsteinia group” in eastern North America is as three species. Nomenclature has also been contentious, with disagreement as to the appropriate basionym (doniana' or ‘parviflora’) to apply to Taxon B (Table 1). Fernald (1935) believed that the application of the epithet 'do- niana' to the small-petaled southern taxa was a misapplication. He argued that the type plate "seems to me a garden-development in which, presumably through unwonted nutrition, the sepals become exaggeratedly large, the petals remaining fairly typical for W. fragarioides.” Teppner (1968) convincingly argued the op- posite case, that the relative size of the sepals and petals are clearly shown and described, and this primary diagnostic characteristic between the two taxa should be taken at face value. A review of the description and plate (which serves as the type) confirms Teppner's opinion, making ‘doniana’ the correct basionym for the southern taxon, should it be recognized taxonomically at the specific level. The plate in Sims (1813) (on which Trattinick's name is based) resembles Taxon B closely and obviously, in that it shows a plant with petals shorter than the sepals, a characteristic that is also stated in the text of both Sims (1815) and Trattin- ick (1823) as a distinctive or diagnostic characteristic. The stated provenance of the plant illustrated (Sims 1813) is pertinent, as well: “the plant, from which our drawing and description were taken, was brought from America, by Mr. Lyons [sic], and purchased at his sale, by Mr. Kent, of Clapton, by whom it was kindly communicated to us.” Ewan & Ewan (1963) show that John Lyon listed this plant in a catalogue in 1812 (as y i, | ic history of Waldes teinia 417 Dalibarda fragarioides), and also listed Mr. William Kent as having purchased plants for 14 £, 18 sh. in 1812. Presumably the plant illustrated the following year was in this lot. Though the location from which Lyon collected the plant cannot apparently be definitely known, the identification of John Lyon as the source of the plant illustrated and described is of interest, since Lyon collected extensively within the southeastern United States range of the small-petaled Taxon B. Thus, it seems very plausible that the plant described and figured as Waldsteinia fragarioides by Sims, and later described and named by Trattinick as the distinct W. doniana, is exactly what it appears to be. Based on our review of generic circumscription, taxonomic distinctiveness, appropriate taxonomic rank, and nomenclature, we hereby effect the transfer of "Taxon B" to Geum at the rank of species: Geum pde AN ) Weakley & Gandhi, comb. nov. Waldsteinia doniana Tratt., R M hia 3:109. 1823 e r ag. 38: pl. 1567 (Lecrorre, designated here). Waldsteinia parviflora Small, Bull. Torrey Bot. Club 25:137. 1898. Waldsteinia fragarioides var. parviflora (Small) Fernald, Rhodora 37:285. 193 Valdsteinia fragarioides ssp. doniana ae ee Zur Kenntnis der Geen Waldsteinia. Diss. Univ. Graz. 1968, nom. invalid., not elfectiv ely os under je N Vienna geg ae ar Article 30.5 states ST “publication on or afier 1 anuary 164 I ] " 1 1 y Tar tainin A 1 of f q Ei ES ] ] Iss 1 a Eech +1 f lI i + L L a L 41 om ] r mr LI y 1 1 - | 1 J cr 11 1 E 1 LI Note 2 suggests fomes of internal evidence as “the ias i an International inis Book e SBN) or a statement of the name of the printer, publisher, ginal p g work was intended to be aie rubia S ns Tepi listributed i ly 50 f his di i ] ical i tions and clearly ive publication (H Teppner pers comm., March foe we ai de conclude that he did t tl j i ts of the Vienna Code, 10 at the subspecific rank. The taxa are distinguishable by the following key. . Leaves trilobed (the sinuses cleft 1/8 to 3/4 of the way to tl irib); petals si tl bout as | the sepais; first-year leaves rather densely pubescent with stiff hairs, these distributed on the veins and on the intervein surfaces; [of a small area at the southern terminus of the Southern Appalachians in n. GA, n SC, and sw. NC) G. lobatu dnd Smedmark k SE ifolola (fully divid 3), | al ically additionally lobed ter than to longer than the pale first year leaves sparsely pubescent with h stiff hairs, these distributed SE or strictly on the veins, the intervein surf g EE | dp EE |] 2. Petals 2,2-4 | (s! tl | tl ), 1-1.5 mm nude terminal leaflet with a distinct pevolule 1 -6r ot Jong, the basal angle of the leaflet acute mostly 60-90” (avg. 75°); [distributed in the Southern Piedmont of AL, GA, KY, NC, SC, and TN] G. donianum (Tratt) Weakley & Gandhi 2. Petals 4-10 mm (longer than the sepals), 2 id | leaflet sessile or with a | lule to 3 long, the hasal angie ovine leaflet SH a dl angle, mostly 75-105? (avg. 90°); [widely distributed in northeastern it j Canada, south to VA, KY, and AR] fragarioides e Smedmark ACKNOWLEDGMENTS We appreciate EE helpful comments ana UM MN of BENE Teppner, Luc pun Jim Phipps, and an E E ae UTR rm A f the taxa involved at the University of North Carolina eba im (NCU). We is gratefully acknowledge the online availability of botanical research resources at botanicus.org (Missouri Botanical Garden), Google Book Search, and Harvard Uni- versity, which greatly facilitated this research. REFERENCES EWAN, J. and N. Ewan. 1963. John Lyon, nurseryman and plant hunter, and his journal, 1799-1814. Trans. Amer. Phil. Soc. 53(2):1-69, | of the Botanical R h Institute of Texas 2(1) 418 FERNALD, M.L. 1935. Critical plants of the upper Great Lakes region of Ontario and Michigan. Rhodora 37:197-341 (pagination interrupted). FerNALD, M.L. 1950. Gray's manual of botany, eighth (centennial) edition. Corrected printing, 1970. D. Van Nos- trand Co., New York, N.Y. GLEASON, H.A. 1952. The new Britton and Brown illustrated flora of the northeastern United States and adjacent Canada. New York Botanical Garden and Hafner Press, New York, NY. GLEASON, H.A, and A. Cronauist. 1991. Manual of vascular plants of northeastern United States and adjacent Canada, second edition. New York Botanical EEN SE KARTESZ, J.T. 1999. A synonymized checklist and atlas | attributes for the vascular flora of the United States, Canada, and Greenland. First Edition. In: Kartesz, J.T and CA Meacham. Synthesis of the North American flora, Version 1.0. North Carolina Botanical Garden, Chapel Hill. McNuL, J, FR. Barre, H.M. Burcer, V. Demouuin, D.L. HawkswoRrH, K. MARHOLD, D.H. NICOLSON, J. PRADO, PC. SiLva, J.E. Skoc, J.H. Wiersema, and N.J. TURLAND (eds.). 2006. International Code of Botanical Nomenclature (Vienna Code). Regnum Veg. 146. A.R.G. Gantner Verlag KG. Pus, J.B. In prep. Waldsteinia. In: Flora of North America Editorial Committee. Flora of North America north of Mexico, Vol. 9. Oxford University Press, New York, NY. Raororo, AE H.E. Artes, and C.R. Beu. 1968. Manual of the vascular flora of the Carolinas. University of North Carolina Press, Chapel Hill. Rvosena, PA. 1913. Waldsteinia. In: N. Amer. Fl. 22(5):398-399. Sims, J. 1813. Curtis's Bot. Mag. vol. 38. London. SMALL, J.K. 1898. Studies in the botany of the southern United States. XIII. Bull. Torrey Bot. Club 25:134-151. SMALL, J.K. 1933. Manual of the southeastern flora, being descriptions of the seed plants growing naturally in Florida, Alabama, Mississippi, eastern Louisiana, Tennessee, North Carolina, South Carolina, and Georgia. University of North Carolina Press, Chapel Hill. SMEDMARK, LEE 2006. Recircumscription of Geum L. (Coluriae: Rosaceae). Bot. Jahrb. Syst. 126:1-9. SMEDMARK, LEE. and T. Eriksson. 2002. Phylogenetic relationships of Geum (Rosaceae) and relatives inferred from the nrITS and trnl-trnF regions. Syst. Bot. 27:303-317. SMEDMARK, LEE T. ERIKSSON, R.C. Evans, and C.S. CamPseLL. 2003. Ancient allopolyploid speciation in Geinae (R ) evidence from nuclear granule-bound starch synthesis (GBSSI) gene sequences Syst. Biol. 52:374-385, Teppner, H. 1968. Zur Kenntnis der Gattung Waldsteinia. Diss. Univ. Graz. TRATTINICK, L. 1823. Rosacearum Monographiae lll. J.G. Heubner, Vienna. THE GENUS ABRONIA (NYCTAGINACEAE) IN COLORADO, WITH NOTES ON ABRONIA BOLACKII IN NEW MEXICO Jennifer Ackerfield William F. Jennings Herbarium, rio e Biology PO. Box 952 Colorado State University Louisville, Colorado, 80027, U.S.A. Fort Collins, Colorado See USA. ABSTRACT Several taxonomic RE on Se peus Abronia in ae and New Mexico are presented. First, Abronia argillosa Welsh & Goodrich is a synonym of A. g Second, Al lt. & Fisher is a synonym of A. fragrans Nutt. ex Hook., and specimens of A. carletonii a New Mexico and me ewe be called. e nealleyi UM ade Abronia EE is a species dis- 1 ] tinct from A. fragrans, as iia Se several fruit and h tics. Lastly, recognition y described A. icd ge es J f A! NUNG (r^ | unwarranted and should be i d y y under A. elliptica. A distril I photographs of fruits of each species. RESUMEN Se presentan | el género Al Colorado y Nuevo México. Primero, Abronia argillosa Welsh & 1 A A 1.1 EC) cC 11 Segundo At H 1.4 TE 1 ce TOS = z 2 E A A j g f e n lz le A letonii de N Méxi Texas deben llamarse A. nealleyi Standl. Tercero, Abronia elliptica es una especie distinta Pi I P p de A. fragrans, | ] terísti lel £ lel hábito. Finalmente, el imi lel ient í / C de dictril 1m de Abronia I I descrita A. bolacl ia de A elliptica The Nyctaginaceae is a taxonomically complex family because of the heavy reliance upon mature fruit for identification (Galloway 2003; Welsh 2003). This is particularly true in the genus Abronia, where vegetative morphology can be highly variable within and among species while fruit morphology remains constant and distinct among taxa. The genus Abronia consists of 20 species, distributed throughout the Southwest, high Great Plains, Mexico, coastal California, and Pacific Northwest (Galloway 2003). Five species have been reported for the state of Colorado: Abronia argillosa Welsh & Goodrich, A. carletonii Coult. & Fisher, A. el- liptica A. Nelson, A. fragrans Nutt. ex Hook., and A. nana S. Wats. var. nana. Abronia bolackii Atwood, Welsh & Heil is a newly described species endemic to San Juan County, New Mexico (Atwood et al. 2002). Abronia argillosa is practically identical to A. elliptica vegetatively, and it is only through analysis of the fruit that identification can be made. Both species are perennial from a branching caudex with elliptic to ovate or suborbicular, entire leaves, and with white or purplish tinged flowers grouped in heads that are subtended by ovate to suborbicular bracts. However, Abronia argillosa has wingless fruit while A. elliptica has winged fruit with the wings dilated and inflated at the top (Fig. 1A,C). Abronia argillosa was described in 1980 (Welsh & Goodrich 1980) as a new species Pas Utah and Colorado restricted to Mancos Shale sub- strates. Examination of the original description of Abronia glabrifolia (Standley 1909) and the type specimen [Colorado: 1878, Wm. F. Flint s.n. (UC)] e that this is deai the same species. The type specimen of A, glabrifolia has the characteristic dark brown, wingless fruit with conspicuous white lines seen in the fruit of A. argillosa (Fig. 1C). The name Abronia glabrifolia predates that of Abronia argillosa by almost 70 years, and should thus be the name applied to all specimens of A. argillosa. William F. Flint was in Colorado at the Los Pinos Indian Agency in 1878 (Flint 1878), where he also collected a type specimen of Senecio flintii Rydb. [Colorado: S.W. Colorado, Wm. F. Flint s.n. (NY)]. Los Pinos was the name for the Ute Indian Agency, located about 25 miles north of Ouray and 11 miles south of Montrose, near the town of Eldredge (Young 1997), on what was then the reservation for the Uncompahgre Ute tribe. This locality is close to the Montrose-Ouray County line. Mancos Shale, the type of habitat known to support A. glabrifolia populations (Fig. 2), occurs in the vicinity. J. Bot. Res. Inst. Texas 2(1): 419 — 423. 2008 Es imm Fic. 1. Abronia fruit (taken at 10x). A. A. elliptica [Colorado: Montrose Co.: P. Valley, Colo 90 lee E ISO ed vet RD: gyp tcroj Celda anfenad 2 May 2004, us E Jennings s.n (CS)]. B. A. fi g ICal 948, H.D. Harrington 4256 (CS)]. C. A. glabrifolia [Col À Plat 1,20 Jun 1981 A lege (CS)]. D. A nana var. nana [Colorado: Mesa Co.: Sinbad Valley, right fork of tl |, 1.9 mi to gyi hill on the N si 27 May 2005, Wm. F. Jennings s.n. (CS)]. The presence of Abronia carletonii in Colorado in particular has been questioned or misinterpreted by numerous taxonomists (Weber & Wittmann 2001). Assumption of its presence was based on the type col- lection made by Mark Carleton in 1891, and attributed to eastern Colorado. Abronia carletonii has not been collected in the state since, but is known to occur in New Mexico and Texas, where it is usually restricted to gypsum soil, very unlike the sandy soil that is found on the eastern plains of Colorado. Examination of the type specimen of A. carletonii [Colorado: Eastern Colorado, 1891, M.A. Carleton 459, (F)] confirms that it is actually a narrow-leaved, trailing form of A. fragrans. The fruit is winged, but the wings are not dilated and are thick and indurate as in typical A. fragrans (Fig. 1B). This form of A. fragrans is seen in other specimens from the plains growing in sandy soil [Colorado. Baca Co.: sand hills on south bank of Cimarron River, 31 Aug 1949, Weber & Anderson 5157 (KANU); Kansas. Finney Co.: ca. 2.5 mi S Garden City, scattered in sand dune area south of Arkansas River, 11 Sep 1989, Brooks & McGregor 19676 (KANU)]. In addition, the specimen was probably not collected in Colorado at all, but in extreme southwestern Kansas, just north of Elkhart, where the only Abronia known to be present is A. fragrans. The name A. carletonii should now be subsumed under A. fragrans, and specimens of A. carletonii in New Mexico and Texas should be referred to as A. nealleyi Standley, as suggested by Turner (2004). In the Holzinger tabulation (Holzinger 1892), collection number 459 is not an Abronia at all, but is shown to be Cnicus altissimus and was collected in Kingfisher County, Oklahoma in August. The specimen 9 A. elliptica A A. fragrans E A. glabrifolia ** A. nana var. nana e ef 4 A à s 8 A 7] = © ce e a E 8 a E e Z e o! A E u e A " O A ` A n O D A^ | a © à 0 A A " En 7 |a HPT oe | gt F AL LU L4 Li P 1 J FL [| L L a I L f al mr FA 0) A elliptica circle; A. fragrans ra triangle; A. A glabrifolia = square; A. nana var. nana = star. labeled collection number 459 of Abronia carletonii likely has the wrong label information, and is probably collection number 352 of Abronia fragrans collected in July in Morton County, Kansas, along the Cimarron River, probably near Point of Rocks north of Elkhart. According to Holzinger (1892), Carleton collected in Oklahoma, Kansas, and Texas, but not Colorado. Carleton began his collection trip in Oklahoma in June, collecting in the Cimarron Valley (collection numbers 202-235). He then continued north to Kansas where he collected until July (collection numbers 236-353), traveling across the southcentral counties of Comanche, Barber, Harper, and Sumner before going north to Sedgwick County. He continued westward from Wichita making collections in Reno and Stafford counties, and then in Seward, Stevens, and Morton counties in extreme southwestern Kansas. Carleton then traveled back to Oklahoma and Texas in July and August (collection numbers e ALA). Some taxonomists consider Abronia elliptica to be with, or a variety of, Abronia fragrans. In particular, Welsh (2003) includes A. elliptica within a broad: concept of A. fragrans. However, he misquotes Galloway (1975) as saying that the fruit of A. elliptica is 2-winged with a groove between the wings, and that of A. fragrans is 2-5-winged but not especially grooved. In fact, Galloway (1975) states that A. elliptica is 5-winged or 2-winged with the wings folded together, with dilated wing tops on the fruits that are not folded. Galloway (1975) further states that A. fragrans is deeply grooved or narrowly winged, with thick wings that are not inflated at the top. The fruits on the periphery of the heads in both A. fragrans and A. elliptica are often contorted, with 2 wings folded together, while the fruits in the middle of the head are 5-winged. Careful analysis of the fruit as well as habit shows that A. elliptica and A. fragrans are indeed different species, as shown by the following characters. First, the fruit of A. fragrans is wingless or winged, with hard, rather indurate wings that are flat and lacking dilations (Fig. 1B). The fruit on the periphery of the heads is 41 422 t tani i Texas 2(1) often wingless in A. fragrans while within the central portion, fruits are winged. In contrast, A. elliptica has winged fruit with papery thin wings that are inflated and dilated at the top (Fig. 1A). Second, A. fragrans is generally a prostrate and spreading or semierect plant with more robust stems 3-5 mm wide, and with leaves that are ovate to triangular or lanceolate, especially on the upper part of the stem. Abronia elliptica, however, is generally erect or sometimes ierect with narrower stems typically 2-3 mm wide, with ovate to elliptic-oblong leaves. In addition, the distribution of A. elliptica in Colorado is scattered in the western counties. In contrast, A. Jaunes is common on the eastern peus but is also found in Chaffee County near Buena Vista. It is int ties with a few records from Montezuma County and one from Mesa County (Fig. 2). Abronia nana var. nana in Colorado is! only from gypsum outcrops in the Sinbad Valley in extreme western part of the state, on the Mesa-Montrose Eun) ime "his is the easternmost edge of its distribution range. It is a very distinctive species with acaulescent stems, and the entire plant is usually extremely viscid glandular. The fruit of A. nana var. nana is conspicuously viscid glandular and also winged. The five wings are broad, flat, and not dilated at the top (Fig. 1D). Abronia bolackii Atwood, Welsh & Heil is a newly described species of Abronia endemic to gypsum soil in Ge Juan County, New Mexico (Atwood et al. 2002). It is delimited from A. elliptica based on the presence fewer flowers per head (15-25 opposed to 25-75). However, the fruit and habit of A. bolackii are identical to that of A. elliptica. Specimens of A. elliptica with rhizomes have been collected as far north as San Juan County, Utah [Utah: San Juan Co.: E base of Comb Ridge just N of Hwy US 613, 1 Jun 1999, L. Yeatts 4251 (COLO). Richard Spellenberg (pers. comments) has also observed rhizomatous A. elliptica plants throughout its range from northcentral Arizona and northwestern New Mexico to northeastern Utah and L er including western Colorado. Welsh (2003) st that gnition of the occasional ts (of A. elliptica, included within his description of A. fragrans) at any taxonomic level seems enel. In addition, several specimens of A. elliptica have as few as 19-24 flowers per head [Colorado: Moffat Co.: ca. 2.5 mi E of mouth ai Irish Canyon, 7 Jun 1983, Baker & Kennedy 83-32 (CS); Colorado: Mot- fat Co.: Junction of Green and Yampa rivers, 8 Jul 1945, H.D. Harrington 1480 (CS); Colorado: Mesa Co: Rabbit Valley, sandstone outcrop in sandy soil, 21 May 2004, Wm. F. Jennings s.n. (CS)]. Finally, the range of A. bolachii is well within the range of A. elliptica. Consequently, the status of A. bolachii as a species appears unjustified, and should be included in synonymy under A. elliptica. The following key includes the common name, scientific name, synonyms (if applicable), ecology and/ or distribution, elevation range, flowering time, and occurrence on the eastern or western slope of Colorado (designated by an E or W, respectively) for each species. KEY TO THE SPECIES OF ABRONIA IN COLORADO Ia 1 | r * Eet qae 1. Plants acaulescent, glandular, with 5 broad wings that are flat and not dilated: leaves to 2.5 cm in length, conspicuously viscid glandular A. nana S. Wats. var. nana DWARF SAND-VERBENA mmon on gypsum outcrops in the Cie Valley, known in Colorado from Mesa and Montrose cos., 5400-5800 ft. Apr-May. W. 1. Plant | Le ling and | bent to EE fruit unwinged or winged, dilated or flat, glabrous to villous or glandular atthe top, but not | ly viscid; leaves 1.5-12 cm in length, glabrous to glandular, but not t densely SO. 2. Fruit all wingless, dark | to gray-brown ith pi Il, appressed WOENE but PRENN aeos | without a distinct reticulate vein pattern; fl glal to s y glandular on the u lobes; bracts obovate to ovate or ellipti | led at the ti A. glabrifolia Standl. IPS CLAY Y SAND-VERBENA. [=A. argillosa Welsh & Goodrich]. Found on clay soil of the Mancos Shale formation, known in Colorado from Garfield, Mesa, eS ane Rio Blanco cos, Se 6800 ft. May-Jun. W 2i Fruit usuall ly winged ti ingl the periphen white, greenish, o hite lines but tl |, usi ua usando andar peca at the top, with a conspicuous reticulate vein pattern ti g y to densely glandular HI J throuchout; bracts ovate to obovate or elliptic with rounded or more often acuminate or mucronate + Milas | Aicts|| | dE | thie] an | t “the peripheral d ips. 3. GENEE ori dee ing or sometimes narrow and lanceolate on the upper part of the stem; bracts ovate to inesplanceolaté with acuminate tips ragrans Nutt. ex Hook. FRAGRANT SAND-VERBENA. ouis in sanay sell especially enn on nea eastern plains, but a eat Fakt (Mesa ne Montezuma), aid "HOO May-Aug(-Sep). EW. 3. Fruit winged i lilated (inflated) and expand y some what spreading leaves ovate, elliptic, or sometimes orbicul T ts o ate to obovate, with rounded or acuminate tip A. elliptica A. Nels. WESTERN EE delo on gypsum, clay, or sandy soil, known in Colorado from the western counties ta, Dolores, Eagle, Garfield, Mesa, Moffat, Montezuma, Montrose, Ee Rio Blanco, and San Miguel), 4500-6500(-8200) ft. Apr-Jun. W. ACKNOWLEDGMENTS 1 1 : (TZ ANTT T) NM “2 We would like to thank the University of California herbarium (UC), University of K and the Field Museum (F) for the loan of Abronia specimens, and the University of Colorado Herbarium (COLO) for access to their specimens and data. Specimens from the Colorado State al Herbarium (CS) were also examined for this study. We would also like to thank Richard Spellenl an anonymous reviewer for comments on the manuscript. REFERENCES ArwooD, N.D., S.L. WasH, and K.D. Heit. 2002. A new species of Abronia (Nyctaginaceae) from San Juan County, New Mexico, U.S.A. Novon 12:167-169 FLINT, W.F. 1878. Juniperus occidentalis in Colorado. Bot. Gazette 3(10):87. GALLOWAY, L.A. 1975. Systematics of the North American desert species of Abronia and Tripterocalyx (Nyctag- inaceae). Brittonia 27: 328-347 GALLOWAY, L.A. 2003. Abronia. In: Flora of North America North of Mexico Vol. 4. Oxford University Press. Pp. HOLZINGER, J.M. 1892. |. List of plants collected by C.S. Sheldon and M.A. Carleton in the Indian Territory in 1891. Contr. U.S. Natl. Herb. 1(6):189-213. STANDLEY, PC. 1909. The Allioniaceae of the United States, with notes on Mexican species. Contr. U.S. Natl. Herb. 12:303-389. Turner, B.L. 2004. Biological status of the names Abronia carletonii and A. nealleyi (Nyctaginaceae). Phytologia 86:42-45 WEBER, WA and R.C. Wittmann. 2001. Colorado flora: eastern slope. University Press of Colorado. WELSH, S.L. 2003. Abronia. In: A Utah flora. Print Services, Brigham Young University. Pp. 446-448. WELSH, S.L. and S. GoobricH. 1980. Miscellaneous plant novelties from Alaska, Nevada, and Utah. Great Basin Naturalist 40:78-80. Young, R.K. 1997. The Ute Indians of Colorado in the Twentieth Century. University of Oklahoma Press, Norman. 424 BOOK REVIEW NIGEL SMITH, RODOLFO VASQUEZ, AND WALTER H. Wusr. 2007. Amazon River Fruits: Flavors for Conserva- tion. (ISBN 978-9972-2974-2-7, pbk.). Amazon Conservation Association, Missouri Botanical Garden Press, PO. Box 299, St. Louis, Missouri 63166-0299, U.S.A. (Orders: www.mbgpress.org, mbgpress@ mobot.org, 1-314-577-9534, 1-314-577-9594 fax). $35.00, 272 pp., color figures and photographs, 9.5" x 8". Introduction, Anacardiaceae, Annonaceae, Apocynaceae, Arecaceae, Celastraceae, Chrysobal- , Euphorbiaceae, Fabaceae, Gnetaceae, Lecythidaceae, A AT 1 J Contents.—Preface, anaceae, Clusiaceae, ae Convolvulaceae, Costaceae, Dicha Malpighiaceae, Malvaceae, Mel , Memecylac eae, M , Moraceae, Muntingiaceae, Myrsinaceae, Myrtaceae, Ola- caceae, Passifloraceae, Polygalaceae, Polygonaceae, Rubiaceae, Sapindaceae, Sapotaceae, Simaroubaceae, Solanaceae, Theophrastaceae, Urticaceae, Verbenaceae, Bibliography Notes, References, Photo cre This bool | it lers by ding the attenti f all thei uni 1.4 ]b in cover o wrapped in a mixed matte and glossy cover weighs nicely in the hand, while photographs depicting col its an iE eid: ` Ss TS 41 ER 1. 1 a 1 a E gee, ee 11 CMT E E “1 1 a 1 an ii + EE IJ Ada AAA E > : £i H H f ALLA”, 3 ) k habitats face. DI J L 3l ] Over 100 i j Peru, representing only a fraction of the Seere eee species of wild and domesticated Gei consumed by people in this region. | follow a standard format throughout the book, with Species accounts are organized by the | y to which they belong and the scientific names listed at the ur of the page and common name(s) listed below, organized by the country to which they pertain 4 +l 1 1 4.1 LaTi ] J 1 4 1 ] «1 E ] f 41 EA E E D + r o text. F AS | Jé f+} 11 Pal + Jee E J; hirti } ] lli ti and disper- J DH D Ir Ge Eé ) I sal, 1] jm Es "a L a J: ] «1 1 1 ] 1; Ff, » | 1 E srt from those eaten only locally ( e 8- Psychotria EE eek a huayo”), to tl ported to local markets (e.g., Passiflora nitida, Passifloraceae, I ing the int ional kets (e.g., Theobroma cacao, chocolate) E E t4 +L, + {+l z + 11 sën 4 J l ] 4 1 ‘1.1 1 : 1 Tes i + 1 e : til à " 1 TA E M 1 The authors cater to a eh naturalist audience that is interested in learning about Amazonian LA and conservation eck a ien dd Modus EEN one Re = ES? easily HRS to as Sua walk through a P I es ac- " ] 1 L +1 1 1 el ina e more compl lerstanding of both tl ironmenta ] and E landscape in Amazonia Tiana F. Franklin, M.S. tomen Science, Botanical Data Specialist, Botanical R h Institute of Texas, 500 East 4th Street, Fort Worth, Texas 76102-4025, U.S. J, Bot. Res. Inst. Texas 2(1): 424. 2008 FURTHER TRANSFERS OF GLANDULAR-PUBESCENT SPECIES FROM CHENOPODIUM SUBG. AMBROSIA TO DYSPHANIA (CHENOPODIACEAE) Sergei L. Mosyakin Steven E. Clemants M G. Kholodny Institute of Botany Brooklyn Botanic Garden nal Academy of Sciences of TM 1000 Washington Ave. 2 EISE Street, Kiev, 01601 UKRAINE Brooklyn, New York 11215 U.S.A. ABSTRACT I (T J.M Black) M kin & Cl t m silla (Hook f.) Mosyal in & Cl ts, D. truncata ae iei OE Ane & Clemants, D. cha (PG. Wilson) Mosyakin & Cl ts, D. andicola (Phil ) Mosyakin & CI ts, D.H & Clemants, D. dunosa (L.E. Simón) Mosyakin & CI ts, D. obl lata (S )M kin & Cl ts, D. retusa ane ance) M ki Cl (Aellen) M kin & Cl ts, D. tomentosa (Th JM 1 EE D M Jas C147] t ,D 1 = rH I DM syal re VE data | T D pl il.) Mo 1-3 £ El + D nepalensis (Colla) Mosyakin & Bee D. procera (Hochst. ex Moq.) Mosy: akin & Clemants, D. dissects (Moq) Mosyakin & Clemants, D. mandonii (S. Watson) Mosyakin & Clemants, D. stellata (Standley) Mosyakin & Clemants, D. congolana (Hauman) Mosyakin & Clemants, and D. minuata (Aellen) Mosyakin & Clemants Key Worps: Chenopodiaceae, Chenopodium, Dysphania, nomenclature, new combinations, taxonomy RESUMEN Se proporen | iguient bi i Dvsphani 2] i i i ] CI li I g. Ambrosia: r oO Paes Y r Lei D i I (T (IM Black) M kin & Cl ts D pusilla nM le fM y alin €y Cl ts, D. truncata PG Wilson) vali in & Clemants, D. saxatilis (EE: Wilson) Mosyakin & iani, D. andicola (Phil) Mosyakin & Clemants, D. burkartii DE Mosyakin & Clemants, D. dunosa (L.E. Simón) E ee & Gees D. oblanceolata EE FI & piano) D. D retusa (Juss. ex Moq.) Mosyakin & Cl ts, D (A y ants, D. venturii (Aellen) Mosyakin & Clemants, D. I i (Hook. £) Mosyakin & Clemants, D. microcarpa (Phil) Mosvakin & Clemants, D. nepalensis (Colla) Mosyakin & Clemants, D. procera uw ex "i Ph akin & Md D. dissecta (Moq.) Mosyakin & Clemants, D. mandonii (S. Watson) Mosyakin & Cl t y) y x Cl , D. congolana (Hauman) Mosyakin & Clemants, y D. minuata (Aellen) Mosyakin & Clemanes INIT RODUCI MUN The genus Chenopodium L. (Chenopodiaceae) in its traditional wide circumscription has long been sus- pected to be an unnatural (polyphyletic or paraphyletic) union of taxa belonging to different phylogenetic lineages. Consequently, many segregate genera and infrageneric taxa of various ranks were described and accepted within this group 9r various authors (see an overview in Scott, 1978). The genus Dysphania R. Br. during its turbulent t an latural history also experienced numerous shifts: from a single genus of a separate family Dysphaniaceae (Pax & Hoffmann 1934) or a representative of Illecebraceae or Caryophyllaceae— to a mere section of Chenopodium s.l. (Aellen 19302, 1930b; see also di ion in: Eckardt 1967; Mabry & Behnke 1976; Scott 1978; Wilson 1983, 1984; Mosyakin & Clemants 2002, and references therein). The present article continues a series of our publications (Mosyakin 1993; Mosyakin & Clemants 1996, 2002; Clemants & Mosyakin 2003) on Chenopodium Ge s.l. ana its Ne Earlier (Mosyakin & Clemants 2002) we justifie di the t y placed in Chenopodium L. subg. Ambrosia A.J. Scott, commonly called Rus HEHGOIDEER ed to as glandular chenopods, to the genus Dysphania, and validated new t | preparat ion for t] t of the genus in Flora of North America north of Mexico (Clemants & Mosyakin 2003). At that time we made 18 new combina- tions, including 11 at the species level, 4 at the section level, and 3 at the level of subsection. Nomenclatural changes were essentially limited to taxa occurring in North America (both native and introduced), leaving a number of glandular chenopods yet to be formally transferred to Dysphania. J. Bot. Res. inst. Texas 2(1): 425 — 431. 2008 426 t tani i Texas 2( The t j ] latural decisions t landular cl 1s withi expanded Dysphania were further discussed in the Dr. Sci. dissertation of Mosy akin 2003). The genus Dysphania circumscribed so as to include all glandular taxa of Chenopodium has been accepted for the — of Chenopodiaceae in Flora of North America north of Mexico (Clemants & Mosyakin 2003) and Flora of China (Zhu Gelin, Mos- yakin & Clemants 2003). At least one additional transfer of a species from Chenopodium to Dysphania (D. pdseudomul inis (Murr) EE Sr Lambinon) has been made (Verloove & Lambinon 2006) since then. Here we validate new tions in Dysphania for other species not transferred to that genus earlier. We refrain here from mal ific conclusion on phylogenetic relationships within Dysphania. However, judging from a thorough dni os of relevant literature, our herbarium and field experience, studies of mor- phology and biogeography of glandular chenopods, we firmly believe that Dysphania in the new circum- scription is a natural phylogenetic unit distinct from Chenopodium s.s. Since 2002 it was partly confirmed by molecular phylogenetic studies. DISCUSSION 1 1 £13; mcm A some extent a deep pl phy ] t] it between at least three Results of recent branches (clades) containing taxa py Hu pae in Chenopodium s.]. (Kadereit et al. 2003, 2005). Species of Chenopodi |. appeared scattere dint g ee EES TE and III. The segregation of bet tl of Chenopodium s.l. T Chenopodium, Blitum (L.) I. Hiitonen, ant mo AJ Scott Scott, 1978). In particular, Kadereit et al. (2003, p. 976) indicated: “In Chenopodieae III, a close e of the Eurasian species Chenopodium botrys and Teloxys aristata, and of the Australian species Chenopodium cristatum and Dysphania glomulifera, is supported by the presence of multicelluar glandular hairs (type 8 in Carolin 1983). By that character, they fit into subgenus Ambrosia (Scott 1978; Simón 1996). <...> These results confirm the proposal of Mosyakin and Clemants (2002) to transfer Chenopodium subg. Ambrina [sic! —S. M. & S. C] to Dysphania <...>.” In a study of the origin and age of Australian Chenopodiaceae Kadereit et al. (2005, p. 74) indicated that "[rlecently, Mosyakin and Clemants (2002) used morphological characters to separate subg. Ambrosia from Chenopodium, and included it in Dysphania. This last view is strongly sup- ported by our results.” Other options alternative to our wide circumscription of Dysphania would be to recognize several gen- era, for example, Teloxys Moa. (including only Chenopodium aristatum L. = T. aristata (L.) Moq., or also other related taxa), Roubieva Moq. (mainly an American group, which may be recognized either in the original sense of Moquin-Tandon (1834), or with addition of other American taxa), the Australian group previously placed in Chenopodium sect. Orthosporum R.Br., and Australian Dysphania in its traditional circumscription. These groups, corresponding mainly to sections in various classification schemes of Chenopodium s.l., seem to be atual Geer EEN of such groups as genera, at Bus at our peccat level of knowledge, has | to paraphyly or yet uncertain phy]! f many taxa. In our opinion, à slightly Minna circumscription of Dysphania (including related SE Gd Se E ge non-Australian taxa) ione be also inconvenient, because it will I diagnostic characters di ishing the resulting Broads! SE from other standhilat taxa. y we strongly prefer to place 1 " 1 + H 1 EE y] ez ———MÀ [e] +- 1 NBA EEN For avoiding such -— andi for maintaining tl all glandular chenopods in Dysphania, which is characterized by its SE ciones and also by other morphological and biochemical characters extensively discussed in literature (Scott 1978; Carolin 1983; Wilson 1983; 1984; Palomino et al. 1990; ad 1993; cedi p de Bonzani et al. 2003 etc.). ] = R 1 1 result ing from In the present article we validate 21 new the transfers of glandular-pubescent species to Dysphania. A moderately narrow —€— concept is applied, which is mainly consistent with prevalent taxonomic treatments of “glandular chenopods” and species of Chenopodium s.l. We admit that many species of the considered groups are in need of further revisions and more detailed studies, but that should not preclude making available new combinations in Dysphania for AA L2 JJ | A Al | 4.2 a n L H 427 d E f W them. In addition to our experience and herbarium studies of these taxa, we especially consulted with and relied upon the basic taxonomic publications by Aellen (19302, 1930b, 1933, 1973), Aellen and Just (1943), Wilson (1983, 1984, 1987), Simón (1995, 1996, 1997), as well as other relevant works (Voroshilov 1942; Scott 1978; Carolin 1983; Weber 1985; Palomino et al. 1990; Kühn 1993; Schwarzová 1993; Uotila 1997, 2001). VALIDATION OF THE NEW COMBINATIONS 1. Dysphania sect. Orthospora (R. Br.) Mosyakin & Clemants, Ukrayins'k. Bot. Zhurn. 59:382. 2002. Several names of putative hybrids between Australian taxa belonging to ape sect. EE were published (Aellen 1933, 1961; Wilson 1983, 1984). However, since their t and their hybrid origin is mostly in need of confirmation, we refrain from making here any new combina- tions for hybrids. Dysphania melanocarpa (J.M. Black) Mosyakin & Clemants, comb. nov. Basionym: Chenopodium carinatum R. Br. var. melanocarpum J.M. Black, Trans. & Proc. Royal Soc. South Australia 46:566. 1922. Chenopodium melanocarpum (J.M. Black) J.M. Black, Trans. & Proc. Royal Soc. South Australia 58:173. 1934 Dysphania pusilla oe f.) Mosyakin & Clemants, comb. nov. B Chenopodium pusillum Hook. £., Handb. New Zealand Fl. 1:231. Dysphania truncata (PG. Wilson) Mosyakin & Clemants, comb. nov. Basionym: Chenopodium truncatum PG. Wilson, Nuytsia 4:177. 1983 Dysphania saxatilis (PG. Wilson) Mosyakin & Clemants, comb. nov. Basionya: Chenopodium saxatile PG. Wilson, Nuytsia 4:179. 1983 The last species was described by Wilson (1983) from Western Australia as C. saxatile. The author of the species noted that “Isluperficially this species would appear to be related to the 4-merous species in the genus Dysphania (e.g. D. rhadinostachya) but in seed character it is obviously different, being very similar to Chenopodium pumilio and its allies of the section Orthospora. Yet even here its placement is uncertain since the other species of this section have flowers arranged in compact glomerules, have one stamen, and normally a 5-merous perianth." (Wilson 1983, p. 180). Wilson also compared his species with members of the Chenopodium botrys group (which evidently differ from C. saxatilis in having 5-merous perianth and horizontal seeds) and section Margaritaria (which has whitish subglobose, as well as glandular, hairs, and some other distinctive characters, see Brenan 1956), and concluded that *it appears that C. saxatile does not fit satisfactory into any of the presently recognized groups." P.G. Wilson refrained from describing a new monotypic section and instead expanded the circumscription of Chenopodium sect. Orthosporum to include his species. Pending further research, we keep the species in Dysphania sect. Orthospora. 2. Dysphania sect. Adenois (Moq.) Mosyakin & Clemants, Ukrayins'k. Bot. Zhurn. 59:382. 2002. In our previous puplicanans (Mosyakiii & Clemants 2002; Clemants 2 ME 2003; Zhu Gelin et al. 2003) we mentioned that opinions vary widely regarding the proper st in this section, with some authors ae Es a few species, including ext ly polymorphic Chenopodium ambrosioides L. with infraspecific taxa, \ t] l (up to 12) ly ci il (Aellen 1929, 1961, 1973; Aellen & Just 1943; fotos diy 1942: Simón 1996, 1997; tom 1997). Gam ance the same author (notably Aellen) in different periods of his work was of differing opinions regarding the number of species related to C. ambrosioides. We believe that taxa in this group differ from each other not less than many generally recognized species of Chenopodium s.l. (subg. Chenopodium), especially in the C. album group. Numerous infraspecific entities were recognized in C. ambrosioides and some other variable taxa. In the present article we are concerned with the species-level taxonomy, and thus for the time being refrain from making any new infraspecific and infrageneric combinations. 428 | lof the Botanical R h Institute of Texas 2(1) Dysphania andicola (Phil.) Mosyakin & Clemants, comb. nov. Basin: Ambrina andicola Phil., Anales Univ. Chile, 1. Mem. Ci, Mem. Ci. Lit. 91:422. 1895. Chenopodium ambrosioides var. andicola (Phil.) Aellen, Feddes Repert. Spec. Nov. Regni Veg. 26:37. o Dysphania burkartii (Aellen) Mosyakin & ae comb. nov. Basionwm: Cl lium ambrosioides L. subsp. burkartii Aellen, Feddes Repert. Spec. Nov. Regni Veg. 26:37. 1 Ch tii (Aellen) Vorosch.. Bot. Zhum. S.S.S.R. 27(3/4):46. 194 Dysphania dunosa (L.E. Simón) Mosyakin & Clemants, comb. nov. Basionym: Chenopodium dunosum L.E. Simón, Bull Mus. Natl. Hist. Nat., B, Adansonia 10:407. 1988 [1989]. ee oblanceolata (Speg.) Mosyakin & Clemants, comb. nov. Basion: Chenopodium ambrosioides L. var. oblan- ceolatum Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 7:137. 1902. Chenopodium oblanceolatum (Speg.) Giusti in Correa, Fl. e (Colecc. Cient. 8) 4a:122. 1984. p retusa Ce ex END Mosyakin & Clemants, comb. nov. Basionvm: Chenopodium retusum Juss. ex Moq. in ., Prod. 13(2):73. 1 Dysphania sooana (Aellen) Mosyakin & Clemants, comb. nov. Bastonya: Cl li Aellen, Acta Bot. Acad Sci. Hung. 19(1/4):5. 1973 (as "sooianum"). Dysphania tomentosa (Thouars) Mosyakin & Clemants, comb. nov. Basionym: Chenopodium tomentosum Thouars, Fl. Tristan d'Acugna 38. 1811 "SE venturii (Aellen) Mosyakin & Clemants, oe nov. Buc sl ambrosioides L. subsp. venturii Aellen in Aellen & Just, Amer. Midl. Naturalist 30:51. 1943. C} h., Bot. Zhurn. S.S.S.R. 27(3/4):46. 1942 ie name: Russian description). Chenopodium venturii allen) Cabrera, Revista Invest. Agric. 11:384. 1958. 3. Dysphania sect. Roubieva (Moq.) Mosyakin & Clemants, Ukrayins'k. Bot. Zhurn. 59:382. 2002. Dysphania bonariensis Kee f. ) Mosyakin « » o Com. nov. Basionyet: Roubieva bonariensis Hook. f. in Benth. &H DI 3: man & Irigoyen, Anales Mus. Nac. Hist. Nat. Buenos Aires 32:90 1923, comb. illeg., non Ten. 1833. Chenopodium Gs di in Engl. & Prantl, Nat. Pflanzenfam. 16c:491. 1934. Dysphania microcarpa (Phil.) o & Clemants, comb. nov. Basionym: Roubieva microcarpa Phil., Anales Univ. Chile (Santiago), Mem. Cient. Liter. 91:423. 1 4. Dysphania sect. Botryoides (C.A. Mey.) Mosyakin & Clemants, Ukrayins'k. Bot. Zhurn. 59:383. 2002. 4a. Dysphania sect. Botryoides (C.A. Mey.) o Er Clemants subsect. Botrys (Aellen & Iljin) Mosyakin & Clemants, Ukrayins'k. Bot. Zhurn. 59:383. 2002. The name of this subsection within the genus Chenopodium is often incorrectly cited as a combination Che- nopodium subsect. Botrys (W.DJ. Koch) Aellen & Ijin. Since the name published by Koch is illegitimate, the above subsectional name should be attributed to Aellen and Iljin, who provided its description only in Russian, but also included a reference to the earlier pre-1935 publication and description by Koch, which makes their subsection validly published. Dysphania nepalensis (Colla) Mosyakin & Clemants, comb. nov. Bastion: Chenopodium nepalense Colla, Herb. Pede- mont. 5:571. 1836 Chenopodium multiflorum Moq. in DC., Prodr. 13(2):75. 1849. This little-known Central Asian lly resembles D. schraderiana in its habit, but differs in having weakly keeled perianth Se with ple non-glandular hairs at the keel (Uotila 1993, 1997). Another species closely related to D. schraderiana was ER transferred from Chenopodium to Dysphania as D. pseudomultiflora (Murr) Verloove & Lambinon (= C! foetidum Schrad. subsp. pseudomultiflorum Murr, C. schraderianum Schult. var. pseudomultiflorum (Mur) Allen, C. pseudomultiflorum (Murr) Uotila) (Verloove & Lambinon 2006). Uotila (2001) indicated that this species of South African origin differs from AA n.º F £l D Ne | Lt H P L H 429 C. schraderianum in its more branched habit, leaves with more lobes and teeth, keeled but not cristate tepals with subsessile glands and long hairs, and smaller seeds. Dysphania procera (Hochst. ex Moq.) Mosyakin & Clemants, comb. nov. Basionym: Chenopodium procerum Hochst. Mog. in DC. Prod. 13(2):75. 1849. 4b. Dysphania sect. Botryoides (C.A. Mey.) Mosyakin & Clemants subsect. Incisa (Standley) Mosyakin & Clemants, Ukrayins'k. Bot. Zhurn. 59:383. 2002. Dysphania dissecta (Moq.) Mosyakin Y Clemants, comb. nov. Basionym: Ambrina di ta Moq., Chenop Monogr. Enum 40. Chenopodium dissectum (Moq.) Standley, N. Amer. Fl. 21:26. 1916. Dysphania mandonii (S. Watson) Mosyakin & Clemants, comb. nov. Basiowvw: Teloxys mandonii S. Watson, Proc. Amer. Acad. Arts. Sci. 9:91. 1874 (as “mandoni”). Chenopodium mandonii (S. Watson) Aellen, Feddes Repert. Spec. Nov. Regni Veg. 26:169. 1929. Chenopodium incisum Poir. var. bangii Murr, Bull. Herb. Boissier 4:991. 1904. Chenopodium graveolens Willd. var. bangii (Murr) Aellen, Verh. Naturf. Ges. Basel 41:107. 1931. 5. Taxa yet unplaced in sections of Dysphania epe stellata (Standley) Mosyakin & Clemants, comb. nov. BASIONYN: o stellatum S. Watson, Proc. Amer. cad. Arts Sci. 18:146. 1883. Meiomeria stellata (S. Watson) Standley, N. Amer. Fl. 21:7 This morphologically deviant pee es E: = ponent segments (described from Coahuila, Mexico) was placed by Standley (1916) in a separate ley, and then transferred to Chenopodium sect. Meiomeria (Standley) AJ. Scott (Scott 1978). Relationships of this little-known taxon with other representa- tives of Dysphania remain insufficiently known. Dysphania congolana (Hauman) Mosyakin & Clemants, comb. nov. Basionru: Chenopodium glaucum L. var. congola- num Hauman, Bull. Jard. Bot. État. Bruxelles 19:447. 1949. Chenopodium congolanum (Hauman) Brenan, Kew Bull. 11(1956):166. 1956 "epe minuata See SEH & Clemants, comb. nov. Basionym: Chenopodium minuatum Aellen, Acta Bot. cad. Sci. Hung. 19(1/4):8. 1 Dysphania congolana, an African species, was placed by Brenan (1956) in Chenopodium sect. Margaritaria Brenan, as C. congolanum; later Simón (1996) added D. minuata to this section (as C. minuatum). ACKNOWLEDGMENTS This article and our previous publications on Dysphania were initiated during our work on treatments of Chenopodium and Dysphania for the Flora of North America (FNA) and Flora of China (FC) projects, and during several visits of the first author to the Missouri Botanical Garden, St. Louis (MO) and the Brooklyn Botanic Garden, NY (BKL). The authors are grateful to ln SEH of nes institutions and herbaria for their kind help and cooperation. Special thanks are due to l n the FNA and FC projects (listed alphabetically): Ihsan Al-Shehbaz (MO), Peter Hoch (MO). Nancy Morin A Point Arena, California), Peter Raven (MO), James Solomon (MO), Nicholas Turland (MO), and James Zarucci (MO), who facilitated our research. We are also grateful to Thomas Borsch (Bonn, Germany), Helmut Freitag (Kassel, Germany), Gudrun Kadereit (Mainz, Germany), Aleksandr Sukhorukov (Moscow, Russia), Zoya Tsymbalyuk (Kiev, Ukraine), Pertti Uotila (Helsinki, Finland), and other colleagues with whom we discussed various issues related to taxonomy, morphology and geography of Chenopodiaceae. Thanks are due to Barney Lipscomb, Donald Pratt, and two anonymous reviewers for their valuable comments on the manuscript. REFERENCES AELLEN, P. 1930a. Die systematische Stellung und Gliederung der R. Brownschen Gattung Dysphania. Bot. Jahrb. Syst. 63:483-490. 430 | | of tl tanical Insti Texas 2( AELLEN, P. 1930b. Eine neue Sektion der Gattung Chenopodium (Sect. Tetrasepaia). Bot. Jahrb. Syst. 63:490—492. ALLEN, P. 1933. Die Arten der Sect. Orthosporum der Gattung Chenopodium L. Verh. Naturforsch. Ges. Basel. 44:308-318. AcLLEN, P. 1961. Chenopodiaceae. In: G. Hegi. Illustrierte Flora Mitteleuropa. Bd. 3. T. 2. Aufl. 2. Lehmann Verlag, München (Reprinted in 1979: Paul Parey Verlag, Berlin & Hamburg). Pp. 533-747. AcLLEN, P. 1973. Formenkreis von Chenopodium L. Sect. Ambrina (Spach) Benth. & Hook. und Sect. Nigrescentia Aellen. Acta Bot. Acad. Sci. Hung. 19(1—4):1-12. AELLEN, P. and T. Just. 1943. Key and synopsis of the American species of the genus Chenopodium L. Amer. Midl. Naturalist 30(1):47—76. Bonzani, N.E., G.E. Bareoza, M.A. Bucarri, and L. Ariza Espinar. 2003. Morpho-histological studies in the aromatic species of Chenopodium from Argentina. Fitoterapia 74:207-225. Brenan, J.PM. 1956. A new section of the genus Chenopodium from Africa. Kew Bull. [Volume of 1956] 11:165-167. Caroun, R.C. 1983. The trichomes of the Chenopodiaceae and Amaranthaceae. Bot. Jahrb. Syst. 103:451-466. CLEMANTS, S.E. and S.L. MosvakiN. 2003. Dysphania R. Brown; Chenopodium Linnaeus. In: Flora of North America Editorial Committee, eds. Flora of North America north of Mexico. Vol, 4. Oxford University Press, New York & Oxford. Pp. 267-299. Eckanpr, T. 1967. Vergleich von Dysphania mit Chenopodium und mit Illecebraceae. Bauhinia 3:32/-344. Gust, L. 1997. Chenopodiaceae. In: Flora fanerogamica Argentina. Fasciculo 40. Proflora, Conicet, Cordoba. Kapererr, G., T. Borsch, K. WeisinG, and H. Freirac. 2003. Phylogeny of Amaranthaceae and Chenopodiaceae and the evolution of C,-photosynthesis. Intl. J. Pl. Sci. 164:959-986. Kaap, G., D. Gorzex, S. Jacoss, and H. Freitac. 2005. Origin and age of Australian Chenopodiaceae. Organisms, Diversity & Evol. 5:59-80. KOHN, U. (WITH ADDITIONS av V. BITTRICH, R. CAROLIN, H. FerrAc, LC Hence, P. UoriLa, and PG. WiLson). 1993. Chenopodiaceae. In: K. Kubitzki, ed. The Families and Genera of Vascular Plants. Vol. 2. SEN verag PA Pp. 253-281. Masry, TJ. and H.-D. Behnke. 1976. Betalains and P-type sieve-element plasti y ti of Dysphania R. Br. (Centrospermae). Taxon 25:109-111. Moauin-TANDON, A. 1834. Descriptions de plusieurs nouveaux genres de Chenopodées. Ann. Sci. Nat. Bot. 1:289-294. MosvAKIN, S.L. 1993. An outline of a system for Chenopodium L. (species of Europe, North and Central Asia). Ukrayins'k. Bot. Zhurn. (Ukr. Bot. J) 50:71-77. Mosvakin, S. L. 2003. Taxonomy, phytogeography and genesis of the family Chenopodiaceae Vent. Dr. Sc. Dis- sertation. M. G. Kholodny Institute of Botany of the National Academy of Sciences of Ukraine, Kyiv (Kiev). [In Ukrainian, with English and Russian summaries.] MosvAKIN, S.L. and S.E. CLEMANTS. 1996. New infrageneric taxa and combination in Chenopodium L. (Chenopo- diaceae). Novon 6:398-403. MOSYAKIN, S.L. and S.E. CLEMANTS. 2002. New nomenclatural combinations in Dysphania R. Br. (Chenopodiaceae): taxa occurring in North America. Ukrayins'k. Bot. Zhurn. (Ukr. Bot. J.) 59:380-385. PALOMINO, G., M. SEGURA, R. Bye, and P. Mercano. 1990. Cytogenetic distinction between Teloxys and Chenopodium (Chenopodiaceae). SouthW. Naturalist 35:351—353. Pax, E A. and K. Horrmann. 1934. Dysphaniaceae. In: A. Engler, H. Harms. Die natürlichen Pflanzenfamilien. 2 Aufl. Vol. 16c. Berlin: Duncker & Hunblot. Pp. 272-274. Schwarzová, T. 1993. Beitrag zur karyotaxonomie und evolution mancher Arten der Gattung Chenopodium L. (Sektion Botryoides und Ambrina). Biologia (Bratislava) 48:421—428 Scott, AJ. 1978. A review of the classification of Chenopodium L. and related genera (Chenopodiaceae). Bot. Jahrb. Syst. 100:205-220. Simón, L.E. 1995. Essai sur l'histoire paleobiogéographique du genre Chenopodium L. sous-genre Ambrosia A.J. Scott. Biogéographica 71:127-142. AA | m Jl en A Al | AB H n L H 431 d zd £ P SIMÓN, L.E. 1996. Notas sobre Chenopodium L. subgen. Ambrosia AJ. Scott (Chenopodiaceae). 1. Taxonomía. 2. Fitogeografía: áreas disjuntas. Anales Jard. Bot. Madrid 54:137—148. SIMÓN, L.E. 1997. Variations des charactéres foliares chez Chenopodium subgen. Ambrosia sect. Adenois (Cheno- podiaceae) en Amérique du Sud: valeur taxonomique et évolutive, Adansonia 19:293-320 STANDLEY, PC. 1916. Chenopodiaceae. In: N. Amer. Fl. 21(1):1-93. New York. Vona, P. 1993. Taxonomic and nomenclatural notes on Chenopodium in the Flora Iranica area. Ann. Bot. Fennici 30:189-194. Lora, P. 1997, Chenopodium. In: LC Hedge et al. 1997. Flora Iranica: Flora des Iranischen Hochlandes und der Umrahmenden Gebirge. Persien, Afghanistan, Teile von West-Pakistan, Nord Iraq, Azerbaidjan, Turkmenistan, ed. KH. Rechinger. No. 172. Chenopodiaceae. Akademische Druck- u. Verlagsanstalt, Graz. Pp. 24-59, (ona, P. 2001. Taxonomic and nomenclatural notes on Chenopodium (Chenopodiaceae) for Flora Nordica. Ann. Bot. Fennici 38:95-97. VERLOOVE F. and J. Lameixon. 2006. The non-native vascular flora of Belgium: a new nothospecies and three new combinations. Syst. Geogr. Pl, 76:217—220. VonosHiLOv V.N. [WonoscHiLov; VoroscHiLov] 1942. Overview of species of Chenopodium L. sect. Ambrina (Spach) Hook. fil. Bot. Zhurn. S.S.S.R. 27(3/4):33-47. [In Russian] WEBER WA 1985. The genus Teloxys (Chenopodiaceae). Phytologia 58:477-478. WILSON, E.G. 1983. A taxonomic revision of the tribe Chenopodieae (Chenopodiaceae) in Australia. Nuytsia 4:135—262. WILSON, PG. 1984. Chenopodiaceae. In: A.S. George, ed. Flora of Australia. Vol. 4. Australian Government Publish- ing Service, Canberra. Pp. 81-317 WiLson, PG. 1987, Generic status in the Chenopodiaceae. Australian Syst. Bot. Soc. Newsl. 53:78-85. ZHU GELIN (CHU GE-LING), S.L. MosvAKIN, and SE CLemants. 2003. Chenopodiaceae. In: Wu Zhengyi and PH. Raven, eds. Flora of China. Vol. 5 (Ulmaceae through Basellaceae ). Science Press, Beijing & Missouri Botanical Garden Press, St. Louis. Pp. 351-414, 432 | tani i Texas 2( BOOK REVIEW ROBERT W. LEBLING AND DONNA PEPPERDINE. 2006. Natural Remedies of Arabia. (ISBN 9781905299027, hbk.). Stacey International Publishers, 128 Kensington Church Street, London, W8 4BH, United Kingdom. (Orders: www. stacey-international.co.uk, info@stacey-international.co.uk, +44 207 221 7166, +44 207 792 9288 fax). $55.00, 210 pp., color T photographs glossary, indexes, reference list, 10 3/4" x 12 1/2". This | i Ith of information on th gnosy of the ens of Panai T iun Yemen, Oman, and E United Arab Emirates. It d ibes in detail 62 herbs, S ies: oils, ] er substance ith factoids, historical nuggets, traditional uses, academi h notes, and family TOES In particular, the family remedy p f h pl ] df q listributed throughout ian Peninsula in 2002. The question- naire.. nis own to explain how E .use various herbs, spi d otl I in natural healing.” a additional t ] Lun | E > plant wa The substances described in detail are: aloe vera, alum, anise, arak, arta, arugula, asafetida, banana, basil, black seed, caraway, cardamom, castor oil plant, chamomile, cinnamon, cloves, coffee, coriander, cress, cucumber, cumin, dates, fennel, fenugreek, frank- incense, garlic, germander, ginger, hasawi rice, henna, herbal waters, hibiscus, honey, incense, lavender, lemon balm, lime, mahaleb cherry, mangrove, mastic, mint, myrrh, nakhwa, neem, nettle, petroleum, pomegranate, purslane, rose, saffron, sage, samh, sarcocol, rE 4 sidr, tarthuth, thyme, desert truffles, turmeric, walnut de Minis (sheeh), wormwood (bu'aythiran), and yarrow. The following , apple of Sodom, black nig , blond psyllium, caper, caralluma, cleome, colocynth, corn poppy, desert bugloss, dogbane, oe neum gum iun harmel, aon jericho rose, lemon grass, milkweed, myrtle, rat-tail plantain, safflower, senna, simp per, smooth sow Bee spurge, iere ane Weieen 7 1: 1 EI E «1 1 | J x H AT A4:3.3.11 T2 ALILLA LA DRA de + Le » ing natu skin care, and cosmetics, sella as uses z oe and mekab Geen This section is presented in bullet — SCH -—À cumcu GE A Recipes section I ^i ov e L is also included, with 29 pages ofi j for the preparati f traditional drinl alada: soup, jam, dips, breakfast, desserts, main j e - ds a 1 = El A 1 1; E WË 1 e i + The bool lud ith the followi j Glossat of Herbal EA, Tum Index of SEH Ailments, Bibliography & NETA ae by Latin Name, and Index by Dd iod The Inde ] handy quic AT. 4 1p J: A 1.3 SU tz, ct £..1 1 E d ses 1 ] y) 1 1 f. 1: ftr «41 Allin all, d g I y own. < :1 H 1 f ¿1 13 T “111 dias l Will UK otal fascinating duelo; Mi Brooke Stegen. Botanical estar Institute of Te Texas, 500 East 4th Street, Fort Worth, Texas 76102- 4025, U.S.A. J. Bot. Res. Inst. Texas 2(1): 432. 2008 NEW COMBINATIONS IN THE PANARCTIC VASCULAR PLANT FLORA Reidar Elven David F. Murray National Center of Biosystematics University of Alaska Museum of the North Natural History Museum, University of Oslo 907 Yukon Driv POBox 1172 Blindern, NO-0318 Oslo, Norway Fairbanks, Alaska 99775-6960, U.S.A. reidar.elvengnhm.uio.no fidímguaf.edu ABSTRACT Thi rty eigh binati le in the families Asteraceae , Brassicaceae, Caryo[ I yll , Equisetaceae, Ericaceae, Fabaceae, Gentianaceae, NN Papaveraceae, Poaceae, Poly "gonaceae, P get Oe Rosaceae, g The RESUMEN Se] j ] binaci las familias Asteraceae , Brassicaceae, Caryophyll , Equisetaceae, Ericaceae, Faba- ceae, Gentianaceae, Orchidaceae, Papaveraceae, Poaceae, Polygonaceae, Pot , Ranunculaceae, Rosaceae, y Saxifragaceae. 1-15 is Ti Eu L L E Eu A Panarctic Flora id oc See has been in progress since 1999 (see Nordal & Razzhivin 1999) ly among botanists tsin Canada, Iceland, Norway, Russia, and U.S.A. Its main aim is to produce a checklist of all arctic vascular plants with critical notes on distribution, chromosome numbers, and with proposes for an bas names anaes on consensus or d eae anne BEE To reach a g y ap plied in Russia, North America, and d | Europe Alignment has meant new bi iri aai: The authors of new combinations are I. Alsos (Longyearbyen, Norway), R. Elven (Oslo, Norway) A.A. Korobkov (St. Petersburg, Russia), D.F. Murray (Fairbanks, U.S.A.), V.V. Petrovsky (St. Petersburg, Rus- sia), O. Rebristaya (St. Petersburg, Russia), P. Schónswetter (Vienna, Austria), H. Solstad (Oslo, Norway), N.N. Tzvelev (St. Petersburg, Russia), and B.A. Yurtsev (St. Petersburg, Russia, deceased). a rense I subsp tre (Wahlenb.) Schónswetter & Elven, comb. et stat. nov. Basionym: Equisetum arvense L. var. alpestre EN El A 296. 1812. Tee N SCANDINAVIA (“Lapland”). Type not traced (Moberg & Nilsson 1991). Equisetum boreale auct., non Bong., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 2:174. 1833. Equisetum arvense L. var. boreale auct., non (Bong.) Rupr., Hist. Stirp. Fl. Petrop. 91. 1845. Equisetum arvense L. subsp. boreale auct., non (Bong.) A. Lóve, Náttúrufredingurinn 18:101. 1948 An arctic—alpine race of Equisetum arvense L. has in recent years been accepted as subsp. boreale (Bong) A. Love (e.g., by Ollgaard 2000). Schónswetter et al. (2001) argued that the name “boreale” is Eege for the race in question. It is based on a Bongard plant from Sitka in soutl Alaska req morph within subsp. arvense, differing from the arctic and high alpine plants in many morphological aiies The next available name is Wahlenberg's var. alpestre, and a combination as subspecies is proposed here. Potamogetonaceae Stuckenia ait caus (Pers.) Borner subsp. borealis (Raf.) Tave len & me. can nov. BASIONYM: Potamogeton borealis Raf., Med. Repos., ser. 2, 5:354. 1808. Tre: CANADA: Q : St. Lawrence PLE geton filij Pers. var. borealis (Raf) H. St. John, Rhodora 18:134. 1916. Stuchenia filiformis (Pers.) Börner subsp. alpina auct., non (Blytt) R.R. Haynes, Les & M. Král, Novon 8:241. 1998. Northern North American plants of Stuckenia filiformis (Pers.) Börner differ from the main race by their broader leaves and more congested inflorescences. Such plants have been considered by authors (e.g., Hul- J. Bot. Res, Inst. Texas 2(1): 433 — 446. 2008 434 tani titute of Texas 2( tén 1968a; Cody 1996, 2000) as subsp. or var. borealis, based on Rafinesque's Potamogeton borealis from the St. Lawrence River region. Les and Haynes (1996) synonymized Potamogeton borealis with the Norwegian Potamogeton marinus L. f. alpinus Blytt, Norges Fl. 1:370 (1861), described from southern Norway, and they raised f. alpina to subspecific rank as the accepted name for the North American subspecies. Haynes et al. (1998) transferred Potamogeton filiformis to the genus Stuckenia, which we accept. Haynes and Hellquist (1998) reported subsp. alpina as restricted to North America and perhaps northeastern Asia, in spite of it being described from Norway. Blytt's f. alpinus has not been lectotypified. The syntypes (O!) from the six sites listed in the protologue are all from southern Norwegian mountain valleys. They have not been studied by Les, Haynes or Hellquist. Les and Haynes (1996: 390) stated, when they synonymized Norwegian ‘alpina’ and North American ‘borealis’, with PROS. for the former, that they had not seen type material (P) of ‘borealis’. Neither have we, but when i y of Blytt's Norwegian f arma 5 Mol Moni American paian we Ge Hen different. We consider the N lant tane f European S as an EE race peda, whereas we consider the North American plant an aceptable race. We the North American name "borealis" for the northern North American plants. This taxon is characterised by two differential characters as given by Haynes and Hellquist (for subsp. filiformis vs. subsp. alpina, slightly modified here): leaves 0.2-0.4 mm vs 0.5-0.8(-1.0) mm broad, and infrutescence strongly interrupted and extending up to 4 cm vs more compact to somewhat interrupted and extending , not only up to 2 cm. Orchidaceae In the Panarctic Flora Checklist we accept PIRE n ie and Lysiella Rydb. as genera different from Platan- thera Rich. We know of no thorough phyl y of the Platanthea group, a; ina nici of Orchidinae and some Hal j , Bateman et al (2003) found ma (Limnorchis) hyr ipl main group of Platanthera spp. Acceptance of Limnorchis necessitates two new ani bination Limnorchis aquilonis (Sheviak) Rebrist. & Elven, comb. nov. B Platanthera aquilonis Sheviak, Lindleyana 14:193 1999 Limnorchis huronensis (Nutt.) Rebrist. & Elven, comb. nov. Basionim: Orchis huronensis Nutt., Gen. N. Amer. Pl. 1:288. 1818. Platanthera huronensis (Nutt.) Lindl., Gen. Sp. Orchid. Pl. 288. 1835. Poaceae Calamagrostis purpurascens R. Br. subsp. laricina (Louis-Marie) Elven, comb. et stat. nov. Basionym: Cal- amagrostis purpurascens R. Br. var. laricina Louis-Marie, Rev. Oka Agron. Inst. Agric. 20:153. 1946. Tree: CANADA: Quzsec: Ungava, iviere aux Melezes (Larch R), 57?42'N, 68°25'W, Dutilly & Lepage 14602 (noLoryre: CAN). Calamagrostis laricina (Louis-Marie) Louis-Marie, Naturaliste Canad. 85:70. 1958. Calamagrostis poluninii T.J. Sorensen, Meddel. Grønland 136, 8:11. 1954. Tire: GREENLAND: Kiagtut in Tunugdliarfik, 15 Jul 1947, Sørensen 188 (HOLOTYPE: C). Greene (1980, 1984) recognized two races in the very widespread Calamagrostis purpurascens: var. purpura- scens from northern Siberia to northern Greenland (and recently found also on Svalbard) and var. laricina restricted to northeastern Canada and southern and western Greenland. The latter race is characterized by being short-rhizomatous and by having awns not distinctly exserted from the spikelet. Sorensen's descrip- tion of C. poluninii in 1954 was made without reference to (and probably while unaware of) Louis-Marie's earlier description of var. laricina in 1946. Greene seems to have been the first to compare the types and recognize that the two names apply to the same plant. The southern and western Greenland plants named C. poluninii are an extension of the northeastern North American C. purpurascens var. laricina. As this is a major geographical race, the rank of subspecies is preferable to variety, and a new combination is proposed. Koeleria pyramidata (Lam.) P Beauv. subsp. seminuda (Trautv.) Elven, comb. nov. Basion: Koeleria cristata Pers. var. seminuda Trautv., Trudy Imp. S.-Peterburgsk. Bot. Sada 5, 1 [PL Sib. Bor.]:138. 1877. Tyre: SIBERIA: northern Sakha (Yakutia), EI DRA Al Le “ zs Al ab "X. 435 "Ad Lena inf. prope pag. Ajakit,” 28 Jul 1875, Czekanovski (LecToTYPE: LE, designated by Tzvelev, Zlaki SSSR 276. 1976.). Koeleria serninuda (Trautv.) Gontsch. in Kom., Fl. URSS 2:331. 1934. Koeleria cristata Pers. subsp. seminuda (Trautv.) Tzvelev, Novosti Sist. Vyssh. Rast. 7:71. 1971 Koeleria cristata is an illegitime, superfluous name and must be replaced by K. pyramidata (Lam.) P. Beauv. Koeleria seminuda is here recombined as a subspecies within K. pyramidata. Puce iim Lap Ge Scribn. & Merr. subsp. neoarctica (A. Love & D. Lóve) Elven, comb. et DOV. BASIONYM: P Li neoarctica A. Love & D. Love, Bot. Not. 128:499. 1976. Tere: GREENLAND: Sydostbugt, Jul 1880, : C). Pu wis neoarctica (A. Love € D. Lóve) Bócher et al., Grønlands PL. ed. 3:293. 1978 [without basionym citation]. Puccinellia phry, ees (Trin.) Scribn. & Merr. “Greenland type” sensu Sørensen (1953). Within the polymorphic, circumpolar Puccinellia phryganodes, Sorensen (1953) informally named four enti- ties as “types”: a fertile, tetraploid “Beringian type” (2n = 28) and three asexual types: a triploid “Greenland type” also widespread in arctic North America (2n = 21), a tetraploid “Spitsbergen—Novaya Zemlya type” restricted to these islands, and a “Siberian type” ranging from northeastern Norway (triploid) Maie un arctic Russia and Siberia to ica (tetraploid). Serensen (1953) d trated anatomical and differences in addition to the ploidy differences. We prefer to consider Sgrensen's types major UM cal races, i.e., subspecies, of a morphologically coherent species phryganodes. Subsp. neoarctica is a sterile stoloniferous race that reproduces by detached shoots Puccinellia phryganodes (rin) Scribn. & Merr. subsp. sibirica (Hadač & Á. Löve) Elven, comb. nov. Ba- A Love & D. Lóve subsp. sibi č & A Lóve in A. Love & D. Lóve, Bot. Not. 114:36. 1961. Tyee: RUSSIA: “alge Sinus Warnek,” 14 Jul 1907, O S). Phippsia vil] (And JÁ Love & D Love subsp. sibirica (Hada? & A. Love) A. Love & D. Love, Bot. Not. 128: 501. 1976. Puccinelli les (Trin.) Scribn. & Merr. subsp. asiatica Tzvelev in Tolm., Fl. Arct. URSS 2:186. 1964 [in error for "sibirica"]. Puccinellia Gigs (Trin.) Scribn. & Merr. “Siberian type” sensu Sprensen (1953) Tzvelev (1964) made a mistake in the basionym, replacing “sibirica” with “asiatica” when publishing Puc- cinellia phryganodes subsp. asiatica. At the rank of subspecies, the epithet “sibirica” (Hadat & Love in Love & Lóve 1961) has priority over “asiatica” (Tzvelev 1964). Vahlodea latifolia (Hook.) Hultén subsp. paramushirensis (Kudó) Elven, comb. nov. Basionym: Deschampsia atropurpurea (Wahlenb.) Scheeie var. n P ia Kudé, J. Ee IL. i i Hokkaido Imp. Univ. 11:71. 1922. Tree?: ee Para- mushir 1, Urafutoyama, 1920, ioni 5787. Vi W b.) Fr Hulté Kamtchatka 1:111. 1927. Ve Erioblastus flexuosus Honda, J. Fac. Sci. Univ. Tokyo, sect. 3, Bot. 1:143. 1930. Described from Japan. Vahlodea flexuosa (Honda) Ohwi, Acta Phytotax. Geobot. 2:33. 1933 is (Kudô) Roshev. in Kom., Fl. URSS 2:242, 750. 1934. The small genus Vahlodea (a segregate of Deschampsia) is disjunct in three very distant regions: around the North Atlantic, the northern Pacific, and in southernmost South America. From one to four species or subspecies have been proposed. Tzvelev recognized, besides the South American V. magellanica (Hook. f.) Tzvelev, three northern species: V. Se M ii ia pd V. onem The amphi-Atlantic V. atropurpurea s. str. is fully allopatric with the two Pacifi ically from both of them. The most evident difference between the mainly ees Asian plant Y feos or V. POramusmténsis and the North American one (V. latifoli that the Asian plant has hai merican plant has glabrous leaves. They also ice in senal habit, but there are or intermediates, especially in southern Alaska. Soreng et al. (2003) recognized only one species with two subspecies: subsp. atropurpurea (including both V. latifolia and the South American V. magellanica) and subsp. paramushirensis. The solution chosen for the Panarctic Flora Checklist is two northern species: one amphi-Atlantic (V. atropurpurea) and one amphi-Pacific (V. latifolia), the latter with two subspecies (subsp. latifolia and subsp. paramushirensis) inasmuch as the entities intergrade. This necessitates the new combination. Polygona Bistorta Eer (Willd. ex Spreng.) V.V. Petrovsky, D.F Murray & Elven, comb. nov. Basionys: Polygonum ellipticum Willd. ex Spreng., Syst. Veg. 2:253. 1825. Described f Siberi bistorta L. subsp. ellipticum (Willd.) V.V. Petrovsky in Tolm., Fl. Arct. URSS 5:176. 1966. Bistorta major Gray subsp. elliptica (Willd. ) Soják, Preslia 46:152. 1 The widespread Bistorta officinalis Delarbre s. lat. is disjunct tane and arctic. The northern montane and arctic plants differ from non-arctic European Bistorta officinalis s. str. in several characters, and they should be accepted as different race(s) or species. Freeman and Hinds (2005) accepted the northwestern North American and northeastern Asian plant as B. plumosa (Small) Greene. Petrovsky (1966), treating the plants for arctic Russia as races of Polygonum bistorta, assigned the plants from northeastern Europe and northern Siberia (Taimyr) to subsp. bistorta (= subsp. officinalis) and those from northeastern Sil (Anabar T ne to the Chukchi Peninsula to subsp. ellipticum. He considered the North American si ibsp. pl sa as different from both of these. We agree with this division of the arctic Russian material but not ih the application of the names. Tzvelev and Yurtsev (pers. comm.) assumed the "elliptica" name to belong to the western entity. Polygonum ellipticum was described from non-arctic parts of the Lena River area. In the Arctic, the limit between the western and the eastern entities runs through the Taimyr, but further south in Siberia and the Russian Far East this otherwise western arctic entity occurs in the mountains all the way eastward to the Okhotsk Sea. We therefore recombine Polygonum ellipticum within Bistorta and accept it as the name for the northern Russian and northwest Siberian plant. We consider the northeast Siberian and northern Russian Far East plants as indistinguishable from the North American Bistorta plumosa, in spite of a suggested dif- ference in chromosome numbers. Caryophyllacea xus i ducis (Cham. & Schltdl.) Bocquet dus furcata (Raf.) VV. Petrovsky & Elven, comb. et stat. OV. Basionym: Silene furcata Raf., Autik. Bot. 28. 1840. Tyre: CANADA: Hudson pp area: type not located. Melandrium sen (Raf.) Hultén, Acta Univ. Lund., n. s., sect. 2, 40, 1:702. 1944. [printing date not given, but probably early 1944] or: (Raf.) Hadaé, Skr. Svalbard Nordishavet 87:34. 1944. [printed 04 May 1944]. S pes (Raf.) Hultén, Acta Univ. Lund., n.s., sect. 2, 40, 1:702. 1944. [comb. illegit] Silene involucrata (Cham. & RE Ue is a Lir aa species. We M it with s major races, a northern Europea subsp. tenella (Tolm ) Bocquet, D S 1 North American subsp. involucrata, and an arctic circumpol ar subsp. furcata. Bocquet (1967) included both the gara northern European and northwestern Siberian plants and the arctic circumpolar plants within Silene i ta sul involucrata, an approach followed by Kurtto (2001) and Morton (2005). i We find the races distinct at the level of subspecies. In three areas of co-occurrence (in the lower Lena River area of northern Siberia, in eastern Chukotka, and in northern Alaska—northwestern Canada) they are fully distinct in nature. Subspecies furcata then becomes the most widespread of the races. Silene furcata Raf. is not typified, but this race is the only one present in the Hudson Bay—Labrador area from where it was described. A subspecific combination based on this name is proposed. This was also the opinion of Tzvelev (2000). Silene soczavana (Schischk.) BO var. macrosperma (A.E. Porsild) VV. Petrovsky, D.E Murray & Elven, omb. e AE stat. nov. Basionym: M Porsild, Rhodora 41:225. 1939. Tree: ALASKA: Norton Sound, Unalakleet, 63°52'N, 106°20'W, A.E. See & RT Porsild 1147 eas CAN). Lychnis macrosperma (A P. a J.P Anderson, Iowa State Coll. J. Sci. 20:250. 1946 (A.E. Porsild) Hultén, Circumpolar PL 2:326. 1971. G i (A.E. Porsild) Tolm. & a in a Ge Ara URSS 6:115. 1971. Silene uralensis (Rupr.) Bocquet BEEN porsildii Senger Candollea 22:27. 1967. } sild Bocquet (1967) treated Porsild's Melandrium macrospermum as subsp. porsildii of the widespread Silene ura- «1 lensis, as a central Alaskan mountain endemic, whereas he accepted the as a separate species. The tetraploid chromosome number, morphology, ecology, and geography SUD inclusion of M. macrosperma within S. soczavana, not within S. uralensis. According to the material we have inspected, Silene soczavana var. macrosperma is a widespread taxon from eastern Chukotka and the north- ern Bering Sea islands northeast to the Alaskan North Slope and south and southeast to Kodiak Island and northern British Columbia. ri DRA M | E + Lea Al D dir Él 437 Silene villosula (Trautv.) V.V. Petrovsky € Elven, comb. nov. Basionvm: Lychnis ajanensis Regel var. villosula Trautv., Acta Horti Petrop. 10:498. 1887. Tre: RUSSIA: northern Sakha (Yakutia), “ostia fl. Lena, Tonoldo [??]," Kol. [Koelreuter?] 4711 (LE). Lychnis sibirica L. subsp. mE (Trautv.) Tolm., Trudy Polyarn. Komiss. 8:118. 1932. Sofianthe villosula (Trautv.) Tzvelev, Novosti Sist. Vyssh. Rast. 33:97. 200 The evidence of ER et al. iun strongly supports inclusion of the Lychnis sibirica L. group in Silene. his group co entities that reach the Arctic (and some more southern entities). Oxelman et al. 2001) accepted two of them as species: Silene linnaeana Vorosch. and S. samojedora (Sambuk) Oxel- man. They did not consider the third entity, “villosula,” in their analysis. Oxelman (pers. comm.) is open to accepting “villosula” as a third species. Tzvelev (2001) had already treated the three entities as species, but in the genus Sofianthe. Silene violascens (Tolm.) VV. Petrovsky & Elven, comb. nov. Basiony: Gastrolychnis violascens Tolm., Fl. Arct. URSS 1971. Tyre: RUSSIA: Sakha (Yakutia), “ad flumen Daaldyn, in ditione fluminis Marcha, fluminis Viluj confluentis,” 16 Jul 1957, A. Lukitcheva (HoLoTYPE: LE?). ?Lychnis uniflora Ledeb., Mém. Acad. Imp. Sci. St. Pétersbourg Hist. Acad. 5:536. 1815. Tire: RUSSIA: “Herb. Ledeb. In Sibiria transbaicalensi, lect. am. Tilesius" (HoLoTYPE: LED. ?Gastrolychnis uniflora (Ledeb.) Tzvelev, Bot. Zhurn. (Moscow & Leningrad) 85(11):101. 2000. Silene violascens is related to S. uralensis but differs in morphology, habitat, and range. It is mainly a Siberian forest species. It is here recombined within Silene as the evidence of sim et al. 2001) makes it niei difficult to retain the genus Gastrolychnis. We also consider the type of lychnis uniflora (Lyc! which Tzvelev (2000) considered a priority name for Gastrolychnis EEN a small “plant of S. wels, This will not influence priority in Silene as there is an earlier S. uniflora Roth 1794. Ranunculaceae up x spitsbergense (Hadaé) Elven, comb. nov. Basionim: R | itsbergensis Hadaé, Stud. Bot. Cech. 5:3. Tre i, Longyearbyen, 16 Jul 1939, E. Hadaé 260 (HOLOTYPE: O). A e Ranunculus pallasii Schltdl. var. minimus Rupr., Fl. Samojed. Cisural. 18. 1845. Tyre: RUSSIA: northeastern European Russia. Ranunculus pallasii Schltdl. var. spetsbergensis Nath., Kongl. Svenska Vetenskapsakad. Handl., n.s., 20, 6:21. 1883. Type: NORWAY: Svaibard. LA V DE Lod I bard, from where it was first bd it occurs fairly frequently in large stands and nearly always in the absence of one or both parents (C. pallasii is very rare on Svalbard). It is obviously not a case of occasional hybridisation. Dispersal is assumed to be mainly or entirely by shoot fragments carried by birds. The iden- tification with RADICE s var. minimus was made Ki Tolmachev (1971). Hada?'s R is was not explicitly based on Nathorst's var. spetsbergensis and was given with a separate type. Bregen s name, spelling and type have priority for a variety, whereas Hadaé's name, spelling and type have priority for a species, see Jalas (1988). The combination Coptidium x spitsbergense (Hada?) Hadac in Á. Lóve & D. Lóve, North Atlantic Biota and their History 218 (1963), is illegitimate as the genus Coptidium was not validly published until 1994. Coptidium (Ranunculus) spitsk is a seed-sterile triploid hybrid Coptidium lapponicum x pallasii. On Sval- Ranunculus oe ed dee Glen (Wahlenb.) Elven, comb. nov. Basionym: Ranunculus acris L. var. pumilus Wahlenb., Fl. Suec. 1:351. 1824. Tire: SWEDEN: Lycksele lappmark, aa? at En d ” 03 Aug1948, O. Rune 200/48 iia UPS, S designated d Nurmi in Jonsell, Nordic J. Bot. 20:520. 2001, Nilsson 1991:298, no g yl is L. subsp ilus (Wahlenb.) A LED Lor: Taxon 34:164. 1985. ?R bus? inosiformis Seli Trautv., Increm. Fl. Phaenog Ross. 1:21. 1882. We accept Ranunculus subborealis Tzvelev (the name replacing R. borealis Turcz. in a northern context) as a species separate from R. acris L There can be two different interpretations of the name “pumilus” and two different meanings derived from them. The older meaning is Nordic and based on Ranunculus acris var. alpha Wahlenb. 1811, formal- ized as R. acris var. pumilus Wahlenb. 1824. Nurmi's typification assigns this name to a Scandinavian alpine subglabrous race more closely related to R. subborealis than to R. acris. Intermediates are frequent between subsp. pumilus and other races of R. subborealis but not between subsp. pumilus and R. acris. 438 The other meaning is Russian and based on R. acris var. pumilus “Wahlenb. ex” J. Fellm., Bull. Soc. Naturalistes Moscou 8:271. 1835. If Fellmann really based his variety on Wahlenberg's var. alpha, then it is homotypic and a superfluous combination antedated by Wahlenberg 1824. Thus the Nordic meaning as established by the typification of Nurmi is also valid for Fellmann’s variety. If not, var. pumilus sensu Fell- mann is a later homonym. From the descriptions in several Russian floras, the application of the name var. pumilus J. Fellm. in Russian areas is different from that of the name var. pumilus Wahlenb. in Nordic areas. R lus subborealis Tzvelev subsp. villosus (Drabble) Elven, comb. et stat. nov. Basionym: Ranunculus acris L. f. villosus Drabble, Bot. Soc. Exch. Club Brit. Isles 9:472. 1930. Ranunculus acris L. subsp. borealis (Regel) Nyman var. villosus (Drabble) S.M. Coles, Watsonia 8:240. 1971 This race connects morphologically with Ranunculus subborealis s. str. in northeastern Fennoscandia, with transitions in the Norwegian provinces of Troms and Finnmark and in northern Sweden and Finland. Transitions towards R. acris s. str. are rare. Subspecies villosus is the dominant entity of the R. acris group in the Faeroes and Iceland and a major entity in western and northern Scandinavia. Papaveraceae Papaver labradoricum (Fedde) Solstad & Elven, comb. et stat. nov. Basionym: ee nudicaule L. subsp. radicatum var. (gamma) labradoricum Fedde in Engl., Pflanzenreich IV-104(40):377. 1909. Tier D: “Flora Groenlandiae boreali- occidentalis, Gebiet des Umanakfjords (70—71? N. Br.)," 9 Aug 1893, E. Vanhöffen 35 (04) Fa B 10 02679991). Papaver radicatum Rottb. subsp. labradoricum (Fedde) Fedde in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 17b:120. 1936. Papaver lapponicum (Tolm.) Nordh. subsp. labradoricum (Fedde) Knaben, Blyttia 16:78. 1958 The eastern Canadian and Greenlandic Papaver labradoricum resembles P. lapponicum (Tolm.) Nordh. and P. radicatum Rottb. and has been considered as a subspecies of either one of these (e.g., Fedde 1936; Knaben 1958, 1959a, 1959b). The assignment to P. radicatum has been justified by morphology (Kiger & Murray 1997), that to P. lapponicum by sharing the same octoploid chromosome number (2n - 56), whereas P. radi- catum is decaploid. In an unpublished study by Solstad et al., based on Structure analysis of AFLP DNA markers, P. labradoricum is a distinct entity (species) which joins comparatively closely with a Beringian species (P. cf. detritophilum) out is M TE from the Atlantic European P. radicatum and the amphi-Atlantic P. lapponicum. Several res are also shared with the Beringian species, but P. detritophilum is hexaploid (2n = 42) A separon is further supported by the low fertility found by Knaben (1959b) in crosses between P. labradoricum on one hand and P. lapponicum and P. radicatum on the other, at the level of crosses between species according to Knaben. Papaver labradoricum is probably an allopolyploid with a complicated parentage, as are P. lapponicum and P. radicatum. Brassicaceae Arabidopsis petraea (L.) VI. Doron. subsp. septentrionalis (N. Busch) Elven & D.E Murray, comb. n Basionvm: Arabis septentrionalis N. = EE Mater. ins Glavn. Bot. Sada RSFSR 3(3-4):10. 1922. Tree: RUSSIA: in Arabis petraea (L) Lam subsp tent lis (N. Busch) European Russia (Novaya Tolm. in Tolm. & Blumenthal, Geet Bot. Muz. 23:204. 1931. Cardaminopsis septentrionalis (N. Busch) O.E. Sula in Engl. & Prantl, 1936. Nat. Pflanzenfam., ed. 2, 17b:541. O'Kane and Al-Shehbaz (1997) proposed Arabidopsis parada s. A as one SE EE subspecies of Arabidopsis lyrata (L.) O'Kane & Al-Shehbaz. Tolmachev (1975) id terial as two subspecies of Arabis petraea (L.) Lam. Doronkin in Malyschev and Peschkova (1994) accepted only a collective Cardamin- opsis petraea (L.) Hiitonen. We have studied herbarium material of the three proposed entities (“petraea,” "septentrionalis," and *umbrosa") and have also seen all three in the field in northern Europe, Siberia, and Alaska-Yukon. They are allopatric to slightly parapatric, with *umbrosa" in northwestern North America and northern-northeastern Asia (slightly overlapping with the next), "septentrionalis" in northern Asia and northeastern Europe, and “petraea” in northwestern and central Europe. There are other, non-arctic entities in the group. We are able to recognize the arctic entities on morphological criteria, and transitional forms have not been documented. Arabidopsis petraea s. lat. also differs in some morphological characters from A. lyrata—kamtschatica, without documented intermediates. The choice is between treating the arctic taxa as Fl FRA Al La D nm Al D ti” El 439 three species or as three subspecies of A. petraea. Pending a combined molecular and morphological inves- tigation of the entire aggregate applying molecular markers more suitable to specific/subspecific variation in recently evolved taxa, we propose to treat them as three subspecies Arabidopsis petraea (L.) VI. Dorof. subsp. umbrosa (Turcz. ex Steud.) Elven & DE Murray, comb. nov. Bastonyu: Arabis umbrosa Turcz. ex Steud., Nomencl. Bot. 2:118. 1824. Tree: RUSSIA: Siberia: “in sabulosis ad fl. Irkut,” Turczaninow (HOLOTYPE: MW). Arabis petraea (L.) Lam. subsp. umbrosa (Turcz. ex Steud.) Tolm., Fl. Arct. URSS 7:97. 1975. Cardaminopsis petraea (L.) Hiitonen subsp. umbrosa (Turcz. ex Steud.) Peschkova, Fl. Tsentr. Sib. 1:396. 1979. Arabis arenicola (Richardson ex Hook.) Gelert var. pubescens (S. Watson) Gelert, Bot. Tidsskr. 21:290. 1898. Tree: CANADA: Hudson Bay, York Facto Arabis media N. Busch, Bot. Mater. Gerb. Glavn. Bot. Sada RSFSR 3(3-4):11. 1922. Tree: RUSSIA: eastern Siberia (LE). Cardaminopsis media (N. Busch) O.E. Schulz in Engl. & Prantl, Nat. Pflanzenfam., ed. 2, 17b:541. 1936. The two entities subsp. septentrionalis and subsp. umbrosa are morphologically close but, as far as we have observed (herbarium, field in northern Sakha), they are without intermediates in the area of sympatry in northern Siberia. Saxifragaceae The phylogenetic analyses of Soltis et al. (1996, 2001) show that there are two major branches of genera in Saxifragaceae, and that a large segregate of traditional Saxifraga L., i.e., Micranthes Haw. (s. lat.) together with, e.g., Chrysosplenium L., belongs in a different branch from Saxifraga s. str. The molecular evidence makes it difficult to retain Micranthes within Saxifraga, and (like Brouillet & Gornall 2007) we accept it as a separate genus for the Checklist. Several combinations within Micranthes have been made in earlier times, and Brouillet and Gornall (2007) made the combinations they needed for their treatment for Flora of North America. This treatment does not cover the Asian taxa, and also for a few North American taxa we prefer another rank than chosen by Brouillet and Gornall. Micranthes hieraciifolia (Waldst. & Kit. ex Willd.) Haw. subsp. longifolia (Engl. & Irmsch.) Elven & D.E Murray, comb. nov. Basionw: i via ii d nicae dis Ge & Kit. ex Willd. f. longifolia Engl. & Irmsch. in Engl., Pflanzenreich IV-117(67):24. 1916. T kchi wrence Bay, “St. Lorenzbusen,” Chamisso (B), “Lútke's Hafen," A. & A. Krause 163 (B), syntypes; Saxifraga hieraciifolia Waldst. & Kit. ex Willd. subsp. longifolia (Engl. & Irmsch.) Jurtz. & V.V. Petrovsky, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 85, 6:101. 1980 TY TC C T A T3 Saxifraga hieraciifolia Waldst. & Kit. ex Willd. subsp. czul Chrtek & Soják, Preslia 53:71. 1981. F East, "Prov. Magadan, regio Tschukotsky, trajectus jugi Maia vicin. 32 km ...,” 2 Aug 1967. T. Derviz-Sokolova & T. Voronova (scri PR). Micranthes hieraciifolia is a polyploid to highly polyploid ¢ (2n = SE T ee species where Zhmylev in Elven et al. (2007) stated that: "Thi gical plasticity, not correlated with the chromosome numbers". We 7 not agree. Large parts of the maei from northeastern Se and Alaska, i.e., Beringia, deviate from the non-Beringian material in several features. We have surveyed North American and some northeastern Asian material (ALA, O) and find that the characters reported for subsp. longifolia vs subsp. hieraciifolia mostly hold, e.g.: leaves narrow, long-petiolate and acute with little pubescence vs broader, more short-petiolate See EE to subobtuse and more hairy; stems with little pubescence vs distinctly pubescent; petal as | ls vs pink and much shorter and narrower than sepals; and sepals purple vs green isvith purple dota The lloressenós of ssp. longifolia is also more interrupted when fully grown, but there is overlap in this character. These characters largely ned SEWER two E waned ian ag subsp. longifolia as narrowly amphi-Beringian and subsp. common co-occurring with subsp. longifolia in the amphi-Beringian ed Subsp. longo ia is thereby fully sympatric with subsp. hieraciifolia, at least in an arctic context. There is also a suggested ploidal difference. All reports of comparatively low chromosome numbers (2n = 56 and 63) are from northeastern Yakutia and Chukotka where these numbers are reported together with the prevalent much higher numbers everywhere else in northern and arctic regions (2n - 100-120). From Alaska, only highly polyploidal counts are reported (2n = >100 and 112), but these reports are partly from localities outside the range of subsp. longifolia as we have recorded it. 440 t tani i Texas 2( The names subsp. longifolia and subsp. czukczorum were published one year apart (1980 and 1981), and we assume that they represent the same taxon and that the authors were unaware of the other publica- tion. On the American side, subsp. longifolia is common in western Alaska north and east to the western Brooks Range (on the north coast as very rare east to Demarcation Point) and to the Kobuk and Yukon river valleys. In southwestern and south-central Alaska it is the exclusive race, e.g., in the Goodnews Bay area, Alaska Peninsula, and on Kodiak and smaller offlying islands. Micranthes lyallii (Engl.) Small subsp. hultenii (Calder & Savile) Elven & D.E Murray, comb. nov. Bastonym: Saxifraga lyallii Engl. var. hultenii Calder & Savile, Canad. J. Bot. 38:418. 1960. Tyre: CANADA: Brrrish CoLumeia: Queen Charlotte Islands, Moresby Island, 20 mi S of Moresby Camp, 29 Jul-2 Aug 1957, Calder, Savile & Taylor 23106 (HoLotyre: DAO). Saxifraga lyallii Engl. subsp. hultenii (Calder & Savile) Calder & R.L. Taylor, Canad. J. Bot. 43:1393. 1965 fe De mae E ls Brouillet and Gornall SE did not include the prop 1 northern tions. There is a gap in t d lations in the eastern cordilleras geben lyallii) and those in the western and northern Condilleras (subsp. hul tenii). We follow Calder and Savile (1960) and Calder and Taylor (1965) in acceptance of two major geographical races, i.e., subspecies in the Checklist concept. Micranthes merkii (Fisch. ex Sternb.) Elven & D.E Murray, comb. nov. B ifi kii Fisch. ex Sternb., Revis Saxifrag., Suppl. 1:1. 1822. Tree: RUSSIA: the Transbaikal area, “In humidis montis Jablonoi Chrebet, ” Redowsky (HoLoTvpE?: LE). Micranthes merkii is a distinct Asian species, assigned by Zhmylev in Elven et al. (2007) to its own sect. Merkianae. ios nelsoniana (D. Pon Small subsp. aestivalis (Fisch. & C.A. Mey) Elven & D.E Murray, comb. Saxifi } Mey., Index Sem. Hor t. Petrop 1:37. 1835. Tyre: RUSSIA: S Siberia, Altai, 1817. Herb. mE (LECTOTYPE: LE), s Sipli inskvy (1976). Mi t} tivalis (Fisch. & C.A Mey.) Small, N. Amer. Fl. 22, 2:145. 1905 Saxifraga nelsoniana (D. D Small sul BECHER (Fisch & CA Mey.) D e Webb, Feddes Repert. 69:154. 1964. Micranthes nelsoniana (D. Don) Small var. als (Fisch. & C.A. Mey) Gornall & H. Ohba in Brouillet & Gornall, J. Bot. Res. Inst. Texas 1:1020. 2007 Brouillet and MOM (2007) recombined Micranthes nelsoniana var. aestivalis. This Asian plant is considered by some Russian aut ors as a species (i.e., Rebristaya 1984), by others as a major geographical race, i.e., a subspecies (Malyschev 1997: Zhmylev in y Elven et al. 2007) due to occurrence of intermediate plants in the zone of overlap. We accept the latter proposal and make the necessary combination. Micranthes nelsoniana (D. Don) Small. subsp. insularis (Hultén) Elven & D.F Murray, comb. nov. Bastonw: L. subsp. insularis Hultén, Svensk Bot. Tidskr. 30:524. ni Type: A ge Aleutian Islands, Carlisle Island, 15 Jul 1932, E Hultén 6591 ( S) iana D. Don subsp. i ltén, Bot. Not. 126:494. 1973. Saxi- fraga insularis (Hultén) ES Novosti Sist, EN Rast. 13:139. 1976. Micranthes "— (D. Don) Small. var. insularis (Hultén) Gornall & H. Ohba in B & Gornall, J. Bot. Res. Inst. Texas 1:1020. 2007. Brouillet and Gornall (2007) recombined Hultén’s Saxif tata subsp. insularis (and Siplivinsky's Saxi- fraga insularis) as Micranthes nelsoniana var. insularis. This’ is an amphi-Pacific plant in very oceanic regions from Sakhalin, = Kuriles, Kamtchatka, and the Aleutian Islands to the very tip of the Alaska Peninsula, mostly well out t] of M. nelsoniana s. str. (see Hultén 1968). Hultén (1968) reported intermediates towards subsp. EE in the zone of overlap on the Alaska Peninsula, Kodiak Island, and the Seward Peninsula. In our opinion, this is a case of subspecific variation, and we recombine accordingly. Micranthes porsiidiana (Calder & Savile) Elven & D.E Murray, comb. nov. Basiony: Saxifraga punctata L. subsp. porsildiana Calder & Savile, Canad. J. Bot. 38:429. 1960. Tree: ipai: EHE Columbia, ea bale at Azouzetta die i Hwy, 4 Aug 1954. a s dii sid 1 Ru ( ifrag D. Don subsp. ; Hultén, Bot. Not. 126:494. 1 porsildiana (Calder & Savile) Jurtz. & VV. eee Bot. Zhurn. (Moscow PR 58:1339. 1973. Micranthes porsildiana was treated by Hultén (1968) as a subspecies of Saxifraga punctata auct. (= Saxifraga (Micranthes) pena as was subsp. insularis. Brouillet and Gornall (2007) did not include or mention this entity. Micranthes ] has alarge overlap in range with M. nelsoniana in Alaska and eastern Chukotka. ic fl 441 Within these regions, we have several times in Alaska and Chukotka observed the two growing in close proximity, sometimes nearly mixed in the stands, without any evidence of intermediates. Zhmylev (1995) assigned M. nelsoniana and M. porsildiana (as Saxifragas) to different series, respectively, series Aestivalis Zhmylev and series Reniformes Zhmylev. Rank as two different species is merited. Micranthes redofskyi (Adams) Elven € D.E Murray, comb. nov. Basionvm: 5 s, Nouv. Mém. Soc mp. Naturalistes Moscou 3:241. 1834. Tree: RUSSIA: N Yakutia, "Fl. Lenam," Adams ex Me Trinius 2911 (LEcTOTYPE?: MW). Micranthes redofskyi is a diploid (2n = 20) belonging in sect. Micranthes together with at least three assumed close relatives: the seed-reproducing, amphi-Atlantic M. stellaris (L.) Galasso, Banfi & Soldano (2n - 28), the also seed-reproducing, Pacific North American M. ferruginea (Graham) Brouillet & Gornall (2n = 20), and the bulbil-reproducing, arctic circumpolar M. foliolosa (R.Br.) Gornall 2n = 40—64). All four species are mor- phologically distinct, but M. foliolosa has been assumed (Zhmylev in Elven et al. 2007) to be: *... a complex of 2-3 entities formed as a result of polyploidy of pseudoviviparous races of Saxifraga stellaris, S. redofskyi an inherited from the parental entities. Arctic plants of Micranthes foliolosa are morphologically much closer to M. redofskyi than to M. stellaris or M. ferruginea. An unpublished investigation based on AFLP DNA mark- ers and comparing these species (A. SEH Salzburg) connects M. foliolosa sampled throughout the Arctic closely to M. redofskyi (and not to M. stellaris). The more floriferous populations of M. foliolosa are found in he Beringian regions, with lower chromosome numbers than the majority of the plants, overlapping in range with M. redofskyi and close to the range of M. ferruginea. Micranthes redofskyi is restricted to northeastern Asia from northern Yakutia eastwards to Chukotka Rosacea Dryas mS Juz. EE hookeriana (uz. 2 juitzs comb. nov. Bastonym: Dryas hookeriana Juz., Izv. Glavn. Bot. Sada SSSR 28:325. 1929. T "Rocky Mountains,” Hooker (LE, ex Herb. Fischer, Ledebour). Dryas octopetala L. subsp. hookeriana (Juz.) Hultén, Acta Univ. Lund., n.s., sect. 2, 42(1):1046. 1946. Dryas hookeriana is recombined as a Cordilleran subspecies of the Asian and Beringian D. punctata. They share the adaxial leaf surface punctate glands, the abaxial leaf surface red, stipitate glands, and the absence of feathery hairs of the D. octopetala type. Potentilla anserina L. subsp. yukonensis (Hultén) Soják ex Elven & D.E Murray, comb. nov. Basionym: Potentilla yukonensis uin Acta aa Lund., n.s., sect. 2, 42(1):1033. 1946. Tree: U.S.A. Alaska: Palisades, 14 Jul 1902, Hollick (HoLOTYPE: db., N. Amer. Fl. 22(4):354. 1908. Potentilla egedii Wormsk. subsp. yukonensis (Hultén) Hultén, El NY). T Ark. Bot. ser. 2, ION. 1968. t] t North American E race , differing from degree as these d ] Treatment Potentilla anserina subsp. yukonensis is an i the other races (subsp. anserina, egedii, pacifica) to the same rank i iate, eit] 1 | fi bspecies. We T chosen the latter course esch + due to the frequent occurrence of intermediates in zones of contact. The name Potentilla yukonensis is a no- men novum as the epithet “subarctica” is inapplicable at species level due to Potentilla subarctica Rydb., N. Amer. Fl. 22(4):347. 1908. The combination P. anserina subsp. yukonensis has already been used as a nom. nud. in a key by Soják to section Pentaphylloides (Soják 1994-70). Fabaceae Hedysarum boreale Nutt. subsp. dasycarpum (Turez.) pe e See & Elven, comb. et stat. nov. Bastonvm: 3. 1840. Tyee: RUSSIA: Sakha (Yakutia). Hedysarum dasycarpum Turcz., Bull. Soc. Imp. ? We consider the nortl tern North American Hedysarum machenzii and the northeastern Asian H. dasy- carpum as allopatric subspecies of a more widely circumscribed H. boreale. The former is already combined as H. boreale Nutt. subsp. mackenzii (Richardson) S.L. Welsh. Ericaceae ` t (Stokes) Harmaja subsp. decumbens Elven & D.F Murray, comb. nov. Basionw: Ledum palustre L. var. ota Aiton, Hort. Kew. 2:65. 1789. Tyre: CANADA: Hudson Bay (HOLOTYPE:?). Ledum decumbens (Aiton) 442 t tani titute of Texas 2( bones ex Se Nomencl. Bot., ed. 2, 2:20. 1841. Ledum palustre L. subsp. d } dis ) tes i o 4:8. 1930. , Ann. Bot. Fenn. 27:203. 1990. Nom for Led ododendron t t (Stokes) H ja subs; ( ja) G.D. Wallace, Madroño 39:77. 1992. Nom. illeg. Harmaja (1990) considered this entity a species and coined the name Rhododendron subarcticum, with Le- dum palustre var. decumbens as its typonym, because a combination as Rhododendron decumbens would be a homonym (due to Rhododendron decumbens D.Don ex G.Don, Gen. Hist. 3:846. 1834). Wallace (1992) con- sidered it a subspecies, as we do, and recombined it as subsp. subarcticum, probably unaware that the epithet decumbens, even if not available for a species, is available and has priority for a subspecies. We accordingly make a recombination on the priority name. Vaccinium uliginosum L. subsp. (Kom.) Alsos & Elven, comb. et stat. nov. Basionvm: Vaccinium vulca- m Kom., Repert. Spec. Nov. Regni Veg. 13:236. 1914. Tyre: RUSSIA (Far East) Kamtchatka, "in alpibus peninsulae Kamtschatka circa lacus Kunotzkoje," 19 Aug 1909 and 02 Sep 1909, VL. Komarov (LE). Vaccinium uliginosum L. subsp. microphyllum (Lange) Tolm. var. vulcanorum (Kom.) Jurtz., Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 85(6):105. 1980. Vaccinium uliginosum L. is polymorphic as described by Young (1970). Komarov's Vaccinium vulcanorum is, according to Tolmachev and Yurtsev (1980), a prostrate-pulvinate dwarf shrub with small, imbricate, rotundate leaves persisting in winter, and globose berries. In a phylogenetic analysis by Alsos et al. (2005), the “vulcanorum” entity is a central node from which both an amphi-Pacific and a circumpolar clade emerge. Both phylogenetic and morphological evidence support the taxon vulcanorum as a race, preferrably as a subspecies. Gentianaceae Gentianopsis barbata (Froel.) Ma subsp. raupii (A.E. Porsild) Elven, comb. nov. Basonvm: Gentiana raupii A P Porsild, Sargentia 4:60. 1943. Tyre: CANADA: Northwest Territory, Mackenzie District, Mackenzie River between Wrigley and Blackwater R, A.E. Porsild 6571 ISLOTE CAN). Gentianopsis detonsa (Rottb.) Ma subsp. raupii (A.E. Porsild) Á. Lóve & D. Lóve, Bot. Not. 128:517. 1976. Genti ii (A.E. Porsild) Iltis, Sida 2:134. 1965. Gentianella detonsa (Rottb.) G. Don subsp. raupii (A.E. Porsild) J.M. Gillett, Ann. SE Bot. Gard. 44:217. 1957. The entities of the Gentianopsis detonsa—barbata group in northern North America are still insufficiently studied, in spite of two revisions (Gillett 1957; Iltis 1965). They are rare taxa, restricted in their ranges and often disjunct. The “raupii” entity has | considered a separate species or a subspecies of Gentianella / Gen- tianopsis detonsa. Hultén (1968b) accepted Gentiana raupii as a separate species in northwestern Canada but said that it is “similar to G. barbata, but with fewer stem leaves; calyx rounded at base; corolla tube broadly obconic, the lobes broad, rounded". We support that view and hereby assign "raupii" to Gentianopsis barbara as a regional race. It occurs mainly along the Mackenzie River, and the entire range is within the glaciated areas. This is an entity that at some time (most probably postglacially) was able to differentiate in isolation from the main body of Gentianopsis barbata. Asterac Achillea alpina L. subsp. multiflora (Hook.) D.E Murray & Elven, on E ME nov. Bastonvm: Achillea multiflora Bor.-Amer. 1:318. 1833. Type: CANADA: "Throughout the wo y, klin,” (HOLOTYPE: 5 ia multiflora (Hook.) Tzvelev in Jurtz., Fl. Arct. URSS 10:107. 1987. The North American representative of the Achillea alpina complex has been treated in most northwestern North American works (e.g., Hultén 1968a; Cody 1996, 2000) as A. sibirica Ledeb. Both names, alpina and sibirica, are based on Asian plants. Trock (2006) applied the name A. alpina without discussion for the North American plant. Russian authors have mostly treated the entities of the A. alpina complex as several species. Rank as largely allopatric subspecies of Achillea alpina is the better solution. The northwestern North American plants differ from the northeastern Asian ones in some quantitative features: capitulae more numerous, ligules smaller, and leaves slightly more dissected. We hereby recombine A. multiflora as a subspecies of A. alpina. The Achillea alpina complex has been considered part of the segregate genus Ptarmica (e.g., Yurtsev ri ] AA Al kh: el HET | D ti 443 1987). Acceptance of Ptarmica as a separate genus is untenable according to phylogenetic evidence (Guo et al. 2004) Artemisia kruhsiana Besser subsp. alaskana (Rydb.) D.F Murray & Elven, comb. et stat. nov. Basony: Artemisia alaskana Rydb., N. Amer. Fl. 34(3):281. 1916. Tyre: U.S.A.: ALASKA: Yukon River, Nulato-Nowikakat, 23-27 Jul 1889, LC. Russell (HoLomee: US-417571) 1 T vith 41 +l Aciqn Hultén (1968b) compared thwestern North American Artemisia A. kruhsiana and considered them related but specifically distinct. Artemisia “o was described with three allopatric to a SE in northeastern Ásia E 1987). Korobkov (pers. comm.) indicated that A. alaskana lly closest to A. kruhsia bsp. multisecta (Leonova) Korobkov. We consider A. alaskana to be des enough to A. kruhsiana to be best considered a subspecies of the latter (see also Tkach et al. 2008 wherein A. kruhsiana and A. alaskana were very close in the phy logeny). In our opinion, the closest relationship is found with the Chukotkan subsp. kruhsiana. There is, however, an ap- preciable gap between the westernmost geed occurrence of subsp. alaskana and the easternmost Asian occurrence of subsp. kruhsiana, as th bsent from all of Chukchi Peninsula. We therefore propose subsp. alaskana as a fourth subspecies gi A. kruhsiana. Packera w (Greenm.) Á. Löve & D. ee subsp. wrangelica Qurtz., Korobkov & V.V. ere Ju Korobkov VV Petrovsky, comb. n SD wrangelica Jurtz , Korobk D & VV. io in uc Fl. Arct. URSS 10:216. 1987. Tyre: RUSSIA: Russian Far East, See Island, “pars centralis, in cursu superiore fl. Neizvestnaja (Incognita)," 22 Jul 1971, VV Petrovsky & O.D. Schteinberg (HOLOTYPE: LE). Transfer of Senecio hyperborealis to Pachera necessitates the new combination. Pachera hyperborealis s. str. is diploid (Packer 1972, northwestern Canada, 2n 2 46; Yurtsev & Zhukova 1972, northeastern Asia, 2n = 48). Subspecies wrangelica, on the other hand, is hexaploid (Zhukova & Petrovsky 1987, Wrangel Island, 2n = 138). This high number is more in accordance with the wide range of numbers found in the closely related P. heterophylla (Fisch.) E. Wiebe (= Senecio cymbalaria Pursh, Senecio resedifolius Less.). Scorzoneroides autumnalis (L.) Moench EE pratensis (Hornem.) Elven, comb. nov. Basionym: Apargia pratensis Hornem., Hort. Bot. Hafn. 2:785. 1815. I ) Rchb., Fl. Germ. Excurs. 1:253. 1831. EH autumnalis . var. pratensis (Hornem.) W.D J. Koch, dn Fl. Germ. Helv. 418. 1837. Leontodon autt subsp. | ) Gremli, Excursionsfl. Schweiz, ed. 3:242. 1878 lis (L.) Moench var. pratensis (Hornem.) PD. Sell, El Great Brit. Irel. 4:531. 2006. Leontodon autumnalis L. var. taraxaci (L.) Hartm., Handb. Skand. Fl., ed. 4:252. 1843 (Basionym: Hieracium taraxaci L., Sp. PL, ed. 2, 2:1125. 1763). The new combination is necessitated by the transfer of Leontodon ae L. to Mie EE Scorzoneroides Moench (cf., Sell & Murrell 2006). The provenience and characters of I tensis is not known to us, but the customary use of the name pratensis in northern Europe is for Tue Blast considered here. The Nordic plants currently assigned under var./subsp. pratensis include Linnaeus' Hieracium taraxaci and are different from the central European alpine plants. Solidago multiradiata Aiton subsp. arctica (DC.) Korobkov & Elven, comb. et stat. nov. Basionya: Solidago virgaurea L. var. (epsilon) arctica DC., Prodr. 5:339. 1836. Type: U.S.A: ALASKA: the Aleutian islands, “Unalaschka,” and possibly RUSSIA: Russian Far d Chukchi Peninsula: *ad sinum Sancti Laurentii." ee multiradiata Aiton var. arctica (DC.) Fernald, Rhodora 17:4. 1 pacta Turcz., Bull. Soc 13:73. 1840. Probably a nomen novum for var. arctica as a species. Solidago multiradiata Aiton is widespread and polymorphic. The plants around the Bering and Chukchi seas in eastern Chukotka and western and southwestern Alaska constitute the westernmost extension of the range for S. multiradiata and differ in several quantitative features. Hulten (19682) accepted a variety, var. arctica. Russian authors (e.g., Korobkov 1987) have instead treated it as a separate species, 5. compacta. We prefer to see S. compacta or var. arctica as a regional race, i.e., a subspecies. It might be uniformly tetraploid (2n = 36), as indicated by Lóve and Lóve (1975), but both diploid and tetraploid counts are known from S. multiradiata s. str. REFERENCES ALSOS, |.G., T. ENGELSKJON, L. GIELLY, P. TaBerLet, and C. BROCHMANN. 2005. Impact of ice ages on circumpolar molecular diversity: insights from an ecological key species. Molec. Ecol. 14:2739-2753. Bateman, R.M., PM. HOLLINGSWORTH, J. PRESTON, Y.-B. Luo, A.M. Prioceon and M.W. Chase. 2003. Molecular phylogenetics and evolution LO: and selected Habenariinae (Orchidaceae). Bot. 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Geraniaceae-Scrophulariaceae. "Nauka, Leningrad. Trock, D.K. 2006. Achillea Linnaeus. In: Flora of North America Editorial Committee, eds. Flora of North America north of Mexico. 19. Magnoliophyta: Asteridae (in part): Asteraceae, part 1. Oxford Univ. Press, New York- Oxford. Pp. 492-494. TzveLev, N.N. 1964. Puccinellia Parl. In: Tolmachev, AL, ed. Flora Arctica URSS. Il. Gramineae. "Nauka; Moscow & Leningrad. Pp. 178-208. TzveLev, N.N. 2000. On the genus Gastrolychnis (Caryophyllaceae) in European Russia. — Bot. Zhurn. (Moscow & Leningrad) 85(11):99-102. [In Russian.] Tzvetev, N.N. 2001. De generis tribus Sileneae DC. (Caryophyllaceae) in Europa Orientali. Novosti Sist. Vyssh. Rast. 33:90-113. [In Russian.] WaLLacE, G.D. 1992. Ledum in the new Jepson manual and a new combination for Ledum in Rhododendron (Ericaceae). Madroño 39:77. Young, S.B. 1970. On the taxonomy and distribution of Vaccinium uliginosum. Rhodora 72:439-459. Yunrsev, B.A. and PG. ZHukova. 1972. Tsitotaksonomicheskaya kharakteristika endemichnykh rasteniy gornogo severo-vostoka Azii. Bot. Zhurn. (Moscow & Leningrad) 57:50-63. Zhukova, PG. and VV. Petrovsky. 1987b. Chromosome numbers and taxonomy of some plant species from the northern Asia regions. Bot. Zhurn. (Moscow & Leningrad) 72:1617—1624. [In Russian HEUCHERA WOODSIAPHILA (SAXIFRAGACEAE), A NEW SPECIES FROM THE CAPITAN MOUNTAINS OF NEW MEXICO Patrick J. Alexander Biology Ee MSC 3AF Mex New co State University Las Cruces, SE Mexico 88003, U.S.A. Pe S O h ABSTRACT " n E 1 o r JH a Tuas phil H J DN ] E 41 nm pit AA a Cy: 1 r^ ty, New Mexico. This HM is similar to thos f TT 1 pe ri pu 1] 1 pe Cyli Js. 1 dicm 1 igl ly ] I 1 J 1 r the subsection. H iaphila i lilv di i if her Heuc) in tl 1 y its greenish hite t tal E bie erect a 2 e 75 mm long. le is EH from dp pP 1 popul f d granitic tal tl tl Peak, between 2,550 and RESUMEN Se describe una especie nueva, Heuchera ]siaphila (Saxifrag ) de las Capitán Mountains en Lincoln County, Nuevo México Tras : = :1 1 1. I7 1 rr.) Lal 1 4 Sal A ^l " "m : 1 1 1 HE 1 $ H LI 1 Sie Lavy J: t [zs D J IY 1 1 na [1 D 1 de blanco d ] sp 1 tos de 2.75 a 3.75 mm de largo. S 5lo de d lect taludes graníti d | 145145 3 y te de C pitá Peak, | ] d 2,550a 2,900m The Capitan Mountains are the most northeastern range in the greater Sacramento Mountains complex, a group of mountain ranges in Lincoln and Otero counties of south-central New Mexico which also includes Sierra Blanca, Carrizo Mountain, and the Sacramento Mountains proper. These mountain ranges include the highest point in the southern half of New Mexico, Sierra Blanca Peak at 3,649 m, and contain large areas over 2,500 m elevation. There are no similar mountain ranges nearby. The nearest peak exceeding 3,000 m is South Baldy in the Magdalena Mountains, ca. 190 km to the west-northwest, while the nearest peak equalling Sierra Blanca's height is Tesuque Peak in the Sangre de Christo Mountains ca. 320 km to the north. Consequently, there is a well-developed high-elevation flora in the Sacramento Mountains com- plex that is isolated from other areas with similar floras. This area contains some of the most southeastern populations of typical members of the Rocky Mountain flora, such as Picea pungens Engelm. and Abies bifolia A. Murr. and is home to 21 endemic plant taxa. Most botanical exploration in the area has been conducted in the Sacramento Mountains proper and on Sierra Blanca. The Capitan Mountains and Carrizo Mountain, both granite laccoliths on the northern end of the Sacramento Mountains complex, are comparatively dif- ficult to access and distant from centers of botanical research. Their floras remain poorly known. It is thus no coincidence that the Capitan Mountains yielded the most recently described of the endemics of the Sacramento Mts. complex, Astragalus kerrii Knight & Cully (Knight & Cully 1991), as well as the presently described species. The endemic flora of the Sacramento Mountains complex has traditionally included a Heuchera, H. wootonii Rydb. of section Parvifoliae (Small & Rydberg b Ge & A d Rosendahl et al. 1930). This species, however, has recently been reported iption is uncertain (New Mexico Rare Plant Technical oun 1999; B. Shipes, personal communication). With the pron - described H. woodsiaphila, the Sacramento Mountains complex gains an endemic Heuchera at the sam that it risks losing one. J. Bot. Res. Inst. Texas 2(1): 447 — 453. 2008 448 | f Texas 2(1) Ge T ENE GE EA RNS en ap SCH Ep ape e a P D x D Fic. 1. Heucl side view. EES woodsiaphila BJ ee sp. nov. (Figs. 1-3). Tre: U.S.A. New Mexico. Lincoln Co.: NE us of capitan Trail, 33° 35' 37"N 105° 15! 31.5"W elev. 9200 ft, steep NE slope, duff boulders in Bee forested als 17 Jul 2006, PJ. Alexander 390. (HoLoTYPE: NMC; IsoTYPES: UNM, ARIZ, ALA, MO, DUKE). Dedh lis a Heurt icola E. Nels. (Idaho, 1 1 EI t. Holochloa Nutt. subsect. Cyl Montana. Oregon) a qu EM Lu inn cordate ee viscido carenti, et bracteis superis brevioribus (a 5 mm). Differt a ] A LGL aa, Heuchera cetera in Nev t s viridi-albas ve Herbs, perennial, acaulescent from an elongate, DEE? caudex; leaves with stipules lanceolate, 5-8 mm long, adnate to the petioles + 2/3 their length, entire, glandular-puberulent, ciliate distally with whit- ish, gland-tipped trichomes to 2 mm long; petioles of the larger leaves ascending, 4-8 cm long, glandular- puberulent and sparsely villous with white, minutely gland-tipped, multicellular hairs to 5-8) mm long; blades green, sub-reniform to broadly ovate, the base cordate, the larger leaves 2.5—4 cm wide and 23.5 cm long, shallowly 5-lobed, with + 25-30 broad teeth, these mucronate, the whitish mucros to 1 mm long, Alexander, A ies of Heuct f New Mexi 449 be 2 | Lia fF Bai J F Lat £11 E Zap | can be found at http /Ipolyploid. L.net/Heuchera/. minutely gland-tipped, secondary teeth often reduced to mucros on the lateral NN ofthe pus teeth, margins also with minute, translucent, gland-tipped projections, surfaces gl pulverulent, also spar- ingly villous on the veins adaxially, with white, gland-tipped hairs to 2 mm long; flowering stems 8-20 cm high, subscapose, glandular-puberulent throughout, usually with 13 sterile bracts (or these absent), the sterile bracts with green blades, resembling the basal leaves, 4-15 mm long by 4-15 mm wide, 3-5 lobed, lobes of the larger bracts crenate-dentate, the teeth acuminate to short-mucronate, surfaces glandular- pulverulent; inflorescences 3—4 cm long, secund or subsecund, flowers born in small cymules of 1-4 flowers, peduncles mostly + 1 mm, the lower often elongate to 7 mm, the pedicels 1-2 mm; bracts of the inflorescence variable, the lowest resembling the leaf-like sterile bracts, the rest linear, 3-5 mm long, pale green or whitish at anthesis, often becoming pinkish with age, glandular-puberulent, the stipules represented by E aristate ës or absent, margins entire or distally fimbriate, bractlets of the cymules becoming 11 talous, regular or nearly so, 5.25—6 mm long at anthesis, the hypanthium " 1. 5mm SEN videns gentis from the inferior portion of the ovary; sepals 5, equal, greenish-white to cream at anthesis, the tips sometimes becoming pinkish in fruit, oblong with obtuse i P e 75 mm long, densely glandular-puberulent on both surfaces; stamens 5, arising from the | ightly below the bases of the sepals, + 2 mm long, gently incurved, persisting until after E of the capsule, the filaments + 1.5 mm long and tke anthers + 0.7 mm long, deeply cordate, orange, red or purplish at anthesis; gynoecium of 2 carpels, at anthesis with the inferior portion of the ovary hemispheric, 2-2.5 mm wide and + 1.25 mm long, the styles and beaks of the carpels separate for + 2 mm, the last 1-1.5 mm of this com- prising the true closed styles, styles included with capitate stigmas slightly wider than the styles; capsules ovoid with a nearly hemispherical base, 5.5-6.5 mm long, the beaks of the carpels splitting longitudinal ly on their interior surfaces, the styles slightly curved outwards, equaling the sepals or exserted to 1 mm, the true closed styles 1-1.5 mm long above the beaks of the carpels; seeds dark brown, narrowly ellipsoid, + 0.75 mm long by + 0.3 mm wide, the surfaces echinulate. 450 Contour interval is 1.000 ft. Lowest contour = 6,000 fi. Ft. Stanton Lincoln“, | vid sa N SC Mi SI L^ Downs T ^ De 2 LF | «th AN Er Ld = | L J A À ] g A I g I Etymology.—The specific epithet refers to the close association of this species with Woodsia plummerae Lemmon, which is ubiquitous at populations of H. woodsiaphila and is often the most abundant vascular plant in the SHOE vicinity. Phenology.—Specim n llected in July and August include both mat fruit d flowers at anthesis Geography m habitat —Heucl tly known only from Capitan Peak at the eastern end of the Capitan Mountains in Lincoln County, New Mexico (Fig. 3), where it has been observed from 2,550 to 2,900 m in elevation along the Capitan Peak Trail, Lincoln National Forest Trail #64. It is found at locally moist sites on north or northeast, stable, forested granitic talus, usually under cover of Pseudotsuga menziesii (Mirb.) Franco or Abies concolor Lindl. & Gord. and in association with Woodsia plummerae. These moist sites are many, but small and scattered. Much of the forest in this area has burned recently, and H. woodsiaphila appears to be equally abundant in burned and unburned areas. Distinction from other Heuchera. Heuchera woodsiaphila is very similar to members of Heuchera section Holochloa Nutt. subsection Cylindricae Rydb. Like other members of subsection Cylindricae, it has elongate (5-8 mm in the subsection), white or whitish, actinomorphic or slightly zygomorphic, apetalous flowers with campanulate hypanthia and long (2.5—6 mm in the subsection), erect sepals. Heuchera woodsiaphila is widely disjunct from other members of the subsection, which are present in the northwestern United States and adjacent Canada. The nearest known populations of H. cylindrica Dougl. are in northeastern Nevada more than 900 km to the northwest. Taxonomy of subsection Cylindricae is uncertain. The subsection includes up to 9 taxa at varietal to specific ranks, all of these taxa apparently intergrading to a greater or lesser extent (Calder & Savile 1959; Alexander, A i f Heuc! f New Mexi 451 Rosendahl et al. 1936). In comparatively recent floras of the northwestern United States and adjacent Canada, subsection Cylindricae is generally treated as two species: H. chlorantha Piper and a highly polymorphic H. cylindrica (e.g., Douglas et al. 1991; Hitchcock & Cronquist 1973). Based on the most recent monographic treatment of subsection Cylindricae (Calder & Savile 1959), H. woodsiaphila is most similar to H. saxicola E. Nelson and H. cylindrica Dougl. vars. cylindrica and orbicularis (Rosend. et al.) Calder & Savile. It differs from the former in lacking dense, viscid pubescence, in having leaves with cordate bases, and in having smaller bracts within the inflorescence. Heuchera woodsiaphila differs from the above-mentioned varieties of H. cy- lindrica in having shorter scapes (8—20 vs 22—70 cm) and usually having sterile bracts with well-developed blades. Although Calder and Savile (1959) do not discuss characters of the androecium or gynoecium in depth, these are described in detail by Rosendahl et al. (1936). Following the latter treatment, H. woodsiaphila also differs from the known members of subsection Cylindricae in having shorter anthers (0.7 vs. 0.9-1.5 mm) and longer true closed styles (1-1.5 vs. < 0.5 mm). To aid in distinction of this species from others in New Mexico, a key is provided below. KEY TO THE HEUCHERA SPECIES OF NEW MEXICO 1. Flowers pinkish to red. 2. Stamens about equaling or slightly exceeding the hypanthium; petals much shorter than the sepals or sent H. sanguinea Engelm. tals both equaling or exceeding the sepals E! Hypanthium 1 mm ono or less at its shortest | yles strongly-exserted, exceeding the sepals by at least 1 mm H. rubescens Torr. 3; BLUE 1.5 mm long or more at its shortest point; styles only slightly exserted, o] the sepals by less than 1 mm pulchella Woot. & Standl. Flowers white, cream, or greenish white 4. Hypanthia deep, campanulate; petals erect and shorter than to equaling the sepals, or absent; sepals whitish basally with greenish tips or whitish throughout, erect basally and spreading apically or erect throughout. 5. Petals absent; sepals 2.5 mm or longer; scapes 8-20 cm H. woodsiaphila PJ. Alexander 5. Petals present; sepals 2 mm or less; scapes 25 cm or more 6. o glandular-puberulent, the longest hairs < 0.2 mm; true closed styles present, > 0.5 m lon ovomexicana Wheelock 6. E glandular- S the longest hairs 0.3-0.6 mm long; true closed styles « 05 mm long or apparently absen H. glomerulata Rosend. et al. 4. Hypanthia shallow, saucer-shaped; petals spreading, usually longer than the sepals, rarely equaling them; sepals green throughout, spreadin 7. Petioles hirsute; plants of cole Otero, and Catron Counties. H. wootonii Rydb. 7. Petioles merely puberulent; plants of north-central New Mexico H. parvifolia Nutt. ex Torr. & A. Gray Although production of new comprehensive keys for subsection Cylindricae best awaits revision of the group and is thus beyond the scope of the present work, the key of Calder and Savile (1959) can be modified to accommodate H. woodsiaphila by replacing the first half of couplet “2” with the following: 2. Well- developed bracts of panicle 4.0-15.0 mm long and 2.5-15.0 mm wide, excluding cilia, often green and leaflike; leaf blades EORSpIEMOSY viscid Or nor L FN UNE “| ] I " il [| £ | 2a. Leaf blades with many y ; g bro ES EE rarely slig ntly cordate H. saxicola E. Nelson 2b. Leaf bl g t not viscid, not il and plant particles; leaf I trongly cordate H. woodsiaphila P.J. Alexander PanATYPE. U.S.A. New Mexico: Lincoln Co.: Capitan Peak, 13 Aug 1995, S. Chaddie 135 (NMC). Other specimens.—Of the other major regional herbaria only UNM and UTEP have specimens of Heuchera from the Sacramento Mountains complex. On viewing the collections of these herbaria no additional speci- mens of H. woodsiaphila were found. Discussion.—Although a conspicuous component of the flora at Capitan Peak, H. woodsiaphila was still unknown when I stumbled upon it while hiking the peak in 2006. Although I quickly found S. Chaddie's 452 1995 i (also from Capitan Peak but misidentified as H icana) and determined it was the same A as m» new collection, I could not identify these plants. Discussion with other botanists in the state, notably Robert Sivinski, quickly made it clear that this was not a species known from New Mexico but left open the possibility that it was a species well-known elsewhere. In the comprehensive monograph of Heuchera by Rosendahl et al. (1936), H. woodsiaphila clearly falls into Heuchera section Holochloa subsection Cylindricae. Although E deiere eg is readily distinct from taxa of subsection Cylindricae as they are described in the of Rosendahl et al. (1936) and Calder and Savile (1959), the confused nature of this sube don makes it difficult to ascertain the limits of the considerable variation found in both the named taxa and their various hybrids. The proper status of H. woodsiaphila is thus difficult to determine at present. Although it could be argued that H. woodsiaphila is better treated as a variety of H. cylindrica, hiding varia- tion within species tends in practice to obscure it or give the impression that it is unimportant. Further, this would not in any way reduce the taxonomic uncertainty in subsection Cylindricae, but would merely maintain the same uncertainties at a different taxonomic level. Unfortunately an adequate resolution to this problem must await a modern revision of the BUDE. Although H. Isiaphila i imil hol llyt | f subsection Cylindricae, the pos- sibility that this species may en bom hybridiza done kiun New Mexican Heuchera and be only convergent with members of subsection Cylindricae should not be excluded. Leaves of H. woodsiaphila are identical in appearance to those of some specimens of the polymorphic H. rubescens, a member of section Rhodoheuchera R. B. & L. subsection Rubescentes Rydb. and, although difficult to distinguish in technical Ea characters, seem subjectively rather different from those of subsection Cylindricae. Furthermore, most char- acteristics in which this species differs from New Mexican members of section Holochloa, including the long sepals, short scapes, and well-developed, leaf-like sterile bracts of the scape, can be found in H. rubescens. Molecular phylogenetic analyses will probably be necessary to ascertain the affiliations of H. woodsiaphila with any certainty. The highly restricted distribution of H. woodsiaphila is also noteworthy. The apparent limitation of this species to high-elevation, forested talus may explain this limited distribution. On the Capitan Peak Trail, H. wootonii occurs in similar habitat to H. woodsiaphila but at lower elevations (1,950 to 2,400 m). On trips to the otherwise geologically and floristically similar mountains nearby, Sierra Blanca and Carrizo Moun- tain, I could find neither high-elevation, forested talus nor H. woodsiaphila. Instead, on these tains the talus is either bare or has only low, tangled vegetation and H. wootonii occurs to at least 3,000 m. In these mountains, H. wootonii is often found near Woodsia neomexicana Windham or W. phillipsii Windham. Here W. plummerae, which is typically present with H. woodsiaphila, is conspicuously absent. Both the Capitan Mountains and nearby Carrizo Mountain are poorly explored botanically, however, and H. woodsiaphila may well have a broader geographic or ecological distribution than is presently known. ACKNOWLEDGMENTS I thank Richard Spellenberg for assistance and motivation in preparing the manuscript, Katy Levings for creating the illustration, Guy Nesom for translating the diagnosis into Latin, Ana Lilia Reina-Guerrero for vetting the Spanish translation of the abstract, and Robert Sivinski for helpful discussion regarding the distinctness of H. woodsiaphila. Larry Mellichamp, Barbara Greene Shipes, Charles B. McDonald, and Jane Mygatt provided helpful review comments. REFERENCES CALDER, J.A. and D.B.O. Save. 1959. Studies in Saxifragaceae-l. The Heuchera cylindrica complex in and adjacent to British Columbia. Brittonia 11:49-67. DoucLAs, G.W., G.B. StraLey, and D. Meneer. 1991. The vascular plants of British Columbia: Part 3 - Dicotyledons (Primulaceae through Zygophyllaceae) and Pteridophytes, p. 177. British Columbia Ministry of Forests, Van- couver, British Columbia. Special Rep. Ser. No. 3. Alexander, A DH £u L E M Aa H 453 HitcHcock, C.L. and A. Cronauist. 1973. Flora of the Pacific Northwest. University of Washington Press, Seattle, Washington. KNIGHT, PJ. and AC Cous 1991. A new species of Astragalus (Fabaceae) from southeastern New Mexico. South W. Naturalist 36:198-200. New Mexico RARE PLANT TECHNICAL COUNCIL. 1999. New Mexico rare plants. Albuquerque, NM. New Mexico Rare Plants Home Page. http://nmrareplants.unm.edu (Latest update: 6 December 2007). RosENDAHL, CO. F.K. Butters, and O, LakeLa, 1936. A monograph on the genus Heuchera. University of Minnesota Press, Minneapolis, Minnesota. Minnesota Stud. Pl. Sci. No. 2. SMALL, J.K. and PA. RYoBERG. 1905. Saxifragaceae. In: North American Flora. Vol. 22. L.M. Und | and N.L. Britton, eds. New York Botanical Garden, Bronx. Pp. 81-158. 454 tani itute of Texas 2(1) BOOK NOTICE LAURENCE M. Harpy (ed.). 2008. Bulletin of the Museum of Life Sciences, No. 14: Freeman and Custis Red River Expedition of 1806: Two Hundred Years Later. (pbk.). Museum of Life Sciences, Louisiana State University Shreveport, One University Place, Shreveport, Louisiana 71115-2399, U.S.A. (Orders: www.lsus.edw/lsusmus, lhardy@lsus.edu, 1-318-797-5338, 1- 318-797-5222 fax). 368 pp., 7" x 10". Contents: Preface (by the editor) D. O R Celebrati F ten Expedition— Dan Flores (University of Montana) 2): Plant Elan and rol of the West Gulf Coastal Plain: An Overview— Barbara MacRoberts and Michael MacRoberts The Promi f Pine in the U West Gulf Coastal Plain During Historical aa Mei Es Bragg m Forest A 4). West Gulf Coastal Plain Pata Ecology: Ice Age to Present —Michael Mach 5). The First People of the Red River: The Caddo Beloreanid After ] Custis—Timothy K. Perttula (Arch. & Envir Consul. LLC), Dayna Bowker Lee (NSULA), aná Robert Cast (Caddo Nation) Tl Ad All Lilo E e E eg PI d DEL 80 i A River Drainages—Malcolm Vidrine (LSU Eunice) Tiscta Ar 1 1 East D: Di] TI oy +1 E T ee + +1 2 Ré f «I T D 1c T + L D d ] Custis Expediti Jeffrey S. Girard, Robert C. Vogel, and H. Edwin Jackson (Northwestern State Univ.) 8). The Fishes of the Lower Red River: 1806 Account of Peter Custis and Present-Day Ichthyofauna— Neil Douglas (Univ. of Louisiana ) Monroe) and Jan Hoover (US Army Corps of Engineers The A bahi J D til Tiha E A E I I Changes to the Present—Laurence M. Hardy (LSUS, Museum of Life Sciences) ). Birds of the F IC Exped 1806 to Present—James Ingold (LSUS) 11). The Dy ics of Louisiana's Past and P tM lian F , in Relati the F ] Custis Expediti f 1806—Brad McPherson (Cent y College) 12). Charting Louisi Mapping & Mi pti den? Wiese alicia ew Orleans Collection) 13). Mapping of the Red River After fi 1 Gary Joi U 14). Gate i Address Making of a History—Dan Flores (University of Montana) 15). Ecology and History of the Red River Raft—Frank J. Triska (US Geological Survey, Menlo Park, CA) 16). Writing a History for Exploration: What became of Thomas Freeman and Peter Custis?— Peter J. Kastor (Washington Univ., St. Louis) Paai | H J. Bot. Res. Inst. Texas 2(1): 454, 2008 THREE NEW SPECIES OF IXORA (RUBIACEAE) FROM THE STATE OF TOCANTINS, BRAZIL Piero G. Delprete Federal University of Goiás, Campus Il Institute of Biological Sciences - ICB-1 e of General oe y 74001-970 Goiánia, Goiá Ee c ABSTRACT uring the preparation of the Rubiaceae treatment for the Flora de Goiás e Tocantins, HERR undescribed species of Ixora were found, all understory shrubs from the state o il). Ixora congestiflora D f two collections, one from the Parque Estaduai do Cantáo, just north of the Ilha ids Bananal, and the other from He state of Maranháo, near the Tocantins River, in semideciducus and gallery forests with Ámazonian influence. Ixora g Delp ] ee mm two Geer de northern Tocantins, near the Araguaia River. Ixora irwinii Delprete is | by a few collecti Tocantins. The tl peci | described and ill ted. A key to the native species of Goiás and Tocantins is included. KEY WORDS: Ixora, Ixoroideae, Rubiaceae, Tocantins, Brazil, Amazon Basin RESUMO Durante a EH do NM das Aa E a iie de Goiás e Tocantins, foram las tré pécies de Ixora, sendo cl | AA PRE my | Ta.) z 1 - 1 Tocantins g por duas coletas, uma no o Parque eege do Cantáo, ao norte da Ilha do Bananal, Estado de M háo, perto do Rio Tocantins, em A y Than] PR 1 k | p li- ge no norte e o de Tocantins SES do Rio Araguaia. Ixora irwinii Delprete é conl porp let florestas d Tocantins. Ás t D I ilustradas. Uma chave para as espécies nativas dos Estados EN Goiás e Tocantins é incluída. PALAVRAS CHAVE: Ixora, Ixoroideae, Rubiaceae, Tocantins, Brasil, Bacia Amazónica. The state of HE was ad from the state of Goiás in 1988, and since then the Brazilian government encouraged its econom t through the creation of new cities and new roads, and by stimulating immigration from other Prom states. For example, Palmas, the state capital and the newest city in Brazil, in 2007 has already grown to more that 200,000 inhabitants, and many other new urban agglomerations have been founded throughout the state during the last decade. As for the vegetation, the southern por- tion of the state is located within the Brazilian Savannah (Cerrado Biome). Whereas, the central-northern portion of the state is an area of transition between the Savannah and s Amazon Basin, with a floristic composition from both biomes and unique to this vegetation. Thi larly true for the floristic composition of the gallery forests geg water courses flowing northwards, which act as migratory routes connected with the Amazon Basin. This region continues to be botanically little-known, and only recently has received some attention through several botanical expeditions organized by local institutions. At the same time, the entire state is under rampant anthropic pressure due to aggressive and extensive farming, construction of new state roads leading to remote areas, and massive human migration that took place dur- ing the last two decades. In addition, even the potentially protected areas are not safe from this ongoing environmental destruction, as it is common to see large herds of cows grazing among the vegetation of the state and national parks present in this state. During the preparation of the Rubiaceae treatment for the Flora de Goiás e Tocantins, three interesting collected in the forests of the central- species of Ixora were encountered. All of them are understory shru bs northern portion of the state of Tocantins. After consulting all the available literature dedicated to South J. Bot. Res. Inst, Texas 2(1): 455 — 462. 2008 ` D fal Dos D in LI E do nd £Tawae 274 d 456 American I (Muller Argoviensis 1875, 1881; Standley 1937; Steyermark 1967; Delprete 2003, 2007, sub- mitted; Delprete & Cortés-B. 2007; Delprete et al. 2005; Taylor & Steyermark 2004), and comparison with the specimens present in herbaria with large Brazilian collections (see acknowledgments), it was concluded that they are, in fact, new to science. One species, Ixora congestiflora Delprete is known from two collections, one from the Parque Estadual do Cantáo, just north of the Ilha do Bananal, and the other from the state of Maranháo, near the Tocantins River, in semideciduous and gallery forests with Amazonian influence. The second species, Ixora araguaiensis Delprete is also known from two collections from northern Tocantins, near the Araguaia River. The third species, Ixora irwinii Delprete, is known from a few collections from gal- lery forests in northern Tocantins. With these three new species, described and illustrated E and a few —— lenar unpublished ies (Delprete, submitted), the number of Neotropical s] f PI 4 4 45, ipsis is the same number earlier estimated by Delprete (2003). 1. Ixora congestiflora Delprete, sp. nov. (Fig. 1). Ty: BRAZIL: Tocantins: Mun. Caseara, Parque Estadual do Cantão, transecto 11, floresta estacional semidecídua, 09°18'00"S, 50%01'57"W, 15 Jan 2000 (fl), PE. Nogueira & M. Richter 692 (HOLOTYPE: IBGE; isorvee: NY). a ae did in Arg. as similis EE EE et paure, differt bracteis subtendentibus nullis (nec bracteis ), et corollis 13-14.5 mm longis (nec 7.5-8 mm longis). Shrub to 2 m tall; branchlets glabrous. Stipules besally connate, 3.5—8 x 3-4 mm, base ovate, 1.5-4 mm long, arista 2-4 mm long, glabrous outside, densely sericeous-pubescent and intermixed with col- leters inside. Leaves opposite, petiolate; petioles 5-15 mm long, glabrous; blades obovate to oblanceolate, (3.5-)7.3-12.5 x (1.22)3—4.5 cm, base acute or decurrent, apex obtuse to acute, or acuminate (acumen del- toid, 5-7 mm long), subcoriaceous, glabrous throughout; secondary veins 12-18 each side, subsecondary veins parallel, about the same number as the secondaries. Inflorescences terminal, subpedunculate, very compact, reduced-trichotomous, glabrous throughout, 21-30-florous; the three branchlets 1-2 mm long, ending with three 2-3(-4)-florous cymules; bracts subtending the cymules ovate-acuminate, 1.5-2 mm long; bracteoles 1 or 2 each flower, ovate, 0.5-0.7 mm long, glabrous. Flower buds with tube narrowly- cylindrical, and fusiform in upper part (lobed portion). Flowers sessile. Hypanthium obovoid, 0.5-0.7 mm long, glabrous. Calyx cupular; tube 0.5-1.4 mm long; lobes unequal, narrowly triangular, 0.5-1.2 mm long, glabrous. Corolla hypoerateriform, 13-14.5 mm ee El es cream-white internally; tube narrowly-cylindrical, 8—8.5 x 0.7-0.9 mm, glal y-lanceolate, 5—6.5 x 1.5-1.8 mm, apex acuminate. Stamens gen else among ER Seet lobes; aliens oblong, ca. 3 mm long. Style exserted just above the mouth, 9.5-10 mm long; style branches narrowly oblong, 0.7-1 mm long (not fully expanded). Fruits unknown. Distribution and ecology.—Known only from two collections, as a shrub growing in semideciduous seasonal forest in the Parque Estadual do Cantáo, near the Araguaia River, in the municipality of Caseara, and the other from the state of Maranhão, in gallery forest near the Tocantins River (on the other side of the state of docu and an area soon to Be submerged because of the construction of a T vations. imilar to Ixora Sege Mull. nci (Müller Argoviensis 1875, 1881; Delprete 2007), because of the Become paucifl infl but differs from it by the absence of subtending bracts (vs. subtending bracts foliose, oblong-elliptic, 5-12 x 23 mm in I. bracteolaris), floral buds fusiform in the upper adi (vs. ovoid in the upper part), and corollas 13-14.5 mm long (vs. 7.5-8 mm long). Addit i: BRAZIL: M Mun. Estreito, left margin of Tocantins River, mouth of Rio Feio, 140 m, 06°44'15"S, 47°29' 26""W, 14 Jan 2008 (fl), G.P. Silva & G.A. Moreira 12598 (CEN, UFG). 14 2. Ixora Asaguatensis DE Preta, Bp nov. I (Fig nd Tyre: BRAZIL: Tocantins: Mun. Pium, Ilha do Bananal, P Araguaia, P Amazonia, 09°50'57"S, 50°11'31"W, 190 m, 26 Mar 1999 (fb, M.A. da Silva, ). R.C. Mendonca, E. Cardoso, A D. dos Santos, X G. Sousa, N.R. Oliveira & J.T. dos Santos 4164 (noLoTYE: IBGE; isorvee: NY T 7 app hel + = sr H "wl Ee à EX 1 1 det a sp 4 AE : ni Frutex 1 m alta statura arboris ad 20m des. inflorescentis 12—18-floris (nec 45-150-floris), t cylindrico), venis secundariis foliorum utrinque 10-15 (nec 15-40). X 11 J 4 + f Eas] 11 457 458 SE < jm Se A KR pst w^ pe ) ds E = m 3 q = Sa = bh ea : E so E BA, = = la q = £ KA. c E E E [^ " ^ “a N = y = VG t M.A. da Silva et al. 4164, NY). Delprete, TI i f | f Brazil 459 Shrub ca. 1 m tall; branchlets glabrous. Stipules basally connate, 3.5-8 x 3-4 mm, base ovate to triangu- lar, 1.5-2.5 mm long, arista 1.5-2.5 mm long, papyraceous, glabrous outside, densely sericeous-pubescent and intermixed with colleters inside. Leaves opposite, petiolate; petioles 7-15 mm long, glabrous; blades oblong-elliptic to oblong-oblanceolate, 7.5-11 x 1.7-3 cm, base acute to decurrent, apex obtuse to acute, or shortly acuminate (acumen, when present, deltoid, 3-5 mm long), subcoriaceous, glabrous throughout; secondary veins 10-15 each side, subsecondary veins reticulate (rarely subparallel) and 2-4 for each sec- ondary vein. Inflorescences terminal, pedunculate, corymbose, erect-pubescent, with 2—4 pairs of lateral branches, 12-18-florous; branchlets 5-18 mm long, ending with three 3(4)-florous cymules; bracts subtend- ing the cymules oblong-acuminate, 1.5-3 mm long; bracteoles 1 or 2 each flower, narrowly oblong, 0.5-1.5 mm long. glabrous. Flower buds with tube cylindrical, and ellipsoid in upper part (lobed portion), pale yellow. Flowers sessile or pedicellate; pedicels (when present) 1-3 mm long. Hypanthium obovoid, 1-1.3 mm long, glabrous or sparsely puberulent. Calyx cupular, 0.3-0.5 mm long, truncate or undulate, rarely with shallowly triangular lobes, glabrous. Corolla hypocrateriform, 7,5-8 mm long, white; tube gradually narrowing toward the middle, 3-3.5 mm long, 1-1.3 mm wide at base and at mouth (0.5-0.7 mm wide at middle portion), glabrous outside, sparsely pubescent at mouth internally; lobes elliptic-ovate, 4.5—5 x 1.7-2 mm, apex obtuse or round. Stamens exserted among the corolla lobes; anthers narrowly oblong, 1.8-2 mm long, acuminate. Style exserted well above the mouth, 6—7 mm long; style branches lanceolate, ca. 2 mm long. Fruits unknown. Distribution and ecology. — Known only from the type material and an additional collection, near the Araguaia River, as an understory shrub growing in sandy soil, in northern Tocantins. Observations.—To my knowledge, Ixora araguaiensis is unique within the genus because of its bifid Supe, as Ixora is SE EE by stipules with one arista. In several other species, older k up irregularly into two units, but this is not the case in I. araguaiensis, because its stipules have pe aristae (sometimes basally connate) since very early stage (Fig. 2B-C). However, the consistency of this character within the species remains to be confirmed, as it is currently known from only two collections. Conservation.—A rare species collected in the Parque Nacional do Araguaia, in the municipality of Pium. Although it was collected inside a national park, this species should be included in the category with imminent threat of extinction, due to the presence of large herds of cows grazing on the vegetation of this park, especially during E season (May—September). Etymology. —TI pithet of this taxon is dedicated to both the river and the national park where it was pad vations.—Thi ies is similar to Ixora brevifolia Benth. (Delprete 2003, 2007) because of the terminal, corymbose to peda inflorescences, but differs by being a shrub ca. 1 m tall (vs. tree to 20 m tall in I. brevifolia), with 12—18-florous inflorescences (vs. 45—150-florous), corolla tube gradually narrowing toward the middle (vs. cylindrical) and leaf blades with 10—15 secondary veins each side (vs. 15—40 veins each side). ^dditi ined: BRAZIL: Tocantins: Araguaia River, right margin, várzea vesetation, 12 Aug 1978 (fD, N.T. Silva 4800 (NY). 1 D: E z AC + A A sa ID a] 3. Ixora irwinii Delprete, sp. nov. (Fig. 3). Tres: BRAZIL: Tocantins: 1 km S of Araguaína, at Rio das Lontras, 300 m, 15 Mar 1968 (fr), H.S. Irwin, H Maxwell & D.C. Wasshausen 21221 (HoLotypE: UB; isotype: NY) “1.2. EI I ` 4-1 . 1 - 11 Pic € Hu DI Ixorae SE limi: ex Schult. & Schult. f. si I ibus), laminis foli 4.5-11.5 cm jonsis (nec 18-30 cm logia); venis secundariis iride 10-15 (nec 20-25). thachidi glabra ( d hirsuta), corollis 9-10 mm longis (nec 15-17 mm). Shrub 0.8—1 m tall; branchlets glabrous. Stipules basally connate, 3.5-8 x 3-4 mm, base ovate, 2-3.5, arista 1.5—2.5 mm long, glabrous outside, densely sericeous-pubescent and intermixed with colleters inside. Leaves opposite, petiolate; petioles 7-15 mm long, glabrous; blades elliptic, oblong-elliptic to obovate, 4.5—11.5 x 1.5-4.5 cm, base acute to decurrent, apex obtuse to acute, or acuminate (acumen deltoid to nar- rowly triangular, 5-12 mm long), subcoriaceous, glabrous throughout; secondary veins 10-15 each side, TL lexas 2(1) > da fT 460 R Flower bud CO H.S. Irwin et al. 21221, NY) Fis. 3. f IFS 5n627.F£ Li [A má UA Delprete, Tk | j f | f Brazil 461 H A A 4 H CH " $ 1 , 4 Heal and al ] 1 ] J T , sessile Or pedimculate, carymibiose to short-paniculate, glabrous throughout, with 2—3 pairs of lateral branches. (27—-)30—75-florous; branchlets 7-18 mm long, ending with three 5—7-florous cymules; bracts subtending the cymules ovate-acuminate, 2-3 mm long; bracteoles 1 or 2 each flower, ovate, 1.5-2.5 mm long, glabrous. Flower buds with tube narrowly cylindrical, and fusiform in upper part (lobed portion). Flowers sessile or subpedicellate; pedicels (when present) to 1 mm long. Hypanthium obovoid, ca. 1 mm long, glabrous. Calyx cupular; tube ca. 1 mm long; lobes 4, unequal, linear to deua 0. J 1.5 mm long, glabrous. Corolla hypocrateriform, 9— SR mm long, dd ea cream-white int ly-cylindrical, 4-4.5 x 0.5-0.7 mm, glal ly-lanceolate, 4.5—5 x L 7-2 mm, apex acuminate. Stamens exserted and reflexed ono the atoll lobes; anthers oblong, 2.8-3 mm long. Style exserted just above the mouth, 5.5-6 mm long; style branches narrowly oblong, 0.5-0.7 mm long (not fully expanded). Fruits subglobose to oblong-ellipsoid, subdidymous, 6.5-7 x 5-5.5 mm, glabrous, passing from red to vinaceous to almost black at maturity. Distribution and ecology.—Known only from a few collections as an undestory shrub growing in gallery forests of northern Tocantins. Conservation.—A rare species in need of urgent protection because the few collections known are from areas without formal protection or from farm land. Etymology. —The specific epithet of this taxon is dedicated to Howard Irwin, indefatigable collector who worked for the New York Botanical Garden, organized and carried out the project Flora of the Brazilian Planalto during 1964-1975, and is the first collector of this species. Additional specimens examined: BRAZIL. Tocantins: Faz. Neto, Rio Agua Boa, 8°12'52"S, 48°09'57"W, 12 Jan 2001 (fl), S.F. Lolis et al UFG); Mun. Tanqueira, floresta de galeria, 8°11'49"S, 48?08'48"W, 16 Dec 2001 (fl), S.F. Lolis et al. 357 (HTO, UFG); Mun. Paraíso do Tocantins, Fazenda Uberaba, 10°09'36"S, 48°55'32"W, 14 Dec 1999 (Fl), Neves et al. s.n. (HTO 7405). Taxonomic observations: This species is similar to Ixora pubescens Willd. ex Schult. & Schult. f. in Roem. & Schult. because of the terminal, paniculate inflorescences and flower buds fusiform in the upper part (lobed portion), but differs by having glabrous vegetative branchlets (vs. puberulent in I. pubescens), leat blades 4.5—11.5 cm long (vs. 18-30 cm long) with 10-15 secondary veins each side (vs. 20-25 secondary veins each side), glabrous rachis (vs. densely hirsute), and corollas 9-10 mm long (vs. 15-17 mm long). KEY TO THE NATIVE SPECIES OF IXORA IN THE STATES OF GOIÁS AND TOCANTINS, BRAZIL 1. Inflorescence very compact, with branches 1-2 mm long I. congestifiora 1. Abi not as compact, with branches more than 7 mm long. . Hower buds oblong-ellipsoid in upper part (tip round to obtuse). S Sue ca.1m tall leaf blades with 10-15 secondary veins on each side; inflorescence 12-18-florous; rona LUPE Qrdaduall y Fiat TO WITIQ toward the midale l. araguaiensis 3. Tree to 20m tall leaf blades with 15-40 secondary veins on each side; inflorescence 45-150-florous; corolla tube cylindrical 2. BONNER Sues Ron in Bet part (tip AE o P | 5 cm long, with 10-15 secondary veins on each side; . breviflora rachis g'aorous, corollas 2 10n Tit ong L irwinii | f blades 18-30 cm long and with 20-25 secondary veins on each 3405 rachis densely hirsute: corollas 15-17 mm long I. pubescens ACKNOWLEDGMENTS This research was realized during a fellowship for Visiting Scientist from the National Council for Scientific and Technological Development (Conselho Nacional de Desenvolvimento Científico e Tecnológico - CNPq) of the alan Government (grant 309885/2003-5), at the Federal University of Goiás (UFG), with the coordination of Vera Lúcia Gomes-Klein (UFG). My gratitude also goes to Rob van Aubel (U), Hendrik Ryp- kema (U) and Kimberly Watson (NY) for help in finding and sending relevant taxonomic literature, and to Lubbert Westra (U) for the Latin diagnoses and reviewing the first draft of the manuscript. The directors 462 and curators of the following herbaria are kindly acknowledged for loan of material, sending digital images, and/or providing working space during my visits: BR, CPAP, CTES, F, G, G-DC, HTO, IBGE, K, MBM, NY, NX, R, RB, S, U, UB, UFG, UFMT, and US. Finally, I am much obliged to Petra De Block (BR) and Joseph Kirkbride (USDA) for the constructive comments that much improved the text and the scientific content of the manuscript. REFERENCES Detprete, PG. 2003. Revision and typification of some species of Ixora (Rubiaceae) from central and southern Brazil. Sida 20: 1471-1480. Deterete, PG. 2007. Ixora (Rubiaceae). In: M.G.L. Wanderley, G.J. Shepherd, and A.M. Giulietti, coords. Flora fanero- gámica do Estado de Sáo Paulo, Editora Hucitec, Sáo Paulo, Brazil. Vol. 5:354-358, fig. 12K-R. Deerere, PG. Rubiaceae, Part I: Introduction, key to genera, and genera from A to L. In: MJ. Jansen-Jacobs, ed. Flora of the Guianas. Kew Publishing, Royal Botanic Gardens, Kew, Richmond, Surrey, U.K. (submitted) Devprete, PG. and R. Corrés-B. 2006 [2007]. A synopsis of the Rubiaceae of the states of Mato Grosso and Mato Grosso do Sul, Brazil, with a key to genera, and a preliminary species list. Rev. Biol. Neotrop. 3(1):13-96. Detprete, PG, LB. Sam, and R. Kuen. 2005. Rubiáceas, Volume II - Géneros de H-T: 20 Gardenia até 46. locoyena. In: A. Reis (Editor), Flora Ilustrada Catarinense. Herbário Barbosa Rodrigues, Itajaí, Santa Catarina, Brazil. Pp. 345-842. MULLER Arcoviensis, J. 1875. Rubiaceae brasilienses novae. Flora 58:449—459. MULLER ARGOVIENSIS, J. 1881. /xora (Rubiaceae). In: C.P. Martius, A.G. EicHLER & l. URBAN (Eds.), Flora Brasiliensis. F. Fleisher, Lipsia, Germany. Vol. 6(5): 57-75, 458-460, figs. 9-10. SrANDLEY, PC. 1937. Ixora (Rubiaceae). Studies of American plants — VIII. Field Mus. Nat. Hist., Bot. Ser. 17: STEYERMARK, J.A. 1967. Ixora (Rubiaceae). In: B. MAGUIRE AND COLLABORATORS, Flora of the Guayana Highlands — Part VII. Mem. New York Bot. Gard. 17:341-353. Taylor, C.M. and J.A. STEYERMARK. 2004. Ixora (Rubi ).In: J.A. Steyermark, PE. Berry, K. Yatskievych, and B.K. Holst, eds. Flora of the Venezuelan Guayana. Missouri Botanical Garden Press, St. Louis, MO, USA. Vol. 8:628-631. LASIAMBIX DOMINICENSIS GEN. AND SP. NOV., A EUDICOT FLOWER IN DOMINICAN AMBER SHOWING AFFINITIES WITH FABACEAE SUBFAMILY CAESALPINIOIDEAE George O. Poinar, Jr. Kenton L. Chambers Department of Zoology Department of Botany and Plant Pathology Oregon State University Oregon State University Corvallis, Oregon 97331, U.S.A. Corvallis, Oregon SE JUSA Alex E. Brown 629 Euclid Avenue Berkeley, California 94708, U.S.A. ABSTRACT Lasiambix dominicensis gen. & sp. nov., represented by six floral specimens at different stages of development, is described from Tertiary Dominican amber. The genus is characterized by small, perigynous, bilaterally-sy trical fl cup-shaped hypanthium, 5 ovate-deltate spas à ge early-deciduous petals, 3 short, incurved stamens, each taled to the pandil cup opposite a sepal, and a superior, simple pistil attached ET within the hypanthium, the style positioned excentrically opposite the 1 bet 2 of the sepals. I f ne a short, thick stipe and rugose surface. The combination of floral it ition in Fak bfamily Caesalpinioideae, but with uncertain tribal affinity. pere] i Key Wonps: amber, Dominican Republic, Caesalpinioideae, eudicot flower, Fabaceae, Greater Antilles, Tertiary RESUMEN E I -1 T a us J sos = & sp nov ] E i £1 ] FRE ra tard ee | 11 , del o I r f] r bar terciario d ini El periginas de copa, 5 sépalos ovado- deletes 5 pétalos es, eege caducos, 3 estambre cortos, incurv ados, adheridos a la copa del opuesto a los estambres y r h: co ii los sépalos, v un Lan dde NL unido | | hipanto I I INTRODUCTION Amber is well known for its preservative qualities of both animal and plant remains (Poinar 1992, Poinar & Poinar 1999). Amber flowers, maintained in three-dimensional form, are often preserved with such detail that they can be assigned to extant genera (Spahr 1993). Such fossils can be useful in paleontological reconstructions (Poinar & Poinar 1999). The majority of described amber flowers originate from Baltic deposits (Spahr 1993); however legume (Dilcher et al. 1992; Poinar 1991) and palm (Poinar 2002a, 2002b) flowers have been described from Do- minican amber, and there are also descriptions of Early Cretaceous flowers in Burmese amber (Poinar & Chambers 2005; Poinar et al. EE During an on; ti T remains in Dominican amber, 6 flowers in 5 separate pieces of amber were TRENT "wich could be linked through the 1 ion of an identical cup-shaped, puberulent hypanthium with deltate-ovate, spreading-recurved sepals. One flower, with petals present, is probably at a pre-anthesis stage; 3 flowers, with petals absent, are at or near anthesis; and 2 flowers show early fruit development, with the SE of Es pee i» d in in EXPO The Esse Diana preservation of the of their unique features. flowers and clarity of t! These flowers, which are described Bes as a new gege Bees a combination of characters that favors an assignment to Fabaceae subfamily Caesalpinioideae. . Bot. Res. Inst. Texas 2(1): 463 — 471. 2008 MATERIALS AND METHODS All 5 pieces of amber containing the fossil flowers originated from mines in the northern mountain range (Cordillera Septentrional) of the Dominican Republic, between the cities of Puerto Plata and Santiago. Dating of Dominican amber is controversial, with the youngest proposed age of 20-15 mya based on foraminitera (Iturralde-Vincent & MacPhee 1996) and the oldest as 45-30 mya based on coccoliths (Cépek in Schlee 1999). Most of the amber is secondarily deposited in turbiditic sandstones of the Upper Eocene to Lower Miocene Mamey Group (Draper et al. 1994). The 6 flowers are similar in the morphology of their floral cup (hypanthium) and are assumed to have originated from t] lant species. The amber piece containing the flower assigned to catalogue number Sd-9-154A (Holotype) is el in outline, measures 13 mm in diameter, 9 mm in greatest thickness and weighs 6 grams. The amber piece containing specimen Sd-9-154B is roughly triangular in shape, measures 12 mm along the sides, 7 mm at the base, is 3 mm thick and weighs 3 grams. The third amber piece (Sd-9- 154C) is rectangular, measures 8 mm by 7 mm along the sides, 2 mm in thickness and weighs 2 grams. The fourth amber piece (Sd-9-154D), which contains the specimen with the petals, is semicircular in outline, measures 17 mm in greatest length, 9 mm in greatest width, 7 mm in greatest thickness and weighs 1 gram. The fifth amber piece (Sd-9-154E), which contains the specimen with the fruits, is trapezoidal in outline, measures 17 mm in greatest length, 12 mm in greatest width, 8 mm in greatest thickness and weighs 1 gram. All specimens are deposited in the Poinar amber collection maintained at Oregon State University. DESCRIP1ALON Lasiambix Poinar, Chambers & Brown, gen. nov. Tyre species: Lasiambix dominicensis Poinar, Chambers & Flowers small, bilaterally sy trical, perfect (bisexual), with a cup-shaped | thi lyx isosepalous, sepals 5, deltate-ovate, Tee ECH or valvate; corolla isopetalous, pas 2: hos ee stamens 3, arising from rim of hy] opposite 3 adjacent sepals, fil hort, linear, strongly incurved, anthers bilocular, dorsifixed, introrse; y superior, sessile at base jdm dudum cup; es le ER ind exceeding stamens, positioned icall ite t] ics simple, Bot enlarged; fruit (immature obovoid, pen: to a short, thick stipe, MS deciduous. Donum dominicensis Poinar, Chambers is DIN. GEN nox. (Figs. 1-9). Tire: ie DOMINICAN REPUBLIC: t 03, (HOLOTYPE: OSC-cat. no. Sd-9-1544A, Poi ] mm Flowers 3.6-3.7 mm in diameter, bilateral, pedicel none; hypanthium cup-shaped, densely tomentulose een pubes eni within, 1.7-2.1 mm long, the E 14-2.2 mm wide, bearing a distinct rim to ate-ovate, densely tomentulose on both surfaces, spreading with E tip, 0.9-1. b» mm long, L 0- E 6 mm at widest point; petals erect, early-deciduous, 1.3-1.9 mm long, 0.8-1.1 mm wide, oblanceolate, bluntly Ge at Ea Geet Put od ciliate; thial rim, linear, stamens short, incurved, slightly exserted from hypanthium, t thickened at point of insertion, glabrous, the anthers positioned horizontally over the mouth of hypanthium, bilocular, slightly dorsifixed, ovate, blunt-tipped, cordate at base, 0.4-0.7 mm long, glabrous, dehiscence introrse by lengthwise slits; ovary sessile in hypanthial cup, lightly pubescent, style linear, 0.6-1.0 mm long, pubescent proximally, stigma terminal, undivided, not enlarged; fruit (immature) obovoid, probably 1-few-seeded, 4.8-4.9 mm long, 3.7-3.9 mm wide, the short, thick stipe mostly enclosed in the hypanthium, surface thick-textured and rugose (as preserved), style not persistent. Etymology.— Genus name from the Greek “lasios,” woolly, and “ambix cup, referring to the tomentulose hypanthium that unifies the 6 specimens. Species name derived from the Dominican Republic, the country Livy Lili where the fossils were found. Poinar et al 3 fes 1-3. leger omnimus gen. ang pn nov. 1 Ceale har D 970 mm A oC. fi APARA PF 465 style. Scale ZS =0. 82n mm. n o? m 327 lo ee IVE VIR CJ A 4FA£F SE LÀ 466 tani i Texas 2( be, 4—7. Lasiambix dominicensis gen. and sp. nov. 4. id view e accession Wel i P Scale bar= 0 idi mm. 2 nin of stamens in sped- men n Sd- 9- Dia arrow points t to style. = bar 0.28 m I | ebar — 0.77 mm. 7. | g ing fruit; note st t base ( ). Scale bar = 0.57 mm. DISCUSSION +1 1 las 1; iio us] Fa : : s single ——— MÀ — dA in The floral features that s illustrations (Figs. 1-7). BS at different Manes of dle the flowers have a — n la hypanthium, with respect to shape, calyx lobes, and pubescence. All have a truncate base where they broke off of their inflorescences. One specimen with an immature fruit (Fig. 7) retains a pair of stamens attached TC iary eudicot fl 467 Poinar et al., F a E ET mr Sé ud é Ge b a < ES {qed D vw? E q Y NAS ge ho Wé d / EAS DOM v SUN dli die $5 4 Ne he Kt ita, us ary D ax et UN 4 i R * 4 kr e a AI * ~ we E t ‘ Ae WES E SE d » St 3 Es ud Y f fees E et: ore JN 4i, of Va Ze Lä dy j SED CNN IC PS ESL EN Ur S ed EA SÉ, A sep Dip ANN da bi ^ -c 1 1] p. ais: Daf Ee Es, 1 D win ah b Mts y LN SE Pre ALY pe N.N An A mite O od vr EN A El X ^ om MN Wé AS us Än Oe Ve AA MOTO Y PCS. YAT Ag Tar aa BI wé rs -h EE: dem FAA A eh Es A Ry S ANY ex 2 BN E jar ES Wa ^ E wen Pä ss NL. = a br ety. de E ER EE E Es m S e ie, SA, à rire. UNE ES ez TM GC R “a EP a Asia e d. e ap q s Ki I Zë e tea to! br titu Ri i! 3 ka + A & - KK EN E “o > Pa 4 d z E (423 +38 > "e 1 ^ A? x a MESS: B d ad. ES A aa H l E ^ A SS v» d ite ag (5 = A ir Ai es LM tric ft ef AE? TT 46, ES) PEA A df ergi d vi Ze MRE SAS Ze ev vt EE A PALMA IW Me VLL VUE es ae a vr m Nc end RA ez 2 VA KC ACC Dr IRAM E TIE SÉ helyg MÉ > rÀ D KL - e? la KÉ SS te ka £ X x v CH A = 3 AAN A geg é AS - za S Sar Cae H Ka VR MA sni PLE / Urs ps Kik t) PN E Done Ch 217 P 23 IS > / $ Y Wat Di ore, ^ua E PE 8 diii ; Si Mff dy Je Y tut £ AME (t & SC ae, uet dee E TH OO Ab A Ñ 7 JA 2 Ving A a E ^4 SE E Jt (Uk d BZW Ge > De TAVA Emr Wa ains e, A MAI AS FATAN VN Ku Vj Hy A AN Ee E dn Wi V fe Vt EG E: TERA H ils) ^ ae Tae X alle ba, E 4 Lc La 4, s I 3 ul Lë D KR ÉS BSc I CS gef Ze kä : - p fr d ENS SAUNA SEN ag ang > TS CAD $US TAT Aeg PX Ce EEN SS tS po a et mas" ^ ^ qoo. i SW Ia "t. ZZ EE $- WEG i a A EA y A. Ne Rakh ee O "Se d E Ae 3 > 1% eamm. fL ty ENTES AOS "A Mos EE LARA fes SSA: Ch E RARUS M eere Seng AE ^ x A ipo c ps = ATI a Ree heel Ze. eat, wi E cope A^ E Ac E d "cb SA eg AE EE bad TRES = ! q s ARES EA 7 Es fu A A AR =p? d 5 = Je ef oT Zeg SE b tens Oyo eat Qk un fee mo, LX Se TOROS OSS.) Py Ao RAN e ma NS OF LON Y plu he E DES AG PN AS ES AE A VP ED NEAR SS E SALAS FT P Jah ée a ESL LT E Y 4 DE SIE T SED IAS wend Ké EE ee Se MN ERRE SIA n VW SE d Geet? A Pr ^ Nd AS A wge c ut ura PY AAN ey d Hi no iu DS + ZC SS EEN MO KAK ʻi Ga A PATO áp. ivi YO EL T V INN Metas - ou M T LET -£ de A ANSIA KSE Ve: M dh vi: E Tee KE ade AN d ke E (Oé, We EN dy Y EE EN ANS 9 PON M DIN AN. NO ^ii) 2x Su 4 RAY G AN dte ^ Jn WI Gr Fics. 8-9. Holotype of Lasiambix dominicensis gen. and sp. nov. ( ion # Sd-9-154A). 8. Top view. Scale bar = 0.57 mm. 9. Oblique view. Scale bar = 0.57 mm. 468 | | of the Botanical R h Institute of Texas 2(1) opposite the calyx lobes. The specimen in early flower (Sd-9-154D, Fig. 6) has all 5 petals visible, and one is detached at the base, illustrative of what we term an early-deciduous corolla. The petals are clearly seen when the specimen is viewed from the top as well as the sides. The center of this flower is unsuitable for study due to an accumulation there of apparent insect remains and frass. The sharp hypanthial rim on the 3 flowers at or near anthesis shows where the petals presumably were attached, alternate with the sepals. It remains ambiguous whether the sepal lobes were imbricate or valvate in the bud, perhaps because of ditfer- ent degrees of spreading and separation in the samples at hand. Because no slits or pores are evident on the back, sides, or tips of the anthers, we suggest an introrse dehiscence, which is supported by observations of the adaxial side of one strongly inflexed stamen. The nature of the mature fruit is uncertain, although from the shape and thick-textured exocarp of the 2 specimens here (Fig. 7), a non-compressed, l- or few-seeded drupaceous fruit is suggested. Although atypical for members of the Fabaceae, several extant genera of the Cassieae have similar indehiscent, drupaceous fruits. Dialium s.l. is characterized by small fruits consisting of one seed surrounded by a soft, but somewhat dry, edible aril (Gentry 1993). Few-seeded, ovoid, fleshy- or thick-walled fruits are not unusual in Fabaceae subfamily Caesalpinioideae (e.g. Lewis et al. 2005). This fruit-type, together with a well- EES hypanthium, a superior simple pistil, regular corolla, reduced stamen number, and a ium and pistil, are what lead us to suggest assigning the fossil 1 d to this family anid ny The caesalpinioid legumes are extremely diverse in floral characteristics, including numbers of stamens, presence or absence of a hypanthium, numbers and form of petals, length of filaments, form of anthers and stigmas, shape and structure of fruits, floral symmetry, modification to unisexuality, etc. The compendium by Polhill and Raven (1981), with its taxonomic treatments of the relevant tribes (Cowan & Polhill 1981; Irwin & Barneby 1981; Polhill & Vidal 1981), was E ina Sie for modern genera with the particular combination of features seen in the fossils. Illust f genera were examined in Taubert (1894), Hutchinson (1964), and Lewis et al. (2005). No examples have been found that correspond in all respects with Lasiambix, but its described traits are present separately in numerous caesalpinioid genera. A cup-shaped, externally puberulent hypanthium occurs frequently in the subfamily. Bilateral sym- metry (zygomorphy) is also common, and this accompanies a reduction to three stamens in such genera as Tamarindus, Macrolobium, Gilbert lron, Elizabetha, CES Dialium, etc. As discussed by Tucker (1998, 2002) there are parallel evolutionary trer dè | of stamen number in groups of related genera in separate tribes (her examples were Detarieae-Amherstieae and Dialiinae). From the basic number 10, stamens may be reduced to as few as 1 (some Dialium spp.), and during development, some of the stamen primordia that form early may cease their growth and become rudimentary or staminodial (e.g. Gilbertio- dendron, Macrolobium; Tucker 2002). Except for Dialium and Apuleia, the 3-stamen genera mentioned fall into the Amherstieae (including Macrolobieae) clade, as defined by analysis of the chloroplast trnL intron (Bruneau et al. 2001). This group is also recognizable in the rbcL analyses by Doyle et al. (1997) and Kajita et al. 2001). However, the corolla evolution in these particular genera leading to loss of all but one petal to form an adaxial vexillum (e.g. Cowan 1953) , the 2-many-seeded, mostly dehiscent pods, plus the usual absence of a hypanthium, argue against a relationship to Lasiambix. Cynometra, also in this clade, is discussed below. Dialium and Apuleia, of subtribe Dialiinae, show some blance to Lasiambix in their zygomorphic, few-stamened flowers and their few-seeded, indehiscent fruits. According to Irwin and Barneby (1981, p. 101) Dialium “can be defined accurately and rigorously by its pod, a sort of indehiscent, 1- or 2-seeded nut, sometimes strongly, sometimes scarcely compressed laterally, consisting of a crustaceous exocarp and a pulpy, when ripe mealy endocarp enveloping the seeds." In their key (p. 100) the fruit is termed “drupa- ceous." The flowers of Dialium, a pantropical genus, are hypogynous and have only 1 or 2 petals and usually 2 stamens. The related Apuleia, of South America, possesses a hypanthium and 3 stamens, but its pistil is exserted, not sessile, its perianth is 3-merous, and its pod is compressed and coriaceous, not ovoid. Despite bix dominicensis, V—— E "S 469 = H d Poinar et al., | their few-seeded, indehiscent pods, neither of these few-stamen genera of Dialiinae suggests a close affinity to Lasiambix. Other possible relationships of the fossils are with genera having regular flowers and more numerous stamens but with a similar type of few-seeded fleshy fruit. One such example is Vouacapoua of tribe Caesal- pinieae (Lewis et al. 2005). This lar flowers with 10 short stamens, but its hypanthium, sepals, and short petals much sale bio and its fruits are round, thick-walled, 1-seeded, and dehisce by a single furrow (Lewis et al. 2005, p. 161; Polhill & Vidal 1981, p. 89). The genus is presumed to have affinity to the Peltophorum group of tribe Caesalpinieae (Bruneau et al. 2001; Haston et al. 2003), but as yet the DNA support for this connection is weak. A similar type of fruit occurs in the large genus Cynometra of tribe Detarieae (Lewis et al. 2005, p. 89). Its pod is described as thick, turgid, rugose to verrucose (rarely smooth), 1-seeded, and regularly in- dehiscent with a spongy pericarp (Hutchinson 1964, p. 235; Dwyer 1958); it lacks a floral disc, its ovary is usually hairy, and its style may be eccentric in the hypanthium (Dwyer 1958, p. 322). However, it has 10 long-exserted stamens, a shorter hypanthium, and a usually showier perianth than in Lasiambix (Dwyer 1958). Other differences, besides these, are that its sepals are glabrous adaxially (except C. cubensis Rich.) and its filaments are united very briefly at the base. Modern-day Arcoa, of Hispaniola, has turgid, 1-seeded, indehiscent fruits (Lewis et al. 2005, p. 132), but its flowers differ considerably from Lasiambix in being unisexual, bearing ca. 12 stamens, PA p an exserted pistil with capitate stigma (Herendeen et al. 2003). Another Caribbean genus, Stahli Caesalpinieae), has fleshy, 2-seeded, indehiscent fruits, regular 5-merous perianth, imbricate sepals, eg well-dev pel. dorsally puberulent petals (Hutchinson 1964, p. 235; Lewis et al, 2005, p. 145). It differs from Lasiambix principally in having 10 erect, connivent stamens with filaments ely woolly in the lower half. Placement of Stahlia near the large genus Caesalpinia and its segregates was established by the molecular and morphological studies of Simpson et al. (2003). The examples mentioned, falling into several distinct clades within the subfamily, illustrate the paral- lelisms in floral and fruit morphology that Ge it difficult to Mg a EE definitive placement of Lasiambix. Alarge amount of recent phylog h, both 1 molecular, was reviewed by Lavin et al. (2005), to estimate the time frame of early ven of this and other subfamilies of Leguminosae. Earlier estimates of a Late Cretaceous origin of the family and a “basal” position for Caesal- pinioideae (e.g. Polhill et al. 1981; Herendeen et al. 1992), are no longer accepted as “conventional wisdom. Rather, the fossil record supports an origination in the Paleocene, allowing some 60 my for diversification within the family (Lavin et al. 2005). In molecular phylogenetic studies, with best resolution provided by the matK chloroplast gene, divergence of 6 early-formed lineages of caesalpinioids was found, along with the similarly old crown nodes of many papilionoid and mimosoid clades. Caesalpinioids are viewed as a paraphyletic grade from which stem the mimosoids and papilionoids, but because of the very early, rapid branching of the ancestors of modern clades, the caesalpinioid grade “harbor(s) neither the oldest diversi- fication nor some other quality of legume LÀ (Lavin et al. 2005). If our suggested placement of Lasiambix is correct, it offers a well-preserved example of several special- ized floral characteristics of Tertiary-age caesalpinioid legumes. These include the loss of all but 3 stamens and accompanying bilateral symmetry of the flowers, an ins of peas at anthesis, and presence of few- seeded, fleshy, probably indehiscent fruits. What polli ted by flowers with short, incurved stamens and no apparent nectar disc, is difficult to m Perhaps Seia was produced at the base of the ovary, which was not visible to us. In this case small bees might have been able to enter the flowers and remove pollen from the arched, introrse anthers. The ecology of the plants is likewise unknown, other than that they grew in proximity to Hymenaea in moist tropical forests such as those described by Poinar and Poinar (1999). Such habitats are opes for many EE of modem: day Caesalpinioideae. Note: In the illustrations, becau e orientation of | , we have not tried to suggest a placement of the stamens as either abaxial, as in Macrolobium et al. (Cowan 1953) or adaxial as in Dialium et al. (Tucker 1998). 470 J loft tanical h Institute of Texas 2( ACKNOWLEDGMENTS The authors thank Richard Halse, Aaron Liston, and Gerald Carr, Botany Department, Oregon State Uni- versity; John L. Strother, University Herbarium, and Margriet Weatherwax, Jepson Herbarium, University of California, Berkeley; Robert Thorne, Rancho Santa Ana Botanic Garden; Lawrence Skog, Smithsonian Institution; Peter Endress, Institute for Systematic Botany, University of Zúrich; and Boris O. Schlumpberger, Systematic Botanical Unit, Ludwig-Maximilians-Universitát, Múnchen, for discussions concerning the place- ment of these fossil flowers. We are also grateful to Steven Manchester, Roberta Poinar and two anonymous reviewers for comments on earlier versions of the manuscript. REFERENCES Bruneau, A. E Forest, PS. HERENDEEN, B.B. KLITGAARD, and G.P. Lewis. 2001. Phylogenetic relationships in the Caesalpin- ioideae (Legum inosae ) as inferred from chloroplast trnL intron sequences. Syst. Bot. 26:48/-514. Cowan, R.S. 1953. A taxonomic revision of the genus Macrolobium (Leguminosae-Caesalpinioideae). Mem. New York Bot. Gard. 8:257-342. Cowan, R.S. and R.M. PolHILL. 1981. Detarieae and Amherstieae. In: Polhill, RM. and PH. Raven, eds. Advances in legume systematics: part 1. The Royal Botanic Gardens, Kew. Pp. 117-142. Dit cHER, D.L., PS. HERENDEEN, and F. Hueser 1992. Fossil Acacia flowers with attached anther glands from Dominican Republic amber. In: PS. Herendeen and DL. Dilcher, eds. Advances in legume systematics: part 4: the fossil. record. The Royal Botanic Gardens, Kew. Pp. 33-42. Doyte, LL J.L. Dove, J.A. BALLENGER, EE Dickson, T. Kasma, and H. OHAsHI. 1997. A phylogeny of the chloroplast gene rbcL in the Leguminosae: taxonomic correlations and insights into the evolution of nodulation. Amer. J. Bot. 84:541—554. Draper, G., P. Mann and J.F. Lewis. 1994. Hispaniola. In: S. Donovan and T.A. Jackson, eds. Caribbean geology: an introduction. The University of the West IndiesPublishers' Association, Kingston, Jamaica. Pp. 129-150. Dwyer, J.D. 1958. The New World a of Ge mu ipud Bot. Gard. 45:313-345. Gentry, AH 1993. A field guid ly plants of Northwest South America (Colombia, Ecuador, Peru) with element notes on bert taxa. The University of Chicago Press, Chicago. Haston, E.M., G.P. Lewis, and J.A. Hawkins. 2003. A phylogenetic investigationof the Peltophorum group (Caesalpin- ieae: Leguminosae). In: Klitgaard, B.B. and A. Bruneau, eds. Advances in legume systematics: part 10: higher level systematics. The Royal Botanic Gardens, Kew. Pp. 149-159 pot > >» j] ` +l | I HERENDEEN, PS., G.P. Lewis, and A. Bruneau. 2003. Floral morphology in C | g testing phyly of the "Umtiza clade" Int. J. Pl. Sci. 164(55):5393-5407. HERENDEEN, P.S., W.L. Creper and D.L. Di.cHER. 1992. The fossil history of the Legumi hyl tic and biogeo- graphic implications. In: Herendeen, PS. and DL. Dicher eds. Advances in legume systematics: part 4. The fossil record. The Royal Botanic Gardens, Kew. Pp. 303-316. HUTCHINSON, J. 1964, The genera of flowering plants. Dicotyledones. Volume 1. Oxford University Press. Irwin, H.S. and R.C. Barnesy. 1981. Cassieae. In: Polhill, RM. and PH. Raven, eds. Advances in legume systematics: part 1. The Royal Botanic Gardens, Kew. Pp. 97-106. ITURRALDE-VINCENT, M.A. and RIDE MacrHee. 1996, Age and Paleogeographic origin of Dominican amber. Science 273:1850-1852. Kauta, T, H. OHASHI, Y. TATEISHI, C.D. BAILEY, and J.J. Dove. 2001. RbcL and leg phylogeny, with particular reference to Phaseoleae, Millettieae, are allies. Syst. Bot. 26:515-536. Lavin, M., PS. HERENDEEN, and M 2005. Evolutionary rates analysis of Leguminosae implicates a rapid diversification of lineages during the Tertiary. Syst. Biol. 54:5/5-594 Lewis, G., B. Schrire, B. MACKINDER, and M. Lock, eds. 2005. Legumes of the world. The Royal Botanic Gardens, Kew Pomar, JR., G.O. 1991. Hymenaea protera sp. n. (Leguminosae, Caesalpinioideae) from Dominican amber has African affinities. Experientia 47:1075-1082. tfl 471 Poinaretal Lasi j ini i f F wë i d Pomar, JR, G.O. 1992. Life in amber. Stanford University Press, Stanford. Pomar, JR, G.O. and K.L. CHAMBERS. 2005. Palaeoanthella huangii gen. and sp. nov. an early Cretaceous flower (Angiospermae) in Burmese amber. Sida 21:2087-2092. POINAR, Jr, G.O., K.L. CHamBers, and R. BuckLey. 2007. Eoépigynia burmensis gen. and sp. nov, an early Cretaceous eudicot flower (Angiospermae) in Burmese amber. J. Bot. Res. Inst. Texas 1:91-96. POINAR, JR, G.O. and R. Pomar. 1999. The amber forest. Princeton University Press. Princeton. Pomar, Jr, G.O. 2002a. Fossil palm flowers in Dominican and Mexican amber. J. Linn. Soc. Bot. 138:57-61. PoiNAR, JR, G.O. 2002b. Fossil palm flowers in Dominican and Baltic amber. J. Linn. Soc. Bot. 139:361-367. Hot, RM. and P Raven (eds.). 1981. Advances in legume systematics: part 1. The Royal Botanic Gardens, Kew. PoLHILL, R.M. and J.E. Vina. 1981. Caesalpinieae. In: Polhill, RM. and PH. Raven, eds. Advances in legume systemat- ics: part 1. The Royal Botanic Gardens, Kew. Pp. 81-95. POLHILL, RM, PH. Raven, and CH. SriaroN. 1981. Evolution and systematics of the Leguminosae. In: Polhill, RM. and P.H. Raven, eds. Advances in legume systematics: part 1. The Royal Botanic Gardens, Kew. Pp. 1—26. Scher, D. 1990. Das Bernstein-Kabinett. Stuttgarter Beitr. Naturk., C 28:1—100. Simpson, B.B., L.L. LARKN, and A. Weeks. 2003. Progress towards resolving the relationships of the Caesalpinia group Caesalpinieae: Caesalpinioideae: Leguminosae). In: Klitgaard, B.B. and A. Bruneau, eds. Advances in legume systematics: part 10: higher level Dites The Royal Botanical Gardens, Kew. Pp. 123-1. SPAHR, U. 1993. Systematischer katalog graphie der Bernstein-und Kopal-Flora. Stuttgarter Beitr. Naturk,, B 195:1—99. TAUBERT, P. 1894, Leguminosae. In: Engler, A. and K. Prantl, eds. Die natúrlichen Pflanzenfamilien. Ill. 3. Wilhelm —, Engelmann, gee is 70- no Tucker, S.C. 1998, Floral genera Petalostylis, Labichea, a series in f'oral Feed Amer Bot. 85: 184—208. TUCKER, S.C. 2002. Comparative floral ontogeny in Detarieae (Leguminosae: Caesalpinioideae). 2. Zygomorphic taxa with petal and stamen suppression. Amer. J. Bot. 89:888-907. nd Dialium (C | inioid Cassieae), 472 Jou BOOK NOTICE CanLos Ropiricuez RAMÍREZ. 2007. Chipinque: Testigo de la Naturaleza. (ISBN 978-970-9715-42-2, hbk.). Fondo Editorial de Nuevo León, Zaragosa 1300 sur, Edificio Kalos, Nivel C2, Local 202, CP 64000 Mon- terrey, Nuevo León, Mexico. (Orders: www.fondoeditorialnl.gob.mx, (81) 8344 2970 y 71). $150.00, 351 pp., color photographs, 10.5" x 14”. SE text, nnd table style book with hundreds of wonderful color ma ba E the ecology and natural beauty of terrey, Nuevo León, Mexico. The table of contents is listed bel 1. Introducción 2. Historia: el devenir de Chipinque 3. Chipinque: el parque ecológico 4. Entorno natural: hidrología, geología y clima 5. Ecosistemas de Chipinque (Matorral submontano, Matorral rosetófilo, Bosque de encino, Bosque de pino, Bosque mesófilo de montaña) le Chipinque (1 tos, Arácnidoa, Reptiles, Aves, Mamíferos) 6. F 7. Biblio ala 8. Agradecimientos J. Bot. Res. Inst. Texas 2(1): 472. 2008 KRUSCHKE NAMES IN NORTH AMERICAN CRATAEGUS (ROSACEAE): A CORRECTION AND CLARIFICATIONS J.B. Phipps Ihe uo of Western Ontario ment of Biology 11 b » ichmond St. N London, Ontario N6A 5B7, CANADA jphipps@uwo.ca In a recent paper (Phipps 2007) I inadvertently stated an incorrect collection date for the holotype of Cra- taegus pisifera Sarg. It should be 28 September 1901 rather than 26 July 1901 as recorded in Phipps (2007). The remainder of the commentary on C. ddp in ot Ge EE however. With respect to the same paper pecialist at Harvard University, Kanchi Gandhi, indicated " me hat I was in error in piis two lectotypes, as the names with which I was dealing had never been validly published. Since 1 January 1958, in order to be validly published, names must have a type indicated and a type must be a single specimen (see Vienna Code Art. 57.1, 8.1 and 8.2). However, the two names below were designated with a “type” representing specimens from different gatherings. Thus, in order to validate Kruschke's names, holotypes rather than Tees i: have been designated. Accordingly, the two names are validly published below with ap] ate It may also be noted that were I to have ascribed the names to myself the Citations of the relevant names would have been ‘J.B. Phipps’ but I retain ‘Kruschke, also permissible, therefore accepting Kruschke as the author since he provided the validating Latin diagnoses (Art. 46.2) 1. Crataegus mollis (Torr. & A. Gray) Scheele var. incisifolia Kruschke, var. nov. Te: U.S.A.: Wisconsin: Rock Co.: Avon, along Sugar River, 27 May 1947, E.P Kruschke K-47-8 (HoLorYPE: MIL; ISOTYPE: A). Comment.—Kruschke (Milwaukee Public Mus. Public. Bot. 3:128. 1965) cited two collections from May and September 1947, respectively. 2. Crataegus schuettei e var. gigantea pees var. nov. Tyre: U.S.A. Wisconsin: City of Milwaukee, north side. 16 Sep 1942, E.P Kruschke K-42-288 (HoLoTYPE: MIL; 1 E: A) Comment.—Kruschke (Milwaukee Public Mus. ET Bot. 3:75. 1965) cited three collections from Sep 1942, May 1943, and Feb 944, respectively. ACKNOWLEDGMENTS I thank Kanchi Gandhi (GH) and John McNeill (E) for nomenclatural advice. REFERENCE Puipps, J.B. 2007. Kruschke names in North American Crataegus (Rosaceae). J. Bot. Res. Inst. Texas. 1:1011-1014, J. Bot. Res. Inst. Texas 2(1): 473. 2008 £Tauae H1) 474 BOOK NOTICE Davip W. ROUBIK, SHCKO SAKAI, AND ABANG A. Hamm (eds.). 2005. Pollination Ecology and the Rain Forest, Sarawak Studies, Ecological Studies Series, Vol. 174. (ISBN 0-387-21309-0, hbk.; ISSN 0070- 8356). Springer Science Business Media, Inc., 233 Spring Street, New York, New York 10013, U.S.A. (Orders: www.springer.com, service-nyOspringer.com, 1-800-SPRINGER). $139.00, 307 pp., 79 il- lustrations, 12 color plates, 6" x 9 1 Contents: Preface Acknowledgments Contributors ge P n iid e n and Pollination in Rain Forests—David W. Roubik 2. The Canopy Biolog Methods, and Merit—Takakazu Yumoto and Tohru Nakashizuka 3.Soil- elated. Floristic GE in a Hyperdi e Di | I t—Stuart J. Davies, Sylvester Tan, James V. LaFrankie, and Mat- thew D. Potts 4.Plant Ao EN dës Geen Flowering in a Mixed Dipterocarp Forest—Shoko Sakai, Kuniyasu Momose, Takakazu , Abang A. Hamid Karim, Tohru Nakashizuka, and Tamiji Inoue ht in Lambir Hills National Park Rhett D. Harrison 2 6.The ee Pollinator QUA in a Lowland Dipt Kuniyasu M 1 Abang A. Hamid Karim (Apidae, Meliponini) T hi N itsu and Tamiji Inoue D 8. Ho ce in Bornso— Davi W. Roubih 9. Beetle Pollination in Tropical Rain Forests—Kuniyasu Momose 10. Seventy-Seven Ways to Be a Fig: Overview of a Diverse Plant Assemblage—Rhett D. Harrison and Mike Shanahan 11. Ecology of Traplining Bees and Understory Pollinators—Makoto Kato 12. Vertebrate-Pollinated Plants—Takakazu Wee 13.1 Predat f Dij ps | Mi Nak , Takao Itioka, Kuni) M , and Tohru Nakashizul 14. Diversity of Anti-Herbi Def i — Taka Itioka vil ET 15. Coevolution of Ants and Pine d Itino 16. Lowland Trop cal Rain 17. Lambir's forest: The World's Most Di K Tree Assemblage?— Peter S. Ashton 18. Toward the Conservation of Tropical Forests—Tamiji Inoue America: Parallels, Convergence, and Divergence—James V. LaFrankie Appendices Glossary Bibliography Index J. Bot. Res. Inst. Texas 2(1): 474. 2008 THOMAS WALTER TYPIFICATION PROJECT, V: NEOTYPES AND EPITYPES FOR 63 WALTER NAMES OF GENERA D THROUGH Z Daniel B. Ward Department of Botany University of Florida Gainesville, Florida 32611, U.S.A. ABSTRACT "mi C ri sm e fT FOO. 0. 1 1 E nA fl = 1 1 1 dd 1 r^ 1: J en H Thomas Walter, ( J, ) I 5 H o H 1 1; 1 . £n £ix7_l. ? H 1 1 POE: ES + e? are without types. Sq E Q I y] pityp RESUMEN Thomas Walter, el autor de la Flora Caroliniana (1788), no dejó un herbario. Muchas de las especies que describió de las Carolinas y Georgia no tienen tipo. Se seleccionan aquí especimenes que se su ponen representan 63 de las especies de Walter, para servir como neotipos o epitipos. This number of the Thomas Walter Typification Project is a continuation of a report begun previously (Ward 2007c). Here, an additional 63 of Walter's species are addressed, with selection of appropriate neotypes or epitypes. Although the number of Walter neotypes, as treated in this and previous reports of the Project, have been many, even now not all Walter names have been examined. No effort has been made to typify most of the numerous Walter names that are now remembered only in synonymy. And those names not represented by suitable materials in the herbarium of the present participating institution (GH) will be addressed in a subsequent Dd This report and its i iate predecessor (Ward 2007c) select types for 106 Walter names. Within this number, 101 were selected as neotypes, and 5 as epitypes. The largest proportion, or 98, were of South Carolina collections, 22 of them from Berkeley County; 4 were from Georgia, and 2 each from North Carolina and Florida. A third, or 37, bore a total of 43 annotations confirming the label identifications. The typifications are presented here in the format used previously (Ward 2007a, 2007b, 2007c), in alphabetical sequence, using the names given them by Thomas Walter. LYPIFICALTIONS WALTER'S NAME: Daucus divaricatus Walter (p. 114) MODERN NAME: Spermolepis divaricata (Walt.) Raf. Common in eastern SC. Identified (from Walter's diagnosis) by Mathias & Constance (1944: 72). Spm. 40-C, a wispy fragment, was labeled *Daucus" by Fraser, and annotated as "divaricatus Walt." by A. Gray, though one wonders what he saw that was recognizable. Boufford & Wood 23862, 27 May 1988 - GH, from Wadmalaw Island, s. of Charleston, Charleston County, South Carolina, is here selected as NEOTYPE for Daucus divaricatus Walt., basionym of Spermolepis divaricata (Walt.) Raf. Known IsSONEOTYPES: MO, NY. WALTER'S NAME: Delphinium carolinianum Walter (p. 155) MODERN NAME: Delphinium carolinianum Walt. Almost unknown in SC (a single county), frequent in central GA. Fraser/Walter 40-B [1787] - BM, a now- unidentifiable fragment labeled *Delphinium" by Walter and “Carolinianum” by Fraser, was cited by Warnock (1981: 48), as “Type: South Carolina, within fifty miles of Berkeley Co., T. Walter s.n. (BM)? The cited specimen is worthless for identification purposes. Radford 22407, 11 May 1957 - GH (annot. Michael J. | Bot Rac Inct Tavac 2(1): A75 — 486. 2008 476 J lof the Botanical R h Institute of Texas 2(1) Warnock 1979), from 4.5 mi SE of Clarks Hill, McCormick County, South Carolina, is here selected as EPITYPE for Delphinium carolinianum Walt., in support of Warnock's typification. Known IsOEPITYPES: CA, FLAS, GA, NCU, NY. WALTER's NAME: Dianthera ovata Walter (p. 63) MODERN NAME: Justicia ovata (Walt.) Lindau Frequent on the SC coastal plain. There is no specimen. Godfrey & Tryon 545, 14 Jul 1939 - GH nodi Martha Meagher 1972), from Andrews, Georgetown County, South Carolina, i ] lected Dianthera ovata Walt., basionym of Justicia ovata (Walt.) Lindau. Known IsSONEOTYPES: BH, CA, CAS, MO. WALTER'S NAME: Diodia teres Walter (p. 87) MODERN NAME: Diodia teres Walt. Common throughout. There is no specimen. o & Tryon bs i UE 1939 - GH, from Georgetown, Georgetown County, South Carolina, is here selecte | Walt. Known ISONEOTYPES: NY, US. WALTER'S NAME: Doronicum acaule Walter (p. 205) MODERN NAME: Árnica acaulis (Walt.) BSP. Common in eastern SC. No specimen. Hunnewell 12681, 30 Mar o — (annot. Beet eee p Laken aalantard ne Artan tar Dianvearniciaum neni from near Goose Creek, Berkeley County, South Carolina, is Walt., basionym of Arnica acaulis (Walt.) BSP. WALTER'S NAME: Eriocaulon anceps Walter (p. 83) MODERN NAME: Lachnocaulon anceps (Walt.) Morong Common on SC coastal plain. Walter's description was identified by Kral (1966: 319). There is no speci- men. Godfrey & Tryon 1210, 7 Aug 1939 - GH, from 6 mi S of Columbia, Lexington County, South Carolina, is here selected as NEOTYPE for Eriocaulon anceps Walt., basionym of Lachnocaulon anceps (Walt.) Morong. Known ISONEOTYPES: NY, US. WALTER'S NAME: Ervum erectum Walter (p. 187) MODERN NAME: Galactia erecta (Walt.) Vail Frequent in eastern SC. No specimen has been identified. Godfrey 147, 10 Sep 1939 - GH, from 5 mi 5 of Andrews, Georgetown County, South Carolina, is here selected as NEOTYPE for Ervum erectum Walt., basionym of Galactia erecta (Walt.) Vail. Known IsoNEoTYPES: BH, DUKE, F, MO, NY, US. WALTER's NAME: Eryngium integrifolium Walter (p. 112) MODERN NAME: Eryngium integrifolium Walt. Frequent on the SC coastal plain. Sp. 42-D was numbered “716” by Fraser, then labeled “Eryngium nov." by Walter, with “Integrifolium” added by Fraser. The specimen is of inadequate, scarcely identifiable quality. Wilbur & Webster 2861, 31 Aug 1950 - GH, from 6 mi E of Summerville, Berkeley County, South Carolina, is here selected as NEOTYPE for Eryngium integrifolium Walt. Known ISONEOTYPES: MICH, NY, US. WALTER's NAME: Erysimum pinnatum Walter (p. 174) MODERN NAME: Descurainia pinnata (Walt.) Britt. Infrequent on SC coastal plain, unknown inland. The label of spm. 43-A bears “Erysimum” in Walter's hand, and “Pinnatum” in Fraser's; it also carries Fraser's 3-digit number. Although Walter saw this speci- men, better material would have been available to him in the field. Godfrey 3515, 17 Apr 1938 - GH, from Brookgreen Gardens, Georgetown County, South Carolina, is here selected as NEOTYPE for Erysimum pin- natum Walt., basionym of Descurainia pinnata (Walt.) Britt. Known ISONEOTYPES: US. Ward, bg nn YA FP " LE ai L LF A | SA E m Z 477 ZE vi V vi -F WALTER'S NAME: Ethulia uniflora Walter (p. 195) MODERN NAME: Sclerolepis uniflora (Walt.) BSP. Frequent in eastern SC. Spm. 15-F ens to be this (even though it atypically bears 2 heads). The label (“Ethulia Uniflora”) is in Fraser's hand. S th evidence Walter saw or used this specimen, Ahles 53495, 25 May 1960 - GH, from 3 mi SW of Moncks OIM AE Oy County, South Carolina, is here selected as NEOTYPE for Ethulia uniflora Walt., basionym of t iflora (Walt.) BSP. Known ISONEOTYPES: CA, FLAS, GA, MICH, NCU, NY, US, USCH. WALTER'S NAME: Eupatorium incarnatum Walter (p. 200) MODERN NAME: Fleischmannia incarnata (Walt.) King € H. Robins. [= Eupatorium incarnatum Walt Rare in SC (but incl. Berkeley Co.). Spm. 46-D was identified as Eupatorium incarnatum by Fernald & Schubert (1948: 227), but not called type. The specimen is poor, and bears only the label “Eupatorium” in Fraser's hand. Without indication the Fraser specimen was seen or used by Walter, Ahles 34320, 18 Sep 1957 - GH, from along Broad River, Cherokee County, South Carolina, is here selected as NEOTYPE for Eupatorium incarnatum Walt., basionym of Fleishmannia incarnata (Walt.) King € Robins. Known ISONEOTYPES: GÀ, NCU, NY. WALTER'S NAME: Eupatorium linearifolium Walter (p. 199) MODERN NAME: Eupatorium hyssopifolium L. [- E. hyssopifolium var. calcaratum Fern & Schub.] Frequent throughout. Walters description ("foliis linearibus integris subverticillatis...”) exactly matches the linear entire mostly verticillate leaves of E. hyssopifolium. Spm. 44-B is the lanceolate serrate mostly opposite- aco variant of E. DNO Or E SEN Short. ER Fernald and Schubert (1948: 226-227) mistakenly a ed pe of Eupatorium linearifolium, which they treated as da distinct bon. E, ign E The Ee bears the label Seen in Walter's hand, butt ridence it had been used by him. A specimen that matches Walter's description, Godfrey e ion 1614, 10 Aug 1939 - GH (annot. Robert K. Godfrey 1973), from 3 mi W of Bonneau, Berkeley County, South Carolina, is here selected as NEOTYPE for Eupatorium linearifolium Walt. [= Eupatorium hys- sopifolium L.]. WALTER'S NAME: Festuca octoflora Walter (p. 81) MODERN NAME: Festuca octoflora Walt. [= Vulpia octoflora (Walt.) Rydb.] Common throughout. There is no specimen. Boufford 12781, 13 Apr 1974 - GH, from Columbia, Richland County, South Carolina, is here selected as neoTYPE for Festuca octoflora Walt. WALTER'S NAME: Galega spicata Walter (p. 188) MODERN NAME: Tephrosia spicata (Walt.) Torr. & A. Gray Common throughout. Spm. 49-B may be the specimen referred to by Wood (1949: 292) as the type of Galega spicata (^GH - photograph of Type in Herb. Walt."). But since Wood cited only the herbarium, not a specific specimen, leaving uncertainty which specimen was intended, unambiguous typification is justified. Godfrey & Tryon 1310, 7 Aug 1939 - GH, from 14 mi S of Columbia, Lexington County, South Carolina, is here selected as nEoTYPE for Galega spicata Walt., basionym of Tephrosia spicata (Walt.) Torr. & A. Gray. Known ISONEOTYPES: CA (“1311”), NY, US. WALTER's NAME: Hibiscus aculeatus Walter (p. 177) MODERN NAME: Hibiscus aculeatus Walt. Common in eastern SC. No specimen. Wiegand & Manning 1959, 10 Jul 1927 - GH, from 3 mi S of Lake City, Florence County, South Carolina, is here selected as neoTYPE for Hibiscus aculeatus Walt. Known ISONEOTYPES: BH. 478 | tanical Insti Texas 2(1) WALTER's NAME: Hibiscus coccineus Walter (p. 177) MODERN NAME: Hibiscus coccineus Walt. Not presently known in NC, SC, or GA (reported from GA by Small 1933), apparently unknown n. of Duval Co., FL. An old (undated) specimen has been seen from Charleston, SC (GH), perhaps from cultivation. Spm. 58-E is clearly this; it bears Fraser's number “674” (an indication of mid-season collection, thus it could not have been obtained on Fraser's spring trip into southern Georgia). It was labeled "Hibiscus" by Walter, with “Coccineus” added by Fraser. Though spm. 58-E could be claimed a lectotype, the possibil- ity that Walter may have had seen it near Charleston, his market town, and his failure to note Fraser's specimen as a species of his own making, makes that status unlikely. Blanchard 173, 14 Jul 1968 - GH, from Fla. 16, 0.4 mi E of St. Johns River, between Orangedale and Green Cove Springs, St. Johns County, Florida, is here selected as NEOTYPE of Hibiscus coccineus Walt. Known ISONEOTYPES: CAS. WALTER' NAME: Hydrastis caroliniensis Walter (p. 156) MODERN NAME: Trautvetteria caroliniensis (Walt.) Vail Nearly absent from SC, but common in western NC; probably a Fraser discovery. Spm. 1-E is of poor quality, and was mislabeled “Actea racemosa?” by Walter. Although Walter may have known this species only by way of Fraser's collections, his failure to recognize spm. 1-E as a species he himself had named makes unlikely his having used it in his writing. Thus, rather than designating spm. 1-E as lectotype, Curtiss 30, Jul [1892?] - GH, from grassy *balds" of Roan Mtn., Mitchell County, North Carolina, is here selected as NEOTYPE for Hydrastis caroliniensis Walt., basionym of Trautvetteria caroliniensis (Walt.) Vail. Known ISONEOTYPES: BH, F, GA, NY, PH, US. WALTER'S NAME: Hyoseris biflora Walter (p. 194) MODERN NAME: Krigia biflora (Walt.) Blake Absent from SC, frequent in western NC. Identified by Blake (1915: 135). No specimen in the herbarium. Cronquist 4349, 27 Apr 1947 - GH, from near Flint River, 4 mi SE of Woodbury, Upson County, Georgia, is here selected as NEOTYPE for Hyoseris biflora Walt., basionym of Krigia biflora (Walt.) Blake. Known 1so- NEOTYPES: FLAS, GA, MO, NY, US WALTER'S NAME: Hypericum denticulatum Walter (p. 190) MODERN NAME: Hypericum denticulatum Walt. Common in eastern SC. No specimen. Wiegand & Manning 2018, 13 Jul 1927 - GH, from 3 mi S of Bonneau, Berkeley County, South Carolina, is here selected as ngoTYPE for Hypericum denticulatum Walt. Known ISONEOTYPES: BH. WALTER'S NAME: Hypericum petiolatum (p. 191) MODERN NAME: Triadenum walteri (Gmel.) Gleason [= Hypericum walteri Gmel.; 2 Triadenum petiolatum (Walt.) Britt.] Frequent throughout. Not Hypericum petiolatum L.; Walter's Hypericum petiolatum was a later homonym (1763 vs. 1788) and thus illegitimate. But Gmelin's replacement (1791) was based on Walter's name and description. No specimen has been identified. Duncan hie 20 SC nari GH, from 4.2 mi E of Lafay- ette, Walker County, Georgia, is here selected Hy walteri Gmel. [2 Triadenum walteri (Gmel.) Gleason]. Known ISONEOTYPES: BH, FLAS, GA, US. WALTER'S NAME: Ilex decidua Walter (p. 241) MODERN NAME: Ilex decidua Walt. Frequent throughout. Spms. 61a-H and 612-J appear to be this, the first labeled “Ilex decid” by Walter, the second “Ilex Decidua" by Fraser. Both are of poor quality. Godfrey & Tryon 652, 17 Jul 1939 - GH, from the Santee River floodplain, 3 mi NE of Pineville, Berkeley County, South Carolina, is here selected as NEOTYPE of Ilex decidua Walt. Known IsoONEOTYPES: BH, CA, CAS, DUKE, F, MO, NY, PH, US. Ward, "T TA Da. A ] SES a! IF F1 | [| ] Hr £ rm Z 479 WALTER'S NAME: Ilex myrtifolia Walter (p. 241) MODERN NAME: Ilex myrtifolia Walt. [= Ilex cassine L. var. myrtifolia (Walt.) Sarg.] Common in eastern SC. Spm. 61a-D was labeled “Ilex Myrtifolia” by Fraser and has been annotated as "type" (by J.E. Dandy?), but is a small twig of poor quality. Spms. 61a-E, 61a-G, and 61a-I appear to be the same species, and are even poorer. Since spm. 61a-D appears not to have been published as type, Bozeman 11368, 27 Aug 1967 - GH (annot. Ross C. Clark 1994), from Francis Marion National Forest, 1.6 mi N of Honey Hill, Berkeley County, South Carolina, is here selected as NEOTYPE of Ilex myrtifolia Walt. Known ISONEOTYPES: MICH, NCU, NY. WALTER'S NAME: Iris tripetala Walter (p. 66) MODERN NAME: Iris tridentata Pursh Frequent on SC coastal plain. Not Iris tripetala L.f. (1782). There is no specimen. Though Walter's name is usually treated as a synonym of Iris tridentata, Leonard & Radford 1599, 2 Jun 1968 - GH, from George- town County, South Carolina, is here selected as NEOTYPE for Iris tripetala Walt. Known ISONEOTYPES: CA, FLAS, GA, NCU, NY. WALTER'S NAME: Ischaemum secundatum Walter (p. 249) MODERN NAME: Stenotaphrum secundatum (Walt.) Kuntze Infrequent in eastern SC. Identified by Hitchcock (1905: 55) from Walter's description. There is no specimen. The species in the Southeast consists of two variants (“demes”), one native, one introduced (Sauer 1972). Ahles 15602, 27 Jun 1956 - GH (annot. J.D. Sauer 1971), from roadbank, U.S. 21, Beaufort, Beaufort County, South Carolina, is here selected as NEOTYPE for Ischaemum secundatum Walt., basionym of Stenotaphrum secundatum (Walt.) Kuntze. Known ISONEOTYPES: GA, NCU, NY. WALTER's NAME: Kalmia hirsuta Walter (p. 138) MODERN NAME: Kalmia hirsuta Walt. Rare in SC (5 counties). Spm. 62b-C, a nearly bare twig, was labeled “Kalmia Hirsuta Nova” by Fraser. Southall & Hardin (1974) referred to a specimen on page 62 as the "type." Their typification has been corrected (Ward 20072) to lectotype. The specimen, however, is unidentifiable without the label and serves no useful purpose as a type. Bell 3956, 29 Jun 1956 - GH (annot. J.E. Ebinger 1972), from S.C. 631, 1 mi SE of Hampton County line, Jasper County, South Carolina, is here selected as EPITYPE, in support of Southall & Hardin's typification of Kalmia hirsuta Walt. (as corrected). WALTER'S NAME: Leontodon carolinianum Walter (p. 192) MODERN NAME: Pyrrhopappus carolinianus (Walt.) DC. Frequent throughout. This may be spm. 64-F, which was labeled “Leontodon novum" by Walter, and "Carolinianum" by Fraser. It is very poor. A specimen with cauline leaves, Wiegand & ciii: 3518, 11 Jul 1927 - GH, from 4 mi S of Kingstree, Williamsburg County, South Carolina, is] Leontodon carolinianum Walt., basionym of Pyrrhopappus carolinianus (Walt.) DC. Known ISONEOTYPES: BH. WALTER'S NAME: Lilium Catesbaei Walter (p. 123) MODERN NAME: Lilium catesbaei Walt. Infrequent on SC coastal plain, but a striking plant and likely known to Walter. Spm. 64-E is this, al- though labeled “Lilium philadelphicum" by Walter. Since the specimen is of poor quality, and appears not to have been recognized by Walter, Godfrey & Tryon 1046, 2 Aug 1939 - GH, from 11 mi N of Georgetown, Georgetown County, South Carolina, is here selected as NEOTYPE for Lilium catesbaei Walt. Known tsoNEo- TYPES: BH, CA, CAS, DUKE, F, MO, NY. 480 ) loft tanical h Instit f Texas 2(1) WALTER'S NAME: Linum striatum Walter (p. 118) MODERN NAME: Linum striatum Walt. Infrequent throughout. No specimen. Godfrey & Tryon 637, 17 Jul 1939 - GH (annot. C. M. Rogers 1960), from Santee Canal, 5 mi W of Pineville, Berkeley County, South Carolina, is here selected as NEOTYPE for Linum striatum Walt. Known ISONEOTYPES: CA, NY, US. WALTER'S NAME: Ludwigia arcuata Walter (p. 89) MODERN NAME: Ludwigia arcuata Walt. Very rare in SC (3 counties, but one is Charleston, Walter's market town). No specimen appears to be this species. But Peng et al. (2005: 345) cited spm. 66-D as “holotype” of Ludwigia arcuata, apparently in gross error! (The cited specimen appears to be L. pilosa, and was so labeled by Walter.) With the expec- tation that the error will be acknowledged and effort undertaken to correct the erroneous typification via conservation, Radford 11411, 8 Sep 1967 - GH, from Millettville, Allendale County, South Carolina, is here selected as NEOTYPE for Ludwigia arcuata Walt. Known ISONEOTYPES: CA, CAS, FLAS, GA, MICH, NCU, NY, USCH WALTER'S NAME: Ludwigia glandulosa Walter (p. 88) MODERN NAME: Ludwigia glandulosa Walt. Frequent in eastern SC. No specimen has been identified. Wilbur 74177, 15 Aug 2001 - GH, from Red Bluff, Horry County, South Carolina, is here selected as NEOTYPE for Ludwigia glandulosa Walt. Known ISONEOTYPES: DUKE. WALTER'S NAME: Ludwigia suffruticosa Walter (p. 90) MODERN NAME: Ludwigia suffruticosa Walt. Frequent in eastern SC. No specimen has been identified. Wiegand & Manning 2200, 10 Jul 1927 - GH (an- not. P.A. Munz 1942; C-I Peng 1981), from 6 mi N of Kingstree, Williamsburg County, South Carolina, is here selected as NEOTYPE for Ludwigia suffruticosa Walt. Known ISONEOTYPES: BH. WALTER'S NAME: Lycium carolinianum Walter (p. 84) MODERN NAME: Lycium carolinianum Walt. There is no specimen. A coastal plant, at times reported for SC, but only on the basis of Walter's listing, and a note in Elliott. Presently unknown in SC; almost unknown in GA (Camden Co., the southeastern corner). Undoubtedly a discovery of Fraser's, but it is difficult to understand how he could have journeyed that far south. Perhaps since his time the range of this cold-sensitive plant has contracted southward. This possibility is supported by Elliott (1816: 1: 200): “I have never seen this plant in its native state, where it is said to be of very humble growth. Found by Mr. Wm. Bartram, in the sali hy marshes of Carolina.” Curtiss 6543, 10 Oct 1899 - GH (annot. C.L. EES E. from mouth of St. Johns River, [Duval County], Florida, is | lected f i arolinianum Walt. Known ISONEOTYPES: BH, CA, CAS, DS, G, MO, NY, UC, US. This collection was noted by C.L. Hitchcock (1932: 239) as: “no type designated, but Curtiss's number 6543 is taken as typically representing this species.” WALTER'S NAME: Melanthium Muscaetoxicum Walter (p. 125) MODERN NAME: Zigadenus muscaetoxicum (Walt.) Regel [= Amianthium muscaetoxicum (Walt.) A. Gray] Frequent throughout. Spm. 58-C was labeled “Helonias bullata?” by Walter, in reference to H. bullata L., a northern species that reaches NC (two counties) but is unknown in SC. Walter did not include H. bullata in his Flora. Though he SHOE AIME muscaetoxicum, which he included as “Melanthium Muscaetoxicum," e tly did not g lant collected by Fraser. Weatherby & Griscom 16486, 1 May 1932 - GH, fem Myrtle Beach, Horry County South Carolina, is here selected as NEOTYPE for Melanthium muscaetoxicum Walt., basionym of Zigadenus muscaetoxicum (Walt.) Regel. Known IsoNEoTYPES: NY. Ward, Thomas Walter typification V: neoty] l epitypes of g D-Z 481 WALTER'S NAME: Melica mutica Walter (p. 78) MODERN NAME: Melica mutica Walt. Frequent throughout. There is no specimen. Griscom 511, 11 Apr 1932 - GH (annot. W.S. Boyle 1943), from Myrtle Beach, Horry County, South Carolina, is here selected as NEOTYPE for Melica mutica Walt. Known ISONEOTYPES: BH, CA, CAS, DUKE, F, GA, MICH, MO, NCU, NY, PH. WALTER'S NAME: Mespilus aestivalis Walter (p. 148) MODERN NAME: Crataegus aestivalis (Walt.) Torr. & A. Gray Rare in SC (3 counties, incl. Berkeley). No specimen. Harbison s.n., 15 Mar & 21 May 1916 - GH, from along Combahee River, E of Yemassee, [Colleton Co.], South Carolina, is here selected as NEOTYPE for Nespilus aestivalis Walt., basionym for Crataegus aestivalis (Walt.) Torr. & A. Gray. Known ISONEOTYPES: NCU. WALTER'S NAME: Nyssa biflora Walter (p. 253) MODERN NAME: Nyssa biflora Walt. [= Nyssa sylvatica Marsh. var. biflora (Walt.) Sarg.] Frequent in eastern SC. Spms. 75-A, 75-B, or 75-C may be this, but are poor. (The first was labeled “Nyssa,” the second “Nyssa Multiflora," both by Fraser, and the third “Nyssa” by Walter.) Little 14304, 15 Jun 1951 - GH, from Hell Hole Bay, S of Jamestown, Berkeley County, South Carolina, is here selected as NEOTYPE for Nyssa biflora Walt. WALTER'S NAME: Obolaria caroliniana Walter (p. 166) MODERN NAME: Bacopa caroliniana (Walt.) Robins. Common in eastern SC. Pennell (1920: 243) stated: “Description sufficiently distinctive.” Pennell (1946) again recognized Walter's description as this species and used this name, but designated no type. No corresponding specimen is apparent. Godfrey & Tryon 948, 26 Jul 1939 - GH (annot. H.S. Wooden 1979), from 1 mi S of Sandinia, Clarendon County, South Carolina, is here selected as neotTYPE for Obolaria caroliniana Walt., basionym for Bacopa caroliniana (Walt.) Robins. Known IsoNEOTYPES: NY, US. WALTER' NAME: Paspalum praecox Walter (p. 75) MODERN NAME: Paspalum praecox Walt. Hitchcock (1905: 42) noted Walter's name now to be “generally acepted by American botanists." Plants of this name are infrequent on the SC coastal plain. There is no specimen. For stability, Godfrey & Tryon 84, 23 Jun 1939 - GH, from Georgetown, Georgetown County, South Carolina, is here selected as NEOTYPE for Paspalum praecox Walt. Known IsONEOTYPES: BH, CAS, DUKE, MICH, NY, PH, US. WALTER' NAME: Pinus glabra Walter (p. 237) MODERN NAME: Pinus glabra Walt. Frequent in southeastern SC. Dayton (1952) and Ward (1962) confirmed Walter's description; Dayton identified spms. 83-D and 83-H as this species. Dayton proposed that these two specimens “perhaps should be considered the type material" of Pinus glabra, but made no designation. Though spm. 83-H was labeled “Pinus” by Walter (and spm. 83-H by Fraser), there is no evidence Walter used it in forming his description. Palmer 35395, 16 May 1929 - GH, from Seneca, Oconee County, South Carolina, is here selected as NEOTYPE for Pinus glabra Walt. Known IsSoNEOTYPES: MO, NY, US. WALTER'S NAME: Pinus squarrosa Walter (p. 237) MODERN NAME: Pinus echinata Mill. Common throughout. Dayton (1952) and Ward (1962) confirmed Walter's description; Dayton identified spms. 83-I (labeled “Pinus” by Walter) and 83-J (unlabeled) as this species. Dayton (1952: 71) suggested, “it is possible that this [spm. 83-I] may be the type of Walter's P. squarrosa.” However he stopped short of designating the specimen as type. A better specimen, with cones, Godfrey & Tryon 1453, 10 Aug 1939 482 ) | tanical h Institute of Texas 2( - GH, from 3 mi W of Bonneau, Berkeley County, South Carolina, is here selected as NEOTYPE for Pinus squarrosa Walt. (= Pinus echinata Mill.). Known ISONEOTYPES: CA, CAS, NY, US. WALTER'S NAME: Polygala cymosa Walter (p. 179) MODERN NAME: Polygala cymosa Walt. Common in eastern SC. Smith & Ward (1976) referred to a “holotype,” but noted “not seen and presum- ably no longer extant." Spm. 84-B, labeled "Polygala" by Fraser, has since been identified as P. cymosa. It is of poor quality, consisting solely of an upper inflorescence. In the absence of evidence that spm. 84-B was seen or used by Walter, Godfrey & Tryon 67, 24 Jun 1939 - GH (annot. Foard 1959), from 15 mi N of Georgetown, Georgetown County, South Carolina, is here selected as NEOTYPE of Polygala cymosa Walt. Known ISONEOTYPES: BH, CA, CAS, DUKE, F, MO, NY, US. WALTER'S NAME: Polygala grandiflora Walter (p. 179) MODERN NAME: Polygala grandiflora Walt. Frequent in eastern SC. There is no specimen. Rodriguez (2003) stated the “Holotipo” to be “Carolina, Walter (BM)," with a symbol indicating the specimen had been lost or destroyed. Since no specimen could be designated, no typification took place. uec 4905, 30 ae 1917 - GH, from Ladies Island, Beaufort, Carteret County, North Carolina, is | oTYPE of Polygala grandiflora Walt. Known ISONEOTYPES: MO. WALTER'S NAME: Polygala polygama Walter (p. 179) MODERN NAME: Polygala polygama Walt. Frequent throughout. No specimen has been identified. Palmer 42407, 19 May 1934 - GH, from Columbia, Richland County, South Carolina, is here selected as NEOTYPE of Polygala polygama Walt. WALTER'S NAME: Polygonum hirsutum Walter (p. 132) MODERN NAME: Polygonum hirsutum Walt. Rare in SC (4 counties, though one is Charleston, Walter’s market town). No specimen has been identi- fied. Ahles 15868, 20 Jun 1956 - GH, from Barton, Allendale County, South Carolina, is here selected as NEOTYPE for Polygonum hirsutum Walt. Known ISONEOTYPES: NY. WALTER'S NAME: Quercus laevis Walter (p. 234) MODERN NAME: Quercus laevis Walt. Common in eastern SC. No specimen. Hill 18860, 21 Sep 1987 - GH, from Bluefield Road, Lexington County, South Carolina, is here selected as NEOTYPE for Quercus laevis Walt. Known ISoNEOTYPES: MO, NCU, NY, USCH. WALTER'S NAME: Quercus lyrata Walter (p. 235) MODERN NAME: Quercus lyrata Walt. Common in eastern SC. No specimen. Spongberg 17209, 14 Sep 1982 - GH, from Perrysburg Landing, WNW of Hardeeville, Jasper County, South Carolina, is here selected as NEOTYPE of Quercus lyrata Walt. WALTER'S NAME: Quercus pumila Walter (p. 234) MODERN NAME: Quercus pumila Walt. [= Quercus elliottii Wilbur] Common in eastern SC. Spm. 90-E is poor, and though it was seen (and labeled, as “Quercus pumila”) by Walter, there is no suggestion that it was used by him. Godfrey & Tryon 1420, 11 Aug 1939 - GH, from 8 mi S of Moncks Corner, Berkeley County, South Carolina, is here selected as NEOTYPE for Quercus pumila Walt. Known 1isoNEOTYPEs: NY, US. Wilbur (2002: 138-140) has argued that Walter's description was de- fective, and that the small shrub should be renamed Quercus elliottii Wilbur. In light of the near-certainty that Walter knew his Quercus pumila (as seen within stone-throw of his grave - D.B.W. obs., Jun 1990), the new name is superfluous. WALTER's NAME: Rhexia petiolata Walter (p. 130) MODERN NAME: Rhexia petiolata Walt. Frequent in eastern SC. James (1956: 216) tentatively designated a specimen, Fraser/Walter 91-D [1787] - BM, marked “Rhexia 723” as “type” of Rhexia petiolata Walt. (The “Rhexia” is in Walter's hand, the “723” in Fraser's.) James' typification has been corrected (Ward 20072) to neotype for Rhexia petiolata Walt. James noted, "Positive identification has not been possible because of pu dris condition of the speci- ” The specimen (91-D) is a bare stem, quite lacking in identifiable feat Wiegand & Manning 2126, da 1927 - GH, from 4 mi S of Florence, Florence County South Carolina, is here selected as EPITYPE, in support of James’ typification of Rhexia petiolata Walt. (as corrected). Known ISONEOTYPES: BH. WALTER'S NAME: Salsola caroliniana Walter (p. 111) MODERN NAME: Salsola kali L. var. caroliniana (Walt) Nutt. Infrequent along SC coast. Spm. 93-F was labeled “Salsola Kali” by Walter, then the epithet was struck and “Caroliniana” added by Fraser. Though only a varietal name is at issue, Godfrey & Tryon 1571, 17 Aug 1939 - GH (annot. H.D. Wilson 1981; S. Mosyakin 1994), from South Island, Georgetown County, South Carolina, is here selected as NEOTYPE for Salsola caroliniana Walt., basioym of Salsola kali L. var. caroliniana (Walt.) Nutt. Known ISONEOTYPES: BH, CA, DUKE, MO, NY, PH, US. WALTER'S NAME: Silene caroliniana Walter (p. 142) MODERN NAME: Silene caroliniana Walt. Frequent throughout SC. Spm. 98-G was labeled “Silene an virginica?” by Walter. Wilbur (1970) referred to comments of J.K. Small and C.A. Weatherby—neither of who personally saw the Walter herbarium—to conclude the specimen was unclear as to its subspecies, and that ^Walter's description is unmistakable and would take precedence over any specimen in the Walter herbarium.” Thus Weatherby 6114, 27 Apr 1932 - GH (annot. R.T. Clausen 1938; Bassett Maguire 1941; R.L. Wilbur 1968 & 1996), from outskirts of Columbia, Lexington County, South Carolina, is here selected as NEOTYPE of Silene caroliniana Walt. Known ISONEOTYPES: DUKE, NY, PH. WALTER'S NAME: Smilax auriculata Walter (p. 245) MODERN NAME: Smilax auriculata Walt. Frequent along SC coast. No specimen is present. Godfrey & Tryon 1169, 5 Aug 1939 - GH, from Myrtle Beach, Horry County, South Carolina, is here selected as NEOTYPE for Smilax auriculata Walt. Known ISONEOTYPES: BH, CA, CAS, DUKE, PH. WALTER'S NAME: Smilax glauca Walter (p. 245) MODERN NAME: Smilax glauca Walt. Infrequent on SC coastal plain, common inland. No speci as found in the Fraser/Walter her] Blake (1918). Blake did find a F with “leaves densely puberulent beneath" in the Der anielle herbarium, Geneva. He noted however that the Fraser specimen was not “in any sense a type of Walter’s S. glauca.” A specimen with leaves smooth beneath, Bell 7168, 12 May 1957 - GH, from Beaver Creek, NNE of Blair, Fairfield County, South Carolina, is here selected as neoryre for Smilax glauca Walt. WALTER'S NAME: Smilax pumila Walter (p. 244) MODERN NAME: Smilax pumila Walt. Frequent in eastern SC. No specimen is present in the herbarium. Ahles 20939, 11 Oct 1956 - GH, from county road, 0.4 mi E of U.S. 21, Beaufort County, South Carolina, is here selected as NEOTYPE for Smilax pumila Walt. Known ISONEOTYPES: GA. 484 | t tanical h Institute of Texas 2( WALTER'S NAME: Staehelina elegans Walter (p. 202) MODERN NAME: Liatris elegans (Walt.) Michx. Common in southeastern SC. Spm. 101-A, a reasonably lete intl el the label “Staehelina” in Walters hand; Fraser added “Elegans.” Since Walter would have known the plant near his home, he need not have used this specimen in forming his description. Boufford & Shi 30292, 25 Sep 2000 - GH, from Halfway Creek road, Francis Marion National Forest, WNW of McClellanville, Berkeley County, South Carolina, is here selected as NEOTYPE for Staehelina elegans Walt., basionym of Liatris elegans (Walt Michx. WALTER'S NAME: Syntherisma serotina Walter (p. 76) MODERN NAME: Digitaria serotina (Walt.) Michx. Hitchcock (1905: 44) found Walter's di ting this and the made the identification on the basis of hese being the only creeping Digitaria “found commonly” in the Carolinas. But, far from common, D. serotina is very rare in SC (2 counties), suggesting Walter's second species may also have been the abundant and somewhat variable D. sanguinalis. There is no specimen. To stabilize the meaning of D. serotina (which is well distinguished elsewhere), Hitchcock 1557, 15 Aug 1905 - GH, from “open low ground,” South Carolina, is here selected as NEOTYPE for Syntherisma serotina Walt., basionym of Digitaria serotina (Walt.) Michx. Known IsSoNEOTYPES: CA, MO, NY. ist: y, but WALTER'S NAME: Syntherisma villosa Walter (p. 77) MODERN NAME: Digitaria filiformis (L.) Koel. var. villosa (Walt.) Fern. Frequent throughout. Hitchcock (1905: 44) accepted spm. 116-C as corresponding to Walter's diagnosis. The specimen is labeled in Walter's hand. But it is poor and unlikely to have been used by Walter in forming his description. Andersen 1408, 1897 - GH (annot. M.L. Fernald 1920), from Oconee County, South Carolina, is here selected as NEOTYPE for Syntherisma villosa Walt., basionym of Digitaria villosa (Walt.) Pers. WALTER'S NAME: Utricularia purpurea Walter (p. 64) MODERN NAME: Utricularia purpurea Walt. Infrequent on the SC coastal plain. There is no specimen. Taylor (1989: 689) stated: "lecto. Barnhart 1916;” but Barnhart's action (1916: 62) was a typification of the segregate genus Vesiculina Raf., not a typification of the species U. purpurea. Godfrey & Tryon 1097, 4 Aug 1939 - GH, from 6 mi NW of McClel- lanville, Charleston, South Carolina, is here selected as NEOTYPE for Utricularia purpurea Walt. Known ISONEOTYPES: BH, CA, CAS, DUKE, F, MO, US. WALTER'S NAME: Viburnum obovatum Walter (p. 116) MODERN NAME: Viburnum obovatum Walt. Frequent in eastern SC. The only possible specimen (spm. 108-H) is a fragment, bearing only Fraser's label, “Viburnum.” Godfrey & Tryon 8215, 24 Jul 1939 - GH (annot. Thomas H. Jones 1978), from 10 mi NE of Moncks Corner, Berkeley County, South Carolina, is here selected as NEOTYPE for Viburnum obovatum Walt. Known ISONEOTYPES: CA, CAS, DUKE, F, MO, NY, PH, US. WALTER' NAME: Vicia caroliniana Walter (p. 182) MODERN NAME: Vicia caroliniana Walt. Frequent in eastern SC, more westward. Spm. 106-D appears to be thi ies. But since Walter labeled it only as “Vicia nova,” it is A it was used in preparing his desceiotiom. Hunnewell 8163, 23 Mar 1922 - GH (annot. Robert Dirig 1990), from Goose Creek, Berkeley County, South Carolina, is here selected as NEOTYPE for Vicia caroliniana Walt. Ward, "Th. m an A A | LE a FR) LT d r1 | [| FH E r 7 485 WALTER'S NAME: Vincetoxicum gonocarpos Walter (p. 104) MODERN NAME: Matelea gonocarpa (Walt.) Shinners Infrequent in SC, nearly absent from the coastal plain. Thus likely a Fraser discovery. Spm. 109-A, marked “Vincetoxicum” by Walter, was identified as Matelea gonocarpa by Drapalik (1970: 76, from photo). But it is sterile, which makes it unlikely to have been of use to Walter, who described the plant in detail (incl. flowers and fruits). A specimen of better quality, Wiegand & Manning 2629, 16 Jul 1927 - GH (annot. Donald J. Drapalik 1974), from Meggett, Charleston County, South Carolina, is here selected as NEOTYPE for Vincetoxicum gonocarpos Walt., basionym of Matelea gonocarpa (Walt.) Shinners. WALTER'S NAME: Viola canina sensu Walter (p. 219) non Viola canina Linnaeus, Sp. Pl. 935. 1753; misap- plied. MODERN NAME: Viola walteri House Infrequent throughout. Not Viola canina L., nor V. conspersa Rchb. When House (1906) realized Walter had EE the Carolina plant he assigned it a new name, V. walteri, but based the new name on Walter's di 1 presumed type. There is no specimen. Hunnewell 12638, 30 Mar 1933 - GH (annot. Harvey E. Ballard 1991), from near Goose Creek, Berkeley County, South Carolina, is here selected as NEOTYPE for Viola canina sensu Walter (basis for Viola walteri House). WALTER'S NAME: Viola villosa Walter (p. 219) MODERN NAME: Viola villosa Walt. Frequent throughout. Spms. 108-D and 108-1 may be this species. Both bear Walter's hand, but only to “Viola” and a descriptive phrase; both are of poor quality. Brainerd (1907) has argued convincingly that Walter's plant was this species (and not V. hirsutula Brainerd, found only inland). Brainerd s.n., 25 Mar 1907 - GH (annot. Landon E. McKinney 1986), from Summerville, Dorchester County, South Carolina, is here selected as NEOTYPE for Viola villosa Walt. WALTER'S NAME: Waltheria caroliniana Walter (p. 175) MODERN NAME: Piriqueta caroliniana (Walt.) Urban Frequent in southeastern SC. No specimen has been identified. Ahles 18262, 9 Sep 1956 - GH, from SC 20, 2.7 mi S of jct. with SC 39, Hampton County, South Carolina, is h elected for Waltheria caroliniana Walt., basionym of Piriqueta caroliniana (Walt.) Urban. Known isoNEorTYPEs: NCU. ACKNOWLEDGMENTS The present report, as expressed previously, would not have been possible without the cooperation of the Harvard University Herbaria (GH), especially Walter T. Kittredge and staff. Robert Dirig and Me- lissa A. Luckow (BH), Andrew Doran (CA), Debra Trock (CAS), Robert L. Wilber (DUKE), Maria Lucia Kawasaki (F), Kent D. Perkins (FLAS), Kelly Bettinger and Wendy B. Zomlefer (GA), Stuart Lindsay and Anton A. Reznicek (MICH), James C. Solomon (MO), Carol Ann McCormick and Alan S. Weakley (NCU), Thomas A. Zanoni (NY), Carrie A. Kiel (PH), George F. Russell (US), and John B. Nelson (USCH) contributed by searching for ISONEOTYPES from within their herbaria. The Spanish abstract was prepared by Christine M. Housel (ABT). REFERENCES BARNHART, J.H. 1916. Segregation of genera in Lentibulariaceae. Mem. New York Bot. Gard. 6:39-64. Blake, S.F. 1918. A variety of Smilax glauca. Rhodora 20:78-80. BRAINERD, E. 1907, The older types of North American violets - I. Rhodora 9:93-98. Dayton, WA 1952. Some notes on United States tree names. Rhodora 54:67-79. Drapauk, D.J. 1970. A biosystematic study of the genus Matelea in the southeastern United States. Ph. D. diss., Univ. of North Carolina, Chapel Hill. 220 pp. 486 ELLIOTT, S. 1816. A sketch of the botany of South Carolina and Ge Ge FERNALD, M.L. and B.G. ScHuBerT. 1948. Studies of American ty tish herbar ia. Part |V: Some species of Thomas Walter. Rhodora 50:190-208, 217-229. GMELIN, J.F. 1791, 1792. Systema Naturae. Leipzig. HitcHcock, A.S. 1905. The identification of Walter's grasses. Missouri Bot. Gard. Ann. Rept 16:31-56. HrrcHcock, C.L. 1932. A monographic study of the genus Lycium of the western hemisphere. Ann. Missouri Bot. Gard. 19:179-374. House, HD 1906. Notes on southern violets - |. Torreya 6:171-173. James, CW, 1956. A revision of Rhexia (Melastomataceae). Brittonia 8:201—230. KraL, R. 1966. Eriocaulaceae of continental North America north of Mexico. Sida 2:285-332. LiNNAEUS, C. 1762-1763. Species Plantarum, ed. 2. 2 vols. Stockholm. Mathias, M. and L. Constance. 1944. Spermolepis. In: Umbelliferae. N. Amer. Fl. 28B:13-295. Pena, CL C.L. Scuuipr, PC. Hock, and PH. Raven. 2005. Systematics and evolution of Ludwigia section Dantia (On- agraceae). Ann. Missouri Bot. Gard. 92:307-359. Dovun, FW, 1920 [1919]. Scrophulariaceae of the southeastern United States. Proc. Acad. Natl. Sci. Phil. 7/1:224-291. PENNELL, FW. 1946. Reconsideration of the Bacopa - Herpestis problem of the Scrophulariaceae. Proc. Acad. Natl. Sci. Phil. 98:83-98. Robricuez, R.R. 2003. Flora de la Republica de Cuba. Fasc. 7(1):37. SAUER, J. 1972. Revision of Stenotaphrum (Gramineae: Paniceae) with attention to its historical geography. Brit- tonia 24:202-222. Smau, J.K. 1933. Manual of the Southeastern Flora. New York. Smith, RR. and D.B. Waro. 1976. Taxonomy of the genus Polygala series Decurrentes (Polygalaceae). Sida 6:284-310. Soumat). RM. and JW. Hardin. 1974. A taxonomic revision of Kalmia (Ericaceae). J. Elisha Mitchell Sci. Soc. Tavlor, P. 1989. The genus Utricularia -- a taxonomic monograph. Kew Bull, Addit. Ser. XIV. WALTER, T. 1788. Flora Caroliniana. London. Warp, D.B. 1962. The genus Anonymos and its nomenclatural survivors. Rhodora 64:87-92. Warp, D.B. 2006. Thomas Walter Typification Project, |. Observations on the John Fraser folio. Sida 22:1111-1118. Warp, D.B. 2007a. Thomas Walter Typification Project, Il. The known Walter types. J. Bot. Res. Inst. Texas 1:407-423. Waro, D.B. 2007b. Thomas Walter Typification Project, Ill. Lectotypes and neotypes for 20 Walter names, as rec- ognized in the Fraser/Walter herbarium. J. Bot. Res. Inst. Texas 1:425-430. Warp, D.B. 2007c. Thomas Walter Typification Project, IV. Neotypes and epitypes for 44 Walter names, of genera A through C. J. Bot. Res. Inst. Texas 1:1091-1100 Waro, D.B. 2007d. The Thomas Walter herbarium is not the herbarium of Thomas Walter. Taxon 56:971—9206. Warnock, M.J. 1981. Biosystematics of the Delphinium carolinianum complex (Ranunculaceae). Syst. Bot. 6:38-54, WaBur, R.L. 1970. Infraspecific classification in the Carolina flora. Rhodora 72:51-65. WiLBUR, R.L. 2002. Thomas Walter's oaks from the coastal region of South Carolina. Rhodora 104:134-150. Woon, C.E. 1949, The American barbistyled species of Tephrosia (Leguminosae). Rhodora 51:193-231, 233-302, 305—364, 369—384. CHROMOSOME NUMBERS FOR SOUTHWESTERN NORTH AMERICAN SPECIES OF ANTENNARIA (ASTERACEAE: GNAPHALIEAE) Jerry G. Chmielewski Department of Biology Slippery Rock University Slippery Rock, Pennsylvania 16057, U.S.A. ABSTRACT Chromosome numbers were determined from somatic material for 30 individuals of Antennaria Gaertner (Asteraceae: Gnaphalieae), ; p 1 : E r. +1 WI MP IS : T1 x D : de E 1 oof e E 4 J RESUMEN Cc 1; 4 za q + ix] Aas m Sy in. AS | A Å aa 4 H £^À Led 1 1; ), que repre t t peci el suroeste de Norte América. I t —À con is es previamente en j , Chromosome ts] ] | unts for North American Antennaria. Cultivation of plants and preparation of somatic rael Gm chromeseme number determinations followed the procedures of Chinnappa (1986). Voucher specimens remain in my possession and duplicates are deposited in SLRO. I collected all specimens; staminate specimens are marked with an asterisk (*). ASTERACEAE Antennaria howellii Greene. 2n = 112. COLORADO: Pueblo Co.: CO 165, Alpine Park NW of Rye, Ch3190; between mileposts 23 and 22, Ch3193. Custer Co.: CO 165, Bigelow Divide, near milepost 8, Ch3200. Antennaria media Greene. 2n = 56. COLORADO: San Juan Co.: US 550, S of Red Mountain Pass, Ch3173; Molas Divide, Ch3177. Antennaria parvifolia Nutt. 2n = 84. ARIZONA: Coconino Co.: US 180, vicinity of Forest Road 760, N of Snowbowl, Ch3154; N of Fort Valley Ranch Road, approx. milepost 222, Ch3155. Rio Grande Co.: US 160, E of South Fork, vicinity of South Fork Rest Area, Ch3183; 2n = 112. ARIZONA: Coconino Co.: Kai- bab Plateau, AZ 67, just S of Jacob Lake, Ch3157. COLORADO: San Juan Co.: US 550, Red Mountain Pass at Red Mountain Creek, Ch3168 (white form), Ch3169 (red form); alpine meadow S of Red Mountain Pass, Ch3174 (white form), Ch3175 (red form); Molas Divide, Ch3176 (white form), Ch3178 (pink form). Mineral Co.: US 160, Wolf Creek Pass, Ch3181, Ch3182*. Custer Co.: CO 165, NW of San Isabel, NW of Beulah Road, Ch3194 (red form), Ch3195 (white form); Bigelow Divide, vicinity of milepost 8, Ch3197*, Ch3198. Antennaria rosea Greene. 2n = 56. COLORADO: Coconino Co.: Kaibab Plateau, US 67, S of Jacob Lake, vicinity of Kaibab Lodge, Ch3160 (red form), Ch3161 (white form); S of Kaibab Lodge, Ch3163. San Juan Co.: US 550, Red Mountain Pass, Red Mountain Creek, Ch3170; S of Red Mountain Pass, Ch3172; Molas Divide, Ch3179. Pueblo Co.: CO 165, between mileposts 22 and 23, Ch3192. Custer Co.: CO 165, NW of San Isabel, NW of Beulah Road, Ch3196; Bigelow Divide, vicinity of milepost 8, Ch3199. ACKNOWLEDGMENTS I thank the Department of Biology, Slippery Rock University for their partial financial support of fieldwork. John L. Strother and Dale E. Johnson provided helpful review comments. J. Bot. Res. Inst. Texas 2(1): 487 — 488. 2008 488 J t tanical hl Texas 2( REFERENCE CHINNAPPA, C.C. 1986. Chromosome numbers in Antennaria (Asteraceae: Inuleae) from western North America. Canad. J. Genet. Cytol. 28:468-475. CHROMOSOME NUMBER OF THEVETIA AHOUAI (APOCYNACEAE: RAUVOLFOIDAE: PLUMERIEAE) WITH DISCUSSION ON THE GENERIC BOUNDARIES OF THEVETIA Justin K. Williams and Julia K. Stutzman! SE of Biological Sciences Sam n State University il Gg 77341-2116, U.S.A. ABSTRACT The mitotic chromosome count (2n = 20) for Thevetia ahouai is the first ted cl t for Thevetia sect. Ahouai. The count together with a previous count in Thevetia sect. Thevetia (also 2n = 20) provides an additional synapomorphy that further supports the monophyly of Thevetia as traditionally recognized. A discussion on the proposal to recognize Thevetia sect. Thevetia as the genus d p Cascabela is provide Key Wonps: Thevetia, Apocynaceae, chromosome number, Cascabela, Cerbera, Plumerieae RESUMEN Fl t Ati itótico (2n = 20) de T! tia al l pri Thevetia sect. Ahouai. Est t otro previo en uai: sect. Thevetia d 2n = =a) e una sinapomori adicional que apoya la monofilia de Thevetia como se ha Se aporta una pror Thevetia sect. Thevetia como el género Cascabela. Thevetia L. belongs to the A bfamily R lfioideae tribe Pl nd ight species of shrubs occurring from Central Mexico to porten South America (Gensel 1969; williams 1996; Allorge 1998; Endress et al. 2007). According to various specialists in the Apocynaceae, the generic boundaries of Thevetia vary. In the concept of Thevetia sensu K. Schum. (Schumann 1895; Gensel 1969; Williams 1996; Allorge 1998; Alvarado-Cardenas 2004), eight species are sub-divided between two sections: sect. Ahouai K. Schum. with three species—T. ahouai (L.) A. DC., T. amazonica Ducke, and T. bicornuta Mull. Arg.—and sect. Thevetia K. Schum. with five species—T. gaumeri Hemsl., T. ovata (Cav.) A. DC., T. peruviana (Pers.) K. Schum., T. pinifolia (Standl. & Steyerm.) J.K. Williams, and T. thevetiodes (H.B.K.) K. Schum. In the concept of Thevetia sensu Lippold (Lippold 1980; Alvarado-Cardenas & Ochoterena 2007) the three species of sect. Ahouai are retained in Thevetia; the other five species of sect. Thevetia are segregated into the genus Cascablea Raf. Although Lippold (1980) and Alvarado-Cardenas and Ochoterena (2007) segregate Thevetia sensu K. Schum. into the two genera Thevetia and Cascablea, at no point do they argue against the genera's “close morphological relationship" (Alvarado-Cardenas & Ochoterena 2007). In fact, a recent morphological cladistic analysis (Fig. 1, Alvarado-Cardenas & Ochoterena 2007) nests all eight species of Thevetia sensu K. Schum. in a clade supported by six synapomorphies. In short, Thevetia sensu K. Schum. is clearly shown to be monophyletic. Nevertheless, Alvarado-Cardenas and Ochoterena (2007) argue for its paraphyly and choose instead to recognize the two sub-clades of the clade (Fig. 1) as distinct genera: Thevetia and Cascabela. Chromosome numbers have proven useful in resolving generic relationships in the Ay (Van der Laan A Pid Wiens 2007): bs date, 73 of the 179 genera of the Apocynaceae s. sit (subfamilies R lf d Apocvnoidoideae) have been coun ed (Van der Laan & Arends 1985; Goldblatt € Johnson 2003; Williams 2007). one | I me ts É t. Thevetia (T. peruviana; 2n = 20) exist, none have been reported for sect. E The piece paper provides the first reported A MUN count for a species of sect. Ahouai and d the utility of in interpreting t relationship of Thevetia and NEM ip dal nd m fall D arro Dad tenini 30201 ITC 7 4 J. Bot. Res. Inst. Texas 2(1): 489 — 493. 2008 490 J ti tanical h Insti Texas 2( Carissa 2n = 22 Mortoniella 2n=32 Himanathus 2n= 18 Plumeria rubra 2n = 36 Plumeria obtusa [ Allamanda 2n = 18 Skytanthus hancorniaefolius Ke Skytanthus acutus Ánechites x =? Cameraria x = ? Cerberiopsis x = ? —— Cerbera floribunda ——1— Cerbera manghus | 2n = 40 — Cerbera odollam Thevetia ahouai Thevetia bicornuta Thevetia amazonica Thevetia pinifolia Thevetia ovata 2n = 20 Thevetia gaumeri — Thevetia thevetioides —— Thevetia peruviana De 1M E gi lly z x g f“Plumeriege” Ia (f 1. £F n 0 n.L.z 2007) a Thovetin concu € Schum and/ or“Cascabela-Thevetia” dade of Alvarado-Card | Ochoterena 2007. This clade is supported by y hies. b = Thevetia sect. Ahouai Mis Thauof linnald This clade i t | by t ynaj phi c= Thevetia sect. Theveti or 6 blea Lip ld This dade is sup- ported by four synapomorphies. MATERIALS AND METHODS Roots tips and voucher specimens were collected from a greenhouse specimen of Thevetia ahouai housed at the greenhouse of the Department of Biological Sciences, Sam Houston State University (Table 1). The root tips were fixed and analyzed for chromosome number using standard procedures (Raffauf 1964; Van der Laan & Arends 1985). A voucher specimen of the greenhouse plant was made and is preserved in the Warner Herbarium (SHST). RESULTS The format used for reporting chromosome numbers in this article follows that established by Strother and Nesom (1997). A mitotic chromosome number of 2n = 20 was recorded for Thevetia ahouai. Van der Laan and Arends (1985) reported chromosome lengths in the Apocynaceae to be between 0.5-4.0 pm, with the average chromosome length between 1-2 pm. The length of the chromosomes in T. ahouai varied between 13 um, consistent with most other chromosomes in the Apocynaceae. The base chromosome number (x - 10) for Thevetia ahouai is consistent with previous reports of x = 10 for Thevetia (T. peruviana 2n = 20; Gadella 1977; Ugborogho 1983; Van der Laan & Arends 1985; Santhosh & Omanakumari 1997). DISCUSSION Van der Laan and Arends (1985) postulated a base chromosome number of x = 11 for the Apocynaceae s. str. based on its prevalence in the family and on the observation that many of the plesiomorphic taxa pos- sess a base number of x = 11. A base chromosome number of x = 10 is found in four genera representing three of the 11 tribes recognized in the Rauvolfioideae (Endress et al. 2007): Hunterieae (Gonioma E. Mey.), Plumerieae (Cerbera L., Thevetia), and Vinceae (Ochrosia Juss.). Based on tribal circumscription (Endress et al. 2007) along with molecular evidence (Simóes et al. 2007), x= 10 is reconstructed as having evolved independently at least three times in the Rauvolfioideae. A cursory review of chromosome counts for the Apocynaceae (Van der Laan & Arends 1985; Goldblatt & Johnson 2003) reveals that at present the only chromosome ts for genera in the Plumerieae, and thus relatives to Thevetia (2n = 20), are for Allamanda L. (n = 9; 2n - 18), Cerbera (2n = 40), Himatanthus Willd. ex Schult. (2n = 18), Mortoniella Woodson (2n = 32), and Plumeria L. (2n = 36). At present there are no chromo- some counts for the remaining four genera in the Plumerieae: Anechites Griseb., Cameraria L., Cerberiopsis Vieill. ex Pancher & Sébert, and Skytanthus Meyen., and it is suggested here that effort be made to ou count of these taxa. Figures 1 and 2 present cladograms of the Plumerieae constructed from (Alvarado-Cardenas & Ochoterena 2007) and molecular evidence (Simóes et al. 2007), respectively Dipleld counts for the respective genera included in the phylogenies are presented for both Figures 1 and 2. Alvarado-Cardenas and Ochoterena (2007) presented six synapomorphies that described the Thevetia sensu K. Schum. clade (Fig. 1, branch a). The diploid count of 2n = 20 presented here adds a seventh syna- pomorphy. When interpreting their data Alvarado-Cardenas and Ochoterena (2007) state that “(here is still no consensus regarding the question of whether one should recognize one genus with two (sections) or two distinct genera (Lippold 1980), given that Cascabela and Thevetia are sister taxa." We would argue that the consensus in evolutionary systematics is to assign generic boundaries that reflect both monophyly and shared ancestry. The decision to divide a well supported clade into two separate genera may support monophyly; however, it excludes shared ancestry. Without prior knowledge most botanists Wong be un- aware that Thevetia and Cascabela ister taxa that sl Instead, an il more meaningful interpretation of the clade would be that Thevetia: sensu K. Schum. is monophyletic aad support is provided for the recognition of two sections as defined by Schumann (1895). In order to maintain systematic consistency as pertains to current trends in phylogenetic nomenclature we recognize Thevetia sensu K, Schum. and regard Cascabela and all taxa pertaining to the genus as synonyms of Thevetia. 492 J | of the Botanical R h Institute of Texas 2(1) Taste 1. Voucher specimen for the cl ber of Thevetia ahoudi. Taxon Voucher specimen Chromosome number (2n) Thevetia ahouai TEXAS: Sam Houston State University 20 Department of Biological Sciences Greenhouse specimen, 22 Jan 2008 Williams 2008-1 (SHST). a Allamanda 2n=18 Plumeria | 2n ^ 36 Thevetia | 2n 20 Cerbera manghas 2n — 40 Cerbera venenifera J f£. E r KEN i Fic p) T P | | | A A | I J f"Plumerieae" | ACKNOWLEDGMENTS We thank Tami Cook for providing us with access to her digital light microscope, and Bob Rhodes for mixing the Carnoy's solution and aceto-orcein. Sibyl Buceli and an anonymous reviewer provided valuable editorial comments. REFERENCES ALLORGE, L. 1998. Les Thevetia, pag des succulentes. Succulentes 21(1):23-32. ALVARADO-CARDENAS, L.O. 2004. Apoc) Flora del Valle de Tehuacan-Cuicatlan 38:1—57. ALVARADO-CARDENAS, L.O. and H. OCHOTERENA. 2007. A phylogenetic analysis of the Cascabela-Thevetia species complex (Plumerieae, Apocynaceae) based on morphology. Ann. Missouri Bot. Gard. 94:298-323. 2007, Enpress, ME and P. Bruyns. 2000. A revised classification of the Apocynaceae s.l. Bot. Rev. 66:1-56. ENDRESS, M.E., S. LIEDE-SCHUMANN, and U. Meve. 2007. Advances in Apocynaceae: The enlightenment, an introduction. Ann. Missouri Bot. Gard, 94:259-267, GADELLA, T.W.J. 1977. IOPB chromosome number reports LVI. Taxon 26:257-274. GENSEL, W.H. 1969. A revision of the genus Thevetia (Apocynaceae). Masters Thesis University of Connecticut. GoLbBLaTT, P. and D.E. Johnson. 2003. Index to plant chromosome numbers 1998-2000. Monogr. Syst. Bot. Mis- souri Bot. Gard. 94. LippoLD, H. 1980. Die Gattungen Thevetia L., Cerbera L. und Cascabela Raf. (Apocynaceae). Feddes Repert. 91:45—55 Rarraur, R.F. 1964. Some chemotaxonomic considerations in the Apocynaceae. Lloydia 27:288-298. SANTHOSH, B. and N. Omanakumarl. 1997. Karyomorphological studies on two varieties of Thevetia peruviana. J. Cytol. Gene. 32:95-98. waren IC... ; et |! of TI +i I H 403 SCHUMANN, K. 1895. Apocyna In: A. Engler and K.A. Prantl, Die Natürlichen Pflanzenfamilien. Wilhelm En- gelmann, Leipzig. 4(2):109— 189. Simoes, A.O., T. LivsHuLTZ, E. Cont, and M.E. Enbress. 2007. Phylogeny and systematic of the Rauvolfioideae (Apoc- ynaceae) based on molecular and morphological evidence. Ann. Missouri Bot. Gard. 94:268-297. STROTHER, J.L. and G.L. Nesom. 1997 Conventions for reporting plant chromosome numbers. Sida 17:829-831. UcBorocHo, RE. 1983. IOPB chromosome number reports LXXIX. Taxon 32:321. VAN DER LAAN, EN. and J.C. Arenps. 1985. Cytotaxonomy of the Apoc, Genetica 68:3-35. WILUAMS, J.K. 1996. A new combination in Thevetia (Apocynaceae). Sida 17:185-190. WiLLiAMs, J.K. 2007. Documented chromosome numbers 2007: Chromosome number of Laubertia contorta (Apocynaceae: Apocynoideae) and its phylogenetic importance. J. Bot. Res. Inst. Texas 1:431—435. 2007. Md p resch | 494 i tani itute of Texas BOOK NOTICES THE FLORA OF CHINA EDITORIAL COMMITTEE. 2007. Flora of China, Vol. 12. Hippocastanaceae through Theaceae. (ISBN Vol. 12: 978-1-930723-64-1, hbk.). Science Press (Beijing) and Missouri Botanical Garden Press (St. Louis), PO. Box 299, St. Louis, Missouri 63166-0299, U.S.A. (Orders: www.mbgpress. org, mbgpress@mobot.org, 1-877-271-1930). $125.00, 534 pp., 8 3/4" x 11 1/4". From the publishers.—“Eighteen plant families are described in this Tune SE these, Hippocastanaceae comprises the horse d longan, and species iie for ge wood; chestnuts and buckeyes; Sapindaceae, the SE family, i T1 : 1 ] DL o the ne m mainly E g yellow and green dyes; V itaceae tains the grap d ther vines; Tiliaceae includes the pon e lindens, or acus Mal , cotton, and okra; Bom! includes kapok; St li "Wee ae a li O P bro Wë of Camellia, as well as C. sinensis, the source e tea.” Editors Wu Zhengyi, Peter H Di Hong De tate, la is a Gm ica SC M il It includes 18 families, 125 genera, and 1275 species, ich fi d 767 ies (6 d 29 species ( (2.396) are introduced to China.” The families eh are (in dh edd -— Actinidiaceae, M per M Dilleniaceae, Elaeocarpaceae, Hippocastanaceae, Leeaceae, Malvaceae, Ochnaceae, Pentaphylacaceae, Rhamnaceae, EE SE Slad- eniaceae, Sterculiaceae, Theaceae, Bee and iow ub included in in text: Preface, Introductio g Nomenclatural Novelties, Inde d an Index to Families. A large ap of C] dits 35 Administrative Divisi j inted on both the inside front ‘front ] d t] de ] / we bd rr back endpaper. Tue FLORA OF CHINA EDITORIAL COMMITTEE. 2007. Flora of China Illustrations, Vol. 22. Poaceae. (ISBN Vol. 22 illustrations: 978-1-930723-61-0, hbk.). Science Press (Beijing) and Missouri Botanical Garden Press (St. Louis), PO. Box 299, St. Louis, Missouri 63166-0299, U.S.A. (Orders: www.mbgpress.org, mbgpress@mobot.org, 1-877-271-1930). $125.00, 937 pp., 8 3/4" x 11 1/4" [hau edi ignifi , namely, the grasses From the cone —“Volume 22 of t | H 7 " 1.1 1 SN Se 1 L D 1 4 1 id ly ltivated ri , wheat, barley, malze (Poaceae or Gramin 1 If H TE 11 1 + a 11.1 J: or corn (Oryza, Triticum, Hordeum, and Zea). yg g life. Al ft] depicted here, including many woody ba bes. occur E cxi in China. Editors Wu Zhengyi, ba H en 1 Hong D tate, “This RR is the iens in a series of 24 volumes of Flora of China Illustrations. It i f China ND ds Volume 22, which I 1 226 genera with 1795 species in the Poaceae. This vol f ill i lud I I ing l of 1271 species, 24 subspecies, and 50 varieties in 226 Pune E Poaceae.” Also included in the text Preface, Acl ledg ts, Index to Chinese Names, Index to Pinyin Names, Index to dex to Families in the Flora of Chi d the Flora Reipublicae Popularis Si and Published E of s did fri Jl ci f TL TI A 1 E est, Rs WE an 4 1 y: “y [e] i i IT. 1 1 J 1 1 f 1 1 I i EN YI J. Bot. Res. Inst. Texas 2(1): 494. 2008 ANATOMÍA FOLIAR DE ALGUNAS GRAMÍNEAS ALPINAS Y SUBALPINAS DEL EJE VOLCÁNICO TRANSVERSAL, MÉXICO versidad Autónoma de Querét Facultad E Cencias Naturales, qute en Biología as Ciencias s/n, 76230 Juriquilla Querétaro, ME. gomezsauaq.mXx RESUMEN 1 a m of s 1 1 Ir: EN — 1.11 "E 1 1 1l or € EE A D Let: [All C "e EN 4 Ec a A 7] dj A NM 1 à LUE + £3 Ta g p g ] I , de las cuales 13 son hato Co identifi E Pooide y Cloridoide y las vías fotosintéticas C-3 y C-4. Se determinaron nueve caracteres s diagnósticos en a en abaxial y 17 en la estructura interna Ge la lámina foliar. Existen dci anatómicas p p de los p E y loss e A Se E iis es el estrés Ge quer Ec las especies. Asi, en la lámina foli ta el d ollo de fibras, yor, AT 1 pa ES PES es DR 1 DÉI ai. ~ 1 ž ch pO A Z son conspicuas, | , gui) y grosor y El patrón anatómico d peci tra relación estrecha con ] dici ientales donde habitan.y es similar al de las especies de zonas desérticas. ABSTRACT The highest elevations in the Transmexican ' ic Belt contain alpi d subalpi lands with duced extensi d thei habitats are fragile. The leaf bl of twent ies of that in tl lands was s tudied. Thirteen species are pue and iL ae ii anatomical pes Pooid and Cloridoid, and the C-3 ana c- 4 oe oun A. were identified. here are — EE 1 e TNT TES land itl from t! balpi aa ard altitude e is ue idis aA d stress fot e prairie EES Thus, ith i ing altitude the p er in the leaf Kee St m Fiba rrirlloc th larl a al le ara Le 3 the ch] ] d th ] duced with altitud on M Eo ue of the species conspicuous. In contrast, yma and the stomata number are reduced with altitude. 1 1 ] JE 1 FLA Lk Lis LA YY Ail AL r * ux IE 2 EE F INTRODUCCIÓN El Eje Volcánico Transversal es un sistema montañoso que cruza transversalmente México, en su mayor parte entre los paralelos 19? y 20? N. Esta área se sitúa a lo largo de unos 950 km desde la región del Volcán Ceboruco al poniente hasta el Pico de Orizaba al Oriente (Rzedowski 1978). En México, el Eje Volcánico Transversal incluye las prominencias topográficas de mayor altitud que, en sus partes altas albergan man- chones de vegetación alpina y subalpina y el área total ocupada por esta vegetación es reducida y está sujeta a cambios ecológicos. Entre 3500 y 4500 m de altitud se observan comunidades de gramíneas amacolladas, denominadas pastizales o zacatonales alpinos y subalpinos. Los pastizales alpinos se localizan por encima del límite de " vegetación Gd El límite superior de este pastizal se sitúa SIE ODE de did m de SE ER en medio d Rosen: de Pinus L., Quercus L. y Abies Mill. y la cota seus de 4000 m señala la altitud máxima que md el ugue ICONS 1975 y 1978; McDonald 1998). ] e de los centros más ricos, por unidad de área, de especies edemas estrictas. DE de las cuales fueron conocidas para la ciencia hace apenas unos años y, salvo sus distribuciones, no existe conocimiento adicional de estas especies raras. Los hábitats de estas regiones ‘Autor para correspondencia J. Bot. Res. Inst. Texas 2(1): 495 -515. 2008 406 | tani i lexas 2(1) son los más frágiles dada la corta duración de la estación de crecimiento y la lentitud de los procesos de regeneración que siguen a la perturbación (McDonald 1998). Los pastizales alpino y subalpino, en forma natural, están condicionados por ciertos factores como temperaturas bajas, disponibilidad de agua, intensidad y duración de luz solar y nutrientes, entre otros. De manera artificial, están condicionados por la tala inmoderada y los incendios, estos últimos usados para provocar el brote de retoños que sirven de alimento para el ganado. La vegetación alpina-subalpina del Eje Volcánico Transversal es única en su tipo y de alto interés biológico, muestra una composición florística notoriamente distinta a la de otras regiones alpinas del país (McDonald 1998) y, es de esperarse que, las especies de estos Ge posean EE anatómicos distintivos y propios de su hábitat. Los estudios en pocos y en su mayoría han tenido, principalmente, un enfoque — donde n anatomía se EE parcialmente. Así, se han abordado revisiones taxonómicas y descripciones de nuevas especies de los géneros Festuca L., Sporobolus R. Br., Triniochloa Hitchc., Trisetum Pers., Muhlenbergia Schreb., entre otros (González-Ledesma 1991; González y Koch 1994; González-Ledesma et al. 1995, 1998; Hernández y Engleman 1995; Peterson y Herrera-Arrieta 2001; Reyes 2004; Cruz Díaz 2005). Algunos estudios que detallan la anatomía foliar han permitido separar géneros (Valdés-Reyna y Hatch 1995) como Erioneuron Nash y Dasyochloa Willd. ex Rydb., establecer grupos y sub- grupos de las especies | de Eragrostis Wolf (Gómez-Sánchez y Koch 1998), proponer la separación de Swallenia alexandrae (Swallen) Soderstr. & H.F. Decker de la subtribu Monanthochloinae Pilg. ex Potzal (Gómez-Sánchez et al. 2001), delimitar algunas especies de Trisetum (Hernández y Engleman 1995) y de Muhlenbergia (Peterson y Herrera-Arrieta 2001), entre otros. Este estudio describe y analiza la anatomía de la lámina foliar de algunas gramíneas que habitan en nas alpinas y subalpinas provenientes de los principales picos montañosos del Eje Volcánico Transversal, en México. Asimismo, los patrones de los gradientes ecológico y geográfico se comparan con las caracterís- ticas anatómicofoliares de los diferentes taxa. MATERIALES Y MÉTODOS Muestras de especímenes de 21 especies se obtuvieron de los Herbarios CHAPA, ENCB, IEB y QMEX, acrónimos según Holmgren et al. (1990). De cada especie, se estudiaron 1-16 especímenes según la di- sponibilidad de material. Hojas basales maduras se rehidrataron con agua caliente durante 1-5 minutos. De la parte media de la lámina foliar se hicieron cortes transversales a mano libre con navaja de afeitar para observar la estructura interna. 1 con hipoclorito de sodio al 6% (cloro de uso doméstico) para aclarar y suavizar los tejidos y posteriormente se lavaron con agua corriente. La epidermis abaxial se aisló mediante la técnica de raspado directo y las muestras | tes se montaron con jalea glicerinada, sin tinción (Gómez-Sánchez y Koch 1998). La epidermis adaxial no se aisló debido a la presencia de surcos fuertemente BEE P lecc ] ión directa al microscopio para definir y diferencias anatómicas entre especies. Mediante encon se hizo una comparación de los patrones anatómicos entre especies y éstos se compararon con el hábitat. La anatomía foliar se definió con la termi- nología y descripción de acuerdo a Ellis (1976, 1979) con algunas modificaciones según Gómez-Sánchez y Koch (1998). Las figuras de la anatomía foliar de las especies se elaboraron con ayuda de un microproyector o por observación directa al microscopio. En el caso de la epidermis, están orientadas de tal manera que el eje longitudinal de la lámina foliar aparece en posición horizontal en la ilustración y el ápice de la lámina se ubica a la derecha. RESULTADOS Y DISCUSIÓN La anatomía foliar se estudió en 21 especies de gramíneas, seleccionadas por su presencia destacada en los principales picos montañosos del Eje Volcánico Transversal, de las cuales 13 son alpinas y ocho subalpinas (Cuadro 1). La lámina foliar mostró 17 caracteres diferenciales en la estructura interna y nueve en la epidermis Cuabro 1. Especies estudiadas y ambiente en que habitan Especie Ambiente Agrostis bourgeaei Subalpina A. subpatens Subalpina A. tolucensis Subalpina Bromus anomalus Alpina B. exaltatus Alpina e orizabae Alpina C. toluc Alpina schampsia liebmanniana Alpina Ossa californicum Subalpina estuca hephaestophila Alpina : livida Subalpina F. rosei pina F. tolucensis Subalpina Muhlenbergia nigra Alpina M. orophila Alpina M. quadridentata Alpina Peyritschia koelerioides Alpina Phleum alpinum Subalpina Poa annua Subalpina > orizabensis Alpina Triniochloa stipoides Alpina abaxial. Con base en la anatomía foliar se identificaron los tipos anatómicos Pooide y Cloridoide. Las es- pecies se arreglaron en dos grupos: Alpino y Subalpino. En el grupo Alpino se identificaron los dos tipos anatómicos, que corresponden a las rutas metabólicas C, y C,. El grupo Subalpino posee especies con tipo aatómico Pooide y metabolismo C, (Brown 1958; Ellis 1977). Las plantas C, tienen anatomía no-Kranz y las plantas C, tienen anatomía Kranz o de corona (Brown 1977; ME éd ” especies no-Kranz (C,) fijan el CO, DEE mediante la enzima ribulosa 1,5-di- fosfato carboxilasa el primer producto es el ácido 3- so esco (3-PGA), proceso que sucede en las células clorenquimatosas del E Cosa 1975 y 1977, Ellis 1977, Gould y Shaw 1983). Anatómicamente estas plantas muestran una vaina interna de paredes g dea todos los haces Pi o por lo menos EECH EC EH sala cimo a tienes los primarios. Al exterior de esta vaina se sitúa ge vaina Į e clor o tiene pocos y de menor tamaño que los cloroplastos del mesóhlo. El mesófilo est dispuesto irregularmente (Ellis 1977). Las especies Kranz (C,) fijan el CO,, durante las horas de luz, en las células del neo mediante la enzima fosfoenolpiruvato carboxilasa y el primer producto de la fotosíntesis es el ácido oxalacético , ácido málico o ácido aspártico. Estos productos son transferidos a las células de la vaina Kranz donde sucede la descarboxilación y luego la fijación (Brown 1975 y 1977; Ellis 1977; Hattersley y Watson 1976). La anatomía de estas plantas se distingue por posset una o o vainas dee MSN muestran una vaina parenquimatosa (Kranz) alrededor de los haces as del mesófilo. Esta vaina tiene sus células grandes y con paredes más gruesas y tienen en cloroplastos en See cantidad y tamaño que aquellos del mesófilo. El mesófilo es regular alrededor de los haces vasculares (Ellis 1977; Johnson y Brown 1973) y con frecuencia es Biens por columnas de células incoloras. Anatomía foliar de Gramíneas Alpinas.—La mayoría de las especies alpinas habitan principalmente por encima de 3500—4500 m de altitud, aunque llegan a crecer en zonas boscosas y en niveles altitudinales inferiores. Estas plantas son herbáceas, perennes, rizomatosas, desarrollan macollas robustas y fuertes y alcanzan de 30—70 cm de altura. Su presencia es notoria en claros de bosques de Pinus, en lugares pertur- bados, a a orilla E caminos y en pce pie ee grupo de e po ae anatómicos en Se + lámina f foliar que | IV E 5 CO L I LI 498 tani ti Texas 2( eon la insolación es alta, la disponibilidad de agua es baja, las sequías y temperaturas son extremas, oca- y heladas y los vientos son fuertes. En estas zonas, al igual que en las áreas secas o desérticas, ocurre el fenómeno llamado “verano de día, invierno de noche”, ya que durante las horas de insolación se alcanzan temperaturas bastante altas y, al llegar la noche, la temperatura desciende tanto que generalmente el suelo se congela y las plantas no pueden disponer de agua. Esto permite que las plantas muestren rasgos morfológicos que también son propios de aquellas que crecen en ambientes secos y desérticos (Smith y Young 1987). En las zonas subalpinas es común encontrar representantes arbóreos de los géneros Juniperus, Pinus y Abies (Starker 1950) por lo que en estas áreas los vientos no son tan fuertes. A mayor altitud, más desarrollados son los rasgos anatómicos que proporcionan protección contra la evapotranspiración, los da- Dos mecánicos y la insolación, tales como los estomas protegidos, desarrollo de aguijones grandes, micro y macropelos, desarrollo conspicuo de fibras y di tas en trabas y células buliformes en abanico y grandes que intervienen en la involución de la lámina, entre otras. Las especies alpinas exhiben un patrón anatómico distintivo. La lámina y ganchos numerosos, las púas son largas y de base ancha sobre el margen, el esclerénquima está dispuesto en trabas o en hebras de tres células de ancho y 6-8 células de alto, el clorénquima es reducido y las células buliformes son grandes que se disponen en formas diversas, y las células incoloras son conspicuas. Las es- pecies alpinas son de fisiología C,, excepto las tres especies de Muhlenbergia que son C,. Este metabolismo les confiere rasgos anatómicos adicionales que les aseguran su desarrollo en estas zonas. Epidermis abaxial.—Los caracteres anatómicos diferenciales de las especies alpinas se muestran en el Cuadro 2. Todas las especies alpinas desarrollan ganchos en las regiones costal e intercostal y son de tamaño mediano y base ancha (Figs. 1 y 2). Púas más grandes (Figs. 2, 3 y 4) y de base más ancha que los estomas y se desarrollan sobre el margen. Esta presencia de aguijones (ganchos o púas) refleja la continua condición de sequía a la que las especies están expuestas (Ellis 1979; González-Ledesma et al. 1995; Gómez- Sánchez et al. 2001; Mejía-Saulés y Gómez-Sánchez 2001) y, a mayor estrés hídrico, mayor es la cantidad y el tamaño de los aguijones y el grosor de sus paredes. El estrés hídrico es habitual en las zonas alpinas, pues la disponibilidad de agua se reduce por la congelación y la alta evaporación producida por la intensa radiación solar. Así entonces, la presencia de aguijones, en las especies alpinas, proporciona un mecanismo de protección contra el daño provocado por la insolación proyectando sombra sobre la lámina y reduciendo el calentamiento y la excesiva pérdida de agua. A mayor tamaño y frecuencia de los aguijones, mayor es la sombra que proyectan y se reduce la exposición a la sequía a la que estas especies están expuestas. En Poa orizabensis Hitchc. (Fig. 4), Triniochloa stipoides (Kunth) Hitchc. (Fig. 13), Calamagrostis orizabae (Rupr. ex E. Fourn.) Beal (Fig. 10), Calamagrostis tolucensis (Kunth) Trin. ex Steud. (Figs. 2 y 21), Muhlenbergia nigra Hitchc. (Fig. 6) y Muhlenbergia quadridentata (Kunth) Trin. (Fig. 8) los aguijones son más abundantes y de mayor tamaño que en el resto de las especies alpinas. Esto explica que dichas especies predominen en los zacatonales alpinos (Rzedowski 1978) y se desarrollan en los hábitats de mayor altitud. Triniochloa stipoides (Fig. 13) es la única especie con pelos en su epidermis. Estos resultados sugieren que en el resto de las especies alpinas los micropelos y los macropelos no son necesarios ó bien son remplazados por aguijones. Las especies alpinas muestran en la epidermis abaxial, una cutícula más gruesa que aquella que se observa en el grupo subalpino, sin embargo, en ambos grupos esa cutícula contribuye a regular la pérdida ca: " EE RE S qam. È de agua por evapotranspiración. Los estomas, en algunas especies, no son visibles (Figs. 2, 5, 8-12) probablemente porque están trasla- pados y ocultos por los EES une SE y Koch te EE están embebidos en el mesófilo (Salisbury y Rose 1994) y f liminado tra, o bien, porque los estomas están confinados a los surcos adaxiales de la lámina (mes Sánchez et al. 2001). En cualquiera de estos microambientes la temperatura es menor y los estomas están protegidos, por lo que también se reduce la pérdida de agua por transpiración. Los cuerpos de sílice son numerosos y más abundantes en las especies alpinas que en las subalpinas y toman formas de hígado (Fig. 5), de pesa (Fig. 8), son redondos (Fig. 6), cuadrados (Fig.1), altos y estrechos 499 Gómez and Téllez, A sesonuis sesonuls Sesonuls EIERE) SJUSWEPUNJOILA sesonuls SJUSWPPUNJOIA sesonuls sepie sein|o» ap ses] e ses] e 949] əyuəwepunyzod e SESI] 31usugepunjoJg So|eui[2Hue sepaJeg sajuasne sajuasne Ssajuasne sajuasne Saluesaid saquasne SO|9dOJ2IN Saluesne sajuasald sojuasne sojuasne sejuasne sajuasne so[edoJoewW sopeDie|e sopebieye sopeipend A opebiu sopeipend sopuopa! esad SD ap euuo., SeLUOISS SELSIIY c SOJSIA OU SEJ9|IU z SCIDIIY € SOJSIA OU SOISIA OU ap uonisodsig sonndyya SOJSIA OU soondi[e soondije SOJSIA OU SO]SIA OU SEUJOJSI ap CUO ly A OM IS IS IS IS IS IS ap uopenpuaisJg sajuasaid sejuasajd Soluasne saJuasne So]uasaid SO]SIA OU $9|£1502431ul seuofimnby Sojuasne sajuasne sajuasaJd salussaid $31U3S310 S9]U9S910 S3/81503 sauolinby SapjoHa[a0y sapiodys SISUaqDzHo pytydoso pibiu Djbjuopuponb Diipsiufad bopuporun] Dod Dibiaquapnyy Dibiaquajunyy Dibiaquojunyy apadsa/19]3e1e) (UGIDENUNUOD) z avn sesonuls sesonuls Sesonuls sesonuis Sesonuls sesonuls ajueuepunjoud ajustuepungoJd 39JUSuJepunjoJd ajuauJaAs| SJUSUUDAS sesonuls sebJe| se|n|a» ap 31usulsA3| e 249] e Sesi| e SESI] e SESI] e sesi| ƏJUJWƏAƏJ| e SESI] SOIBUIDIUUE SAPaled soluasne Ssajuasne sajuasne sojuasne sajuasne sajuasne sojuasne soado IA sojuasne saluasne sajuasne sajJuasne saJuasne Sojuasne sajuasne so[edoJ2ewW sopuopa) sopebJe|e sopebleje sopebieyje opebly opebiu A opebiu A sopuopal A sopeipeno sopeipend A sopeJpeno SD 9p Puto, SO]SIA OU SOJSIA OU eajly | SOJSIA OU SOJSIA OU eJəjly | SeJallU Z SPLUOISS op uonpisodsig SOJSIA OU SOJSIA OU soondi[e SOISIA OU SOISIA OU soondijo soondi|a seulO1so SP PLO IS IS IS OU OU IS IS ly A DY ep uopemuaialiq sejuasne Soluasald sajuasald sajuasaud sojuosaJd sojuasaJd Sejuasne S9|e1502J91u! seuofinby sojuesaJd $9]U359.d sejuasaJd saJu9saJd saJu9saJd sajuasaJd sajuasaJd $3|£1505 souolinby 19501 Djydo3saDydoy DUDILUDUIQO!H sisuaonjo] aDquzHo snjb)jpxo SN[DUIOUD ponjsaj HITCH Disduipipsag SHSOJÓDUID[D) SIJSOJÓDUID[D) snulolg snuoig apadsa/I12]p4bp) 2721 BP sodianD = $) 9150313141 uoiDai = [y 18505 ugiBal = Dy 'seuid | [9p | pida e| 319 UI] 128Je^ “7 CC? 500 YH = — e P $ A = - " H + b a E e à Co EE = Wu om h . ei e A a e Des- Sg E TESES E d A E CU Leak spotters Chris ee ae A TROP oI a pi uk? RA — > SEH = AS eS Pa um a ee rdi A a E - , ie T DH a SAA A ag EIA MAS RENA E AM m fa 0.3 mm y 1 d 0.1 mm g e E f E. ¡AAN IN AO a eege EE Tur qum... qu wm cÉ UN RADA ARA RAMUS [AO 0 0 a LS SA ¿A E A elis SSC up > PF Rd LI AA ER! Ll em Ls : VS D «MA Ús a <> a +. mar Ne Ge c S Co i SUSAN Voz Matres | teen | + en gt Lo ae. m m, CERES : = a mm a é s A on , a? Ke ALT = IS uu Hr UR PRO e LAA a spese dor soapy eg ee bl “au = m m AM N a a + A LA — wT nis d i "— pum Md SS OR ms 1 3 0.4 mm 4 0.1 mm ee CEET A CREER Wee DEE a Loire E Em " UTE at MT re 0.5 mm 5 b 0.3 mm 6 Fic : —f Er Sin D L ata [| pz D TU [| | D Ir » P Ex 8 (Di y mí 5963, IER) 9 Ful q PA D (Ni z a al. 707, EB). 3. Muhlenbergi hila (Rzedowski 36012 ENCB). 4. Poa orizabensis (Rzedowski 38363, ENCB). 5. Desch Ga lik WM» ENCB). 6. Muhlenbergia nigra (Koch 280, ENCB). «c = célula corta, cl = célula larga, cs = cuerpo de sílice, e = estoma, g = gancho, mi = micropelo, p = púa, rc = región costal, ri = región intercostal. (Figs. 2 y 4), o alargados (Fig. 10). Este incremento en el número de células de sílice sugiere que la lámina de ] : Tas "ES " Pi : “a más alta, y 1 I : 1 Lud a las E mecanismo que contribuye con soporte y regula la temperatura de la lâmina foliar (Kaufman et al. 1985). Las paredes anticlinales de las células largas y cortas son lisas (Figs. 1, 7, 9 y 10), moderada (Figs. 8 y 12) o profundamente sinuosas (Figs. 6, 11 y 13). Las paredes anticlinales sinuosas de las células largas y los cuerpos de sílice numerosos brindan rigidez a la lámina (Ellis 1979) y la protegen también contra daños mecánicos provocados por el viento o los animales. Muhlenbergia orophila Swallen (Fig. 3), M. nigra (Fig. 6), M. quadridentata (Fig. 8) y Festuca hephaestophil d. (Fig. 11) muestran células largas con paredes Gómez and Téllez, Anatomía foliar de algunas gramíneas 501 a be s Sor SS SS AAN a TE edid —— - inicie rum MEN m | : I ET Sm eo AS p E EE COEN 6 “tam So > Cx Da Oo De E ` vi s A O AR eege e Soe, — a iene wok ae Un. DP geet eeng Fun eC unb is ee EE is: Rue vn Dia dad A LT o EE A Ee e z mm: E gor eie 5 Pa PARE = 28 > Eegen 5 À, o = - —— 1 EP E kun Pi 5 E DEI E Lee. ES MI E Ce Ne ere AO E ED = A EPA ss m O) O na rie mo OEA AN me EP Cc = LC on PD a ae a im EEO ee eS erat item, oe tg i GER 3 a anal QUA Su ee D = II [e] AS Ss dd se Lek e Cf e A ( ————À E A Eer mou eo VERA A a cnr MIS e ER aa ae; - + er > unas tor EE (Anda ct ERE O li - EK a KE -— d eu er ER qui i X Eres EE, wm Mr eng VA TR PME CMRI? (T Fm ren ae epa Po e e dier rst mm P EN E 1 AA pU MR E LAT VITE Jerg TI x co E AAA Es [IA TER MORE ERAS u o e tog Exe aA yt 3 aed us cla p e — A A A AO pS Aa petty > ee. em ALA "T =>, qué e EDU A. AA a eU PAX AC A Ror A = a — E) Mee aN VN CRT CS AAA po = = E. E E mp e as ae EE dum OE AAA H 8 et ae “cat Mist” s, m 0.6 mm CP TEARS e p ee O IEA > E ——— —M3 = === — 3d — A A AAA k o E AAA GA CEO E e oi A E CEC == REE EE a, a) a a a FC EM = PEA E === Äert) <= e E S SI EN go] LU Mane We selon o A i n AH duutz e TIET II iail é diii g | assenza AAA oe ss Ln Car ! ` " á a É rar 1 fp oe E eaaa S ri ro oe a ea o TE AAA —— do e m me m FM SER hh EA ae AARMA Ea eba nisso SATA UE 0.2 mm 12 cà —— eo GC ee DS P o A Em Mis, : = a EI [E a RM a LE UI. ria] LES ma Figs. 7-13. E a L LEM EN | | nz El: A 8 Lon 7 n Jl. (R J Fea A.A , ENCB) Q TT L gi ] ap dd r1 Fi (G Af 1914, ENCB) 9 n. LP Le i f - oos | rn d r’ gan’ -ENCB). 1 1n Fal g E’ cl L Tis A “ONA ,TEB). 1 J | Laf (G rp ` García 90, IER) 12. E rosei (Pérez Cálix 26 , IEB). 13. Triniock les (Rzedowski 42438 8, IEB) cc = célula corta, cl = célula larga, cs = cuerpo de sílice, ma = macropelo, LE región costal, ri = región intercostal 3 T ran Dos H In LI Ma dien adis £T BILE df 502 anticlinales moderada o profundamente sinuosas que brindan rigidez a la lámina lo que sugiere que son especies que resisten fácilmente daños mecánicos eS See E viento (Ellis 1979). Estructura interna.— Los iferenciales de la estructura interna que distinguen a las especies alpinas se muestran en el Cuadro 3. El quina es copioso y forma hebras (Fig. 22) o trabas (Figs. 14, 16, 23 y 25) orientadas adaxial o abaxialmente como una extensión de la vaina de los haces vasculares. Algunas veces, el esclerénquima se desarrolla en bandas (Fig. 18 y 26) que recorren la anchura de la lámina. Su presencia sugiere una baja disponibilidad de agua, que es un elemento importante de sos- tén, y que evita la apertura total de la lámina impidiendo que ésta tenga una prolongada exposición a la luz "ow SEH dU Las us hacen las veces ES varillas BEE sostén a la lámina y amortiguan los fil ofrecen a la lamina resistencia mecanica los fuertes vientos, p y evitan su Rea Por el contrario, un desarrollo escaso de id explica la presencia de culmos laxos y hojas delicadas y frágiles, en g ue habitan los o menos adversos (Gómez-Sánchez y Koch 1998; Nava-Rojo et al 2002). Una quilla conspicua provista de grandes y fuertes trabas o hebras de esclerénquima es evidente en.Bromus exaltatus Bernh. (Fig. 14), Triniochloa stipoides (Kunth) Hitchc. (Fig. 17), Poa orizabensis (Fig. 19), Deschampsia liebmanniana (E. Fourn.) Hitchc. (Fig. 22) y Festuca hephaestophila Nees ex Steud. (Fig. 24). Esta quilla proporciona sostén y está asociada al control de la pérdida de agua por lo que sugiere que estas especies están NN a una excesiva EE EE Ha lámina foliar de estas especies, a través de la quilla, manifiesta apertura parcial mostrando lución en forma de “V.” En SS resto de p indi alpinas la lámina t involución en rn de “U” o es convoluta, lo cual completamente a la insolación (Ellis 1976; Gómez-Sánchez y Koch 1998; Mejía- Saulés y messi 2001) y así regula la pérdida de agua. Las células buliformes se restringen a los surcos adaxiales, son conspicuas, en forma de abanico (Figs. 14, 16, 23) o son redondas (Figs. 18, 22 y 24). Las trece especies alpinas, estudiadas aquí, muestran células buliformes. Estas células responden a cambios de turgencia provocando la involución de la lámina, mecan- ismo que regula el paso de luz hacia el mesófilo y la evapotranspiración (Ellis 1979). Estas células no sólo intervienen en la apertura y cierre de la lámina, sino también en s absorción y retención de agua (Tschirch 1882, citado ud Els Vind Así entonces, cuando las células buliformes disminuyen su turgencia provocan enlaláminau gr itando así que la lámina tenga una iva exposición a la luz solar (Ellis 1976, Gómez-Sánchez y Koch 1998). De esta manera, las especies alpinas son beneficiadas porque las células buliformes permiten la involución de la lámina y absorben y retienen agua favoreciendo su super- vivencia en los zacatonales alpinos, mismos que están expuestos a una intensa y prolongada insolación y a sequías extremas. Se | jue atraviesan el 5filo se EECH en Muhlen- £1.41 H 1 E aoe bergia nigra, M. orophila y M. quien m (Figs. 16, 25 y 26). Est asociadas a las buliformes, funcionan como canales facilitando y regulando el See de luz hasta las células de clorénquima (Ellis 1976) o las de la vaina Kranz. Esta regulación, además de incidir en un metabolismo eficiente, evita alteraciones al material genético de los cloroplastos, mismas que pudieran ser provocadas si la lámina tuviése una prolongada exposición a los rayos ultravioleta (Ellis 1976; Salisbury y Ross 1994; Gómez-Sánchez y Koch 1998). Las tres especies de Muhlenbergia (Figs. 25 y 26) estudiadas aquí, manifiestan una anatomía Kranz y metabolismo C,. Estas Mee muestran el mesófilo reducido, regularmente dispuesto alrededor de los haces vasculares, estos últimos, están protegidos por una vaina a manera de corona (vaina del haz) de células grandes donde está contenida la mayor parte de | plastos y la presencia de células b oras es importante (Ellis 1976). Est | las pl tol con éxito las alt por una insolación excesiva (Salisbury y "ipse 1994). Esta est alee es común en ambientes como id de los pastizales alpinos o las zonas secas o desérticas, donde la radiación solar es alta y prolongada. Todas las especies alpinas, excepto Bromus anomalus Rupr. ex E. Fourn. y B. exaltatus Bernh., tienen 503 aJuasny ajuasne ajuasaud 31U3SaJd 3juesne ajuasne ajuasne elno SEJSIA OU sepuopal ODIUEqe sepuopal sepuopal oDIuege oDjuege S3UJOH|Nq Spina? sajuasne saJuasne s3aJuasne sojuasne sajuasne sojuesne sajuasne SBJOJODUI Se[n|g2 SepeAo A sasen Buen salejinbueu saJejnbuen) saJe¡nbuel) SepeAo epuopal saquasne "pe se|nsoo ap PLLC auany əyən} Piaf alan} Piaf Piaf 391U3SNE ‘pe soouns ap pepipunioJg Jembaul Jepnboui ien 6a Jeinbaul Jenboul JI Dau Jejnbaul euimbua1op Jap ojbauy 31uasaud 3]uasaud 3juasaid 3]U9S910 3Ju3asaud 31ueseJd syasaid ` AH aqua onunuoo euumnbugio[5 qe ajuasne qe Op ge pe A qe pe A qe dAH U2 HA 9p ugirdnuaiu| pe ejuasne ajuasne ajuesaud pe ejuasne ajuasne HA e| 9p ugisua1q ajuesne ajuasne ajuasne aluesald 31uesaud ajuasne ajuasne ‘qe 3 ap epueg e1qay ajuasne elqau epueq epueq əzuəsne sjuasne [2150313341 qe 3 ap uornquisig eJqeu 3jussne eJqau eIqay e1qay eqe 3juesne SAH Ue pe 3 ep uomngusid eqen eiqau eJqau eqen egen eqeu eqen SAH Ue qe 3 ap uomngusig eJqau sjuasne eJgau egay 3jussne POS eqen dAH Ue pe 3 ap ugionqussig eqen e1qay eiqau POS eqe1] eqen eqen dAH US qe 3 ep ugionalasiq epuopel epuopal epucpal epuopel epuopal eDeAO epuopal SAH Ue HA 9p PWC epuopsJ EDeAO epuopal epuopal epuopal epeAo epuopas dAH U9 HA 9p PUNO, [BUD |PJ3ua2 EINEN Op qe jenuso |BQuaD eulUuJe| 2] US AH 9p UODISO4 19501 Dydo3sabyday DUDIUUDUIQO! SISU2MJO] aDqpzuo 5n]b)jDXa SN/DUICUD D)n)521 Dnjsaj Disduipipsag spsobpuibjp) SIJSOJÓDUID|D) DT snuio4g epadse/Iapee) A | | 'Zey jap eulea = HA 'souepunoss salejnosea sa»eu = SAH > '€ ouavn 'SOMBUILIA SaIe|nseA sa2eu = JAH 'saJe|noseA sa2eu = AH "euinbuai9[sa = 3 'eixepe = pe 'jeixege = qe à + + =| [ove Fad E 504 azuəsəd ajuasne ajuasaJd ajuasaJd ajuasne ajuasne elmo OoIueqe oDIuege sepuopal oDluege SEISIA OU SEJSIA OU S3UJOH|NQ Sena) sajuasne sajuasne sajuasne PULUN|OD ua eULUN|OD ua euuinjoo ua seJo|o»ut SÉIS? sepuopeal Sepero saJuasne sepuopal PDPAO SepeAo Se]|11505 ap euo. Glaf Glaf 3juesne aJuasne ayan] e1301] pe ONS [ep pepipunjoJg sejnBaul Jenbaul Jepnbaui Jejnba) Jejnba! sena ewnbuaiop [ep ojbauv ajuesne a}uasaid 391U3SSIO ajuasne 3juesald sjuasne — AH anus onunuoo euinbusio|> pe A ge ajuasne aquasne pe A qe qe pe A qe dAH Us HA ap vondn1a1u| ajuasne pe A qe pe A qe ajuesne ajuasne ajuasne HA PI ep ugisua1x3 ajuasne sjuasne ajuasne ajuasne ajuasald ajuasne ‘ge 3 op epueg ojuesne ajuasne ajuasne ajuasne epueq 3JU3sne |91509/3141 Ge 3 ep uomnguusig eqen eiqau eqen eqen eJqau eqen SAH us pe 3 ep ugionquisiq esgay eqen eqes eqen elqau eqen SAH ua qe 3 ep uoionguisiq eqe eqəy ajuasne eqen elqeu eqeil dAH Ua pe 3 ep UONNquasig eqen egen egay eqen e1qay eqen JAH ue qe 3 ep ugionquisi(] epeAo epuopel epeAo epuopal epeAo epuopal SAH Ua HA ap ewo epuopal epuopaJ epuopal epuopal epeao epeao dAH Ua HA ap euo ¡Es GIVÉEN [24JU3) IER [nus |[e3ua2 eup] e| US AH 9p uopisog sapiouajaoy saprodns SISUIQDZLLO pjiydoso Dubju pjDjuapuponh DIpsyufad Dojiporun] Dod Dibiaquojunyy pibiaquayyny pibiaquayyny apadsa/I3ppIe) (UQDENUNUOS) E ouavnm) LE a4 || A A r £..1* | D at n ZE, SSC aff NATI NA, pe Pred pe Err WW. Le AE ETA A AS KF LL E NIC PCT 5 RÄ CHAP! ARTE Witte T, Yn SH. VOD SA PAAD 5 SS VIN Walt lia a A HN Zä tam Nu à oa itte s BD LEO CUN Dor T fo XC: oe ate Vë mx, ARO Md atis et oes a eras d o e A pee e d (e = AE e D LJ r a " F — CO LA A " Dv » M j e n SA Sa Së Wë iat th fi / da, Kai di =-: ) > Dei? Oé (echt, EX Y E © Mo SEL "a9 e a, EPA e a s a Déi ND | Feta Ned G Tte E TS ANS MNT ¿a AE Zen, KE ie Pi: G d * fr - " m D ae eat MANN E Et e ro» CAT $ ‘ar À DM hii 4 qdo = LES mia a eg WE AO D EEN a WE, EE > sa L^ ef, nef F PA A LES ee Ach) AO on mw z om REEL e “alo qp um dre Boro E, d AA AS eiert H amoo IA OMS ERA AAA "Té d E79». C Ei mor ^ P Ze at LJ 5 ag Ke Kéi ZS Pus 7 IUS 2 d -— wo. WEE SE E D SE ? ` Fics.14—20. Estructura int le la lámina foliar de especies alpinas. 14. Bromus exaltatus (Díaz Pérez 5963, 1EB). 15. Festuca rosei (González et al. 26, IEB). 16. Muhlenbergia orophila (H 325, IEB). 17. Triniochloa stipoides (Rzedowski 42438, 1EB). 18. Calamagrostis orizabae (Ventura 1824, IEB). 19 Poa orizabensis (Koch 75519, ENCB). 20. B lus (Rzedowski 29299, ENCB). cb = células bulif , ci = células incoloras, cr = clorénquima regular (radial), cre lorá i i lar ( lial), eb lará ] : hiad lov | . tah , L : i e : hvs = haz vascular secundario, q = quilla. hojas más estrechas y delgadas que las subalpinas. Esto sugiere que hay una relación estrecha entre la anchura y el grosor de la lámina y la exposición continua a ambientes secos, aspecto que coincide con Red- man (1985, citado por Gómez-Sánchez y Koch 1998) quién determinó que la anchura y el grosor de la hoja están asociados a la resistencia a sequía. Bromus anomalus y B. exaltatus (Figs. 14 y 20), como las demás especies alpinas, desarrollan un es- clerénquima copioso que forma trabas abajo y arriba de los haces vasculares, tienen láminas estrechas, células buliformes dispuestas en forma de abanico y poseen aguijones de paredes gruesas y lignificadas 506 Se CC ty O d PTE. Ae D Ü Ate A jr Ha tos A E eso 5 4 TL omy A KA PELIS ci ter "VIE! AAA WC A ee mets arr. die SESS a ae a n A Hm ze e, A [] esti ats + + 7 tree woe WAS DS ar, CA 22> fon, ‘an wt te, «ie mes vw RE oa mn go D E uy pé ERA e m Sech : 0.3 mm AS E E PEX-- LEE PLC O E ei a HO" Dé CR SO Pr Ti d ny * "NALE Ta us f ow ata - br 0, ES v ond 3h a) EIA AA ES e E A, a MAAN, AL SP TZ CELUM AP A Ser + 4 T E ` 2.4: ta Sa ST Zë ver o A E Ak, a $ 2. ^ EY ei: e "de E Lë P A UE AS Jr vr: ae Te à $e MEAT: ISE ej ao A UR Ze Ee A T am. E. Pp aa A te i,» rez “SP e bw ug ^ (oc 1% EL AS A H^ ERA KSE EX 4 Rh GA $ rhet] a e PE Ena E, A e AS ee t5 - SE ta eh» wl. bar D “ E sl" re EE m Le TEA PRO Cl zd wm de La KE) bes 21-26. Est 2 SW a A | [| Lz H "dë | J . la! 31 st PAR? |] A ($ a E 85, IEB) 33 D E J $ A) L] (Ki i 96269, ENCB) 73 n j L^ ES f A | (R J | 11933, IEB) IA Fr P L L A Lo (G £p y F E 151, IEB) DE AA. LE L 9 a 7 LEN | de ¡Miranda ot al 204 IFR} ae Muhlont H H "y r 7rr IER) ph E PR | Lert, A ralidlacincalarac rra || E E) H 1 (no radial), ck = corona Kranz, eh lerénqui hebra, eb lerénqui banda, et lerénqui tral en su estructura interna. No obstante, su epidermis abaxial ( en las especies subalpinas. Estos rasgos anatómicofoli e Figs. 1 y 7) es semejante a la que se observa eren que, aunque estas especies predominan en lugares altos, cuentan con adaptaciones que les permite sobrevivir a las adversidades del tiempo de las regiones intermedias de los bosques y los pastizales. A 4 Ju Js E J : Lal Eet Aonripe habitan ] J en los bosques d posible localizarl e Pinus, Pinus-Quercus y Quercus, desde 1500 hasta 3000 m de altitud, a 11 D unque también es damente 10-40 cm de altura, no obstante, algunos individuos llegan a m de aproxima- edir hasta 90 cm ó más. La lámina Gómez and Téllez, Anatomía foliar de algunas gramíneas 507 foliar de estas especies no desarrollan los mismos | d tección que | pecies alpinas. Sobre la epidermis, los ganchos tienen su base mediana ó más ancha "m los estomas y las púas son cortas. En el margen, los ganchos son cortos y de base angosta ó están ausentes en Agrostis bourgeaei E. Fourn. y Phleum alpinum L. En su estructura interna se identifica el esclerénquima escaso y dispuesto en hebras, a veces se observan pequeñas trabas de 2 a 4 células de ancho "n de los Gees Keser SE el clorénquima es abundante con arreglo irregular y las células buliformes están son pequeñas, a veces son redondas. Todas las especies subalpinas, aquí "v carecen de "m incoloras y de la corona Kranz manifestando una anatomía no-Kranz y metabolismo C,. Epidermis abaxial.—Los rasgos anatómicos observados en la epidermis abaxial de las especies sub- alpinas se describen en el Cuadro 4. El grupo de especies subalpinas, tratadas aquí, contrasta fuertemente con el grupo alpino en la mayoría de sus rasgos anatómicos. En la epidermis de estas especies subalpinas los ganchos son escasos y de tamaño menor (Fig n Agrostis tolucensis Kunth y Festuca tolucensis Kunth (Figs. 28 y 29) poseen peq dio en la región costal. Poa annua L. (Fig. 31) exhibe escasos aguijones en E margen de n lámina y, en otras especies los aguijones están ausentes Asimismo, las especies subalpinas carecen de micropelos bicelulares y macropelos. Estos rasgos sugieren que las especies subalpinas no están expuestas a fuertes vientos o heladas frecuentes, como ocurre con las alpinas, por lo que no requieren de una fuerte protección. El tamaño pequeño de los ganchos y la ausencia de micropelos y macropelos hace suponer que estas especies no requieren evitar la pérdida de agua pues los árboles de los bosques les proveen de sombra, tem- peratura y humedad estables (Rzedowski 1978) por lo que la evapotranspiración, a través de los estomas, es baja (Salisbury y Ross 1994). La ausencia de aguijones (en la mayoría de las especies), micropelos y macro- pelos sugiere que estas especies no están expuestas a sequía (Gómez-Sánchez y Koch 1998; Nava-Rojo et al. 2002), o al menos no a condiciones tan adversas como sucede con las especies alpinas. Esta condición contrasta con la presencia de aguijones, micro y macropelos en especies alpinas y también en aquellas de zonas desérticas. Los estomas están dispuestos en una o varias hileras (Figs. 30 y 31). En ocasiones, los estomas no están visibles, probablemente porque están cubiertos por los escasos aguijones, están inmersos en el mesófilo y éste se eliminó durante el proceso de aislamiento de la epidermis, o bien están confinados a los surcos de la lámina. En la mayoría de las especies subalpinas, los cuerpos de sílice son escasos y/o pequeños y se observan en forma de hígado (Fig. 33), alargados (Figs. 27, 28 y 32) y redondos o cuadrados (Fig. 34). Este carácter está asociado a las paredes anticlinales de las células largas lisas (Figs. 27, 31 y 32) y ambos rasgos, permiten que las láminas sean laxas y menos rígidas que aquellas de las especies alpinas, lo que coincide con la opinión de Ellis (1979). Estructura interna.—| en el Cuadro 5. El esclerénquima es escaso y se desarrolla en hebras, — trabas de apenas 2-3 células de ate (Figs. 36 y o o en peda (Fig i ere desarrollo MÀ EH del EE contrasta con el POY Ca: a sl al : Tan bens Anan AA + PENAL OSA I sugiere q protecion contra «daños Eege Las especies subalpinas exhiben clorénquima abundante EEN ES manera ca y sus ecules generalmente son grandes (Figs. 38-40). Las células del mesófilo g É una buena absorción, retención y disponibilidad de agua. De acuerdo con el criterio de Ellis (1976), la ausencia de costillas y surcos en la lamina de especies como Dissanthelium californicum (Nutt.) Benth. (Fig. 41) o Poa annua L. (Fig. 42), o costillas y surcos leve- mente pronunciadas como en Festuca tolucensis Kunth (Fig. 37), se asocian con la apertura completa de la lámina lo que sugiere que estas especies no están expuestas a la radiación solar por periodos prolongados o la intensidad de la luz solar no es extrema. Festuca livida (Kunth) Willd. ex Spreng. (Fig. 35) y las tres especies de Agrostis (Figs. 38, 39 y 40) muestran surcos y costillas pronunciadas, no obstante, la cantidad lo TE AN lexas Z(1) £T a rd 508 Sesonuls Sesonuls sesonuls sesonuls ejueujepunjoid = ejueujepunjo)d ajuawepunjold ajusuwepunjold sepie; sena) ap SesI| e 39A9| e 3A9| e sonal e 249] SeSI| pel Ses|| so|eulpouue sepaJeg sajuasne sajuasne sajuasne sajuasne saquasne sajuasne Ssajuasne sejuasne SO|9dOJ IA sajussne sajuasne sajuasne saluasne Sajuasne Ssajuasne sajuasne sajuasne sojadoJe IA sopuopal sopebieye sopebieje Á SOPeJpen) opebiy opebiu sopebue|e sopebue|e sopebue|e SD ep ewo SEI sejany z SO]SIA OU Sau | SOJSIA OU Blat | elo | eJo|u i seulojsa ap uonisodsig soondifo sopeJpeno SOJSIA OU soondi|e SOJSIA OU SO2naye soondi[a sopeipen» SeulO]s2 9p LUJO, OU IS IS IS IS ou IS IS IH A DY 9p uopenpueiajig sajuasne sejuasne sejuasne soluesaid Ssajuasaid sojuesaJd SoquasaJd saJu3saJld S3|e1s02J31u! seuofinDy Sojuasne sequasne Sejuasne səpuəsad saju3asald Sajuasne SsajuasaJd sajuasne S9[e1s02 sauo(inby pnuup unuidip DpIAI| sisuaonjo) OTT suajpdgns sisua2njo} Jabinoq bod unajid Dn]524 DMI594 untjaujupssip spsoiby 5isoiby snsouby əpədsə / Japeie) E 1915031314] ugiDal = [M 21503 uoi re Ua ‘QD [IS op sodiano = $) asa oe | IP Sa19126J6 “y OVaYND e A | A A Li [s ERRE ARA a Mim GU UA dd LT (per LED FEVER E s AW CG p aq, elt nas T3 SIX KA reas yn E Et AEN IA rm Mera BO EE E pep — ensi RR ces. cmm. Emm > rr i n ae e PERE E SEH geen — e e = LE C oc VIO RI eege E | bibi ttd Qe tg emm am : Es Sege ees SE e Vestib od being rr (en Ca eeneg? ria | Kee , "I iae RR. p Zen Siemens Se ege Ee T ERE d =N E o = MES ies [i I ORIO T fN eer | indus peer gen Leederen Gëtter ge de p E apa, dir c aa ep AAA A x = = el ^a na M — ee A eee = El vba di oF a cs oy, T.U E kV"? ERN Eer Cj Sc Ge de LT ai r^ mul: "i ai Og x, i deg E EE POSE. ? E Dom 0.3 mm 33 CS 0.3 mm 34. CS Figs. 27-34 Fnid ic al ial dala limi Gelli J H hale! 77 À m g a (R 4 Le 37853, IER) 2 Ag i Fen L: 23315, ENCB) 20 F Fi i D (G Zi J Ve 206, IEB) 1 DL | m Fn. i Lear 107, ENCB) 21 n / Fr 17, IER), A ra) L ri ín i eegne a IER) 22 FX: i ae if. In J ki 20174, ENCB) 24 F. A Pn. (G L4 | ) EJ r I sa i, 32. Ag | ( IEB). cs = cuerpo de silice, g = gancho. STE! 3a1uasaud aluasaid Sjuasne ajuasne ajuesne aluasne aJuasne aluesne e| mo sepuopal OSIUEqe odiuege oDjueqe sepuopal odJuege sepuopa) sepuopal seuuoJng senap sajuesne sajuasne sequasne sequasne saquasne sajuasne sejuasne saJuasne SeJOJODUI senan epeao epejpeno sajuasne sajuasne PDPAO JeinDuen epeao A Jejnbueum A seinbuewy epeao Se||l]502 ap euuoJ ajuasne ajuasne 21361] oen) Glaf Ei Si sant ‘pe ODINS jap pepipunjojd Jena Jejnbaui ien BaN Je nba! Jepnbaui Jenboui aen 6a] Jejnbaui eujmnbugJop rep ojbauy 3JUISDIO Slussaid ajuasaid auəsəd Sjuasaid 3]U3saJd 3juesald aquasald AH enue onunuoo euuinbuaJlo[5 pe A qe ajuasne peAge qe qe qe |eixeqe qe dAH Ue HA 9p ugmdnuaiu| ajuasne ajuasne 3]u3sne ajuasne ajuasne ajuasne Sluesne ajuasne HA 8| 9p ugisuaxq ajuasne ajuasne ajuasne ajuasne aluasaid ajuasne ajuasne ajuasne ¡e1xeqe q ap epueg aquasne ajuasne ajuasne ajuasne epueg eJqau ajuasne aJuasne |e3502J81Ul qe 3 ap uopngquisiq ajuasne elqay ojuasne ajuasne 3juesne eJqau egay eiqəy SAH Ue pe 3 ep ugmngustq ajuesne eiqau ajuasne e1gay eqen eJqau eJqau eJqau SAH Ue qe 3 ap ugmnquisig eqen ajuesne equi 3JU3sne aluasne eJqau eJqau eqein dAH ua pe 3 ep UQDNQuIsIg eqei} e1q3y eqen eiqau eqen eIqeu elqou eqen JAH Ue qe 3 ep uogpngquisiq epuopaJ epuopaJ epuopal epuopal epuopel epuopal epuopal epuopal SAH Ua HA 9p ewo eDPAO epuopel epeao epuopal Ppuopal epuopal epuopal epeAo dAH Ue HA 9p euo [eus ¡e11U9) ¡nues [2/11U9) IEN ¡211ua) qe DIEN eure; e| ua AH ap uopisog Unomaojibo pnuup uwnuidjo DIM] SIsuaonjo] suazodgns sisuaonjo) rapabunoq unioujupssig bod (Dal Dn]5oj DJS] snsoJby snsoiby spsoiby apadso/1a)e1e) 510 ZY op euie^ = HA 'souepunoss saJeIn2seA sadey = SAH 'soueuJud S92EU = dAH 's9Je|n2SPA Sa5eu = AH 'euinbus!apso = 3 '[eixepe = pe 'jeixeqe = qy ‘seuldjeqns sanadsa se) ap Luau, ein3onaisa e| ue sopeAJasqo saJeiuaJallp saJa12eJe) € ouava» Gómez and Téllez, PA hh i | E TOR LN ren Oe: É as 3 SUR f- S A + pat SES yt 4 TEN = Se KEE? Se A IES: ah KS Zar am Kai ^ Se NI. NS as Um ER rm AAA Zë 6 A LI Y O SE Di V Mte - oo I d TS Pass [ YA. Zéien eh $ Data +04 Ge Si an -È AY (ës prio M A d ^ PT t“ 5 ¿Gal b AS eT pr peto e BOSSA À - YE OPA a 4 AAA) d > KA - SR x : Gi * F te nS LJ A W LA P t Ee EE E DE 01) ^J ew + di LE Y, * + Lé yor + ". Cé $ En é 8 + M D pt e ki " a a b a e Te ant d "AN N en [3 A O Ka + k wei d Ka m o^) Herr) Ee = D Np DA E Ge Su AJA “3 rè, pat Fea Kl A ESE T. ML A Eer E E e NEE 222789, ETE Beta fo Ee sw ae GL e SE H Be Mages. GAR LO * A? Ze 2 .... cb Fics, 35-42. Estructura interna de la lámina foliar d | bal T" livida (Ménd. 188, IEB). 36. Phi [pinum (Reeder & Reeder 5374, ENCB). 37. Festuca tolucensis (González y V IER). 38. Agrostis bourgeaei (Rzedowski 25949, ENCB). 39. Agrostis subpatens (Rzedowski 36013, IEB). 40. Agrostis tolucensis (Rzedowski 41921, ENCB). 41. Dissantheli lifornicum (Rzedowski 22848, ENCB). 42. Poa annua (Morones 052, IEB). cb = células bulif , cre = dorénquima irregular ( lial), eb énqui banda, eh lerénqui hebra, et = esclerénquima en traba, q = quilla. de mesófilo es abundante, las células epidérmicas son grandes y el esclerénquima es escaso, lo que indica que las especies disponen de buena condición hídrica. Agrostis subpatens Hitchc., Dissanthelium californicum y Poa annua (Figs. 39, 41 y 42) tienen una quilla, sin embargo, en la primera especie la quilla está provista de una pequeña hebra de esclerénquima, en la segunda el esclerénquima está ausente y, en la tercera, la quilla tiene solamente un pequeño cúmulo de células de esclerénquima. En los tres casos, la quilla solamente es un elemento que da forma a la lámina y proporciona un ligero sostén (Ellis 1976). 511 512 t tani i Texas 2( Phleum alpinum (Fig. 36) y Poa annua (Fig. 42) muestran células buliformes dispuestas en abanico y ocupan la mitad o más de la mitad del grosor de la lámina. No obstante, las células buliformes en la mayoría de las especies subalpinas son de menor tamaño que aquellas de las especies alpinas, lo cual sugiere que la involución de la lámina ocurre en menor grado que en las especies alpinas, puesto que no están expuestas a una pérdida excesiva de agua. En contraste con el grupo alpino, las especies subalpinas estudiadas carecen de células incoloras y tienen anatomía no-Kranz y metabolismo C,. Esta condición también es típica en especies que crecen en climas templados y presume que las especies subalpinas reciben luz en cantidad e intensidad adecuadas (Ellis 1976; Hattersley y Watson 1975; Nava-Rojo et al. 2002). Dissanthelium californicum (Fig. 33), Festuca livida (Fig. 34), F. tolucensis (Fig. 29) y Phleum alpinum (Fig. 30) tienen, en su epidermis, células largas con paredes anticlinales profundamente sinuosas y numerosos cuerpos de sílice como el grupo de especies alpinas. No obstante y como el resto de las especies subalpinas, carecen de aguijones o cuando existen son cortos y de paredes delgadas poco lignificadas, en su estructura interna el esclerénquima es escaso y el mesófilo es conspicuo, mismo que se dispone de manera irregular y sus células son grandes, aumentando así el grosor de la lámina lo que sugiere una presencia de humedad estable (Ellis 1976). Festuca tolucensis muestra una combinación de caracteres tanto de especies alpinas como de las sub- alpinas. Esta especie exhibe, en su estructura interna, pequerias trabas abaxiales de esclerénquima que conectan a los nas veseulates primarios, el clone niquama esta names GES y sus células son grandes di te, la involución de la lámina tiene forma de *U" y, las púas y ganchos sobre las caras EE y banal son Brandes y de paredes delgadas (Fig. 37). Sin embargo, el esclerénquima es escaso y ausente de la superficie adaxial de la lámina, los aguijones tienen paredes con escasa lignificación y son cortos como en las especies subalpinas pt habitan "ines Pd an ome eden de caracteres nije Ft 1 M A tadaa A ems A PA PA los pastizales M y su P en claros de be cues de pino-encino (Rzedowski 1978). Situación similar ocurre con Phleum alpinum, especie con algunos caracteres de las especies alpinas como el esclerénquima en trabas abaxiales y adaxiales que conectan a los haces vasculares con la epidermis, células buliformes en abanico y paredes anticlinales de las células largas de leve a fuertemente sinuosas (Figs. 30 y 36). No obstante y como las especies subalpinas, carece de cuerpos de sílice o son escasos, los aguijones son cortos y de paredes delgadas, el esclerénquima es escaso, los surcos y costillas adaxiales son apenas notorias y la lámina está expuesta y tiene mayor amen y ined Existen dif ] especies q en | tizal las que habitan en el interior de los bosques Las pac desitrollam y mecanismos SE Ge protegen c contra 7, f] aan erlarida en ci i MT 1 nia D diversos daños y j foliar g | avansa en altitud, ] dias 4 [A345 A ] ` «T —€——— P m las altas montañas, de manera que éstas desarrollan cáracieres monies. que les permiten permanecer con éxito en esas altitudes y condiciones ecológicas. A partir de un bosque de encino y hacia el pico de una montaña, a mayor altitud mayor es la agresión ambiental y el desarrollo de fibras del esclerénquima se in- ] son conspicuas, la anchura y el crementa, el tamaño de las células buliformes es mayor, las células 1 grosor de la lámina es menor, el clorénquima y la cantidad de estomas se reducen, y los aguijones aumentan su tamaño y el grosor de sus paredes. Especies, como Muhlenbergia nigra, M. orophila y M. quadridentata tienen anatomía Kranz y una fisiología alterna, la C, Estos rasgos les permiten tolerar las distintas condiciones extremas (Ellis 1977 y Salisbury y Ross 1994) que prosperan en estos ambientes de alta montaña. Dadas las condiciones tan especiales que imperan en los ambientes de alta montaña, es posible que otras especies alpinas y subalpinas tengan distintos patrones anatómicos a los descritos aquí, lo cual podría confirmarse estudiando un número más amplio de especies. por TE HI : A " NE JA E J || H 513 APÉNDICE 1 ESPECÍMENES REVISADOS i E Fourn. MÉXICO. Ixt pal tacié peri tal dei tige IÓN y f de Z iapan, 8 km al EM Río E S. D. Koch 75328 (ENCB) | tald a Zoquiapan, B em al sur de río as. uae 186 (ENCE? Tialmanalco: La Ciénega, región de Menos Cuatas S. Acosta 961 (ENCB). Ayucan: 3 km al NW de San Luis Ayucan, Acosta 35955 lowski 38021 ioe mm Hitchc. SE TEEN la Joya de Alcalican Rzedowski 36013 (IEB). A | s kung la Joya de Alcalican, Castellanos s.n. (ER) Se Cerro Teotepec, Rredowsk bd EE B.D ITO CEDERE Le leg. Ves Magi an Puerto de las Cr ices, H 98 (IEB). MICHOACÁN rro del Burro, Ee g Si de Chincua, M Mei (ER) B E. Fourn. HIDA Zempoala: SEH de Ds Pitos, culis 277 (ENCB), 5 km al i de e Atizapán, Rea 20921 (ENCB). Bromas See Gs MÉXIC g Puebla, 8 km, al S de rio Frio, camino 5, Vega 288 (ENCB) HIDALGO. El Chi le Las Vent (si de Pachuca), Vera 777 (ENCB). Calamagrstsorabas (Rupr. ex E. Fourn) Beal. Ventura 1824 (IEB) (sin registro de lugar de colecta). Calamagrostis tolucensis (Kunth) Trin. ex Steud. México. Ixtapaluca: cerro Tláloc, Flores 288 (IEB). San Mateo: a 2.4 km de la desviación al Nevado de Toluca, Deg et dl. on (IEB). Tlalmanalco: la Ciénega, región de la cabeza del Ixtaccíhuatl, Rzedowski 37689 (ENCB). Santa Ana de alrededores del Llano de la Horca, Rzedowski 31668 (ENCB). POEDER 23 km ns Río Frío, barranca entre supercarretera y carretera libre Meer Geier Ge E B) Edel Cerro ros serranía deg Ajusco, ied 12626a (ENCB). E. Fourn.) Hitchc. MÉXICO. Tlal ano Grande, arriba de san Rafael, Rze- dowski 38266 (ENCB). La Ciénega: e de renas Cuatas, de la cal lel | ihuatl, Rzedowski See EB), eH pie SW dei Ixtaccíhuatl, 4 N Rzedowski 20163 (ENCB MICHOACÁN. San Laguna e cerca ER SCH EE a bci o Tancítaro: above apo on cerro Tancitaro, Ge 585 (ENCB). DISTRITO CO, EE ad ENED) aan a californicum (Nutt.) Benth. MÉXICO. Texcoco: cmo Tages N | | | y y Atkinson 3950 (ENCB). Festuca ! hila N Steud. MÉXICO. Ixtapal tacic 'imental de i ti y enseñanza de Zoquiapan, 8 km alS de RÍO uis a uf et al. 99 (IEB). Festuca livida (Kunth) Willd. ex png González y Hernández 104 (IEB), Méndez 188 (IEB). S ta. Festuca rosei Piper. MICHOA ÁN El Frijol, al N de Santa Juana, Pérez 690 (EB). Festuca tolucensis Kuntl MICHOACÁN. Zi á te alta del cerro San Andrés, Zamudio 5586 (IEB). MÉXICO. Amecameca: la Joya de Alcalican, Cruz 14 (IEB). Muhlenbergia Ee Hitchc. MÉXICO. Amecameca: 10 km al E de EE E el Sule a Tlamacas, Rzedowski s.n. (ENCB). MICHOACÁN. Quiroga: cerro del Tzirate, López 755 (IEB). M S DISTRITO FEDERAL. Cuajimalpa: Ae de las Cruces, Herrera 325 (IEB). mm qua aa (Kunth) Trin asia Ladera E del Popocatépeti, 10 km al W de Santiago Salicinta, Herrera y Cortés 917 (| rqueza, 1 km al S de la Marqueza, Herrera y Cortés 929 (IEB). MÉXICO: a 7.2 km de la ae al Nevado de Toluca, carretera SE Manrique et al. 260 (IEB). MÉXICO. Amecameca, La BO de Alcalican, EECH 1319 (IEB). MICHOACÁN. Quiroga: cerro Boe Tree López 1176 (IEB). MÉXICO. Ixtapaluca: | ío, Vega 135 (ENCB). DISTRITO FEDERAL. Milpa Alta: cerro Tilcoayo, Miranda et al 894 (IEB). Peyritschia koelerioides (Peyr.) E. Fourn. MICHOACAN. Huiramba: parte alta del Cerro del Burro, Rzedowski 41933 (IEB). Phleum alpinum L. VERACRUZ. eters Cofre de Perote, lado N en la base de! macizo de roca que se encuentra en la cima del Cofre de Perote, González et al.136 (ENCB). Poa annua L. MEXICO. La Joya de Alcalican: parte SW del Ixtaccíhuatl, d 25676 (IEB). Amecameca: 10 km al E de Amecameca, Jiménez 76 (IEB). Exp. de Inv. de Zoquiapan, 8 km al S |S de Rio as Koch (Aa TALENCE); HIDALGO: Fpazoyucan: 3 km al SE de Real del Monte, Medina-Cota 1466 (EB). DISTRITO lowski 20381 (IEB), Tlalpan: Cerro del Ajusco, Bojorquez s.n. (IEB). VERACRUZ. Calcahualco: 8 km al O de Calcahualco, Mejía- -Sauléz 1151 (IEB). MICHOACÁN. Pátzcuaro: Joya de Las Navas, Escobedo 669 (IEB). JALISCO. Mazamitla: a 6 km de Mazamitla por brecha a microondas El Montoso, Morones G. Geo (EB). oa BEE nn MÉXICO. Ixtapaluca: Estación Experimental de Inves- tigación y sao de Zoquiapan , o, Koch 75519 (ENCB). Triniochloa stipoides (Kunth) Hitchc. PUEBLA. as de Y | Za a uds 369 (COCA) MICHOACÁN: | See eg km al E de Morelia, sobre la carretera: a Mil Cumbres, ENEE (IEB). MICHOACÁN. Pat carretera Ge Morelia, Díaz 1670 (IEB). MICHOACÁN. Pátzcuaro: Joya de Las Navas, ds ! (IEB). o Jenécuaro: terrenos pertenecientes a El Tejocote, Rzedowski 50574 (IEB). MICHOACÁN. Nahuatzen: Cerros Pelón y Cherán de la Mojonera, e 402 (IEB). Nahuatzen Jii iio cerro La Bandera, cerca de Comachuen, Martínez 752 (IEB). Michoacán: 19 km al E de Querétaro, sobre la carretera a Maravatio. EE E SE (EB) GUANAJUATO, o 3 km al NE de Santa. Rosa. Rzedowski 42044 (IEB). SCHER TARO. Pin | le H nza 2926 (IEB). Pinal de amoles: al S de los Pinos, Carranza 2912 (IEB). VERACRUZ: ise Vigas Los Hermanos, Ventura es PC OAXACA. Sola de Vega: carretera Puerto Escondido, 13 km al S de Sola de Vega, Koch et al 79559 (CHAPA). GUERRERO: Heidoro, 68 km de filo de caballo por terracería a El Paraíso, González-Ledesma et al. 487 (CHAPA). NUEVO LEÓN. Zaragoza: 9 km de Zaragoza por camino de terracería a La Encantada, González-Ledesma y García 683 (CHAPA). 514 | loft tanical h Institute of Texas 2( AGRADECIMIENTOS Agradecemos a los curadores de los herbarios IEB, CHAPA, COCA y ENCB por facilitar las muestras de las especies estudiadas. Robert W. Jones amablemente revisó el resumen en inglés. El Programa Integral para el Fortalecimiento Institucional (PIFI 2003-23-08) otorgó un apoyo a la segunda autora. Este trabajo se desarrolló con un apoyo parcial de la Comisión Nacional para el Conocimiento y Uso de la Biodiversidad (CONABIO) a través del proyecto U031. REFERENCIAS Brown, WM 1958. Leaf anatomy in grass systematics. Bot. Gaz. 119:170-1 78. Brown, WV. 1975. Variations in anatomy, associations, and origins of Kranz tissue. Amer. J. Bot. 62:395-402, Brown, W.V. 1977. The kranz synd and he pen in ge sao Mem. Torrey Bot. Club 23:1-97. Cruz Diaz, J.E. 2005. Revisión taxonómica del género fia (Poaceae: Pooideae) en México. Tesis.de Licencia- tura. Facultad de dS iino: Universidad A Autonoma de iaa Querétaro. Erus RP. 1976, A | | y in the Poaceae. |. The leaf blade as viewed in transverse EH Bothalia 12:65—1 09. Eius RP. 1977. Distribution of the Kranz syndrome in the southern african Eragrostoideae and Panicoideae ac- cording to bundle edd anatomy ane vlog. Agroplantae 9:73-110 Es R.P 1979. A | | parative leaf anatomy in the Poaceae. II. The epidermis as seen in surface view. Bothalia 12:641-671. GouLp, FW. y R.B. Shaw. 1983. Grass systematics. Segunda edición. Texas A&M university press. College Station. Texas, E.U.A. GoMEz-SANCHEZ M. y S.D. Koch 1998. Estudio anatómico comparativo de la lámina foliar de Eragrostis (Poaceae: Chloridoideae) de México. Acta Bot. Méx. 43:33-56 Gómez-SÁNCHEZ M., P. DAVILA-ARANDA y J. VALDÉS-REYNA. a dd anatómico de Swallenia (Poaceae: Eragrostidae: Monanthochloinae), un género monotípico de ^ ica. Madroño 48:152-161. GonzáLEZ-LEDESMA M. 1991. Revisión del género Festuca L. (Gramineae) en el Eje Neovolcánico Transversal. Tesis de Maestría en Ciencias, Colegio de Posgraduados. Montecillo, México GOoNZALEZ-LEDESMA M. y S.D. Koch. 1994, Tres especies nuevas de Festuca (Poaceae: Pooideae) de México. Novon GonzAtez-Levesma M, M. Gómez-SANCHEZ, S.D. Koch y T. Mesta-SauLés. 1995. Triniochloa talpensis (Poaceae: Pooideae), a new species with deciduous leaves from México. Novon 5:36-39. GONZALEZ-LEDESMA M, S.D. Koch y M. GómEz-SANCHEZ. 1998. Two new species of Festuca (Gramineae: Pooideae) from Mexico. Novon 8:147-151. HarrersLer, PW. y L. Watson. 1976. C, grasses: an anatomical criterion for distinguishing between NADP-malic enzyme species and PCK-malic enzyme. Austral. J. Bot. 24:297-308 HERNANDEZ TÓRRES |. y E.M. ENGLEMAN. 1995. Anatomia de la lámina foliar del género Trisetum (Gramineae: Pooideae) en México. Acta Bot. Méx. 31:39-50. HoLmGren, K.P, N.H. Howesp y L.C. Barnett. 1990. Index herbariorum. Part 1. The herbaria of the world. Octava edición. New York Botanical Garden. Nueva York. Johnson, S.C. y WM Brown. 1973. Grass leaf ultrastructural variations. Amer. J. Bot. 60:727-735. KAUFMAN, PB., P. DAYANANDAN, C.l. FRANKLIN Y Y TATEOKA. 1985. Structure and function of silica bodies in the epidermal system of grass shorts. Ann. Bot. 55:487- e McDoNALD, J.A. 1998. Fitog fía e historia T.P, R. Bye, A. Lot, y J. Fa. Eds Diversidad biológica a México, Soe distribución. Instituto de Biología, Universidad Nacional Autónoma de México. México. Pp. 665-685. Mejía-SauLes T. y M. GówEz-SÁANCHEZ. 2001. Primer registro de Triniochloa andina (Poaceae: Pooideae) para la Flora Colombiana. Caldasia 23(2):405-412. lis A en A EEN ac Md EE ei, | D +1 "m: ITI l A " fa Enl! J || H 515 at "i Nava-Rojo A. M. Gómez-SáncHEz y M. GoNzÁLEz-LEpEsva. 2002. Agrostis avenacea (Poaceae: Pooideae) first record for the mexican flora. Sida 20:423-429. | H I CAA LI i H Fry RA L. | L aja A PETERSON, PM. y Y. HERRERA-ARRIETA. 2001. A leaf bl | vey gia ( g Sida 19:469-506, Reyes Gutiérrez, G.B. 2004, Estudi fológi tómico del género Sporobolus R. Br. (Poaceae: Cynodonteae) de la península de Yucatán. Tesis de eno culo de Medicina Veterinaria y Zootecnia, Universidad Autónoma de Yucatán. Mérida Rzepowskl, J. 1978. Vegetación de México. Limusa. México SALISBURY, EB. y CW. Ross. 1994. Plant physiology. Fourth edition. Wadsworth Publishing Co. Belmont, California. Smity, A.P. y T.P. Youna. 1987. Tropical alpine plant ecology. Ann. Rev. Ecol. Syst. 18:137-158. VaLDÉS-REYNA J. y S.L. Hatch. 1995, An anatomical study of Erioneuron and Dasyochloa (Poaceae: Chloridoideae: Eragrostideae) in North America. Sida 16:413-426. 516 BOOK NOTICE M.K. SAUNDERS. 2007. World Checklist of Dioscoreales: Yams and D RAFAËL GOVAERTS, PAuL WILKIN, Their Allies. (ISBN 978-1-84246-200-3, pbk.). Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 ; UU, Nb EE 3AB, United Kingdom. (Orders: www.kewbooks.com, publishing@kew.org). $50.00, 65 pp., 9" Contents: Introduction How to use the Checklist Structure Names Acceptance of taxa Geographical distribution Life-forms Abbreviations References Acknowledgments Burmanniaceae Dioscoreaceae Nartheciaceae I 111 | ically and ically significant order of tyled The most diverse à ] ic t | community levels. Other ¿1 11.1 crys 4 C | and importan member is the yam genus, Dioscor species of Dioscorea provided the chemical basis of EE pui de a oral nbsp Ally families oo the HR largely a a aah ue B ispecialised Nartheciaceae i hical distribution; indi ll synonymy, P a M source guide y g., pathologists, [eJ L d their allies; andy id j , foresters, EEN l } loeist d anti I l g ever publi she £ 41] t LA 4 geneticists, plant breeders, J. Bot. Res. Inst. Texas 2(1): 516. 2008 A NOVEL DESIGN FOR A LIGHT WEIGHT AND DURABLE FIELD PRESS Charles T. Bryson Richard Carter USDA-ARS Herbarium Southern Weed Science Laboratory Biology Department PO. Box 350 Valdosta State University Stoneville, Mississippi 38776, U.S.A. Valdosta, Georgia 31698-0015, U.S.A. charles.bryson@ars.usda.gov rcarter@valdosta.edu ABSTRACT The development of a light weight, washable, durable, and inexy field described and illustrated. A pattern an d protoco | RESUMEN Se d il ilustra un t átil i tal l campo , hecho de material ligero, resistente y barato. I r r D r o D I I d I E D E Oo As long as Botanists have collected plants, new and innovative methods have been developed to dry and preserve specimens that retain as much of the natural integrity of the living plant as possible (cf. Smith et al. 1886). Results of the progress in pressing, drying, and preserving methods are readily apparent upon viewing a series of herbarium specimens covering several generations of botanical collectors. The standard methodology employing newsprint, blotters, corrugated cardboard, and heat has been used for several de- cades to dry specimens, and details about composition, construction, and procedures for drying specimens are provided by a number of authors (e.g., Fosberg & Sachet 1965; Simpson 2006; Carter et al. 2007). Various materials and methods including metal tubes or cans (vascula), cloth sacks, Styrofoam ice chests, and paper and plastic bags have been used to transport specimens from the field to drying presses, Alternatively, heavy and Aa besote Mi presses may be taken directly to the field; however, their use is often prohibited by terrain, dis! d, and adverse weather liti Several field 1 (portfolio) designs have been used over the years, iedig that of Fosberg and Sachet (1965). Haynes 2006) described a field press made from several pieces of cardboard corrugate cinched with a press strap. In the early 1990s, Dr. Anton A. Reznicek, curator of vascular plants at the University of Michigan, constructed a field press of cardboard and duct tape and secured with a cord, which was light weight and easy to use under most field conditions. During a field trip to centra l Arkansas in 1993 with Dr. Reznicek, Dr. Paul Rothrock, and Philip Hyatt, Dr. Reznicek was observed using a cardboard and rope press. Immediately after this field trip, the senior author constructed a prototype of the press described herein out of heavy cloth and rope. Over the years this cloth press has been modified to increase the ease of use and to perfect the overall dimensions. The second author has successfully improvised a variation on Dr. Reznicek’s field press by cutting down corners of a cardboard carton and using the sides and top flaps, still attached, to hold specimens enfolded in newsprint. MATERIALS AND METHODS Instructions and materials for making the Bryson Field Press (Fig. 1) ¡A BCE out two pieces of cotton canvas or similar weight cotton cloth, one piece 1 to 1.5 m long and 0.35 m ud dd to 1.0 m long and 0.5 m wide. 2. Place é E right side down To create hem, turn edges « over 1.5 cm and fold over again, then pin hems to hold in place (Fig. 1, hem inset 4. Clip small triangle at end of each piece of cloth, then fold the miter corners with double folds and pin (Fig. , Corner insets). J, Bot, Res. Inst. Texas 2(1): 517 — 520. 2008 : » - e o A ap e P dU d 4 i L] U U 1 + i 3 Step 4c Velcro = A Fic. 1. Diagram of the Bryson Field Press. 5. Iron hem and corners prior to sewing. 6. Stitch around each hem and corner twice on the inside and twice near the outside edge and tie thread ends. Y Cross pieces with hem side up and cloth right side down forming equal size opposing flaps. 8. Pin the two pieces together. 9. Sew the two pieces tog revious hem stitches where the pieces overlap and tie thread dni One side can be left open to create à pocket for a record book, piece of sturdy corrugated card- ard, plywood, or other field supplies. 10. Pin Velero to the cloth using care to align corresponding pieces on each side. Note that Velcro strips are ached to opposite sides of the cloth on opposing flaps. 11. Stitch around the edge of each piece of Velcro twice and tie thread ends. Strips of Velcro at least 5 cm wide provide better field press closure than narrower strips. 12. Remove all pins and the field press is ready to use. DISCUSSION The field press described herein (Fig. 1) is the result of over 15 years of field testing by the authors and except for its size and more permanent materials is based on Dr. Reznicek's initial concept. Durable, light weight, and washable, it is easy to use in a variety of field situations (Fig. 2). The press can be carried under the arm or attached to a back pack, an all terrain vehicle (ATV), or a saddle. It is also convenient for press- a)? mL 4 HI AP al n fg Am IR | ye al ES sl m || . al a e field press). ing and transporting plants in a boat or canoe where a conventional press is too heavy or bulky. It is useful for pressing large numbers of specimens rapidly and for pressing plants that wilt rapidly or lose floral parts upon harvesting. The capacity of the press varies depending on the length of the press flaps and thickness of plant specimens and nan used to divide Ge SE however, we have stacked more than 100 specimens in a mee field press. The p tt 11 to “breathe” and the drying process to begin. Whereas other materials, EE as ul canvas, ies lined cloth, or heavy plastic were tried in several prototypes, tl ble, because they prevented drying and increased condensation, promoted mold and mildew growth, and caused discoloration of specimens. However, heavy weight polyester or other synthetic fabric may be substituted for cotton to make the field press suitable for field work in tropical areas or environments with constant moisture. Although extra cardboard corrugates, plywood, or felt blotters add weight and bulk, they can be used in the field press. Three heavy pieces of corrugated cardboard (one on top, one on bottom, and one between newsprint and the pressed specimens) provide the lightest weight, while allowing for adequate handling of specimens. Additional corrugates can be used to separate sp from different collection sites or to separate pre-numbered and unnumbered newsprint. We developed a system of pressing the first specimen in pre-numbered newsprint and the du- plicates in unnumbered newsprint. When these are placed into a conventional drying press from the top to bottom of each stack from the field press, the numbered newsprint is removed first from the conventional drying press. Thus, the duplicates can be easily tucked into the folded numbered sheet. Unknown to us, Dr. John Thieret and his students used a similar numbering technique (Haynes 2006). 520 À reasonably adde seamstress can cut out, ps and sew a field press in about two hours. The major expens esin t ftl the fabric and Velcro. Blemished and seconds fabric can be ai to save in ie overall cost. PORA we have noted that lighter colored fabrics absorb less heat from sunlight than darker colored fabrics. Although wider Velcro strips are substantially more expensive, Velcro strips at least 5 cm wide provide better closure than narrower strips. The estimated cost of fabric, Velcro, and thread is $20.00 to $35.00 per press, but may vary depending on the price and weight of the fabric and amount and width of Velcro. The first author experimented with addition of expandable pockets for e ee E weeny stems and fruit and straps or a for SEE the pe in the Ge od to of tl bett utility f but add cost ad bulk to ode simple design described herewith, Specimens are pressed between newsprint and stacked on top of one another with one piece of cor- rugated cardboard on the top and another piece on the bottom of the stack. The press is closed and secured by pressing the Velcro strips on the first set of flaps against those on the opposing set. Next the second set of opposing flaps is closed and secured. Two sets of opposing flaps allow for greater pressure on the stacked specimens, hold specimens in place within the newsprint, and provide pressure on plant parts that were folded over in fitting the specimen to newsprint sheets. With this field press, unlike vascula, cloth sacks, LAM ice EE eus E or plastic bags, specimens are in the exact ud in aa CR were SE lected, facili with field notes. When field work is completed, t j| removed and placed into a standard drying press. While placing the me in the deum RM they can easily be repositioned as needed, e.g. refolding leaves or other plant parts to expose flowers, fruits or other structures. Precautions should be taken to keep loaded field presses as cool as possible and out of direct sunlight. For optimal results, plants should be taken from the field press and placed in drying presses as soon as possible, e.g. at the end of each day. However, the time in the field press can be extended depending upon environmental conditions and the kinds of plants being pressed. Plants such as grasses and sedges may remain in the field press longer than plants that are subject to discoloration or contain high water content, and specimens may be kept in the field press under refrigeration overnight with no discern- able ill effects. Although not led, in unusual circumstances of exigency, we have kept specimens D in a field press for almost a week. ACKNOWLEDGMENTS We thank A.A. Reznicek (MICH) for sharing his idea of the cardboard and rope press, Nancy B. Bryson for sewing and design input for prototypes and the current version of the field press described herein, J. Paige Goodlett for demonstrating use of the press in photos, and John J. Pipoly, I1 and A.A. Reznicek for helpful suggestions. REFERENCES Carter, R., C.T. Bryson, and S. Daagvsuie. 2007. The importance of voucher specimens in documenting research in weed science with notes on their preparation, handling, and storage. Weed Technol. 21:1101-1108. FosserG, FR. and M.-H. Sacher. 1965. Manual for tropical herbaria. Regnum Veg. 39:1-132. Haynes, R.R. 2006. John W. Thieret, a student's perspective (1967-2005). Sida 22:25-31. Simpson, M.G. 2006. Plant systematics. Elsevier Academic Press, San Diego, California. SmITH, J.D., LC MARTINDALE, J.W. CHICKERING, JR, C.E. Bessey, A.W. CHAPMAN, Hl CRATTY, J.J. Davis, C.F. JOHNSON, C.E. Smit, and G. McCanrnv. 1886. Specimens and specimen making. Bot. Gaz. 11:129-134. STRUCTURE AND DIVERSITY OF A RIPARIAN FOREST AT KATETEUR NATIONAL PARK, GUYANA Carol L. Kelloff Department of Botany, MRC166 10th and Constitution Ave, NW O. Box 3/01 Smithsonian Institution, NNH Washington, DC 20013-7012, U.S.A. email: kelloffc@si.edu ABSTRACT TU ua ON T ais 1D 1 e + Jt] - Cal DA: mE [1 1 Eu E ] ] , I g I , Guyana, I $ > AXI “41 D me : : f. " [ Të ] 341 Lil + A e A 1 : letra | to riparian forests (100 — 450 m) I y E I g y x 1 Tea PE J; 1 YIT] f E. y = ] f£, F +1 Dant Diar + ] + +1 H Ts MP tree composition within the park. Approximately 133 species in 33 families comprised 1585 trees with a DBH 2 10 cm. Chamaecrista adiantifolia var. Lire dim and Eperua au ee o x een ides for 26.196 of all trees inventoried in the Of the three subfamilies of Legom inosae, Į Kaieteur. We used two 1 ha plots at each site to compare the dorem at Kaieteur with tl d forest of Kwakwani (Guyana) and Barro Colorado Island (Panama). Of these three, Kaieteur had the highest number of stems o 6%) < cm DBH = Se total week area Gen d / d Statistical analysis of the d Wallaba ¡Epia aaa two plots at Kaieteur clearly indicates a h forest. Although dominated by E. falcata und e adiantifolia va var. pteridophylla, the plots at 1 subdominant tree taxa of the Potaro Flateau. Dal D nT Key Worbs: Barro Colorado Island, Guiana Shield, Kwakwani, RESUMEN El Parque Nacional de Kaieteur localizado a lo largo del curso medio del rio Potaro en Guyana, es un mosaico de hábitats que incluye desde areniscas UE hasta ole ribereños d bin m) con Seen de arenas eege con Sc? bajos contenidos de materia orgánica. C B , se instalaron dos parcelas de 1 ha en el Wallaba ( )-bosque mixto del altiplan Potaro - alos efectos de documentar la diversidad ifica y posición del dosel arbóreo dentro del Me En ambas parcelas se encontraron un total de 1585 individuos con DAP mayor à ado | li n te 133 especies reunidas en 33 familias. Chamaecrista adiantifolia var. nn: y Ena falcata estancas: EE constituyeron el 26.1% del total de los tallos inventariados. De las tres subfamilias de Leguminosas, Caesalpinoideae fue la mejor rep tada en las puse de pond con mas dh ~ E E Lade) E 1 = peus ca el Re de uaa con los bosques de aieteur tuvo el mayor numero de i d iii con DAP menores a los 20 cm, yl d feint "-———" parcelas en | TN es 9 ree en " 11 A 1 1 ERN RR E to L + por E. falcata y C. ned is Var gt t t difi j ] I ición y est t lo] g A ' Jur Ber Ji E d eee +27 1 d. pousse f 1 er a E 1 d 1.41 L + r dominantes del altiplano de Potaro. INTRODUCTION Kaieteur National Park (Fig. 1), the only national park in Guyana, is located approximately 230 km southwest (5° 10' 42" N; 59° 29' 44" W) of Georgetown, the nation’s capital. The park is situated on an upland region of Guyana known as the Potaro Plateau of the Pakaraima Mountains. This plateau is an eastern extension of the largely Venezuelan Roraima sandstone formation that forms the flat table-top mountains known as tepuis. The Potaro River flowing over this plateau has cut a 20 km gorge in the easternmost portion of the Roraima formation. The river eventually plunges 226 meters via a single drop into the gorge. Kaieteur Falls is one of the most spectacular cataracts in the world and is the main geological feature of the national J. Bot. Res. Inst. Texas 2(1): 521 — 545. 2008 [| £ al Dos + ID LJ Me nd ET 522 J Texas 2( park. The park was established in 1929 and encompassed 44 km" to protect the area around the falls, the Potaro gorge, and the surrounding Potaro Plateau. Today the park has been expanded to 627 km! to protect a greater representation of the vegetation around the falls and a larger expanse of the savannas and forests of the plateau. In 1998, the park was incorporated into the National Protected Areas System (NPAS) that is currently being developed as part of Guyana’s ecotourism business. With this NPAS Project, Guyana urgently needs information on the biodiversity of the area before any conservation or management policy can be implemented. Botanical exploration of the mid- to lower portions of the drainage has been limited to the area around the falls and along the Potaro gorge; such exploration has resulted in a checklist of the vascular plants of the area (Kelloff & Funk 1998), but there have been virtually no ecological studies in an attempt to quantify species A In general, tropical forest ] ized as having a high diversity of trees, with some inven- tories recording over 300 species with a diameter at breast height (DBH) > 10 cm per hectare (Gentry 1988; Hubbell & Foster 1986; Valencia et al. 1994). Several hypotheses offer explanations regarding the mainte- nance of high diversity. Huston's (1979, SE ee E SUOMI SUEESSIS that UNE in populations is reached by low rat (Hustor 1979). If the population has a high dumber or high frequency of a Doing species will disappear and be replaced by the faster-growing pioneer species over time thus reducing diversity. In order to overcome this species must have some minimum growth rate to recover from population reductions. In other words, an increase in growth rates during periods of high frequencies of population reduction can actually increase diversity (Huston 1979). Another hypothesis is an ecological response to environmental conditions such as low and high light conditions. By changing the availability of resources, disturbances such as canopy gaps may influence succession. Those species efficient in recruitment quickly move in, and the site becomes 1 with pi (Bazzaz & Picket 1979). The ability to survive under low light conditions and the ap) to achieve high growth under high light conditions is the most significant differ- ence between species diversity (Bazzaz & Picket 1979; Hubbell 1995). It's the survival of the fittest that can utilize the disturbance, increasing the diversity of species. In Guyana, Dicymbe Spruce ex Benth., which is locally known as clump wallaba, produces coppices (suckers) on the healthy parent tree. When Gs parent trees die, these young trees can take advantage of the mature root system of the parent, as well as the light gap, and thus can out-compete other species in the area (Henkel 2003; ter Steege et al. 1993). Not all tropical forests show a high alpha tree diversity, however. Productivity and disturbance vary in the tropics (Richards 1952; Connell & Lowman 1989), and this is evident in the mixed forests of central Guyana where there is a tendency for some species to dominate the vegetation (Davis & Richards 1934; Fanshawe 1952; Johnston & Gillman 1995; ter Steege 1993). Severa lexamp les of this type of forest in Guyana are the wallaba (Eperua Aubl.), mora (Mora Benth.), or clump wallaba (Dicymbe) forests (Davis & Richards 1934; Fanshawe 1954; Henkel 2003; ter Steege 2000b). These forests have a few species that represent a high percentage of the stems and/or basal area. The tropical forest of Guyana contains a wide array of forest types, e.g., mangrove, moist and dry ever- green, montane, seasonal, swamp, and Greenheart forests. In swamp and mangrove forests, edaphic factors such as flooding and soil clearly explain these forest types (Fanshawe 1952). However, in the mixed forests of Guyana, the relationship between soil, hydrology, and forest types is not clear (ter Steege et al. 1993). Generally, classification of forest types has largely been based on climate, soil, and physiognomy (Ducke & Black 1954; Prance 1987), which reference species compositions or have generalized tropical floras at the genus or family level (Forero & Gentry 1988; Maguire 1970; ter Steege et al. 20002). Recent studies used plots to examine patterns of plant diversity, e.g., how species richness (Gentry 1988) or habitat diversity contributes to species diversity (ter Steege 1993; Sabatier et al. 1997; Tuomisto & Ruokolainen 1997). Plot studies produce data at the species level and can provide long-term information on growth, mortality, re- generation, and dynamics of forest trees. Sampling riparian trees of the plateau using a grid system of collecting allows us to (1) describe the Kelloff, Kaieteur National Park, Guyana 523 de J || all Pas rat H Fm L H t A Ene Len il [| LL - L 1 Fic. 1. Kaieteur Falls, 11 524 | t tanical | Texas 2(1) relative abundance and distribution of the trees, (2) provide a framework by which the forest structure of Kaieteur can be compared to other known study sites, (3) help fill in the missing tree taxa in the checklist, and (4) provide a standard by which forest structure and composition can be measured. This paper presents the results of the two, one-hectare (ha) inventories carried out on the Potaro Plateau within Kaieteur National Park. Tree data from two other plot studies were used as a comparison of the tree diversity at Kaieteur. Data from two one ha plots were borrowed from a lowland forest in Guyana near the town of Kwakwani (a Guiana Shield community) and two one ha portions of a 50 ha plot at Barro Colorado Island, Panama (non-Guiana Shield community. In general, plot data from the Guiana Shield and particularly the Potaro Plateau are scarce (ter Steege et al. 20002). One ha plots allow for us to quantify the relationship between tree species as well as among tree species and add to the knowledge of the tree diversity of Potaro Plateau in Guyana. The ultimate goal of this project was to document the species diversity and tree composition of the Wallaba (Eperua)-mixed forests of the Potaro Plateau and to compare the tree species diversity of Kaieteur National Park to those of other plots. METHODS Tt 42. | m TZ. Ls ANITA TDA.1,: . LL P E PP scrub, E riparian forests. The average air temperature for the "— is 26. 5°C (USDA 1974). The average rainfall for the northeast face of the ai escarpment is approximately 3750 mm per year (Fanshawe 1953). Two 1 ha plots blished in the Wallaba (Eperua)-mixed forest of the Potaro Plateau. The study was conducted between 1995 and 1998 with sites selected based on the surrounding vegetation. In both 1 ha plots, some level of previous human disturbance was found (e.g. several large trees were removed for the building of the guesthouse). The first plot (Fig. 2a) is located at the headwaters of the Korume Creek (5º10'42"N, 59°29'44"W). The lower portion of the first plot is seasonally inundated and includes a portion of the creek in which grew a few patches of Thurnia sphaerocephala (Rudge) Hook.f. Along the western side of the plot, the land gradually rises up tl ine towards Muri-Muri savanna. The second plot (Fig. 2b) was established in the upland forest of the plateau between the Korume Creek and Potaro River gorge (5*11'16'N, 59º28'52"W). The terrain is rocky with a slight downward slope towards the northeastern part of the trail. There is no evidence of seasonal flooding as in the first plot. The soils from these two sites were not analyzed, but it was generally observed that they consisted of a course white sand of quartzite with very little peat. These shallow white sand areas of the Pakaraima Plateau (Fanshawe 1954) are associated with the peneplain of the interior of Guyana. The plots in the lowland area of Guyana used for comparison are located near the town of Kwakwani at the Aroaima (bauxite) mining site on the Berbice River, 238 km up the canje! ave from New Amsterdam (5°30'N, 58°W). Data for these plots were kindly provided by James Comiskey (Smi ian Institution Man in the BioSphere Program). The average rainfall recorded for this area is 2400 mm per year. The land is low lying with swampy areas near the river. The sites for the plots were selected within the area designated as "south mine," a proposed area for future bauxite mining. In this area, the shallow layer of topsoil (averaging 1.5 meters in depth) covers ca. nine meters of bauxite that overlays 60 meters of volcanic rock. Both plots were irregular in shape, and the vegetation is classified as non-flooded tropical mixed forest (Comiskey et al. 1994). It was noted that the soil was composed of a mixture of sand and clay with high porosity and high drainage. The soils were not sampled (Comiskey et al. 1993). The second set of plots were part of a 50 ha study site at Barro Colorado Island (BCD located near the Panama Canal at the Smithsonian Tropical Research Institution’s station (9°9'17"N, 79°50'53"W). This for- est is classified as a rich, old growth tropical forest dominated by Bombacaceae and Leguminosae (Condit et al. 1996). The terrain on the island's summit is relatively level with a variance of 28 meters in elevation. The average rainfall for this area is about 2500 mm per year (Condit et al. 1996) with a mean annual tem- perature of 27°C. The soil at BCI is a clay-rich tropical soil with an organic matter content (Yavitt 2000). Data from two of the 50 1 ha plots (designated as Hec21 & Hec37) were selected and kindly provided by Richard Condit for this study. Kelloff, Kaieteur National Park, Guyana 525 99227 W Menzies = Landing O e Së eege JEE th EN o P dd Ge A | E Ea E B so A oo DM. A i pp i | E = d d N | Mile 1 au 42 a 0 500 — 41000 meters ~ A ON Lë H A Ai x" I! m. | pl | A E al ri AN Fal fw a E a? | a Lëba t i Fic. 2 E E FI 526 Established on the central plateau of BCI, the 50 ha permanent plot has a 70 year history of the flora and fauna and over 1500 scientific papers written on various topics (Hubbell & Foster 1992). For more information on Barro Colorado Island, selected readings are: Hubbell and Foster 1992 and Hubbell et al. 995. +1 ah d'H £-11 € «Lond TL thad adanted f, woody vegetation Ol 1 Dallmeier O (1992). The terrain at both sites was uneven, and ti de t t for sl (Durr et al. 1988). The two 1 ha plots at Kaieteur were subdivided into 25 M 20 x 20m quadrats, end all stems with a minimum DBH of 10 cm were measured. Spatial location of the trees within each quadrat was recorded for mapping purposes. Field vouchers were obtained, where possible, for each species encountered. Information was recorded to aid in identifications of the sterile vouchers, e.g. features of the wood, bark, and sap (Polak 1992; Rosayro 1953). Romeo William, tree spotter and guide, assisted in the identifications of trees in the field. Local or vernacular names tore trees E did 1223; di Kee to imi ae DOE ey similar trees. Most vouchers Trees were identified to the species level when pila but were otherwise assigned to morphospecies. Taxonomic nomenclature aaa tHe” SUCCI of Se Plants of the Guianas: Guyana, Surinam, French Guiana” (Boggan et al. 1997). Voucl deposited in the Guyana National Herbarium, University of Guyana (BRG) and at the U.S. National Herbarium, Washington, DC (US). Density, frequency, and dominance were calculated to describe how individual species are spatially distributed within the plot and how these distributions contribute to the community (Barbour et al. 1987). Stem basal area (a) and density of each tree were calculated using a = 0.7854 x (DBH) 2, where a = basal area of each tree in m? (Anderson & Ingram 1989). This measurement is useful in estimating canopy cover of trees because it assumes to take trunk diameter at breast height (DBH) into account. We also calcu- lated species basal area (BA), defined as the sum of all stem basal areas (a) of a given species. Density is defined as the number of trees per hectare, and relative density is the percentage of one species in relation to the total plant diversity. Relative frequency is expressed as a percentage of the number of quadrats that contain a given species. Dominance is an important ecological t. A species that | the most cover or basal area to the community is considered the dominant species. Dominance is calculated as the percent of basal area oh a given species to ota! vn area Mimi et al. E Finally, the relative tion that ity is called its importance value (Balslev et al. 1987). The importance value iudex (IVI) can be calculated by the addition of the relative density, frequency, and dominance for each species and for all trees at each site. High IVI values indicate that a species has a high basal area, frequency, and density at a site or when one or two of these parameters are much higher than other species (Felfili & da Silva 1993). The sum of relative parameters places species in a hierarchical order in the community and is useful for comparisons between different sites (Curtis & McIntosh 1951; Felfili & da Silva 1993). The IVI values can range from 0 to 300, with 300 representing a pure stand of one species (Balsiev et al. 1987). Essas aes PEA x32] ] 1 s ws z + 1. =o 1: +1 of describing best straciuire (Ďutilieul SR eg et i 1998), and raw pee ni ion pios can be too large and | to yield any m u Or data into a space with fewer Ge Es can reflect sample configurations in e space (Gauch & Whittaker 1981). In community ecology, such ordinations can describe some of the strongest patterns in species composition (McCune & Mefford 1999; McCune & Grace 2002). Ecologists have applied metric ordination, which includes principal components analysis (PCA), to analyze vegetation data (Proctor 1967; Debinski & Brussard 1994; Tuomisto et al. 2003). This type of ordination assumes a linear relationship between the variables and is rarely amenable to ecological community data; it can also lead to consider- able obscurity in data interpretations (Goodall 1953). Non-metric ordination methods are considered more powerful for analvzing vegetation data because this method assumes a non-linear relationship between the Kelloff, Kaieteur National Park, Guyana 527 variables and the use of ranked dist tends to li ize t] lationship between the distances measured in environmental space and species space (McCune & Mefford 1999). Species data per plot were ized into data matrices (taxon vs. quadrat) and imported into PC-Ord (McCune & Grace 2002), a multivariant statistical program. Because the data represent quantitative scores per taxon, the original values were log transformed using b; = log(x; + 1), where the value of one is added to each species score to define zero values in the data. Log transformation reduces any dominant effects and normalizes the relative importance of common or rare species (Digby & Kempton 1987; McCune & Grace 2002). To visualize multivariate patterns among the quadrats, non-metric multidimensional scaling (NMS) was conducted using Sgrensen (Bray-Curtis) distances for each plot and on each plot pair. NMS in PC-Ord is based on Mather's (1976) program and the algorithm of Kruskal (1964). For each dataset in this study, the "autopilot" mode in PC-Ord was used. This mode provided a random starting configuration for each run. The program calculates each solution by running 40 runs with the real data and 50 runs with the randomized aa, wa a maximum number of 400 iterations for six axes. The program selected the best solution at each where p < 0.05 for the Monte Carlo test and where that value of the “stress” parameter was t I anes Clarke’s (1993) “rules of thumb” were used to interpret the final stress. Clarke found that most ecological community data will have values with a final stress between 10 and 20. MS provides an optimal ordination technique to analyze patterns in tree species composition for each of the studies. The Anal: result in each analysis was an ordination in two-dimensional space. The two axes show the relationship between species abundance and the axis score. RESULIS Kaieteur National Park, Guyana.—There were 1724 total number of stems 210 cm DBH at Kaieteur in the two study plots. These stems covered a total basal area of 66.34 m?/ha. Plot KF1 at Kaieteur contained 757 stems per hectare, with most taxa b dea Dy a Ld stem. Dicymbe pharangophila R.S. Cowan (swamp wallaba, Leguminosae—Caesal.) produced coppice the base of the trunk that resulted in 65 stems being measured and tagged froin a total of 28 trees. This habit of producing multiple stems from the base was also noted by Whitton (1962) and Henkel (2003) of Dicymbe corymbosa Spruce ex Bentham, another “swamp wallaba” found on the Potaro Plateau. The largest diameter tree recorded for the site was Ocotea sp. (Lauraceae), with a DBH of BN cm. cn EE with a DBH > 60 cm were Pouteria cf. cuspidata (A. DC.) Baehni (Sapot ) and S ; ii Bentham (Leguminosae—Fab.). Sixty-four percent of the stems were ani 20 cm in ees and abner one percent of the trees reached over 60 cm DBH. The two species that had the highest Important Value Index (IVI) were Chamaecrista adiantifolia (Spruce ex Bentham) H.S. Irwin & Barneby var. pteridophylla (Sandwith) H.S. Irwin & Barneby (Caesal.; IVI-20.052) and Eperua falcata Aublet, (Caesal.; IVI-15.427). These two species were found throughout the plot and accounted for 13.4% of the stems. Dicymbe pharangophila (IV1214.679), Ormosia coutinnoi Ducke (Fab.; IVI=10.109), Dicymbe sp. (1V1=10.027) and Pouteria cuspidata (IV1=10.442) ranked next in importance. These species accounted for 32% a ine stems recorded for KF1. | species were not recorded, but it was noted that t sp. and Meno ama spicatum (Aublet) J.F. Macbride) dominated the loners wet areas of the plot. RE Species of Bromeliaceae (Ananas parguazensis L.A. Canargo & L.B. Smith) dominated the dryer soils of the plot. Thurnia spherocephala (Ridge) Hooker f. (Thurniaceae) was quite abundant and grew in the flowing waters of the creek. The second plot (KF2) at Kaieteur had more stems (967) than the first plot. Dicymbe pharangophila was neatly absent from this plot. Chamaecrista adiantifolia var. pteridophylla (95 cm DBH) and Swartzia schom- burgkii Bentham (Fab.; 92 cm DBH) were the largest trees recorded for the site, followed by Elizabetha sp. (Caesal.; 66 cm DBH) and Pouteria cuspitata (64 cm DBH). Stems 20 cm DBH or less accounted for 69.596, whereas only 1.096 of the stems exceeded 60 cm in diameter. Several species ranked high in IVI: Eperua ERA " PA DEE 528 Jou tl tanical Insti Texas 2(1) falcata (1VI=20.048), Chamaecrista adiantifolia var. pteridophylla (19.486), Ocotea spp. (IVI-18.255), and Swartzia schomburgkii (IVI=17.404). When Catostemma commune Sandw. (Bombaceae; IVI 215.082), Ormosia coutinnoi (IVI212.328), and Eperua spp. (IVI215.702) are added, these species account for 40.396 of the total stems EH for "ped KF2. The understory species were not surveyed, but it was noted that Ishnosiphon sp. and M somewhat evenly distributed in the plot. Vriesea splendens (Brongn.) Lem. E DEER was seen along with the spiny bromeliad (Ananas parguazensis) from the first plot but was not as abundant. A broad-leaf aroid grew on tree trunks, and a Rapateaceae (Stegolepis ferruginea Baker f.) dominated the understory at quadrat 14. In both plots at Kaieteur, the Leguminosae had the highest number of individuals per site (815 stems or 47.396 of the plots) most of which were from the subfamily Caesalpiniodeae (607). In plot 1 Bombacaceae ranked as the second most abundant family across all quadrats with 64 stems. Bombaceae, in plot 2, had 77 stems but ranked third after Lauraceae (136 stems). Kwakwani, Guyana.—There were a total of 995 stems > 10 cm DBH at Kwakwani in the two plots, these with a total basal area of 52.13 m?/ha. Five hundred and seven stems were recorded from first plot (ART) at Kwakwani. Only four trees had a second stem measuring over 10 cm DBH. The largest tree on the site was Trymatococcus amazonicus Poepp. & Endl. (Moraceae) at 91 cm DBH. Eschweilera pedicellata (Rich.) S.A. Mori (Lechythidaceae) represented 23% of the total stems (115 stems) in this plot and had an IVI of 33.425. The taxon with the next highest IVI (6.751) was Aspidosperma excelsum Benth. (Apocynaceae) followed by Bocageopsis WER Dee ) R.E. Fr. (Annonaceae; 6.733), Unonopsis rufescens (Baill.) R.E. Fr. (Annonaceae; kii (Fab.; 5.116), Prunus sp. (Rosaceae; 3.857), and finally Cordia sagotti I.M. Johnst. (Garip mace 3.795). These taxa reseed 163 stems or 21% of the stems counted in the site. Sixty three percent of the stems were under 20 cm di a only 2% were over i cm DBH: Me pedicellata (I thid - 88.2 cm), Aspid (Sp ex Benth.) plam NE mass 83.5 cm), Inga pna Benth. MAE 76 Em And Bocageopsis multiflora (Mart.) R.E. Fr. (Annonaceae; 72.8 cm The second plot (AR2) at Kwakwani had only 491 trees of single recordable stems. Aspidosperma excel- sum was the largest tree (91 cm) and Eschweilera pedicellata had the highest IVI value of 63.671 (138 stems, 2896 of the total stems). Two other taxa significant were Mora gonggrijpii (Kleinhoonte) Sandw. (Caesal.; 1V1=36.486) and Aspidosperma excelsum (IVI-15.600). Inga acrocephala Steud. (Mimos.), Mora gonggrijpii, Swartzia polyphylla DC. (Fab.), Eschweilera pedicellata, Chrysophyllum sparciflorum Klotzsch ex Miq. (Sapota- ceae), and Aspidosperma excelsum represented the 396 of the total stems over 60 cm DBH. Fifty-six percent of the total stems were under 20 cm. Lecythidaceae was the most abundant family at Kwakwani, with 254 stems documented for the two plots. Leguminosae was tl ond most abundant with 184 stems (107 of these were Caesalpinioids in site 2). Next in rank were the Annonaceae with 55 stems over all quadrats in ARI and Boraginaceae (34 stems). In site 2 these families represented only 19 and 2 stems. Apocynaceae had more stems (26) in AR2 and ranked third in abundance across the plot. All other families had less stems. Barro Colorado Island, Panama.—1n the two 1 ha plots selected from the Barro Colorado Island (BCT) 50 ha plot, the total basal area of trees > 10 cm was 57.153 m?/ha. Information provided for this study includes only data for the main stem on each plot. No data were available for multiple stems and thus these were excluded from this study. The first plot (designated as Hec21 at BCI) had 418 trees. The tree that had the largest diameter on the site was Hura crepitans L. (Euphorbiaceae; 102.1 cm DBH). Seventy-five percent of the trees had « 20 cm DBH. Two percent of trees had a DBH » 60 cm: Brosimum alicastrum Sw. (Moraceae; 88.5 cm); Chrysophyllum cainito L. (Sapotaceae; 88.5), Jacaranda copaia (Aubl.) D. Don (82 cm) and Tabebuia guayacan (Seem.) Hemsl. (Bignoniaceae; 82 cm); Prioria copaifera, (Caesal.; 82 cm); and Luehea seemannii Triana & Planch. (Tiliaceae; 76.8 cm). The dominant species for Hec21 were Faramea occidentalis (L) A. Rich. (Rubiaceae; IVI = 18.644), Trichilia tuberculata (Triana & Planch.) C.DC. (Meliaceae; IVI = 23.870), Kelloff, Kaieteur National Park, Guyana 529 Prioria copaifera, (Caesal.; IVI = 17.581), and Alseis blackiana Hemsl. (Rubiaceae; IVI = 10.344). These taxa accounted for 139 trees or one-third of the entire tree census. The second plot (designated as Hec37 at BCI) contained 447 recordable trees, 62.6% of which were < 20 cm DBH. Ceiba pentandra (L.) Gaertn. (Bombacaceae) was the largest tree in the plot with a diameter of 209.5 cm DBH. Other species over 60 cm DBH were Ficus costaricana eoe o (Moraceae, 138 cm), Tachigali versicolor Standl. & L.O. Williams (Caesal.; 109.1 cm), Hura ; 99.4 cm), Aspidosperma cruenta Woodson (Apocynaceae; 89.0 cm), Dendropanax arbores (L) Been & Planch. (Araliaceae; 83.0 cm), Brosimum alicastrum Sw. (Moraceae; 75.3 cm), and Cecropia insignis Liebm. (Cecropiaceae; 66.0 cm); Guapira standleyana Woodson (Nyctaginaceae; 75.3 cm); SE EE Pittier (Bombacaceae; 69.8 cm); and Guatteria dumetorum R.E. Fr. (Annonaceae; 63.5 cm). The t species on the site were Faramea occidentalis (IVI = 23.340) and Trichilia tuberculata (IVI=27.239); together, these two accounted for almost 31% of the total countable stems. Other species with high IVI values were two Bombacaceae: Quararibea asterolepis (IVI=14.721) and Ceiba pentandra (IVI-12.455). From the two plots examined at Barro Colorado Island, Rubiaceae ranked the highest for individual stems (217). Meliaceae, with 122 trees, ranked second in the "tree per family" category. The Leguminosae contained only 67 stems, 47 of which were Caesalpinioids. A comparative summary of the most abundant tree species from the six study sites can be found in Table 1. Cintictical analwerc Non of the three study sites. In each plot there were several mm that id a strong relationship with the ordination scores. These species were displayed on the ordination as a joint plot. The radiating lines of the joint plot reflect the direction and strength of that species that is influencing the separation of the quadrats in the plot. Plot KF1 at Kaieteur had 103 species in the analyses. The final stress was 15.68692 for the 3-D solution with 99 iterations. The Monte Carlo test for this solution was p < 0.0196. The first two axes accounted for 56.8% of the variance; the first three accounted for 74.1%. Three patterns are seen in the grouping of quadrats based on species assemblages (Fig. 3A). Dicymbe spp., Chamaecrista adiantifolia var. pteridophylla, Pachira flaviflora (Pulle) Fern. Alnso. (Bombacaceae) form one group, and Chamaecrista apoucouita (Aubl.) H.S. Irwin & Barneby (Caesal.) and Pterocarpus rohrii Vahl (Fab.) form a second group; these two groups have a stronger influence in some of the quadrats whereas a third group composed of Hevea guianensis Aubl. (Euphorbiaceae), Inga gracilifolia Ducke (Mimos.) and species in Meliaceae have a stronger influence on others. Kaieteur plot KF2 had the fewest species (71) in the dataset. The best result for the dataset is the 3-D ordination from NMS with a stress value of 13.30984 with 162 iterations (Monte Carlo p < 0.0196). The variance for the first two axes accounted for 66.496 of the variance; if the third axis is included, 82.596 of the variance is accounted for. Quadrats in the EE are an into speci lages influenced by two groups: the Licania alba (Bernoulli) Cuatrec. (Chry pp., Clathrotropis macrocarpa Ducke (Fab.), Eschweilara spp., Ocotea spp., and Macrolobium iaa Spruce ex Benth. (Caesal.) group; and the Henrettea ramiflora (Sw.) DC. (Melastomataceae), Sextonia rubra (Mez) van der Werff (Lauraceae), 3, Je? ` 1 1: INTRAC) | A +l Tatanta At )/ Was used t each a Inga sp.3, and Catostemma Sandw. (Bombacaceae) in the other (Fig. 3B). Combining the data from the two plots at Kaieteur, the best result from multiple NMS runs was a 3-D solution (stress = 18.09208, p < 0.0195) with a variance for the first two axes ting for 54.696 or 75.096 including the third. The first two axes were chosen for the ordination to best represent the data (Fig. 3C). Axis 1 represents the area of collection and EES in a between the locations of the plots. Axis 2 represents a clear separation between tl an ges at plots KF1 and KF2. Species that contributed to the discrimination in dein were DIOE i a a Eperua rubiginosa Miq. (Caesal), and Catostemma fragrans Benth. (Boml g to discrimination in KF2 were Fperua spp., Sextonia rubra, Henriettea UM and Gesten spp. There was no overlap in species composition between quadrats of these two plots. 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H | ye CRIBACY (E D | CFA’ a £r. | j" J £ a. L A all d Mt J J VJ d 2 i - T E J Li J DH bno l n: | ID«n] et m p TO E ctas add Drs PB avid rog : vw d d af ke a= LI d * =e A e? - LI FL a LM ital h | | Fia E . b » AL I idi Li H Tel k | B PF H LU ët ri 8 al LL. $ | fro al f Le re F * bk e iu Jo | Lih" 4 FAT J A Ll KE? A H ra SA . LI fur L IL Ti I1 n L [Ren] ei gk A a D F d LI = D LI po] e eE re - qn.” D D [Cmal. Fschweilara ssp. [Esp]. Ocotea sop. [Osp] iM lobi lens [Msuli iH H fora IHral, Sextonia rubra [Stu] L Je He Leeds Fr" Ad wl L d ho | f L dr L Je Inna «n 2 (ECH Ka nt ET tu Al ai 4 L: J H lata E Lan La TIPA O VES] nh Va! M PA CO A f L af k d kl | uor E f i E] A I ` Ll NUT al J mm Ai Ez D & V. I DH AR1 "al Eran dh at en all D Lu rl L| Fue f F e | J 1 m J H au. Ai H e £ PUR, aal Sneed? PROS Eë aal Tamil 4 [7.9]: Miranin = Lu M kl T d F E af ki bd Li F bd d Ki Er hvnnlemca IMhvl D : Lët TD J'a al A I Frol H] Pole EP 1 £L ` a «al La /F s al Id oa ZB L eat a F L 4 Li F Lp. T PUE i Kwakwani AR? Fe ch ij lirell iM jini si ifi tl t ial | tt H j ] tl Iculati Tha x A i wë - së FF d wë F L J - A . al AL S fa | J al H A H i TAAL " f LZ TI 3 Pruna en [Pen] Pal I. Jr I: re " L] F - eN we Li wf =F SP D Kai ES aa Se a Evie rag 1 Jn P r " fX aL La H Jag L DI Inte ADT O F, || SI ah r4 V P f T L T rr 5096 of the quadrats between the study sites. 534 | £ ul naa H In L Pa att iT fay 17 RO] 24 drat Bros, wie Cat 1.07 Ars e sth e o % D a $ e Au . P Di 10 Axis 1 Bo) YT desch aiser Bra Curia 1 q m è e ga $ LS e + ^ sex er SDK ° “Acr B | Ahi, a a Abl S a E E $ Gst $ LP . > 6 z+ . e è LÀ ds ae es a: 15 Ads 1 Bel od +. 47 raclrat en. D rt H Q5 4 e E MX 1.5 T y T T T—— F 15 dk 05 15 Ads 1 F A Al but la: [| Ir fFRIAACA I: AP [| E AY DO ful Ia In! E mas HFC?1 la ys gi Las al IG. 1 Li d SS D + Li U L H | D L aus LI. Pads e e | I an rj or TA 1: n Af = | T f T euge wé r? F rj A Sat) 1 Tre a “Tr 1 JA TA Ti al J IF SA Aalt, PPA sl aL al J [Ppal Lp mra P L JF k f Liye“ ir F F d F L 4 F fr) al IA £n J J ures A B4 al Sal LI [| H A IL A a bush Fada de d'W A Më 1 Li T a F T B 3 ne | Ler rir . l E) ` a Feed A sf a. TA cul TT LI L: TALI] te el | vma AA A o PE L 3 I f 3 L 4 f LU dy L d , ef p: A [| Y e D Fe al! al AL Su fe ye a? E al MA L| H AL, L Jag E n real | | p | | les L d P L 4 F a [Hec21 3 E | »Lh d EH J* 4 H HI SA E tgs L £L LPS i Lj LI D Kelloff, Kaieteur National Park, Guyana 535 Ordination was used to reduce the datasets at the Kwakwani study site. Site ARI had 84 species with a final analysis in a 3-D solution of 15.71292 (stress) over 82 iterations (p < 0.0196). The overall patterns between the quadrats indicate some differences in species composition between the plots. The quadrats separated into several groups (Fig. 3D). These Ge were E So ee ie (Palb) R R.E. Fr. (Annonaceae), Cordia sagotii I.M. Johnst. (B h.) Triana (Melastomataceae), Bocageopsis multiflora (Mart.) R. E. Fr. EE and Tuus sp.2 acia diaceae). The first two axes accounted for 53.0% of the variance. Including the third, 72.9%of the variance was accoun ted for LI. DN T5211 y Ind a A A At site AR2, there were 58 species in the matrix were abun- dant throughout the plot. Since a d Wee contributed 1 more than the other a NMS failed to find a useful ordination. Esch llata an i were deleted for remaining calculations. The result was a 4-D solution (stress = 13.80266; p < 0. 0392) for this plot (Fig. 3E). The first two axes ac- counted for 18.7%, and the third added only 34.6% of the variance. These blages were defined by the following species in the plot: Miconia hypoleuca (Benth.) Triana (Melastomataceae), Inga pezizifera Benth. (Mimos.), Prunus sp. al Chl DIDI rodiei (R.H. Schomb.) Rohwer, H.G. Richt. & van der Werff (Lauraceae), Ocotea spp , and À bilis L.f (Euphorbiaceae). Combining the latasets for both plots at Kwakwani resulted in 93 gesat in the analysis. NMS ordination yielded a 3-D solution with a stress of 19.64717 (p < 0.0196) over 326 iterations. The first two axes EE for TR 196 of the variance (65.796 with all three). Fifty percent of the | both plots had species that were similar. Swartzia schomberghii, Prunus sp., Mora gonggrijpii, in BEE Aubl. (Chrysobalanaceae), and d e excelsum were the species that define the g the BCI plots. Eschweilera 1 in both plots (Fig. 3F). At the Barro Colorado Island (BCI) Hec21 plot, a total of 92 species was observed. The final stress for the 3-D solution was 15.82730 (p < 0.0196) in 88 iterations. The first two axes accounted for 49.2% of the variance (73.896 with all three). Figure 4A shows the result of the non-metric scaling. The angle and length of the radiating lines in the joint plot indicate the direction and strength that each species has with the ordination score. Radiating vectors indicated that Trichilia tuberculata, Allophylus psilospermus Radlk. (Sapindaceae), Protium panamense (Rose) I.M. Johnst. (Burseraceae), Faramea occidentalis, Jacaranda copaia, Tetragastris panamensis Kuntze (Burseraceae), and Oenocarpus mapora Karst. (Arecaceae) had the strongest relationship with the ordination scores. The BCI plot Hec37 had 82 species; the ordination had a final stress of 17.47696 over 131 iterations (p < 0.0392). The first two axes accounted for 30.5% of the variance and the first three, 61.1% of variance. The species that contributed to discrimination in this plot were Trichilia tuberculata, Virola sebifera Aubl. (Myristicaceae), Cecropia insignis, Aspidosperma cruenta, Alseis blackiana, Brosimum alicastrum, and Guapira standleyana (Fig. 4B). A total of 114 species in the two BCI plots was observed. The resulting ordination of the combined plots produced a 3-dimensional solution with a stress of 21.77097 (p « 0.0392) in 400 iterations. The vari- ance explained by the first two axes was 33.696 and 58.196 including the third. The quadrats of the two plots overlapped significantly, indicating a similarity of species composition within each plot (Fig. 4C). The species with the strongest relationship in this ordination were Trichilia tuberculata, Adelia triloba Hemsl. (Euphorbiaceae), Oenocarpus mapora and Jacaranda copaia. DISCUSSION Establishing two permanent biodiversity monitoring plots at Kaieteur National Park has provided Guy- ana with the scientific data required for the prioritization of conservation initiatives. This study provides baseline data on tropical woody plant communities to aid in measuring ecological change over time and to help distinguish between natural and human impact. Measuring relative density, relative frequency, rela- tive dominance, and importance index values of individual species provides information on how species 536 | tanical h Institute of Texas 2( are DM Man the forest and how they contribute to the community in which they live. Using a lology f. ing all trees > 10 cm DBH ensures comparability with other study sites and provides a framework ior studying the dynamics of tropical forests. This survey at Kaieteur resulted in 133 taxa representing 1586 trees (1,725 stems = 10 cm in DBH) almost twice the number of trees from the Kwakwani, Guyana or Barro Colorado Island, Panama. The density of trees at Kaieteur were similar to a caatinga forest on white sand at Pico da Neblina in Brazil where 1569 trees were measured in two one- hectare plots (Boubli 2002). The number of stems recorded in each plot at Kaieteur bordered on the upper range of stems per hectare (965 stems har!) found on similar plot studies. This high number of stems were found by Davis and Richards (1933, 1934) in five plots at Moraballi Creek, Guyana (460 to 919 trees per 122 x 122 m plot) where as those of similar studies in the Neotropics for trees 2 10 cm DBH have not been. Sabatier and Prévost (1990), Poncy et al. (1998), and Bordenave et al. (1998) found densities of 473 to 570 stems per hectare at Les Nouragues, French Guiana, and up to 882 stems per hectare in one particular transect (Sabatier & Prévost 1990). Johnston and Gillman (1995) recorded 357 to 742 trees per hectare at the four one-hectare study plots at Kurupukari, Guyana. At the El Caura Forest Reserve, South Venezuela, mean densities ranged from 563-573 trees ha! (Castellanos 1998). In Manaus, Brazil mean densities were 550 stems ha! (Ferreira & Rankin-de-Mérona 1998) Davis and Richards (1934) noted that the forest at Moraballi Creek dominated by Eperua falcata had an extraordinary large number of trees ha! and that Leguminosae was the most abundant family (Whitton 1962). This was also noted by Boubli (2002) at Neblina where Eperua leucantha Benth. accounted for a large percentage of the stems. Guyana’s rain forests can be dominated by one to several species, these often in the same family (Davis & Richards 1933; Fanshawe 1952; Whitton 1962; Henkel 2003). Kwakwani's plots were dominated by Lecythidaceae (30.9%) and Kaieteur's (47%) were dominated by Leguminosae. This is not unique to Guyana as a Lecythidaceae or Leguminosae dominance can be found in many other tropical forests, e.g., Richards 1952; Whitton 1962; ter MAN 1993; Nascimento & Proctor 1994; Henkel 2003. Stem diameters were summarized into two categories (10 cm increments and percentage of stems per class size) in order to facilitate comparisons between the study sites (Table 2). The distribution of tree per class size in the six plots shows a characteristic inverse J-shape (Fig. 5) typical of forests that have been relatively undisturbed in the recent past ER E Mori 1989), with over 5696 of the trees equal to or less than 20 cm DBH. Overall, the stem class size tions from 20.1 cm to 60 cm DBH at Kaieteur were comparatively similar to Kwakwani and BCI in this paper. Total p tage of stems below the 40.1 cm DBH class in each of the three study sites ranged from a low of 86.7% at BCI to 95.9% at Kaieteur plot KF2. Boubli (2002) commented that 46% of trees at Neblina were small in girth while Whitton (1962) commented that in the Wallaba forests of Guyana, very few trees exceed 70 cm in diameter. At Kaieteur only ca. 2% reached the larger diameters with only three species, these all Caesalpinioid legumes, reaching over 30 cm DBH: Eperua falcata, Chamaecrista adiantifolia var. pteridophylla and Swartzia schomburgkii. The total calculated basal area/ha from the three sites in this study ranged from 52 to 66 m? / ha. Kwakwani and BCI were similar to the 53 m*/ha found by Whitton (1962) and Mori and Boom (1987) at Saúl, French Guiana, but higher than those found in the forest plots (27-34 m?/ha) of Central Amazonia (Rankin-de-Merona et al. 1992). Kaieteur’s total basal area (66 m*/ha) was similar to that of Neblina (Boubli 2002, 73 m%/ha). In order to understand the importance value index (IVD of each species and how it contributes to the community one has to look at its relative parameters. Of all the Caesalpiniodeae legumes at Kaieteur two species, Eperua falcata and Chamaecrista adiantifolia var. po e a tota! IVI va of 75. K These m 1 PE species were widely spread throughout the plots, had a In Kwakwani, the highest IVI values (119.5) for both sites came from one species al Lecythidaceae, Eschweilera pedicellata. This species dominated the plots and was higher than the Lecythidaceae at La Fumée Mountain, French Guiana (Mori & Boom 1987). Such high IVI values have only been recorded elsewhere by Gibbs et al. (1980) for Cyclolobium vecchii A. Samp. ex Hoehne (Leguminosae; IVI=82) and Sebastiania klotschiana Kelloff, Kaieteur National Park, Guyana 537 TABLE 2. Stem class size in cm DBH (number of stems/percent of stems) at Kaieteur N Barro Colorado Island, Panama. i, Guyana, and z 10.0 z 20.1 > 30.1 > 40.1 > 50.1 > 60.1 Kaieteur Plot 1 488 / 64.3 147/194 712495 287 3.7 1572.0 8/10 Plot 2 673 / 69.6 201/208 5375.5 19/20 11/11 10/ 1.0 Kwakwani Site 1 3167522 110/21.6 45/89 14/28 7414 13/26 Site 2 2///564 107/217 60 /12.2 207 4:1 12/24 1573.1 Barro Colorado Island Hec21 287 / 68.7 52/124 38 / 9.1 19/46 9/2.1 1373.1 Hec37 7807 62.6 88 / 19.7 36 / 8.1 13/29 14/ 3.1 16/3. 6 Stem Class Size Distribution 700 - | "T N |». EKF2 600 - = TAAA u $ E ART e 500 = EM : = AR2 O f 200 E Hec21 RGE > Hec37 t 300 IEEE ec - e ] m S 200 +- 100 +-- =r - — HÀ 0 H A E == eae en. 10 - 20 20.1 - 30 30.1 - 40 40.1 - 50 50.1 - 60 60.1 + Stem Size DBH (cm) Fic. 5. Distributi (Muell. Arg.) Muell. Arg. (Euphorbiaceae; IVI=119) from his survey in Mugi-Guicu, Brazil. At Barro Colorado Island to species in different families scored high, Trichilia tuberculata (Meliaceae, IVI-51.1) and Faramea occidentalis (Rubiaceae, IVI241.9). When the plot data were examined at the family level, there were 33 families at Kaieteur and 36 at Kwakwani. The number of families in the Guyana plots was lower than the number of families recorded by Balslev et al. (1987) for the floodplain (44) and non-flooded forests (53) of Añangu, Ecuador. It was also lower than the 42 families at BCI. Thirty-three percent of the families were found in all three study sites, with 538 | tani i Texas 2( Taste 3. Dominant famili ithin the tl it itl bined total of over 50% of the stems: Kaieteur (KF), Kwakwani (AR), and Barro Colorado Island (BCI). Numbers shown are percentages of overall st per family found at each site. KF AR BCI Annonaceae = 74 - pocynaceae e 5.2 " Arecaceae = E 5.6 Bombacaceae 8.2 - - Euphorbiaceae - 4,3 a eae 2:5 : Lecythidaceae - 254 d Leguminosa 49.2 18.4 76 Meliaceae 14.1 osaceae 4.] Rubiaceae - - 25.1 E | Annonac. 20222222224] —] [1KF1 = A Apocynac. zz. Ss Em KF2 Bombacac. AR1 uN pa | AR2 Euphorbiac. a Hec21 Leg-M i m o S, : ARA E Hec37 Leg-Fab. Laurac. Lecythidac. Melastomatac. ] Rubiac. = a | Sapotac. e 0 20 40 60 80 100 120 140 160 Number of Stems the dominant families varying among the sites (Table 3). The total number of stems for each family shared by all three sites is compared in Figure 6. Caesalpinioideae dominated the plots at Kaieteur with over 300 stems/ha. Subsequently, it was necessary to leave this subfamily of Leguminosae out of the comparison so that the bars on the graph representing families with less than 50 stems would be displayed. Kelloff, Kaieteur National Park, Guyana 539 69 E KFI KF2 " ARI AR2 40 Hec21 [L]Hec37 Ave DBH (cm) aV aw aV ër ër a éi GC ze e " a > SS ni " 4 SÉ a A Fo PS < SÍ SES S P "y ES $ e "gz p^ d d Plant families De 7 A ye L L e La IRADIER fal H E i, fe It Cal I Lal i, Panama Key to the abbreviations: KF1=Kaieteur, plot 1; KF2-Kaieteur, plot 2; ARI=Kwakwani, site 1; AR2=Kwakwani, site2; Hec21=Barro Col Island , plot 1; Hec37=Barro Colorado Island, plot 2. When the average di ter at breast height of major tree families at Kaieteur, Kwakwani, and BCI were compared (Fig. 7), the DBH ranged from 10.2 cm (Melastomataceae) to 47.7 cm (Bombacaceae). The largest tree recorded among the research sites was an individual of Ceiba pentandra, Bombacaceae, from Hec37 at Barro Colorado Island, with a DBH of 209.5 cm. From the 325 taxa recorded for the three study sites, only 3% (10 species) of all species were shared between the plots. Apeiba aspera, Brosimum alicastrum, and Virola surinamensis were shared between Barro Colorado Island and Kwakwani. Catostemma fragrans, Licania alba, Swartzia schomburgkii, and Tapirira gui- anensis were shared by Kwakwani and Kaieteur. BCI and Kaieteur shared only Pterocarpus rohrii. The only species shared by all three sites was Jacaranda copaia. Thus, there is a substantial lack of overlap in species composition among these tropical tree communities. Comparing genera among sites, only 6% were shared among Kaieteur, Kwakwani and Barro Colorado Island. These were Aspidosperma, Inga, Jacaranda, Licania, Ocotea, Pouteria, Protium, and Swartzia. When pair-wise comparisons were made between the sites, BCI and Kwakwani shared 8.196 of their genera. Kwakwani and Kaieteur Falls shared 6.396 and 5.096 was shared between BCI and Kaieteur. Based on a phytogeographical study of the taxa at Kaieteur, the flora of this area has its strongest affinity (4296) with the Guiana Shield area of South America with only about 1596 of the taxa having a widespread distribution across the Neotropics (Kelloff & Funk 2004). Statistically, non-metric (NMS) ordination on ecological data derived from PC-ORD has helped to describe the vegetation data from the plots at Kaieteur National Park and Kwakwani, Guyana and Barro 540 t tani i Texas 2( Colorado Island, Panama. In all six plots, the first t lescribe the relationshi dissimilarity between the quadrats (in ordinary space) and represent this variation in ordination space Nec & Grace 2002). The quadrats with floristic similarities tend to form loose clusters. In the joint plots these similarities sug- gest trends that can be indicators of microhabitats induced by substrate characteristics, hydrology and/or other edaphic effects in the site. The Janzen-Connell model for tropical tree diversity proposes that seeds and seedlings in proximity of conspecific adults are have a higher mortality rate resulting in recruitments some distance from the parent tree leaving space for colonization by other species (Schupp 1992). Other factors to consider are the ability of the species to disperse its seeds, the competition for light or nutrients, pollinators. Ordination of tree species at Kaieteur separated the 20 x 20 m quadrats in each plot primarily by spe- cies composition. The quadrats were strongly correlated with some of the less dominant tree species such as Eperua rubiginosa, Inga gracilifolia, Bombax eege DES SP. Hevea guianensis in KF1 and Pouteria cuspidata, Clusia sp., Tapirira sp., Licania canella a KF2. Some species such as Hevea guianensis that grows along more inundated soils get Bombax flaviflora of the dryer soils may help explain the variation in the plots. Although the soils of the Potaro Plateau are largely composed of porous white sand with very little nutrients, slope, accumulation of leaf litter, and hydrology can change over relatively short distances (C. Kelloff, pers. obs.). Although ane by Eperua falcata and Chamaecrista adiantifolia var. pteridophylla, the two plots at Kaieteur indicate differences in the forest community over very short dis- tances (Fig. 5). This was not the case at Kwakwani or BCI where analyses indicated some overlap in species composition between the quadrats of the plot pairs at each study site. Although a forest type may be designated based on the dominant species, the forest is not without an array of subdominant or even rare species that have an overall large effect on the floristic composition of the forest. Such changes can occur over distance and time with changes in light (tree falls), seed dispersal, soil composition or by turn-over (David & Richards 1934). An example of one such change is in the Kaburi district of Guyana where the Eperua falcata forests were replaced over time by Dicymbe corymbosa Spruce ex Benth. which expanded its dominance by self-pollarding, thus replacing mature trees with large clumps of slender stems (Davis & Se E and eventually Gi out the other species. The forest at Kaieteur could not be considered a t stand nor could it be considered stri ctly a a Wallaba (Eperua) forest. As seen in plot 2 Chamaecrista adiantifolia var. pteridophylla became the dominate species in that area of the forest. The main family that dominated the forest was the Caesalpiniodeae of the Leguminosae with over 300 stems. The subdominant families range from Lauraceae, Bombacaceae, Clusiaceae, and Fabiodeae. CONCLUSION Plots can be a powerful tool for providing long-term information on forest composition, diversity and struc- tural change. They can then be used to assess changes in the forest over time, and the information gathered from plots can be used to understand how other physical parameters may influence species composition and distribution (Dallmeier & Comiskey 1998). The two plots established at Kaieteur National Park were only a small sampling of the riparian forests of the Potaro Plateau. The information gathered on the tree composition of just one area on the plateau dem- onstrates the diversity and turnover in this forest. The Wallaba (Eperua) —mixed forest plot study at Kaieteur had a larger number of trees per unit area over 10 cm DBH and represented almost twice the stems surveyed in the Mora forests at Kwakwani, Guyana or in the lowlands of Barro Colorado Island, Panama. The plots at Kaieteur were similar to the white sand forest studies by Whitton (1962) at Amatuk, Guyana and by Boubli (2002) at Neblina, Brazil, with their large number of trees and dominance of the family Leguminosae. One hypothesis presented by Torti and Coley (1999) suggested that legumes may be more successful in this region because they have ectomycorrhizal fungi that are good scavengers for nutrients in the otherwise nutrient poor soils and that these fungi suppress saprophytic fungi that are potentially detrimental to this symbiosis. A study by ter Steege and Hammond (2001) suggested that seedlings were compensating for low Kelloff, Kaieteur National Park, Guyana 541 light conditions by providing an internal energy source in the form of large cotyledons. Bulky endosperms sustained larger seedlings and supported juvenile plants when light were insufficient. Although designated as a Wallaba-mixed forest, Kaieteur did not have a SE dominant species such as Eschweilera dior dd that was dominant in the lowland forest of Kwakwani. Studies have shown that Kaieteur | to the Guiana Shield flora (Kelloff & Funk 2004) with an upland element of the Roraima formation. This was noted from this study with via a similarity of Kaieteur to Pico da Neblina in Brazil. Non-metric ee scaling (NMS) methods m E Kee n from li rero pes. researchers ties of vegetation m Ge Beete inierence Statistical analyses al the plots at Kaieteur National Park revealed d the species compo n between the plot pairs. This suggests subtle changes in microhabitats and species dynamics of the subdominant tree taxa on the Potaro Plateau. This study has shown that two 1 ha plots can adequately demonstrate how diverse tree species are in one section of the forest; furthermore, it indicates that sampling was probably too small to capture all of the different habitats represented at Kaieteur National Park. It still remains to be understood why the Caesal- piniodeae and in particular the Eperua and Chamaecrista dominated the forest. Does this indicate favorable ecological condition, the Janzen-Connell effects or can the Caesalpiniodeae legumes with its root nodules thrive better is the nutrient poor soil of the Potaro Plateau? To answer these questions more studies need to be done on the soils of the plateau. Forests on white sand generally have a lower alpha-diversity than those on terra firme (ter Steege et al. 2000a) but noted for their monodominance and high abundance of several tree families, such as the Caesalpiniaceae. These forests also have trees that are locally abundant but globally restricted such as the Greenheart, Chlorocardium rodiei (Rob. Schomb.) Rohwer, Richter, & van der Werff (Lauraceae), in central Guyana. Development or poorly designed logging and mining practices can quickly lead to irreversible damages to the forests and habitats, with eventual species loss or extinction. Analysis of the diversity and the composition on the scale of plots can provide some of the best information needed define protected areas in Guyana (ter Steege et al. 2000b; Kelloff 2003). It is on this scale that we can best understand plant spatial distributions and how radically these can change over a relatively small geographic area. Tue iamen. collected in the plot study at Kaieteur ES Park can be useful for monitoring t assessment. This information, along ral changes of the forest over time | with the “Checklist of the Plant of Kaieteur National Park, Guyana” provides data on the plant taxa found on the Potaro Plateau in the vicinity of Kaieteur National Park. In addition to minimizing habitat damage due to industrial development, these data can be used to provide the framework for conservation efforts in the park as well as for the development of ecotourism. ACKNOWLEDGMENTS There were many people during the course of this research that helped in one way or another. I thank the Smithsonian Institution’s Biological Diversity of the Guiana Shield Program for its financial support dur- ing this project. For work done on the plot study in Guyana, I thank Abelardo Sandoval for his expertise with the surveyor compass; Shawn Lehman, Dyantie Naraine (Centre for the Study of Biological Diversity), Keith David, D. Jafferally, and Jennifer Singh (University of Guyana students), Greg McKee (Smithsonian Institution), and Godfrey Bourne’s 1995 class (University of Missouri-St. Louis) for assistance with data collection and mapping the plots. In Guyana, I also thank the Park Wardens, Michael Phang, L. Gibson, the EPA-Guyana and the National Parks Commission for collecting permits. A special thanks is given to Romeo Williams who was invaluable as a tree spotter and climber. I am also grateful to Raquel Thomas (Iwokrama Rain Forest Reserve) who helped me understand vegetative characters to use in identifying trees in the field. To Margaret and Malcolm Chan-A-Sue, I give many thanks for their friendship, moral support, and excellent flights in and out of Kaieteur. I thank Francisco Dallmeier, James Comiskey (SI/MAB Program) and Richard Condit (STRD for sharing plot data from Kwakwani, Guyana, and Barro Colorado Island, 542 J lof tl tanical h Instit Texas 2( Panama, and Pedro Acevedo-Rodriguez for translating my abstract into Spanish. Special thanks I give to James Comiskey, Erin Tripp, and to all the reviewers that provided helpful suggestions to make this a bet- ter paper. 1 am thankful for staff at many herbaria that I visited for help with identifications: U5, NY, MO, BRG, K, U, P, and the three herbaria in Guyana: BRG, Jenman Herbarium and FDG. This paper was part of my dissertation from George Mason University, Fairfax, Virginia. Special thanks go to my committee: Ted Bradley, Associate Professor of Biology and Committee Director; Barry N. 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Forests and dominant legumes of the Amatuk Region, British Guiana. Caribbean Forest. 23:35-57. Yavitt, J.B. 2000. Nutrient dynamics of soil derived from different parent material on Barro Colorado Island, Panama. Biotropica 32:198-207. 546 BOOK NOTICE RAFAEL GOVAERTS AND DAVID A. SIMPSON, WITH JEREMY BRUHL, TATYANA EGOROVA, PAUL GOETGHEBEUR, AND KAREN WiLson. 2007. World Checklist of Cyperaceae: Sedges. (ISBN 978-1-84246-199-0 pbk.). Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, United Kingdom. (Orders: www.kewbooks.com, publishing@kew.org). $196.00, 765 pp., 9" x 6". Contents: Introduction How to use the Checkiist ure Names Acceptance of taxa Geographical distribution Life-forms Abbreviations Acknowledgments References Cyperaceae Unplaced Names e uri 1 "ae Sol PR T S fis TE Pues | 1 " T] Lx Hei From the publishers.— g y, Cyg ; g y y Į y g importance, especially g l ities in the tropics, where sedges are intensively used. The World Checklist of Cyperaceae is H 1 +] 11 1.11 1 ER | 1 I A Lu EL +1 Cal a 1.1: Aë Jj. a ES s sz) q er [e] Eu metres e 1 I " ] ] 1 1 E y It will raj s Te *z Sw: y e E f. 1 ists, horticulturists, ecologists, L «1 CULICLUELY ALL S 1 LI E ct family. The book J. Bot. Res. Inst. Texas 2(1): 546. 2008 REDISCOVERY OF MIRABILIS HINTONIORUM (NYCTAGINACEAE), A STRIKING FOUR-O'CLOCK ENDEMIC TO THE SIERRA DE COALCOMÁN, MICHOACÁN, MEXICO Mark Fishbein Victor W. Steinmann Department of Biology and Herbarium Instituto de Ecología Portland State University Centro Regional del Bajío egio PO. Box 757 61600, Pátzcuaro, Ste MEXICO Portland, Oregon 97207, U.S.A mfish@pdx.edu ABSTRACT R Ilecting in the Si de Coal an, Michoacan, Mexico, lted in tl li f Mirabilis hintoni ,aspeci that had | li ] f g ] g 1939. The rediscovery i ble I ft] triki fl ftl i which are lin tl by virtue of thei ly limbless, tubular f 1 “firecracker” red and yellow colors: The species is sndedbiediy A 11-51 1 1 q: da (nal RESUMEN En una d a la Sierra ac Ge d México resultó el E de Mirablis hintoniorum, una EE que no primera colecta hecha en 1939. El red notable dado y atractiva por sus flores, | | tubul , con lóbul lucidos y la d illo y rojo. A especie es muy rara y Wee de la Sierra de Coalcomán. From 1931 to 1941, famed plant collector George B. Hinton (1882-1943) conducted extensive botanical exploration, primarily in the southern Mexican states of Guerrero, México, and Michoacán. During this time he made over 16,000 collections, and his specimens have served as the types of more than 360 spe- cies (Hinton & Rzedowski 1975). On July 11, 1939 Hinton secured specimens in the Sierra de Coalcomán, in southwest Michoacan of what is undoubtedly the most striking of all species of four-o'clocks (Mirabilis L.). Despite the dist and showiness of the species, and the several duplicate specimens that were distributed to major herbaria, the species remained “undiscovered” by taxonomists until Le Duc (1992) described Mirabilis hintoniorum Le Duc over 50 years later. In the course of monographic work on Mirabi- lis section Mirabilis, Le Duc came across four duplicates of Hinton’s collection, housed in herbaria at the University of Texas (TEX), Conservatory and Botanical Garden of Geneva (G), Missouri Botanical Garden (MO), and University of California, Berkeley (UC). When she published her monograph in 1995, Le Duc had not encountered additionai collections of M. hintoniorum (Le Duc 1995). In this note, we provide a fuller description and photographs of this remarkable species that had apparently not been recollected until we, too, encountered plants in the Sierra de Coalcomán. Although unremarkable in vegetative morphology, M. hintoniorum is unrivaled among the Heu 60 species in the genus by virtue of the form and color of the flowers. Le D similar species on the basis of glabrous, few-flowered inflorescences and the ely shallow lobes and orange color of the perianth. However, her observations, based only upon examination of few herbarium specimens and Hinton's economical observations on the specimen labels, are not entirely accurate and fail to convey the spectacular impression of the living plants. In particular, the inflorescences are hardly few- flowered and the perianth scarcely orange. On June 28, 2003 we collected Mirabilis hintoniorum for apparently the first time since Hinton encoun- tered the species 64 years earlier. Had we been familiar with Hinton’s collection or Le Duc's description of the species, we would have been wholly unprepared for the appearance of this seeming botanical fireworks display (Figs. 1-3). The flowers, described by Hinton as simply “orange”, are in actuality pendent flames— J. Bot. Res. Inst. Texas 2(1): 547 — 550. 2008 E Jn, re Je Sch d » "nec A de e Fo rum A PAS TS > AS E EF. dotes "an o HE AU Pr A SC "eg D A Ce d Ge e EE a Gs ie KI wf pá = d MAIN j py NE AAA (Gab Ai d + » a y Y d ja y» » x PA Er 1 AA? Lafe Lad | TA L A L As Pe | ge J r | z j Michoacán, Mávirn 1. habit. the perianth tube is bright red, grading into a vibrant yellow with red venation in the limb. However, these features are lost with drying, and the flowers on both Hinton's and our herbarium collections are nearly colorless. In contrast to the "few-flowered" inflorescences (Le Duc 1992, 1995) of Hinton's specimens, those on the plants that we encountered comprise a many-flowered, pyrotechnic cascade. To Le Duc's otherwise accurate description we can add that the plants are erect, bushy, perennial herbs reaching 1 m in height with red stems. In addition to the aforementioned characteristics of the flowers, the stigmas are pink and the anthers red. Like Hinton, we encountered M. hintoniorum only in the Sierra de Coalcomán. Whereas the type local- ity of Villa Victoria is along the Pacific Slope, our initial encounter took place approximately 55 kilometers (airline) away, at the opposite (northeastern) end of the range within the Balsas Depression. We were brought to an abrupt stop by the sight of M. hintoniorum while traveling along the unpaved road from Aguililla at the foot of the mountains to the high-mountain logging town of Dos Aguas. As we approached the unknown marvel, we were dumbstruck with the realization that we were gazing at a Mirabilis dramatically unlike any that either of us had ever seen, the myriad cultivars of M. jalapa L. not excepted. Only a small patch of these robust plants was found at this site, which we documented with several specimens (Fishbein and Steinmann 5113). Later the same day, we observed another small patch on the southwestern side of the range, between Dos Aguas and Coalcomán along the road to Villa Victoria, from which no collections were made. We did not observe the species anywhere else. It is paradoxical that M. hintoniorum, arguably the showiest species in the genus, should have remained so obscure, notwithstanding being a narrow endemic to the Sierra de Coalcomán. Two populations were Fishbein and Steinmann, Mirabilis hintoniorum, rediscovered Fics. 2-3. Mirabilis hintoniorum Le Duc photographed in the Sierra de Coalcomán, Michoacán, México. 2 (top). flowering stems. 3 (bottom). close up of flowers. 350 t tani i Texas 2( found along a moderately traveled, albeit rugged, road that has been traversed periodically by plant col- lectors, including Rogers McVaugh. In fact, we made our collection at the type locality of Ipomoea fissifolia (McPherson) Eckenw. It seems likely that the species is indeed endemic to the Sierra de Coalcomán and it is quite likely rare. Furthermore, it possibly has a very brief flowering span in the early part of the rainy season, and these factors may help to explain not only the few encounters by plant collectors but also the absence of the “firecracker four-o’clock” from our gardens. Exsiccatae. MÉXICO. Michoacán. Distr. Coalcomán: Villa Victoria, alt. 700 m, dense woods, 11 Jul 1939, G.B. Hinton et al. 13909 (HOLOTYPE: TEX; IsoTYPES: G, MO, photo!, UC); Mpio. Mnt ge ae "Des BS ue 18.2 km dee in of La Paz at jet N Aguililla road, 18°46.0°N 102*50.90'W, elev. 15 transition to oaks, with Guazuma, Helioc rpus, Pouzolzia, Oxalis, Gonolol lif Opuntia Cordia, 28 Jun 2003, M Fishbein E VW Steinmann 5113 (ARIZ!, F!, HPSU!, IEB!, MEXU!, MISSA!, MO!) ACKNOWLEDGMENTS We used the Missouri Botanical Gardens w"TROPICOS data (mobot.mobot.org/W3T/Search/vast.html) to examine an image of MO's isotype of Mirabilis TM, We thank Ignacio García, Guy Nesom, and Richard Spellenberg for reviewing the manuscript and providing helpful suggestions. Fishbein's fieldwork was supported in part by the Department of Biological Sciences, Mississippi State University, and Stein- mann’s studies were supported by a grant from the Comisión Nacional para el Conocimiento y Uso de la Biodiversidad to the Instituto de Ecología (account number 20007). REFERENCES HINTON, J. and J. Rzepowski. 1975. George B. Hinton, Explorador botánico en el suroeste de México. Anales Esc. Nac. Ci. Biol. 21:1-114. Le Duc, A. 1992. A new species of Mirabilis (Nyctaginaceae) from Michoacán, México. Sida 15:53-55. Le Duc, A. 1995. A revision of Mirabilis section Mirabilis (Nyctaginaceae). Sida 16:613—648. FLORISTIC COMPOSITION OF SEASONALLY DRY TROPICAL FOREST FRAGMENTS IN CENTRAL BAHIA, NORTHEASTERN BRAZIL Domingos Benício Oliveira Silva Cardoso Luciano Paganucci de Queiroz Herbário da Universidade Estadual Universidade Estadual de Feira de Santana de Feira de Santana (HUEFS) epto. de Ciéncias Biológicas, Herbário da Km 03-BR 116, Campus. 44031-460 Uni idade Estadual de Feira de Sant (AGEFS) Feira de Santana, Bahia, BRAZIL Km 03-BR 116, Campus. 4403 1-460 cardosobot@gmail.com.br Feira de Santana, Bahia, BRAZIL Igueirozquefs.br ABSTRACT The Serra do Orobó M j gei ] lin centra ] Bahia State, northeastern Brazil, and pi f PI imately 7,300 ha = g I " eee phy i g E te 11 TA 1 g 1 1 ] i] L r a that grow the 1 l ] ti (450 to 700 meter) and f g t f sul t ] id f tg i g the summit (700 to 850 meters). À base survey was undertaken at the 1 I 1 fi frag t d al eh Orobó g 1.3 : GE i - : A A total of 615 species within 378 genera and 106 families 597 speres DS 365 Ge e 96 families, while peridophytes comprised 18 species in 13 genera and 10 families. The floristic m diversity 5 I the € zil 141 Ze | fl [41 11 Dees | -1 r i Cal 1 1 1 3 E 65 d e Pi ALEA LAA A al 11 J r A TD A +] Tm 1 1 1 1 P J LILO eCaoUllall) UL y IOLESL e 5 a Serra do Orobó 1 tan ] lizad ] área central del estado de Bahia, gi 4 iental del Brasil, que ocupa I 7 ke? 300 ha r7 J E. 1:£ 1D l TT. I E UT... a 1 $ E (SDTP), incluyendo 1 ed 1l D E 1 1 : . 1 1.3: T E 1 ` z AEN i ) L t 1 FZ X 700 metros, y E g CH 1 1 ] 1 + Laz 1 1 1 r I 2] m 7005 ocr P T 11 1 1 Je mn t ] f buen estado d ión a lo largo de la Serra do Ge Fueron muestreados un total de 615 ee en 378 genera y 106 familias, incl do di taxa de dif famili PN ~ Pd en 365 indi y 96 Ge > "iss pcs Bus ni 18 dn e en 13 genera y 10 ert La diversidad a la f1 al E é E T + a] zl 21. p pa | ry TA 1 H 5 1 + a dall Jal 4 L L de la Floresta Atlánti lAs] A E INTRODUCTION Seasonally Dry Tropical Forests (SDTF) are one of the world’s most threatened ecosystems. Miles et al. (2006) observed that ca. 97% of these dry forest remnants are at risk for biodiversity loss due to a number of different anthropogenic Emel such as dira e On pret change, fire, or conversion into cultivated areas. However, SDTF relatively little attention fro m vationists and ecologists in comparison to humid forests (Pennington et al. 20060). Little is presently known about the floristic diversity of neotropical SDTF although in recent years there has been an increasing interest in this vegetation type (Gentry 1995; Giulietti et al. 2006; Pennington et al. 20062). However, the growing knowledge of the SDTF flora has stimulated the formulation of various hypotheses concerning the historical biogeography of this vegetation (Prado & Gibbs 1993; Pennington et al. 2000, 2004; Prado 2000; Lavin et al. 2004; Queiroz 20062) and the geographical EEN of its com- ponent species, and studies have indicated that SDTF are independent phy ts distributed disjunctly through South and Central America (Pennington et al. 2000; Prado 2000). According to the definition elaborated by Pennington et al. (2000), the neotropical SDTF comprehend different physiognomies, and can include both forests as well as more open vegetation types where shrubs predominate. Terrestrial bromeliads and succulent plants belonging to Cactaceae and Euphorbiaceae are J. Bot. Res. Inst. Texas 2(1): 551 — 573, 2008 552 | | of the Botanical R h Institute of Texas 2(1) common in SDTF areas, as well as an ephemeral herbaceous tapparent only during the short rainy season. Another important aspect characterizing SDTF is their maket; seasonal phenology of flowering and fruiting, with many species demonstrating synchronous flowering at the transition between the dry and the rainy seasons (Machado et al. 1997). The Caatinga domain constitutes the largest and most isolated nucleus of the SDTF (Queiroz 20069). It occupies ca. 800,000 Km? in the northeastern section of Brazil (Ab'Sáber 1974) with a semi-arid climate mainly in the low altitude areas between extensive plateaus and/or mountain ranges (Prado 2003; Queiroz 2006a). Andrade-Lima (1981) noted that the vegetation of the Caatinga domain comprised different physi- ognomies, including areas of open vegetation dominated by shrubs or areas with relatively tall forests. These seasonally dry forests are commonly referred to as arboreal caatinga, dry forests, or deciduous forests, and occur as highly fragmented areas throughout the semi-arid region of northeastern Brazil, principally from Bahia State through northern Minas Gerais State. In spite of the fact that these dry tall forest areas are situ- ated within the Caatinga domain (Rizzini 1979; Andrade-Lima 1981), different workers have indicated that they may in fact be phytogeographically more related to the dry vegetation of the Atlantic Forest domain (Oliveira-Filho & Fontes 2000; Amorim et al. 2005; Oliveira-Filho et al. 2005, 2006), or may represent dis- junct areas of SDTF without any relationship to the humid areas of the Atlantic Coastal Forests (Pennington et al. 2000; Prado 2000). The deciduous and semi- deciduous Desin of See and southeastern Brazil, for example, have been Eeer to have a strong flori hip with the moister sites of Atlantic Coastal Forest and therefore should not be treated as a type of dry tall forest included in the Caatinga domain (Oliveira-Filho & Fontes 2000; Oliveira-Filho et al. 2005). However, Oliveira-Filho et al. (2006) have more recently proposed a wider definition for SDTF that would treat these same deciduous and semi-deciduous forests of the Atlantic Forest domain as another nuclei of SDTF. As such, it is clear that there are still problems to be resolved concerning the nature of SDTF in eastern South America. The delimitation of SDTF is complicated by the fact that Penington et al. (2000) and Prado (2000) did not include humid forests in their analyses, while Oliveira-Filho et e (2006), on s ge nano, included only a very limited sampling of Caatinga areas. Additionally, the | ledg composition of many areas of SDTF in northeastern Brazil is still very incipient. As such, the present study undertook a floristic survey of several fragments of seasonal dry forests in the Serra do Orobó Mountains in central Bahia FS Sa uode GEN of these dry forests will be discussed here in terms of the present und graphy of SDTF. Specifically, we intend to demonstrate that the dry forest framents of the Bera do Orbe Mit s are floristically related to the SDTF floras. To account this goal, we will treat the SDTF as a broader definition (sensu Oliveira-Filho et al. 2006) that encompasses the dry sites of the Atlantic Forest domain as a distinct nuclei from the Caatinga of the Serra do Orobó mountain range with these former nuclei. domain and discuss the I Study Area The Serra do Orobó mountain range is located within the municipalities of Ruy Barbosa and Itaberaba in central Bahia State, Brazil (12°15’ — 12º25'S and 40°19’ — 40?30"W) (Fig. 1). The range occupies an area of approximately 7,300 ha and has a maximum elevation of 1,014 m (BAHIA 2002). The region has a semi- arid climate (SEI 1998), and although there are no systematic rainfall SE for this area, estimates of precipitation in the region place the average annual rainfall at approximately 800 mm (SEI 1998). The aver- age annual temperature is 26 ?C, with an bolas maximum af 33 °C, and an p minimum of 18 *C (SEI 1998). The forest fragments examined have shallow soils that are sometimes covered by a significant layer of leaf litter. These soils are classified in the main group “luvisoil” and present moderate to high pH and nutrient levels (IBGE/EMBRAPA 2001). In addition, they are comparatively more fertile than those of the nearby savannas. The Serra do Orobó mountain range is located within the Caatinga domain of northeastern Brazil (Velloso et al. 2002), and several vegetation types can be found there (Fig. 2) according to general altitudi- nal gradients. A shrubby caatinga with palm trees is most common at lower altitudes, while forests occur Serra do Orobó = d Espinhaco range 181) | cio --18 (land areas above 1000m) E 0 5 nm egen Lee EES -45 -42 ES De 1 Lu! [| Ir 4° fal e LA LZ AA ha! E A FL | J. DS Ai : A A Chat , Brazil at higher altitudes (between 450 and 850 meters). Dry forest fragments and riparian forests are found on the slopes at elevations between 450 and 700 meters. There is a more humid area near the summit on the southern slope (between 700 and 850 meters) harboring a submontane humid forest. Above the forest zone, and surrounding the mountain peaks up to the 900 meters, is a savanna-like vegetation with large popula- tions of Vellozia furcata L.B.Sm. & Ayensu (Velloziaceae). Near the summit, the rock outcrops harbor campo rupestre (rocky field) vegetation. Although the Serra do Orobó has suffered strong anthropogenic alterations in 400 years of European colonization, some forest fragments are still relatively well-preserved. These fragments have extremely high conservation value as they represent some of the last intact fragments of dry forest in northeastern Brazil. Additionally, many new species belonging to different groups, such as the fungi (Neojhonstonia minima, Gusmáo & Grandi 2001), psychodid flies (Psychoda serraorobonensis, Bravo et al. 2006) have been described from this area. Besides, some new species of plants from the campo rupestre vegetation were identified and are in process of publication (Calliandra, Fabaceae-E.R.Souza & L.P.de Queiroz, pers. comm.; two new | || E ak n.a H Inm L I FRI A ET mia ACF 554 t* B Rinarian Fic > ML A E A L H a E J: al e I ri LAA A H d Bahia State, Rrazil A LE g E Li LI r AL A D Interior of af t extendi | | fth tai Ê with Eriotheca cf g lof the humid forest, showing many epiphytes. species of Marcetia, Melastomataceae—A.K.A.Santos, pers. comm.; Mitracarpus, Rubiaceae—E.B.Souza, pers. comm.; Oncidium, Orchidaceae-C van den Berg, pers. comm.; and Sauvagesia, Ochnaceae-D.Cardoso, pers. comm.). In 2002, the forest remnants as well as the campo rupestre vegetation of the Serra do Orobó mountain range were assigned the status of sustainable use protection areas in recognition of their importance to conservation and to biodiversity research. METHODS A floristic survey was carried out from August 2004 to June 2006 in the best preserved sites of semidecidu- ous, riparian, and submontane humid forest fragments. The vegetation classification used here for each for- est fragment follows Veloso et al. (1991) and stresses the distinct physiognomies and ecological conditions observed in each. It was studied 8 fragments of semideciduous forests (2.85 km2), 6 of riparian forests (0.37 km?) and one of submontane humid forest (0.07 km2). During the course of the present study a total of 15 field trips were carried out, each lasting from three to four days. Of all the 15 studied fragments, the submontane humid forest one was the unique where our collect effort was lower due to its difficult access and because it was discovered only recently. All fertile material of vascular plants was collected according to the standard methods described by Filgueiras et al. Fal J A | Q H À e lly J E PRE E FEM! fa La Brazil 555 (1994). In some cases, ue collections were Educ for peres never encountered with flowers or fruits. A total of 1,385 voucherc de and y stored at the Feira de Santana State University Herbarium (HUEFS). In some cases, species that were ay difficult to collect were included in the checklist if they could be positively identified in the field; these are indicated as “Not collected” in Appendix 1. All specimens were identified by the authors with the aid of identified collections stored at the HUEFS and/or by using specific bibliographies. Specimens identified only to genus or family, are clearly labeled in the Appendix. po were sent to specialists in order to Dëse or to confirm He dun Angiosperm families are organized according to APG II (2003) in the floristic list according to Tryon and Tryon (1982). Authors of species are abbreviated according to Bram and Powell RESULTS The vascular flora of the forest vegetation of the Serra in b mountain range comprises 615 species belonging to 378 genera and 106 families (Appendix 1) represented by 597 species in 365 genera and 96 AS thg pa SECHER 18 neces PM DN to 13 EE gie a ee 2 total of 561 taxa 96). However, in som many taxa were identified only to d genera (51 specimens, or 8.3% of he total collection) or family pu (3 specimens, 0.596) due to their taxonomic difficulty. The semideciduous forests demonstrated the greatest species richness (465 spp.), while 141 and 87 species were sampled in the riparian and in the submontane humid forests, respectively. Considering all forest fragments together, the commonest life form was trees (182 species), followed by vines (138), herbs (11D, shr E (109), See (45), and epiphytes (35). The most diverse families were Fabaceae Ge spe- cies), Eupl (38 (24), Orchidaceae (24), Rubiaceae (24), Asteraceae (22), Bi (19), Myrtaceae (18), RE (17), and Rutaceae (15), which together comprised 4896 of the total number of species surveyed in the Serra do Orobó mountain range. Thirty-five families were represented by only a single species. The most species-rich genera were Croton (11 species), Passiflora (8), Senna (8), Eugenia (7), Myrcia (7), Solanum (7), Miconia (6), Bauhinia (5), and Senegalia (5). Several species were identified by different specialists as pang new taxa (Fig. 3). These included: GE nee ee Castro & Rapini "ido: P} tri pala (Fabaceae, Queiroz 2006b), and to the genera C (Fal L.P.de Queiroz, pers. comm.), Conchocarpus (Rutaceae—J.R. Piani pers. ae ee (Commelinaceae—L.Aona, pers. comm.), Eugenia (Myrt- aceae-M.Sobral, pers. comm.), Sinningia (Gesneriaceae—A.Chautems, pers. comm.), Solanum (Solanaceae— L.P.de Queiroz & D.Cardoso, pers. comm..), Standleya (Rubiaceae—J.G Jardim, pers. comm.), and Spermacoce (Rubiaceae-E.B.Souza and E.L.Cabral, pers. comm.). Additionally, of the total number of species collected, 12 were considered to be endemic to the dry forest fragments of Serra do Orobó or to the adjacent dry forest areas. These species are marked by an asterisk in the Appendix 1. In oo to ness new species, other taxa were collected for the first time in Bahia State, including Bulbophyll anum (Orchidaceae), Croton warmingii (Euphorbiaceae), es Oxalis roselata EE (Fig. 3. Other une in D included Cordia blanchetii (Boragi Heteranthia (Solan aceae), and Passiflora bahi ), which were only previously known from their SC material collected in Bahia: as well as lovotilbesis bech and Wullschlaegelia calcarata, both very uncommon sapro- phytic orchids (Fig. 2). The forest fr ae composed of different plant strata. In general there was a continuous canopy 20) meters high camnoce d of Acnid ¿fal (A y TT trelloncic (T thid X Ca. E (23 UE Á ZW C * / 1 T1: 1 mr 1 Li ^ e 1.1 71 FAA ^ H a i , Heis- Centrolobium E (Fabaceae), Cni IT (Olacaceae), Hirtella 1 gla dulosa (Cl bal ) EE cfi Myrsine ferruginea (Myrsinaceae), Peltogyne confertifióra Firad. Piptadenia paniculata (Fabaceae) and Vantanea compacta (Humiriaceae). Emergent trees could reach up to 30-45 meters, and included Eriotheca } NOMAS 556 t tani i lexas 2(1) Fic 1 P PASE Fall £l Pal e L A LGS CNET A AA APA FS A A D Ze Th Cinningin d T E 1 ui Ee = G. Ovalis roselata: H P, F , r - F , wë LI en nov: E Friaenia sn. nov: E Pf LI Li a LI - 1 aw ^ Te ] 1 ylla, E cf. globosa (Malvaceae), Caesalpinia leiostachya (Fabaceae), Cavanillesia gomelleira (Moraceae), Goniorrhachis marginata (Fabaceae), Vochysia sp. (Toyata, and Mandhari m (Sapotaceae). Several vine species, mostly belonging to the families Apocynaceae, Bignoniaceae, Fabaceae, Passifloraceae, and Sapindaceae, occur in the canopy, within treefall gaps, or along the edges of the forest fragments. The subcanopy was dominated by species of Euphorbiaceae, Fabaceae, Myrtaceae, Melastoma- taceae, Rubiaceae, and Rutaceae. The submontane humid forest fragment was the richest area for epiphytes, including Anthurium pentaphyllum, A. scandens (Araceae), Nematanthus albus (Gesneriaceae), Oncidium ciliatum, O. flexuosum (Or- chidaceae), Peperomia sp. (Piperaceae), Tillandsia stricta (Bromeliaceae), and Vriesea simplex (Bromeliaceae). This submontane humid forest fragment also had a dense herbaceous stratum composed of sciophilous herbs such as Paepalanthus spatulathus (Eriocaulaceae), Sinningia barbata (Gesneriaceae), Begonia petasitifolia (Begoniaceae), Dichaea congniauxiana (Orchidaceae), Sobralia sessilis (Orchidaceae), Gibasis geniculata (Com- melinaceae), Ichnanthus grandifolius (Poaceae), and Ruellia affinis (Acanthaceae). rn fe J J | Fi H E M | £ £ £. Ka | P.L” Brazil 557 d d =f The commonest species found at forest edges and in disturbed sites were the trees Albizia polycephala (Fabaceae) and Maclura tinctoria (Moraceae), the shrubs Diospyros inconstans (Ebenaceae), Lantana camara (Verbenaceae), and Senna ples alia EE the vines Arrabidaea cinerea (Bignoniaceae), Chaetocalyx scandens (Fabaceae), Dicell (Malpig ), Macroptilium erythroloma ER ee Periandra coc- cinea (Fabaceae), and the herbs Chaptalia int E ilia fosbergii (Ast RES a ane H. suaveolens eo) tin the best preserved f lorest fragments, such Gall ifoli pau- -d'alho”, Phytolaccaceae) nud espia leiostachya (“pau-ferro”), are used locally as folk Méd cues mala we found evidence of fully grown trees being harvested for their wood, including Copaifera langsdorffi (“pau-d'óleo”, Fabaceae), Brosimum guianense (“oiticica”, Moraceae), Ocotea glomerata ("lóro", Lauraceae), and Hymenaea courbaril. jatobá,” Fabaceae). DISCUSSION Floristic diversity The forest fragments of Serra do Orobó mountain range demonstrated a relatively higher number of spe- cies in comparison to other Brazilian semideciduous forests, even comparing the arboreal component only (Meira-Neto et al. 1989; Oliveira-Filho & Machado 1993; Soares-Filho 2000; Werneck et al. 2000; Jarenkow & Waechter 2001; Moura & Sampaio 2001; Cielo-Filho & Santin 2002; Paula et al. 2002; Silva & Soares 2002; Nunes et al. 2003; Souza et al. 2003; Andrade & Rodal 2004; Toniato & Oliveira-Filho 2004; Rodal & Nascimento 2006; Macedo 2007), or shrubby caatinga (Rodal et al 1998; Lemos & Rodal 2002; Pereira et al. 2002; Alcoforado-Filho et al. 2003; Nascimento et al. 2003; Andrade et al. 2004; Rocha et al. 2004), riparian forests (Vilela et al. 1995; Funch 1997; Passos-Júnior 1999; Stradmann 2000; Ribeiro-Filho 2002; Lacerda et al. 2005), or enclaves of humid forests within the caatinga (locally known as *brejo de altitude") (Tavares et al. 2000; Rodal & Nascimento 2002; Melo & Rodal 2003; Agra et al. 2004; Barbosa et al. 2004). The species richness encountered in the SUDAN humid forest (87 spp.) was surprisingly lower than would normally be expected i in this forest | ] y in comparison with other studies (Mori 1989; Lombardi & Goncalves 2000; Tavares et al. 2000; culo Sobrinho & Queiroz 2005; Amorim et al. 2005; Neves 2005; Martini et al. 2007), and lower also that observed in the dry and riparian forest fragments of the same mountain range. This may be due to the reduced size of the fragment examined, and/or to the relatively low sampling effort in this area. As such, more field trips to the area may reveal the presence of other species not yet recorded. With minor variations, the ten most species-rich families in this survey were the same families identi- fied as important in other SDTF sites, irrespective of the general aspects of the local climate, soil, and forest physiognomy. Similarly, several of these families (usually Fabaceae, Euphorbiaceae, Apocynaceae, Bignoni- aceae, SM AM and Rubiaceae) were important in sand dune areas of the middle Sáo Francisco river in Brazil (Nascimento et al. 2003; Rocha et al. 2004), in caatinga areas on sandy soils (Araújo et al. 1998; Rodal et al. 1998; Andrade et al. POE o a areas on I AN basement rocks (Pereira et al. 2002; Alcoforado-Filho et al. 2003), astern and southern Brazil (Jarenkow & Waechter 2001; Cielo-Filho & Santin 2002; Paula et al. 2002; Silva & Soares 2002; Toniato & Oliveira-Filho 2004), riparian forests in the Chapada Diamantina of Bahia State (Funch 1997; Passos-Jünior 1999: Stradmann 2000; Ribeiro-Filho 2002), as well as in several neotropical SDTF (Pennington et al. 2000; Bridgewater et al. 2003; Linares-Palomino et al. 2003). Fabaceae was the most species-rich family in the Serra do Orobó mountain range as well as in the other sites referred to above. Schrire et al. (2005) presented historical and biogeographical evidence supporting the hypothesis of a long term association between le- gumes and dry forests. Amongst the vines, the families with the greatest numbers of species in the Serra do Orobó (Apocynaceae, Bignoniaceae, Malpighiaceae, Fabaceae, and Sapindaceae) were also encountered in other semideciduous forest fragments (Hora & Soares 2002; Udulutsch et al. 2004). These too similar species richness at the family level encountered throughtout different physiognomies of dry vegetation in 558 Brazil as well as in the Central and northern South America may also be concerned for justifying that the Serra do Orobó mountain range should be included into a broad definition of SDTF. On the other hand, a more restrict evaluation comparing the most species-rich genera may be sugges- tive to stress that within the different physiognomies of SDTF in eastern Brazil, the dry forest fragments of the Serra do Orobó Mountains should be floristically related to the semideciduous forests of the Atlantic Forest domain rather than to the dry shrub-arboreal vegetation of the Caatinga domain. For example, most of the genera comprising the largest numbers of species identified in this survey, such as Croton, Passiflora, Miconia, Myrcia, Eugenia, Solanum, and Bauhinia are better represented in several areas of semideciduous forests (e.g. Soares-Filho 2000; Paula et al. 2002; Silva & Soares 2002; Nunes et al. 2003; Andrade & Rodal 2004; Amorim et al. 2005; Macedo 2007) than in shrub-arboreal caatinga areas (Araújo et al. 1998; Rodal et al. 1998; Lemos & Rodal 2002; Pereira et al. 2002; Alcoforado-Filho et al. 2003; Cardoso & Queiroz, unpublished data). Moreover, the paucity of Cactaceae species in the Serra do Orobó Mountains is another evidence for lack of closer relationship with the Caatinga domain, where this family is usually abundant (see for example Taylor & Zappi 2002; Rocha et al. 2004; Gomes et al. 2006). Endemism Among the 152 arboreal species surveyed in the dry forest fragments in the Serra do Orobó Mountains, 16 (see Append 1) are Loon ott Ee of the SDTF flora (Prado & Gibbs 1993). Besides, the genera 4 il tia (Cactaceae), and Pterogyne (Leguminosae) surveyed here are endemic to the SDTF as a whole (Prado 2000). In addition, another 14 species considered endemic to the caatinga domain occur in the dry forest fragments of the Serra do Orobó Mountains or in other nearby areas, such as Anemopaegma laeve (Bignoniaceae), Apodanthera glaziovii (Cucurbitaceae), Averrhoidium gardnerianum (Sapindaceae), Capparis jacobinae (Brassicaceae), Heteranthia decipiens (Solanaceae), and Senna acuruensis var. caatingae (Fabaceae) (Giulietti et al. 2002). This highlights that the Serra do Orobó Mountains may be indeed related to the other remmants of seasonlly dry forests of the neotropics. The presence of a considerable number of new species (ten) in the dry forest fragments in the Serra do Orobó Mountains may be explained in term of the processes of metacommunities, a model recently put forward for explaining the maintenance of diversity within SDTF (Lavin et al. 2004; Schrire et al. 2005). These authors propose that the biogeography of the succulent biome (which includes neotropical SDTF) can be better understood on a global scale by postulating metacommunities than by proposing historical processes of speciation due to climatic changes or tectonic movements. According to this model, SDTF oc- cur in isolated areas (i.e. much like islands isolated within the continental area), with each area functioning like a local community that is linked to other areas through species exchange based on random dispersal events. The more extreme environmental conditions in SDTF restrict colonization by species originating in most other vegetation formations (Lavin et al. 2004), limiting the rate of immigration into a given local community and resulting in floras that are locally endemic. These processes also result in genera with very few species in local SDTF communities, in contrast with those observed in humid forests (Lavin 2006). For example, the most diverse genera in SDTF (e.g. Senegalia, Croton, Machaerium, Mimosa, Senna, and Zan- thoxylum) usually do not comprise more than ten species in any given forest area (see Werneck et al. 2000; Nascimento et al. 2003; Rodal & Nascimento 2002; Bridgewater et al. 2003; Linares-Palomino et al. 2003; Melo & Rodal 2003; Rodrigues et al. Pid Santos & Kinoshita 2003; Andrade & Rodal 2004; Lemos 2004; Rocha et al. 2004; and this study), while 1 diverse genera in humid forests (e.g. Eugenia, Inga, Miconia, Myrcia, Pouteria, Psychotria, and Solanum) can have from 15 to even 27 species of the same genus occurring in the same locality (see Amorim et al. 2005; Neves 2005; Moreno et al. 2003; Lombardi & Gonçalves 2000; Oliveira & Amaral 2004; Martini et al. 2007). The dry forests of Brazil are distributed in a fragmented and isolated fashion (very much like islands), which has led to the appearance of a significant number of endemic species. For example, 12 species en- countered in this study had distributions limited to the forest fragments of the Serra do Orobó Mountains and adjacent areas. As such, we suggest that the endemism observed in these forests can be explained (on fe [| ) | ri = E | || | E A q A E Ba: Brazil 559 a regional level) by the processes of met ities, in much the same way as species exchange has been proposed on a continent scale (Lavin et al. 2004; Lavin 2006; Schrire et al. 2005). In that context, the forest vegetation of the Serra do Orobó Mountains as a whole could be viewed as a local community that is isolated from other dry forest fragments found in eastern Brazil. Additionally, the recognition that this set of dry forest fragments (together with the Serra do Orobó Mountains) is a regional metacommunity is supported by the presence of disjunct species among the different areas—a pattern that has been similarly observed at a continental scale among the different nuclei of SDTF (Pennington et al. 2006b; Lavin et al. 2004). Cordia blanchetii, for example, has only been observed in the Serra do Orobó Mountains and in another dry forest fragment in Jequié, Bahia State; Passiflora bahiensis has been recorded only in a few collections from an urban forest fragment in Salvador, Bahia; Croton warmingii demonstrates a disjunct distribution within other semi-deciduous forest fragments in a eieiei Gees jorgei and Senna acutisepala (Fabaceae) are known in Bahia only from a few semid forest fragments in the southern part of that state; and Margaritopsis carrascoana (Rubiaceae) occurs in isolated areas of dry vegetation from Ceará State to northeastern Bahia. Floristic relationships Riparian forests.—These forests are closely associated with rivers and are widely distributed throughout Brazil, principally in the central region of the country (Oliveira-Filho & Ratter 1995), although local abiotic conditions may not always be adequate for the establishment of typical riparian forests (Rodrigues 2000). The riverside forests of the an do iud von BE in steep and relatively narrow valleys. As such, g Very great; ana frequently results in intercalation ly found in those forests, such as Cupani ] ), Dictyoloma vandellianum Tamra, Machaerium acutifolium (Fabaceae), Pera pron basa, Piper arboreum (Piperaceae), Shoepfia brasiliensis (Olacaceae), and Vitex aff. klugii (Lamiaceae). However, a majority of the arboreal species encountered in the riparian forests occur only in this vegetation type. Many of the species occurring in the Serra do Orobó Mountains have been reported from riparian forests in the Chapada Diamantina in central ae EUREN Too E SE E E 2000; Ribeiro-Filho 2002) as, for o iii vir- (Clusiaceae), Fi geg contact with tl la thibaudiana cee) Pouteria ion etc Sloanea guianensis (Plasgearpateae): Simarouba amara (Simaroubaceae), Richeria grandis (Euphorbiaceae), Roupala montana (Proteaceae), and Vantanea com- pacta. However, some species observed in the riparian forests of the Serra do Orobó Mountains are widely distributed within the riparian forests of central Brazil, such as Copaifera langsdorffii, Hirtella glandulosa, Ta- pirira guianensis, and the tree-fern Cyathea delgadii (Cyatl ) as noted by Oliveira-Filho & Ratter (1995). These same authors postulated that these riparian forests could act as ecological corridors linking the two principal nuclei of humid forests in South America (the Atlantic Coastal Forest and the Amazon Forest) due to d dense EES EE all of the Cerrado Biome in central Brazil. J ts.—The classification of the SEH ay tees of DOM Brazil is still a t l subject pur Lima (1981) considered t caatinga, and thus part of the Caatinga domain. Prado (2000) and Pennington et al. (2000), based principally on the patterns of distribution of the arboreal species, adopted a similar circumscription, but considered the caatinga and the seasonal dry forests of eastern Brazil as part of a greater phytogeographical unit of the neotropics—the so common referred seasonally dry tropical forests (SDTF)—as distinct from the Atlantic Coastal Forest. This phytogeographical unit would comprise a large variety of vegetation formations, from tall forests in more humid areas to cactus scrub in the EE regions. This controversy can be clearly seen in the great variety of names applied to these diff physiog ies, such as: in Gs EE dry forest t, bosque mi forests, caatinga, mesotrophic, caducifólio, and thorn woodland (Penpüngión et al. 2000). On the other hand, Oliveira-Filho et al. (2006) performed similarity analyses with areas of P d dry forests, shrub-arboreal caatinga, and humid Atlantic Coastal Forest areas, and identified g 360 t tani i Texas 2( affinity between seasonal dry forests and Atlantic Coastal dude areas than with shrub-arboreal caatinga vegetation. Ás such, Oliveira-Filho et al. (2006) proposed a wider ition of SDTF of eastern South America that would include the three major floristic nuclei of dry vegetation formations: the Caatinga, the Chaco, and the seasonal forests of the Atlantic Forest domain, sensu latissimo. An analysis of arboreal and shrub species distributions from semideciduous and humid forest frag- ments in the Serra do Orobó Mountains demonstrated that 106 species (46.596) also occur in areas of the Atlantic Por: domain co Oliveira-Filho et a A ee hrysophyllum g pum (Sapotaceae ) FRA n XA EE amaral E ‘TD Y 1 AS (Fabaceae), and Trichilia hirta Miaa A total p 56 ue 04%) v were common to areas sampled in the Caatinga damain and included A Senna spectabilis (Bibacedó) and Zizi joazeiro (RI ) (Giulietti etal. 2006). Of diesen n Bub. nine of were considered as enden to ehe Castings domain (Giulietti et al. 2002). Additionally, if consider- ing all the 318 endemic species to the Caatinga domain cited by Giulietti et al. (2002), another eight non arboreal or shrub species occur in dry forest fragments of the Serra do Orobó Mountains or in other nearby areas. However, these 17 species make up only 4.596 of the total endemic flora of the Caatinga domain, suggesting that the dry forests of the Serra do Orobó Mountains might not be phytogeographically related to the Caatinga domain, but rather are of Atlantic Forest vegetation. Therefore, these numbers reinforce the hypothesis of Oliveira-Filho et al. (2006) that the semideciduous forests of eastern Brazil are floristically more closely related to the Atlantic Forest domain than to the Caatinga domain. These suggestions are further strengthened by comparisons made between the flora of the Serra do Orobó Mountains and the Serra do Teimoso Reserve (a fragment of Atlantic Coastal Forest in southern Bahia State). The Serra do Teimoso Mountains, like the Serra do Orobó Mountains, have dry vegetation at their lower altitudes but more humid conditions exist at greater elevations, and the flora there is more closely related to the Atlantic Forest domain than to the SDTF sensu Pennington et al. (2000) (Amorim et al. 2005). Among the 252 arboreal species that occur in the Serra do Teimoso Mountains, 52 can also be found in the Serra do Orobó Mountains, including Andira fraxinifolia (Fabaceae), Brasiliopuntia brasiliensis (Cactaceae), Brosimum guianense (Moraceae), Cavanillesia arborea (Malvaceae), Caesalpinia pluviosa, Centrolobium tomen- tosum, Goniorrhachis marginata (Fabaceae), in ae moa O Urera caracasana (Urticaceae), ldin common include: and Zantł | hoifolium (Rutaceae). E Costus spiralis Coca Sinningia barata (Gesneriaceae), Coccoloba decimal (Polygonaceae), Faramea hyacinthina (Rubiaceae), and Conchocarpus macrophyllus (Rutaceae). Additionally, species such as Croton warmingii (Euphorbiaceae), Dimorphandra jorgei (Fabaceae), and Heteropterys macrostachya (Malpighiaceae) demonstrate a disjunct distribution pattern between the Serra do Orobó Mountains and the semideciduous forests within the Atlantic Forest domain. Thus, kills theres Ed ME. D EN 4 +: 1 H qa ] laas A E the Serra do Orobó Mountains band the SDTF nuclei of canal d lóresis min e Aida Forest do- main, additional studies involving a larger number of areas also from the Caatinga domain would be useful in confirming some of the conclusions presented here. APPENDIX 1 Species list of the plants found in semideciduous, riparian, and submontane humid forests of the Serra do Orobó Mountains, Panta State, Braal, ae asterisk before the ecol names EE the endemies PUR Serra do Orobó t dry forest areas an real species of SDTF — (€ to Prado &G Gibbs (2002). Abbreviations for vegetation types: SE semidecidu- ous, RF: riparian, and ShF: submontane humid forests; abbreviations for collectors are: LP=L.P.de Queiroz; C=D.Cardoso; N=F.R.Nonato; H=R.M.Harley; B-H.P.Bautista; P=G.C.P.Pinto; L=].C.Lima; CO=T.A.B.Costa; RC=R.M.de Castro, and S-E.C.Smidt; abbreviations for life forms are: Sub=Subshrub; Epi=Epiphyte. la, Brazil 561 ANGIOSPERMS Acanthaceae Anisacanthus pide Lindau — LP 9383; Sub, SF S e i Ruellia affinis Lindau — LP 10771; C 296; Vine, ShF Ruellia bahiensis (Nees) Morong - C 381; Sub, SF Agavaceae Herreria salsaparilha Mart. — LP 12022; Vine, SF Amaranthaceae Alternanthera brasiliana (L.) Kuntze — LP 9538; Herb, RF Gomphrena demissa Mart. — LP 9801; Herb, SF Amaryllidaceae Hippeastrum stylosum Herb. — C 835; Herb, SF Anacardiaceae Anacardium occidentale L. — LP 9392, 9434; Tree, SF Spondias venulosa (Engl) Engl. — LP 9470, 12028; Tree, SF Tapirira quianensis Aubl. - LP 9778, 9802; C 343; Tree, RF, SF Annonaceae Oxandra reticulata Maas — LP 9788; Tree, SF Rollinia emarginata Schltdl. — LP 9840; Tree, SF Rollinia sylvatica (A.St-Hil) Mart. — LP 9967, 10663; C 1232; e Apocynaceae Asclepias curassavica L. — LP 9551; Sub, SF ° Aspidosperma discolor A.DC. — C 1274, Tree, ShF Aspidosperma parvifolium A.DC. — LP 9816, 12230; Tree, SF ? Aspidosperma pyrifolium Mart. — LP 9785; Tree, SF gh nitidum (Vell) J.F.Macbr. — LP 9876; C 356; Vine - Ti pm Turcz. - LP 9254, 10686; Vine, RF, SF Forsteronia sp. — LP 10830; Vine, SF Forsteronia glabrescens Müll.Arg. — C 1288; Vine, SF Forsteronia pubescens A.DC. — LP 9944; Vine, SF Gonolobus cordatus Malme — LP 10837; Vine, SF Macroditassa adnata (E.Fourn.) Malme — LP 9264; ME RF Mandevilla funiformis (Vell) K.Schum. - C 270; Vin EE scabra (Hoffmanns. ex Roem. & Su K Schum. 4; Vine, SA maritima (Jacq.) Woodson - LP 10643; Vine, SF Oxypetalum erostre E. raan - E 10863; C lom we oF = W e ER SIC 767: Vine, SF Peltastes peltatus (Vell) Woodson - LP 12084; C 828; Vine, Prestonia bahiensis Müll.Arg. — LP 9836; C 1293; Vine, SF Gare coalita (Vell) Woodson - L? 10689; C 378; Vine, SF volfia bahiensis A.DC. — LP 10804, 12247; Tree, insi multiflora Mart. — LP 10692; Vine, SF Secondatia floribunda A.DC. — LP 9896; Vine, SF Tabernaemontana solanifolia A.DC. — LP 9300, 9812; Tree, SF Temnadenia violacea (Vell) Miers — LP 9867; C 347; Vine, RF Araceae Anthurium pentaphyllum (Aubl) G.Don — LP 12051; Epi, RF Anthurium petrophilum K.Krause — C 272; Herb, SF Anthurium scandens (Aubl.) Engl. — C 1275, Epi, ShF Monstera adansonii var. klotzschiana (Schott) Madison — LP 12090, Epi, ShF Philodendron pedatum (Hook) Kunth — LP 9244, 9767; Epi, p Arecaceae Geonoma pauciflora Mart. - LP 9513; Shrub, RF Geonoma pohliana Mart. — LP 12254; Shrub, ShF Syagrus coronata (Mart) Becc. — LP 9846; Tree, SF Aristolochiacea Aristolochia ech Mart. €: Zucc. — LP 9447; C 798; Vine, SF Aristolochia gigantea Mart. € Zucc. - H 53484; LP 10668; C 6; Vine, Asteraceae Asteraceae sp. — LP 9940; Shrub, SF Acanthospermum australe (Loefl.) Kuntze — C 792; Herb, SF Achyrocline satureoides (Lam.) DC. — LP 10752; Sub, SF Acritopappus confertus (Gardner) R.M.King & H.Rob. — LP 12249; Tree, ShF Baccharis calvescens DC. — H 55454; Shrub, SF Chaptalia integerrima (Vell) Burkart — LP 9857; C 358; Herb, SF Chromolaena cf. laevigata (Lam.) R.M.King €: H.Rob. — LP 10678; Shrub, SF Chromolaena morii R.M.King & H.Rob. — LP 10770; Shrub, SF Cyrtocymura harleyi (H.Rob.) H.Rob. — H 55455; P 389; Shrub, SF Cyrtocymura scorpioides (Lam.) H.Rob. — H 53480; LP 10709; Delilia biflora (L.) Kuntze — LP 10737; Herb, SF Elephantopus sp. — LP 9543; Herb, RF Emilia fosbergii Nicolson — C 1284; Herb, SF Erechtites hieracifolia Raf. ex DC. — LP 12239, Sub, SF Eremanthus capitatus (Spreng. MacLeish — LP 12248; Tree, SF Lepidaploa cotoneaster (Willd. ex Spreng.) H.Rob. — LP 9427; Sub, RF Mikania cordifolia (Lf) Willd. — LP 10816; Vine, ShF Mikania elli a s LP 9508; Vine R Mikania obov .— C 1278; Kees SC (Cass.) S.FBlake — LP 9294; Sub, SF Vernonanthura brasiliana (L.) H.Rob. — LP 9554; C 805; Shrub, Trixis antimenorrhoea (Schrank) Kuntze — LP 12278; Herb, SF Begon Begonia jai Brade - LP 9403, 10787; Herb, ShF, SF Begonia lobata Schott — LP 9925; C 247; Shrub, R See petasitifolia Brade — LP 9768, 10751; i RF, ShF Begonia vitifolia Schott — LP 10790; Shrub, ShF Bignoniaceae proc sp. - LP 12024; Vine, SF Anemopaegma laeve DC. — LP 10832; Vine, Pies a een (DC) Bureau — LP 10677; Vine, SF EEN anera Poepp:= LP 9253; C 278; Vine, RF, SF DC.) Baill. ex Bureau - C 1241; Vine, T 562 Arrabidaea lasiantha Bureau & K. Schum. [= Cuspidaria lasian- reau & K Schum) AH. Gentry comb. ined.] - C 280; Vine, SF Cybistax antisyphilitica (Mart.) Mart. — LP 9475; Tree, SF Fridericia speciosa Mart. — C 277; CO 17; Vine, S Handroanthus heptaphyllus (Vell) Mattos — LP 12038; Tree, SF Handroanthus ochraceus (Cham.) Mattos — LP 9969, 9998, 12016; Tree, SF TURA irwinii A. j ipai m : cu Shrub, RF ith —LP 10724; Shrub, SF Lundia cordata (Vell) A.DC. — LP 10773; C 812; Vine, ShF, SF E valida UN eg o B 1236; Ms SF 27 6,1 1282 3 ne SF GE EE we) enon goë 2531 C odi vine; BE LIM vv -= LP 9799, 9901; Vine, SF Tynanthus sp. — LP 10681; C 357; Vine, SF Tynanthus labiatus Cham) Miers - LP 10661; Vine, SF Boraginaceae * Cordia blanchetii DC. - LP 9448; € 770; Tree, SF Cordia curassavica (Jacq) Roem. & Schult. - H 53485; LP 9939; S Cordia superba Cham. — LP 9932; C 383, 1254; Tree, SF Heliotropium angiospermum Murray — LP 9979; Sub, SF Heliotropium procumbens Mill. — LP 12280; Sub, SF Heliotropium transalpinum Vell. - C 367; Sub, SF Tournefortia rubicunda Salzm. ex DC. — LP 9978; Shrub, SF Brassicaceae Capparis brasiliana DC. — LP 9843, 12079; Shrub, SF ° Capparis flexuosa (L.) L. — LP 9930; C 302; Vine, SF Capparis jacobinae Moric. ex Eichler — LP 9968; Shrub, SF Capparis lineata Pers. — H 53483; LP 9934; C 271; Vine, SF ° Crataeva tapia L. — LP 12056; Tree, SF Bromeliaceae Aechmea aquilega (Salisb.) Griseb. — LP 12055; Herb, SF Aechmea bromeliifolia (Rudge) Baker — C 295; Herb, ShF Aechmea nudicaulis (L.) Griseb. — C 327; Epi, RF Billbergia porteana Brongn. ex Beer — LP 9790; Epi, SF romelia pinguin L.— Not collected, Herb, RF Catopsis sessiliflora (Ruiz €: Pav.) Mez — LP 10820; C 829; Epi, ShF Cryptanthus bahianus L.B.Sm. — LP 9793; Herb, SF Tillandsia polystachia (L) L. — LP 10848; Epi, SF Tillandsia stricta Sol. ex Sims — LP 10774, ee = ShF Tillandsia usneoides (L.) L. - Not collected, E Vriesea procera (Mart. ex Schult.f) Wittm. — d e Herb, hF Vriesea simplex (Vell.) Beer — C 288, Epi, ShF urmanniaceae B i Apteria aphylla (Nutt) Barnhart ex Small — LP 10616; C 344; - Gymnosiphon divaricatum (Bentn.) Benth. & Hook.f. — LP 10617; | £a Dos . In LI Par oe a ET Sif), re Cactaceae Brasiliopuntia brasiliensis (Willd.) Haw. — LP 9469; Tree, SF Epiphyllum phyllanthus (L) Haw. — C 332; Epi, SF Pereskia aculeata Mill. — LP 10697; Tree, SF Campanulaceae Siphocampylus imbricatus G.Don — LP 12257, Shrub, ShF Cannabaceae ° Celtis iguanaea (Jacq) Sarg. — LP 9297, 9459; Tree, SF Trema micrantha (L.) Blume — LP 12089; Tree, SF Cannaceae Canna glauca L. — C 814; Herb, RF Celastraceae Hippocratea volubilis L. — LP 9556; Vine, RF Maytenus erythroxylon Reissek — LP 10654; C 784; Tree, SF is eee (Schrad.) Loes. — LP 12042, 12113; ms Ve (A.C.Sm) A.CSm. — LP 9890, 12065; E Chrysobalanacea Hirtella glandulosa — — LP 9539, 9776; Tree, RF, SF Hirtella racemosa Lam. — LP 9544; C 315; Shrub, RF, SF Clusiaceae Clusi G.Mey. — LP 9868; C 808, 827; Tree, RF, ShF Combretaceae Buchenavia capitata (Vahl) Eichler — LP 9780, 12252; Tree, RF, ShF Commelinacea Commelina benghalensis L. — E 348; Herb, RF * Dichorisandra sp. nov. — LP 12115; C 305; Herb, SF Gibasis geniculata (Jacq.) Rohweder — LP 10797; Herb, ShF Connaraceae Connarus detersus Planch. — C 298; Tree, ShF Rourea sp. — LP 12004; Shrub, Rourea doniana Baker — LP 12109; C 291; Shrub, SF Rourea martiana Baker — LP 9894; C 242; Vine, SF Convolvulaceae Evolvulus glomeratus Nees & Mart. - LP 9910; C 286; Sub, SF Evolvulus latifolius Ker Gawl. - N 1027; C 331, 1218; Sub, SF moea 40; C 766; Vine, SF Ipomoea hederifolia L. — LP 10736; Vine, SF Ipomoea wrightii A.Gray — LP 12283; Vine, SF Jacquemontia confusa Meisn. — C 384; Vine, SF Jacquemontia densiflora (Meisn.) Hallier f. — C 9483; Vine, SF Jacquemontia martii Meisn. — LP 9985; C 389; Vine, SF Jacquemontia montana (Moric.) Meisn. — C 319; Vine, RF Merremia cissoids (Lam.) Hallier f. — LP 10639; C 316; Vine, SF Costaceae Costus spiralis (Jacq) Roscoe — LP 9871, 9775; Herb, RF, ShF Cucurbitaceae Anisosperma sp. — LP 12111; Vine, SF Apodanthera sp. — C 380; Vine, SF Apodanthera glaziovii Cogn. — LP 12064; Vine, SF Cayaponia cf. petiolulata Cogn. — LP12069; Vine, SF E Eu DL! , Brazil 563 d sp. - LP 12086; Vine, SF nia subumbellata (Miq.) Cogn. - C 1247; Vine, SF "ia triphylla (Miq.) CJeffrey — LP 10700; Vine, SF Cunoniaceae Lamanonia ternata Vell. — LP 9270, 12092; Tree, RF, SF Cyperaceae Cyperus cf. compressus L. — C 1255; E SE Cyperus laxus Lam. — C 372; Herb, S Rhynchospora E EE ( e LP 9430, 10826; Herb, RE SF Rhynchospora cf. exaltata Kunth — Si RER Herb, HE Scleria bracteata Cav. — H 55452; Herb, H Scleria latifolia Sw. — H 55451; LP SS - RF Dilleniaceae Doliocarpus sp. — LP 9537; Vine, RF Dioscoreacea Dioscorea sp. — ie 9958; Vine, SF Dioscorea altissima Lam. — LP 12088; Vine, SF Dioscorea dodecaneura Vell. — LP 10711, 12279; Vine, SF Ebenaceae spyros inconstans Jacq. - LP 10007; C 256; Shrub, SF Diospyros sericea A.DC. — LP 9247, 10849; Tree, RF, SF Y S Elaeocarpacea Sloanea gege (Aubl) Benth. — LP 9504, 9782; Tree, RF Eriocaulaceae aepalanthus spathulatus Korn. — LP 10768; Herb, ShF Tonina fluviatilis Aubl. — LP 10754; Herb, RF Erythroxylaceae án macrocalyx Mart. — LP 9830, 9902, 12005; Tree, F, SF rom pe Mart. — LP 9936, 10720; C 355; Shrub, R zu phorbiaceae (NACH Arq.) NAIL Arm Shrub, SF par É Acalypha brasiliensis Müll.Arg. — LP 10006, 10683, 10799; Shrub, SAF, SF Actinostemon conco! for (Spreng.) Müll.Arg. — LP 9952; Shrub, SF Actinostemon verticillatus (Klotzsch) Baill. — B 1239; Tree, SF Alchornea triplinervia (Spreng.) Müll.Arg. — LP 10803; Tree, h ShF Bernardia tamanduana (Baill) Müll.Arg. — LP 12047; C 1215; Shrub, SF Cnidoscolus oligandrus (Müll.Arg.) Pax — LP 9451, 9408; Iree, SF Cnidoscolus urens (L.) Arthur — LP 9995; Shrub, SF Croton betulaster Müll.Arg. — LP 9265; Shrub, RF Croton cordiifolius Baill. - LP 12041; C 262; Shrub, SF Croton echioides Müll.Arg. — LP 9961; Shrub, SF Croton hirtus UHér. - C 1256; Sub, SF Croton lobatus L. — LP 9926; C 801; Shrub, RF, SF Croton sincorensis Mart. ex Müll.Arg. — LP 9895; Shrub, SF Croton tetradenius Baill. — LP 9295, 10662; Shrub, RF, SF Croton triqueter Lam. — C 363; Sub, SF Croton urticifolius Lam. — C 1222; Sub, SF Croton velutinus Baill. — LP 9271; Shrub, RF Croton warmingii Müll.Arg. — LP 10005; Shrub, SF Dalechampia brasiliensis e LP 9986; Vine, SF Dalechampia cf. pentaphylla Lam. — LP 9402; Vine, SF Dalechampia scandens Vell. — LP 9903; Vine, SF ae sylvatica S.Moore — LP 10674; C 802; Vine, > Euphorbia comosa Vell. — LP 10806; C 1267; Herb, ShF, S Euphorbia hyssopifolia L. — LP 9717 , Herb, SF Euphorbia insulana Vell. — LP 10796; C 1285; Herb, ShE SF Jatropha palmatifolia Ule — LP 9933; Shrub, SF Maprounea guianensis Aubl. — LP 9761, 9869, 12044; Tree, Manihot sp. — C 310; Tree, SF Manihot cf. anomala Pohl — LP 9904; C 385, 782; Tree, SF Manihot cf. epruinosa Pax €: K. Hoffm. — 387; Tree, SF Pachystroma longifolium (Nees) LM. Johnst. — LP 10659; F ree, Pera glabrata (Schott) Poepp. ex Baill. — LP 12243; C 337, 342; e, RE, ShF Philyra brasiliensis Klotzsch — L 259; Tree, SF Richeria grandis Vahl — LP 10028; Tree, RF — corniculata (Vahl) Müll.Arg. — LP 10632; Shrub, d lessertiana (Baill) Müll.Arg. — LP 9941, 12097, 12792; ine, ShF, SF Tragia volubilis L. — LP 10002; Vine, SF Fabaceae-Caesalpinioideae Apuleia leiocarpa (Vogel) J.F.Macbr. - LP 12063; C 809; Tree, SF Bauhinia sp.1 — LP 9817; Shrub, SF Bauhinia sp.2 — LP 12043; Vine, SF Bauhinia forficata Link — LP 9815; Tree, SF Bauhinia longifolia (Bong.) Steud. — LP 10753; C 364; Tree, SF Bauhinia maximilianii Benth. — C 787; Vine, SF Caesalpinia aff. echinata Lam. — LP 9405; Tree, S Caesalpinia leiostachya (Benth.) Ducke — C 1250; Tree, SF Caesalpinia pluviosa DC. — LP 12264; Tree, SF Cassia ferruginea (Schrad.) Schrad. ex DC. var. ferruginea — C Ci OMM losa (Benth) HSA — LP 9278; Sub, RF Chamaecrista olaa var. grandiflora (Benth.) H.S.Irwin & by — 3,6236; Shrub; SF Copaifera cearensis os Ducke — LP 9586; Tree, SF Copaifera langsdorffii Desf. — LP 10625, 10756; De RE oF Dimorphandra jorgei M.FSilva — C 834; Tree, Goniorrhachis marginata Taub. — LP Se E Sr Tree, SF ° Hymenaea courbaril L. — C 768; Tre Melanoxylon brauna Schott — LP EC Tree, SF Peltogyne confertiflora (Hayne) Benth. — LP 10779, 10831; im 8 Darm alb: P ee, ShF, Peltogyne aff. recifensis Ducke — LP 12013; Tree, SF ° Peltophorum dubium (Spreng.) Taub. var. dubium — LP 9258; C 306, 243; Tree, RF, SF * Phanera trichosepala L.PQueiroz — LP 9413; C 281; Vine, SF ° Poeppigia procera var. conferta Benth. — LP 9914; C 257; Tree, SF 564 ° Pterogyne nitens Tul. - H 53478; LP 10016; C 1239; Tree, SF Schizolobium parahyba (Vell) S.F.Blake — Not collected, Tree, F C ] di (sro UI €] D Darnr nal LP 9251, 9824; Shru Senna acutisepala pn H.S.Irwin €: Barneby - C 1270; ree Senna aversiflora (Herb.) H.S.Irwin & Barneby — LP 10739, Senna hirsuta (L.) H.S.Irwin € Barneby — LP 10672, 10695; Shrub, Senna macranthera var. micans (Nees) H.S.Irwin & Barneby — LP 9292; Tree, SF LN ffe = 1 D Darneal; | mw Ki d = 9555; Tree, SF Senna pendula (Willd.) H.S.Irwin €: Barneby — LP 9453, 9980; Sh ° Senna spectabilis var. excelsa (Schrad.) H.S.Irwin €: Barneby - C 360; Tree, SF EE eier Killip ex 4; Tree, SF Blencherodendron blancheti (Benth) pum. & fad Inga mre SE - LP 9302; C 374; Tree, SF Inga aff. laurina Desv. — LP 9990; C 807; Tree, SF Inga subnuda Salzm. ex Benth. subsp. subnuda — C 289; Tree, SF Inga thibaudiana DC. subsp. thibaudiana — LP 10029; C 252; Tree, Mimosa invisa Mart. ex Colla — LP 9277; Shrub, RF Mimosa tenuiflora (Wild) Poir. — LP 12242; Tree, SF Piptadenia sp. — LP 9255, 12007; Vine, SF Piptadenia adiantoides (Spreng.) Macbr. — LP 9825, 10713; ine s paniculata Benth. — H 53481; LP 10682; s a pta tadenia bahiana G.P.Lewis & M.PLima ; egen contorta (DC.) G.PLewis €: M.P.Lima — LP ree, SE esu (Harms) Barneby & J.G.Grimes — LP 9473; Senegalia sp.1 — LP 12068; Tree, SF Senegalia sp.2 — LP 10657; Vine, SF Senegalia martiusiana (Steud.) Seigler & Ebinger — LP 10729; C 390; Shrub, SF Senegalia riparia (Kunth) Britton & Killip - C 351; Tree, SF Senegalia velutina (DC) Seigler & Ebinger — C 382; Shrub, SF Fabaceae-Papilionoideae Aeschynomene elegans Schul. & Cham. var. elegans — LP 1 ; Sub, SF Aeschynomene histrix var. densiflora (Benth.) Rudd — LP 9293; UD, Andira fraxinifolia Benth. — LP 9284; Tree, RF Bionia coriacea (Ness & Mart.) Benth. — LP 9272; Shrub, RF Bowdichia virgilioides Kunth — LP 9781; Tree, RF Canavalia parviflora Benth. — LP 12275; Vine, SF 1 £ al Centrolobium tomentosum Guill. ex Benth. - H 53496; LP 9828; Tree, SF g (Humb. & Bonpl. ex Willd.) B | — LP 9441, 10013; Vine, SF Centrosema virginianum (L) Benth. — LP 9974; Vine, SF Chaetocalyx scandens var. pubescens (DC) Rudd. - LP 9454; a Vine, SF Clitoria falcata Lam. — LP 9463; Vine, SF Clitoria laurifolia Poir. — " e Sub, RF ratylia sp. nov. — LP 10656; CO 15; Vine, SF Crotalaria holosericea a & Mart. — LP 9426; C 322; Sub, SF Crotalaria micans Link. — LP 10651; Sub, SF Dalbergia decipularis Rizz. & Mattos — LP 12029; Tree, SF Desmodium barbatum (3 Benth. — C 800; Herb, SF Desmodium incanum (Sw.) DC. - LP 10645; Herb, SF Desmodium tortuosum (Sw. DC. — LP 10644; Herb, SF Desmodium uncinatum (Jacq) DC. -C 1248, 1271; Herb, SF Dioclea violacea Mart. ex Benth. — C 352; Vine, Galactia striata (Jacq.) Urb. - LP 10687; C 359; Vine, SF o EE pn - nei o. i PA o & H.C.Lima - LP 10010; Tree, SF ° Machaerium acutifolium Vogel — LP 9433, 10000; C 290; Tree, RF, 5 Macherium aff. gracile Benth. — LP 12020; Tree, gé Machaerium hirtum (Vell) Stellfeld — C 287; Tre Macroptilium erythroloma (Mart. ex Benth.) in - im 10640; ; Vine, Myrocarpus fastigiatus Allemáo — LP 12017; C 1230; Tree, SF Periandra coccinea (Schrad.) Benth. — LP 9993; C 249; Vine, mio, a D H j Ls Ce Sp E ‘LJ \ in J —LP 9920; C 1251; Tree, SF ° Platypodium elegans Vogel — LP 10017, 10667; Tree, SF Poiretia punctata Desv. — LP 10650; C 778; Vine, SF Rhynchosia edulis Griseb. — LP 10641; Vine, SF Rhynchosia melanocarpa Grear — LP 10671; Vine, SF Rhynchosia phaseoloides (Sw.) DC. — LP 9924; Vine, RF Rhynchosia reticulata var. kuntzei (Harms ex Kuntze) Grear -C — 15; Vine, SF Stylosanthes scabra Vogel — LP 10647; Herb, SF Swartzia acutifolia Vogel — LP 9414, 9948; Tree, SF Swartzia apetala Raddi var. apetala — LP 10015, 12073; Tree, SF Vigna sp. — LP 10749, Vine, SAF Vigna candida (Vell) Maréchal, Mascherpa €: Stainier — LP 72; Vine, SF 2 Zollernia ilicifolia (Brongn.) Vogel - LP 12030; Tree, SF Gentianace Irlbachia nen (Aubl) Maas — LP 10634; Sub, RF Voyria aphylla (Jacq) Pers. — LP 10618; C 345; Herb, RF Gesneria Saas albus Chautems — LP 10782, Epi, 4 * Sinningia sp. nov. — LP 12277, DC 2066, Her e barbata (Nees & Mart.) G. cud ~ LP 10795, erb, ShF r J J a. e E. an É A E duc om Le di 1 Brazil 565 Heliconiaceae Heliconia pendula Wawra — C 1272; Shrub, ShF Humiriaceae Vantanea compacta (Schnizl.) Cuatrec. — LP 9921; C 361; Tree, RF lridaceae Trimezia spathata subsp. sincorana (Rav.) Chukr — LP 9771; Herb, RF Juncaceae Juncus microcephalus H.BK. — LP 10757; Herb, SF Lamiaceae Hyptis pectinata (L.) Poit. — LP 10765; Herb, SF Hyptis sidifolia (U'Herit.) Brig. — LP 10003; Sub, SF Hyptis lici (L) Poit. - LP 10638; C 362; Herb, SF m echianum Mill. — H 28446; LP 9997; Herb, SF Salvia H dsl Benth. - H 53497; LP 10792; Sub, ShF, SF Vitex aff. klugii Moldenke - LP 9946, 12015; C 283, 326; Tree, RE SAF SF Lauraceae Cinnamomum sp. — LP los SF Nectandra sp. — LP 12233; Tre Nectandra cf. membranacea un H 53500; LP 10684; C 365; Iree, RF, SF Ocotea glomerata (Nees) Mez — LP 12059; C 795; Tree, SF Ocotea velutina (Ness) Rohwer — LP 10635; C 1292, 1290; e, RES Lecythidaceae Cariniana estrellensis (Raddi) Kuntze — LP 10008; Tree, SF Loganiace Spigelia Ger L. — LP 9774; Herb, RF Spigelia ~ Cham. €: Schltdl. - LP 10750, 12081; Herb, ShF, Stryc. 2 5 sp. — LP 9965; C 388; Vine, SF Strychnos brasiliensis Mart. — LP 9841; C 361; Vine, SF anthaceae Struthanthus polyrrhizus Mart. - LP 10685; Epi, SF Lythraceae Cuphea circaeoides Sm. ex Sims — LP 12282; Herb, SF Cuphea impatientifolia A.St.-Hil.— LP 12226; Herb, SF Cuphea racemosa (Lf) Spreng. — LP 9560; C 813; Herb, RF, SF Pleurophora anomala (A.St-Hil.) Koehne — LP 12281; Sub, SF Malpighiaceae Bunchosia fluminensis SCH LP 9912, 12049; Shrub, SF Byrsonima sericea 7, 9994/ 9385; Tree, RF, SF Dicella bracteosa (A Juss) Gin LP A C 2/5; s SF 994 rlearcmm -a Vine E Heteropterys macrostachya A.Juss. — LP 10780; Vine, ShF, SF Heteropterys perplexa W. R. Anderson - C 336; Vine Mascagnia sp. - C 811; V Mascagnia chasei WR. HEN LP 9390; Vine, SF Mascagnia rigida Griseb. — LP 10655; Vine, SF Mascagnia sepium (A.Juss.) Griseb. — LP 9823; Vine, SF Stigmaphyllon blanchetii C.E.Anderson — LP 9423, 9827; Vine, F RF, dbi sp. ; — LP 9947; Vine, SF Tetrapterys sp.2 — CO 16; Vine, SF odi Mua Mart. - C 235; Vine, SF Malvaceae Malvaceae sp. - LP 10840; C 779; Tree, SF Bombacopsis stenopetala (Casar.) A.Robyns — LP 9291, 10819; Mop Cavanillesia arborea (Willd.) K a — C 2295; Tree, SF Corchorus hirtus L. - LP 1064 , SF Eriotheca cf. globosa A. ie — LP 10030, 10776; Tree, RF, ShF Eriotheca macrophylla (K.Schum.) A.Robyns - C 1237; Tree, SE Guazuma ulmifolia Lam. — LP 9467; C 265; Tree, SF Helicteres macropetala A.St-Hil. - LP 10665; Shrub, SF Luehea candicans Mart. — LP 9458, 9290; Tree, SF Melochia betonicifolia A.St.-Hil. — LP C 366; Shrub, SF Melochia tomentosa L. — LP 9450; Sub, S Pavonia malacophylla (Link & Otto) Sen LP 9509; Shrub, RF Pavonia martii Colla — LP 9853; C 1286; Sub, SF Sidastrum micranthum (A.St.-Hil.) Fryxell - LP 10642; Sub, SF Waltheria indica L. — LP 9245; Sub, RF Wissadula amplissima (L.) R.E.Fr. — LP 9443; C 771; Sub, SF Wissadula contracta (Link) R.E.Fr. — LP 12238; Sub, SF Maran pa sp. — Gm 9929; Herb, RF Maranta bicolor Ker Gawl. - LP 10698; Herb, SF Maranta divaricata Roscoe — LP 9557; B 1260; Herb, RF, SF Marcgraviaceae Schwarizia brasiliensis (Choisy) Giraldo-Cañas — LP 12053; Melastomataceae Clidemia hirta (L) D.Don - LP 10619; Shrub, RF Henriettea succosa (Aubl.) DC. — C 300; Tree, RF Leandra aurea (Cham.) Cogn. - LP 12093; Shrub, ShF Miconia sp. — LP 9766; Shrub, RF Miconia albicans (Sw. Triana — LP 9866; es RF Miconia aff. caudigera DC. — LP 9536; Tre Miconia ciliata (Rich.) DC. — LP 9866; T E RF Miconia mirabilis (Aubl) L.O.Williams — LP 10628; C 246; ree, Miconia rimalis Naudin — LP 9533; Tree, RF Pterolepis glomerata (Rottb) Miq. — LP 9274, 10747; Sub, RF Meliaceae Trichilia sp. — LP 12046; Tree, SF Trichilia emarginata (Turcz.) C.DC. — LP 9964; Tree, SF Trichilia hirta L. — LP 12037; Tree, SF oraceae Brosimum gaudichaudii Trécul — LP 9962; C 275; Tree, SF Brosimum guianense (Aubl) Huber — LP 12062; C 810; Tree, SF * Dorstenia caatingae R.M. Castro ~ LP 10701; Herb, SF 566 Ficus citrifolia Mill. — LP 9468, 10813; C 349; Tree, RF, ShF, SF Ge cyclophylla (Mig) Mig. - LP 12071; C 308; Tree, SF icus ire Kunth & C.D.Bouché - LP 12078; C 354; Tree - F, SAF mi pulchella Schott ex Spreng. — LP 12058; C 978; Tree, SP Maclura tinctoria (L) D.Don ex Steud. - LP 9982; Tree, SF Sorocea hilarii Gaudich. - LP 12057; Tree, SF Myrsinac ra sp. - C 1276; Tree, ShF Myrsine ferruginea (Ruiz & Pav.) Spreng. — LP 10759; C 1246; Tree SF Myrsine guianensis (Aubl) Kuntze — LP 9286; C 350; Tree, RF Myrsine venosa A.DC. — LP 10764, 12245; Tree, SF Myrtaceae Eugenia sp.1 -C 772, 777; Shrub, SF Fugenia sp.2 — LP 9991; Shrub, SF Eugenia sp.3 - LP 12045; Tree, SF * Eugenia sp. nov. - LP 10727; C 393, 1487; Tree, SF Eugenia cf. candolleana SCH Gë e SF Eugenia florida DC. — C 204 Eugenia punicifolia en SR - e 9283, 9796; Shrub, RF, SF Myrcia sp.1 — C 245, 391; Tree, SF Myrcia sp.2 — C 341; Shrub, SF yrcia sp.3 — LP 9949; Shrub, SF yrcia blanchetiana (O.Berg.) Mattos — LP 12104; Tree, ShF yrcia dis (Aubl) DC. — LP 9959, 12006; Tree, RF, SF yrcia rostrata DC. — LP 9762; Tree, RF rcia E (Sw.) DC. - LP 12009, 12074; Shrub, RF, ShF yrciaria floribunda (H West ex Willd.) O.Berg - LP 10703; € 774; Tree, SF Psidium sp. — LP 12026; Shrub, SF Psidium cf. brownianum DC. - LP 12033, 12096; Tree, ShF, SF Psidium schenckianum Kiaersk. — C 330; Shrub, SF = 35.235 = Nyctaginaceae Bougainvillea spectabilis Willd. — C 263; Vine, SF Guapira hirsuta (Choisy) Lundell — LP 9972; Tree, SF Guapira opposita (Vell.) Reitz — LP 10009, 12080; C 1279; Tree, ShF, SF Ochnaceae Sauvagesia erecta L. - LP 10627; Herb, RF Olacaceae Heisteria blanchetiana (Engler) S'eumer — LP 12098; Tree, ShF Schoepfia brasiliensis AIDC. — LP 9542, 10769; Tree, RF, ShF Ximenia americana L. var. americana — LP 10775; C 266, 1238; Tree, ShF, SF Orchidaceae Bulbophyllum sanderianum Rolfe — C 312; Epi, R Campylocentrum micranthum (Lindl) Rolfe — G 9535; Epi, RF Cleistes metallina (Barb. Rodr) Schltr. - LP 10630; Herb, RF Cleistes pluriflora (Barb. Rodr.) Schltr. — LP 12256; Herb, ShF Dichaea cogniauxiana Schltr. — C 1269; Herb, ShF Eltroplectris calcarata (Sw.) Garay €: H.R.Sweet — LP 9424; H Eltroplectris triloba (Lindl.) Pabst — LP 9424; Herb, RF Furystylis actinosophila (Barb. Rodr.) Schltr. - LP 10823; Epi, Shr Galeandra beyrichii Rchb. f. - H 53499; LP 9506; Herb, RF, SF Maxillaria sp. — C 1297; Epi, 5 Notylia hemiticha Barb. Rodr. — LP 9464; Epi, SF Oeceoclades maculata (Lindl) Lindl. - LP 9438; Herb, RF Oncidium ciliatum Lindl. — LP 10798; Epi, ShF Oncidium flexuosum (Kunth) Lindl. — LP 10785, Epi, ShF Pogoniopsis schenkii Cogn. — LP 12255, Herb, ShF Polystachya estrellensis Rchb.f. — LP 10808; Epi, ShF Prescottia oligantha Lindl. — LP 9534; Herb, RF Prosthechea aemula (Lindl.) W.E.Higgins — C 1295; Epi, SF Psilochilus cf. us Sie Rodr. — LP 9507; Herb, RF Rauniella sp. — S 674 Scaphyglottis mds a ce — LP 10818; Epi, ShF Sobralia sessilis icu H 55461; LP 10791; Herb, ShF - LP 10812; Epi, ShF a SE Benth. — LP 10767; Herb, ShF Oxalidaceae Oxalis barrelieri L. — H 53498; LP 12225; Sub, SF Oxalis neuwiedii Zucc. — LP 9988; C 1262; Herb, SF Oxalis roselata A.St-Hil. - LP 10699; Herb, SF -—— Passifloracea Passiflora alata bande — LP 10622; Vine, RF Passiflora bahiensis Klotzsch — LP 10675, 12235; Vine, SF Passiflora cincinnata Mast. — C 314, 1268; Vine, S Passiflora edulis Sims — LP 9875; Vine, SF Passiflora galbana Mast. — LP 9305, 10623; C 1281; Vine, RF, Passiflora organensis Gardner — LP 9288; Vine, SF Passiflora setacea DC. - LP 10011, 9907; Vine, SF Passiflora suberosa L. - C 1244; Vine, SF Phyllanthace * Astrocasia a (Müll.Arg.) G.L Webster - C 1235; Pisanus sp. - LP 12117; Herb, SF Phyllanthus niruri L. — C 1259; Herb, SF Phytolaccaceae Gallesia integrifolia (Spreng.) Harms — C 1257; Tree, SF Microtea paniculata Moq. — LP 12083; Herb, ShF, SF Rivina humilis L. — LP 10648; Shrub, SF Seguieria floribunda Benth. — LP 12261; Vine, SF Piperaceae Peperomia sp. — C 294; Epi, ShF Peperomia cf. glabella (Sw.) A.Dietr. — LP 9460, 10708; Herb, SF Piper aduncum L. - LP 9559; Shrub, RF Piper amalago L. — LP 10004; Shrub, SF Piper arboreum Aubl. var. arboreum — LP 9241, 9466; C 793; Shrub, RF, SF Plumbaginac Plumbago bii L. — LP 12266, Herb, SF ia, Brazil 567 Poaceae Poaceae sp. — LP 9243; Herb, RF emana SEET MU Sdn — E SC p E hn, arctr 2 Herb, ShF Ichnanthus pallens (Sw.) Munro ex Benth. — LP 9783; Herb, RF Olyra latifolia L. — LP 12231; Herb, S GC Ded (Kunth) Ee & Zuloaga — LP 9429, er cron oiK S DC 2063; Herb, SF Streptostachys asperifolia Desv. — LP 12019; Herb, SF Polygalac Bredem ann ne AW.Benn. — LP 10844; Vine, SF a mollis Kunth — LP 9540; Herb, RF Polyga sae lice ea H 55457; Herb, urbani Chodat - LP diee 9975; Sub, RF, SF cea Aubl. — LP 9897; Sub, SF aa dives » o LP 9842, 9893; Vine, SF Securidaca lanceolata A.St.-Hil. & Moq. — LP 9870; Vine, RF Polygonaceae Coccoloba alnifolia Casar. - LP 1071; C 1260; Tree, SF Coccoloba declinata (Vell) Mart. — LP 12031; C 1253; Vine, SF Portulacaceae Talinum t tum (Jaca) Gaertn. - LP 9461; Herb, SF Talinum triangulare (Jacq.) Willd. — C 1266, Herb, SF Proteaceae Roupala montana Aubl. Rafflesiaceae Pilostyles blanchetii (Gardner) R.Br. — — LP 9514, 9923; Tree, RF, ShF H 53495; Fpi, SF Rhamnaceae ° Ziziphus joazeiro Mart. — LP 9831; Tree, SF Rubiaceae ° Alseis floribunda Schott — LP 9905; Tree, SF Amaioua guianensis Aubl. - C 323, 1164; Shrub, RF, SF Borreria latifolia (Aubl) K.Schum. — LP 10744; Herb, SF Chiococca alba (L) Hitchc. — H 53493; LP 12087; Shrub, SF melia sp. — LP 12114; Tree, SF Ge cypselum sp. — LP 10629; C 251; Herb, RF ° Coutarea hexandra Jaca.) K.Schum. — LP 10817; C 1165; , ShF, SF Decio fruticosa (Willd. ex Roem. & Schult.) Kuntze — LP ; C 248; Herb, RF, SF Dede Mn (Willd. ex Roem. & Schult) Bacigalupo & bral - LP 10741; Sub, SF aere umbellata (Spreng.) K.Schum. — LP 10846; Vine, SF Faramea hyacinthina Mart, — LP 9956; Shrub, SF Hamelia patens Jacq. — LP 10021; C 373; Shrub, SF Manettia cordifolia Mart. - LP 10676; Vine, SF Mitracarpus frigidus KSchum. — LP 10761; Sub, SF Mitracarpus hirtus (L) DC. — LP 10742; Herb, SF Molopanthera paniculata Turcz. — LF 9285; Tree, SF Palicourea blanchetiana Schltdl. - LP 9864, 9760, 10621; Shrub, RF Margaritopsis carrascoana (Delprete & E.B.Souza) C.M.Taylor & E.B.Souza - LP 9953; Shrub, SF Psychotria Ee — LP 9246, 10614; Shrub, RF Psychotria chaenotricha DC. — LP 12034; C 264, 261; Shrub, SF Psychotria hoffi ggi (Willd. ex Roem. & Schult.) Müll .—LP 10633; Shrub, RF Randia armata (Sw) DC. - LP 10704; Tree, SF * Spermacoce sp. nov. - LP 10828; C 1294; Herb, SF * Standleya sp. nov. — LP 10835; C 764; Herb, SF Rutaceae Almeidea rubra A. St.-Hil. - LP 10706; C 303; B 1261; Shrub, SF za ih! 2 KAart\) Mallt inks d / 9 LP 9844, 10653; Shrub, SF * Conchocarpus sp. nov. — LP 9845, 10845; C 392, Shrub, SF sis e (ASt.-Hil) Kallunki € Pirani — H ; C 769; Shru Con pis Deo s ea eee -C 786; Shrub, SF Dictyoloma vandellianum A.Juss. — LP 9547; Tree, RE, SF Esenbeckia grandiflora Mart. subsp. grandiflora — C 785; Tree, Esenbeckia leiocarpa Engl. — C 780; Tree, SF Helietta glaziovii (Engl.) Pirani — LP 12052; Tree, SF Pilocarpus riedelianus Engl. — LP 9791, 12067; Shrub, SF rcs spicatus A.St-Hil. subsp. spicatus — LP 10658; C 836 Zan thoxylum acuminatum (Sw.) Sw. — C 267, 783; Tree, SF Zanthoxylum cf. nigrum Mart. — C 1233; Tree Zanthoxylum rhoifolium Lam. — LP 10624; E RF Zanthoxylum tingoassuiba A.St.-Hil. — C 1486; Tree, SF Salicaceae Casearia arborea (Rich) Urb. - H 53489; Tree, 5 Casearia javitensis Kunth — LP 9891; C 334; zm RF, SF Casearia sylvestris Sw. — LP 12110; C 776; Tree, SF Sapindaceae Allophylus cf. laevigatus Radlk. — LP 12050; C 307, 1234; hrub, SF iiie sericeus (Cambess.) Radlk. — H 53488; LP 12070, e, SF mic gardnerianum Baill. — LP 9809; Tree, SF Cardiospermum a var. elongatum (Radlk.) F.A.Barkley - LP 10020; Vin Cupania rigida ae LP 9455, 9835, 10841; Tree, RF, SF Paullinia elegans Cambess. — LP 10670; Vine, SF Paullinia racemosa Wawra — LP 10842; Vine, SF Paullinia revoluta Radlk. — C 773; Vine, SF Serjania sp. — LP 9386; C 386; Vine, SF Serjania fuscifolia Radlk. — LP 9384, 10719; Vine, SF o pernambucensis Radlk. — LP 9249, 12190; Vine, SF Thinouia sp. — LP 9439; C 1245; Vine, SF ilta laevis a LP 9449; Vine, SF Urvillea ulmacea Kunth — LP 9259, 9388; Vine, RF, SF Sapotaceae L L H 9298; C 788; Tree, SF 568 Chrysophyllum rufum Mart. — LP 12048; Tree, SF Manilkara rufula (Miq.) H.J.Lam — LP 12054; Tree, ShF, SF Pouteria sp.1 — LP 9452; Tree, SF Pouteria sp.2 — LP 9407, 12112; Tree, SF Pouteria ramiflora (Mart.) Radlk. - LP 9772; C 233, 775; Tree, Simaroubaceae Simarouba amara Aubl. — Smilac Smilax iia SS 370; Vine, SF LP 9303; Tree, SF Solanaceae ° Brunfelsia uniflora (Pohl) D.Don - H 53494; Tree, SF Cestrum gardneri Sendtn. — LP 10702; Tree, SF Cestrum laevigatum Schltdl. — LP 10024; Shrub, SF Dyssochroma viridiflora Miers — LP 10777; Vine, ShF Heteranthia decipiens Nees & Mart. — LP 10710; Herb, SF Metternichia princeps Mik. — LP 9287; C 258; Shrub, SF Schwenkia americana L. — LP 10745; Sub, SF * Solanum —LP 10850; Shrub, SF Solanum caavurana Vell. — C 377; Shrub, SF Solanum crinitum Lam. — LP 9549; Shrub, SF Solanum depauperatum Dunal — LP 10801; Shrub, ShF Solanum jabrense Agra & M.Nee — LP 9432, 10014, 10838; Shrub, RF, SF Solanum palinacanthum Dunal ~ LP 12236; Sub, SF Solanum paniculatum L. - B 1237; Shrub, SF Styracaceae Styrax camporum Pohl — LP 3541; C 335; Shrub, RF, SF Trigoniaceae s bahiensis E.FGuimar., Miguel & Fontella — LP 9937 SF - 660; C304; ME m N Ere vom 2 E Camrmc C 781; Shrub ~ SF Trigonia nivea Cambess. var. nivea - LP 9957; C 282; Vine, SF oie nivea var. pubescens (Cambess.) Lleras — LP 9252 9798, 9874; Vine, SF Turneraceae Turnera cearensis Urb. — H 53491; LP 9922; C 1220; Shrub, SF Di Urticaceae Cecropia pachystachya Trécul — LP 9770; RC 1031; Tree, RF, SF Urera caracasana (Jacq.) Gaudich. ex Griseb. — LP 12076; ree, Urera nitida (Vell) Brack — LP 9465; Shrub, SF Verbenaceae Aloysia virgata (Ruiz & Pav.) Juss. — LP 9834; Shrub, SF Lantana camara L. - LP 9859; C 1212; Shrub, SF Lantana canescens Kunth — LP 12240; Shrub, SF Lantana fucata Lindl. — LP 10726; Sub, SF Violaceae Anchietea selloviana Cham. €: Schltd'. —C 279; Shrub, SF Hybanthus atropurpureus (A.St-Hil) Taub. — H 53479; Shrub, SF Hybanthus calceolaria (L) Schulze-Menz — C 379; Herb, SF Viscaceae Phoradendron crassifolium (Pohl ex DC) Eichler - C 293; Epi, SF Phoradendron mucronatum (DC) Krug € Urb. - LP 10847; C ; Epi, Phoradendron perrottetii (DC) Eichler — LP 9550; C 328; Epi, RF Phoradendron cf. pteroneuron Eichler — LP 12066; Epi, SF Vitaceae Cissus albida Cambess. — C 250; Vine, RF Vochysiaceae Vochysia sp. — Not collected, Tree, ShF idaceae Xyris jupicai Rich. — LP 10755; Herb, SF PTERIDOPHYTES Aspleniaceae Asplenium auriculatum Sw. — LP 12102; Herb, ShF Asplenium serra Langsd. & Fisch. — LP 10766; Herb, ShF Cyatheaceae Cyathea delgadii Sternb. — LP 9263; Tree, RF Dennstaedtiaceae Lindsaea lancea (L.) Bedd. - LP 9512; Herb, RF Lindsaea stricta (Sw.) Dryand. — LP 9545; Herb, RF Dryopteridaceae Ctenitis submarginalis (Langsd. & Fisch.) Ching - C1273; Herb, ShF Lycopodiaceae Lycopodiella cernua (L.) Pic. Serm. — C 804; Sub, SF Polypodiacea Niphidium crassifolium (L) Lellinger — LP 10805; Herb, ShF Microgramma geminata (Schrad.) Presl — LP 9792, Epi, SF Microgramma squamulosa (Kaulf) de la Sota — LP 10814; Polypodium catharinae Langsd. & Fisch. — n Epi, ShF Polypodium triseriale Sw. — LP 10815; Herb, S Pteridaceae Doryopteris lomariacea Kl. — LP 9398; Herb, SF Schizaea me. cS Sw. - LP 12267; Herb, SF Anemia hirsuta (L.) Sw. — LP 9289; Herb, SF Schizaea elegans (Vahl) Sm. — LP 9266; Herb, RF Selaginellaceae Selaginella muscosa Spring — LP 10809; Herb, ShF Thelypteridaceae Thelypteris salzmannii (Fée) C.V.Morton — LP 9472; Herb, SF e J UA, EN b c "n y £ Lf t." Dake Brazil 569 ACKNOWLEDGMENTS Ges work was supported by the Instituto do Milénio do Semi-árido (IMSEAR/MCT) and grants from the onselho Nacional de Desenvolvimento Científico e dL s i o dao aid 5 ane see 2004-2). The authors would like to thank the f of some plant specimens: F.R.Nonato and J.L.Ferreira (Pteridophytes), LCordeiro and D.S.C arneiro-Torres (Euphorbiaceae), J.G.Jardim and E.B.Souza (Rubiaceae), A.M.Giulietti (Eriocaulaceae), A.Chautems (Gesne- riaceae), C.Correia (Anacardiaceae), M.S.Ferrucci (Sapindaceae), R.M.Harley and F.Franca (Lamiaceae), M.G.L.Wanderley (Bromeliaceae), M. Sobral a. J.G.de Carvalho-Sobrinho (Malvaceae), E.Melo e (Polygonaceae), E.B.Miranda-Silva (Log dP | ), R.P.Oli (P ), T.D.Pennington (Sa- potaceae), M.M.Silva-Castro (Big ), N.Roque & S C.Ferreira (Ast ), M Sant aiid S.F.Conceicáo (Boraginaceae), M.C.Machado (Casaca, A A Conceição (Oxalidaceae), J.R. Pirani (Rutaceae), R.M.Castro (Moraceae), S.Romaniuc (Urticaceae), A NM Amorim and P.Dib (Malpi ), A. Rapini and R.F.Souza-Silva (Apocynaceae), MJ.S.Lemos-Costa (Aristolochiaceae), R. ls and A. K. A.Santos (Melastomataceae), M.F.Agra (Solanaceae), J.A.S.Costa (Begoniaceae), V.L.Gomes-Klein (Cucurbitaceae), T.R.Silva (Verbenaceae), C.van den Berg (Orchidaceae), T.S.Nunes (Passifloraceae), A.Gil (Iridaceae), and L.Aona (Commelinaceae). We are deeply indebted to Municipal government of Ruy Barbosa, SEMARH, Maria de Lourdes Janzen (Lurdinha), and Sr. Edson Pedreira who provided us with great attention during our visits into the study area; to Elvis, Ernani, Daniel, Sr. Messias, and Sr. Elias who helped us during the fieldwork; to Jorge Costa, Abel Conceição, Toby Pennington, and an anonymous referee for reading and commenting on the manu- script; to Ricardo Villas-Boas for helping us produce the map; to Freddy Bravo for translating the abstract into Spanish; and to Roy Funch for improving the English version. REFERENCES Ab SABER, A.N. 1974. O domínio morfoclimático semi-árido das caatingas brasileiras. Editora da Universidade de Sao Paulo, Sao Paulo. Acra, ME M.RV. Barsosa, and W.D. Stevens. 2004. Levantamento Florístico Preliminar do Pico do Jabre, Paraíba, Brasil. In: K.C. Porto, J.J.P Cabral, and M. Tabarelli, eds. Brejos de altitude em Pernambuco e Paraíba: história natural, ecologia e conservacáo. 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The Arboreal component of a dry forest in northeastern Brazil. Braz. J. Biol. 66(2A):479-491. RODRIGUES, LA. DA CARVALHO, AT, Ouivelra-FiLHo, R.T. Borre, and E.A. DA Sitva. 2003. Florística e estrutura da comuni- dade arbórea de um EE WEE em EE MG. Acta Bot. Brasil. 17:71-87. Ropricues, R.R. 2000. Uma discu S iliares, In: R.R. Rodrig d H.F Leitão-Filho, eds. Matas ciliares: conservacáo e recuperacáo. Ed. da Universidade de Sáo Paulo, Sáo Paulo. Pp. 91-99. SANTOS, K. DOS and L.S. Knoshma. 2003. Flora arbustivo-arbórea do fragmento de floresta estacional semidecidual do Ribeiráo Cachoeira, municipio de Campinas, SP. Acta Bot. Brasil. 17:325-341, Scunie, B.D., M. Lavin, and G.P. Lewis. 2005. Global distribution patterns of the Leguminosae: insights from recent phylogenies. In: |. Friis and H. Balslev, eds. Plant diversity and complexity patterns: local, regional and global dimensions. Biol. Skr. 55:375-422. r M | d | f^ H e EE E £ a a FE n.L* Brazil 573 SEI (Superintendência de Estudos Económicos e Sociais da Bahia). 1998. Análise dos atributos climáticos do Estado da Bahia. Série estudos e Pesquisas 38. SSES) Bahia. Siva, LADA and J.J. Soares. 2002. | fi ioldgico em um fragmento de floresta estacional semi- decidua no municipio de Sáo Carlos, SP. Acta Bot. Brasil. 16:205-216. Soares-FitHo, A.O. 2000. Estudo fitossociológico em duas florestas em região ecotonal no planalto de Vitória da Conquista, Bahia, Brasil. MSc dissertation, Universidade de Sáo Paulo, Sáo Paulo. Souza, J.S., F.D.B. ESPÍRITO- A MAT Fontes, A.T. OLiveira-FiLHo, and L. Borezetu, 2003. Análise das Mp dis florísticas e estruturais da de arbórea de um fragmento de floresta semidecídua às margens do Rio Capivari, Lavras-MG. R. Árvore 27(2):185-206. STRADMANN, M.T.S. 2000. Composicáo florística da mata ciliar da foz do rio Capivara e análise quantitativa do estrato arbustivo-arbóreo, Parque Nacional da Chapada Diamantina. MSc dissertation, Universidade Federal da Bahia, Salvador. Tavares, M.C.G., M.J.N. pee li L. MeLo, and MEA "din 2000. Fitossociologia do componente arbóreo de um trecho defl nontana do Parq lógico Joáo Vasconcelos Sobrinho, Caruaru, Pernambuco. Naturalia 25:243-270. TAvLoR, N.P and D. Zapri. 2002. Distribuição das espécies de Cactaceae na Caatinga. In: E.V.S.B. Sampaio, A.M. Giulietti, J. Virgínio, and C.F.L. Gamarra- las ES EH e flora da ls ANS RESTO is es 125. Toniato, M.T.Z. and Eck EK FILHO. di Variations in tree community | a frag of tropical tern Brazil related to different ! disturbance histories. Forest Ecol Managem 198:319-339. Tryon, RM. and AE IRvoN. 1982. Ferns and allied plants with special reference to Tropical America. Springer- Verlag, New York. UpuLutscH, R.G., M.A. Assis, and D.G. Picchi. 2004. Florística de trepadeiras numa floresta estacional semidecidua, Rio Claro — Araras, Estado de São Paulo, Brasil. Revista Brasil. Bot. 27:125-124. VeLLoso, Al E.V.S.B. Sampaio, A.M. Guuem, M.R.V. BARBOSA, A.A.J.F. CASTRO, L.P. DE Queiroz, A. FERNANDES, D.C. OREN, L.A. CESTARO, A.J.E. CARVALHO, F.G.C. PAREYN, F.B.R. SiLva, EE MIRANDA, S. KEEL, and R.S. Gonbim. 2002. Ecorregiões: propostas para o Bioma Caatinga. APNE, The Nature Conservancy do Bn Recife. VeLoso, H.P, A.L.R RANGEL-FiLHo, and J.C.A. Lima. 1991. Classificação da vegetação brasileira adaptada a uma sistema universal. IBGE, Rio de Janeiro. Vigia, EA AT. OuveiRA-FiLHO, DA CarvaLHo, and M.L. GaviLANES. 1995. Flora arbustivo-arbórea de um fragmento de mata ciliar no alto rio Grande, Itutinga, Minas Gerais. Acta Bot. Brasil. 9:87-100. WERNECK , M.DE S., G. PEpRALLI, R. Koenig, and L.F. Giseke. 2000. Florística e estrutura de trés trechos de uma floresta semidecídua na Estação Ecológica do Tripuí, Ouro Preto, MG. Revista Brasil. Bot. 23:97-106. 574 Journa BOOK NOTICE Ana CLAUDIA ARAÚJO, EDGLEY A. César, AND Davip Simpson. 2007. Preliminary List of the Cyperaceae in Northeastern Brazil: Repatriation of Kew Herbarium Data for the Flora of Northeastern Bra- zil Series, vol. 3. (ISBN 978-1-84246-204-1, pbk.). Royal Botanic Gardens, Kew, Richmond, Surrey, TWO 3AB, United Kingdom. (Orders: www.kewbooks.com, publishing@kew.org). $51.55, 32 pp., 11 3/4" x 8 1/4" Contents: Texto em Pe cnl I map Bibliografia Reference Text in English Listagem/Checklist Lista de Exsicatas/List of Specimens e ghtf FERIDA EE fthe Cyr Ps ol : Brazil, p AN ERR Be py d English text, with a Forward by Wm. Wayt Thomas of The New York B ical Garden. F tl blis] “Between 2002 anid 2004, 1932 Cyperaceae specimens ined and the inf ti patriated. This checklist ds 191 species and 24 genera, MIE entries See Kiwi and sorted by state, collector and number. Àn additional exsiccatae list enables duplicates lodged in otl ze ain the correct determination.” J. Bot. Res. Inst. Texas 2(1): 574. 2008 DIVERSITY AND FLORISTIC COMPOSITION OF THE VASCULAR PLANTS IN THE FOREST FRAGMENT IN SOUTHEASTERN RIO DE JANEIRO, BRAZIL Regina H.P. Andreata Haroldo C. de Lima Instituto de Pesquisas Jardim Botánico do Rio de Janeiro 22460-030; R. Pacheco Leão 915, RJ, BRASIL hl Universidade Santa Úrsula/ICBA 22231-040, R. Fernando Ferrari 75, SiL regina.andreata@gmail.com ima@jbrj.gov.br Angela S. Fonseca Vaz José Fernando A. Baumgratz Instituto de Pesquisas IBGE/Instituto de Pesquisas Jardim Botánico do Rio de Janeiro 22460-030, R. Pacheco Leão 915, RJ, BRASIL avaz@jbrj.gov.br Jardim Botánico do Rio de Janeiro 22460-030, R. Pacheco Leão 915, RJ BRASIL jbaumgra@jbrj.gov.br Sheila R. Profice Instituto de Pesquisas Jardim Botánico do Rio de Janeiro 2460-030, R. Pacheco LeGo 915, RJ, BRASIL sprofice@jbrj.gov.br ABSTRACT The diversity and floristic composition of the vascular plants of the Morro de Mouráo, AS the Serra i Tiririca n i Rio de Janeiro State, Brazil (22? 58' 04" S x 43º O1' 17" W), were analyzed. The stud est within the Atlantic Coastal Forest biome. A total of 369 taxa were collected, including 7 Preridophytes, 295 [Dieotsdedons, and 67 forms: arboreal, shrub-arboreal, pe Monocotyledons, belonging to 84 cian families. T p herbaceous. Á map of the area, a f the 1 water balance, lani profiles, soil anal climates of the eastern and western facing slopes e s cua da a uae are Ro es or ee UM and E: Pa sl ] Al DN q e D Ge collectors. RESUMO A diversidade e a composição florística das plantas vasculares foram analisadas no Morro Alto Mourão (22° 58' 04" S e 43º 01' 17" W) situado no Parque Estadual da Serra da Tiririca, oon no estado do Rio de Janeiro, Brasil. A área de estudo compreende um Mata Atlántica. Foram a 369 táxons, sd 7 importante ECH Ge AF “1 Ja Pteridófitas, 295 Di , pert I - > arhmnctiva z 1 dz Anrecenta_cee 111 mang at le] 1 hidri fi ati ] A SA 1 j| inf * ULLFIÃOLI VS EF I 7 0 5 I L toc à lico g PE AEE EE EN tij 1; d Itad I i | la S da Tiririca al se pa ceo | ani / 1 Jami É f id list g dos tá , dados do hábito, floração, frutificação d Ej e dos coletores do material examinado. INTRODUCTION Coastal forests are an important component of the Atlantic Coastal Forest biome. These forests are distributed along most of the eastern coast of Brazil. However, they have been subject to intense degradation within this range, due mainly to their proximity to large urban centers. Detailed information concerning the coastal vegetation of Rio de Janeiro State is relatively limited. This study examines the coastal vegetation of the Morro Alto Mouráo region, including components of the humid coastal forest and the near-shore (restinga) vegetation. The Morro Alto Mouráo is one of the highest pou in Ee Serra da Tiririca Range did m). It is EE in southeastern Rio de Janeiro State between the tal if EL ragments of one of the most highly threatened f Niterói and Maricá. It still Planes: in the world—the Atlantic Coastal Forest. J. Bot, Res. Inst. Texas 2(1): 575 — 592. 2008 [| [| £ al n In LI MP don nd ET 576 tani Texas 2(1) The Serra da Tiririca Range is oriented almost perpendicular to the Atlantic coastline, which results in contrasting climatic conditions on its two flanks. The western slopes are generally exposed to the humid ocean winds. Intense rainfall is also common, principally due to cold fronts arriving from the southern polar region. On the other hand the more sheltered eastern slopes lie in the rain shadow and receive less and lighter rainfall. The first historical references to the area date from the end of the 16th century when the Portuguese crown awarded a ae land title esa to Duarte Martins Mourão. This title granted formal control of the coastal lands | the Lagoa de Piratininga and Pedra de Inoã, in the current municipality of Maricá. ce Human use and occupation of this region is very intense. It was cultived with large numbers of small sugar cane, coffee, and banana plantations, as well as citrus orchards up until the start of the 20th century (Grael et al. 1995 Even though this region was highly disturbed (and is located near large urban centers as Niterói and Maricá cities) the vegetation on the Morro Alto Mouráo is experiencing a gradual but natural regeneration. Currently, plants with ornamental, medicinal, timber value and species of great scientific importance (rare or endemic to the State of Rio de Janeiro and/or the Atlantic Coastal Forest) that are often little known to botanists have been discovered. Preliminary data from this study have been presented at scientific meetings and helped transform the Serra da Tiririca Envi tal Protection Area into the more highly protected Serra da Tiririca State Park (Rio de Janeiro State Law No.1901, of November 29, 1991). This study presents the results of a floristic inventory undertaken in the area around the Morro Alto Mouráo. This inventory examined the diversity and composition of the vascular plants within an area of coastal forest. It is intended to be used to support conservation and degraded areas restoration efforts along Rio de Janeiro State coast. MATERIALS AND METHODS Study Area The Morro Alto Mouráo is located in the Serra da Tiririca Range, which covers a approximately 2700 hectares near Pontas de TUAE and dere Gef 5804" S x id O! p uL (Fig. a The Morro Alto Mourão is partofa pre-Cambrian (Niterói Massi | tl rn shore of the Guanabara Bay. The peak has one ol ¡he highest elevations (369 m) in the Tiririca OUTING. together with Morro do Elefante (or Falso Páo-de-Acúcar) (412 m) and Morro do Telégrafo (250 m) (Pontes 1987). The ground is generally covered by leaf litter although there are also exposed rock areas. The sandy- clay soil has a medium texture, is highly acidic (pH = 4.7-4.8) and has mid-range quantities of calcium (3.5 me/100 mg), phosphorous (12 ppm) and magnesium (1.6 me/100 mg). The aluminum concentration is high (1.0 me/100 mg), and the potassium concentration very high (> 135 ppm). The high acidity and large quantities of aluminum and potassium found in the soils are probably the result of leaching and frequent burning. The western slopes of the Tiririca Range have a sub-humid climate (C2), with an A’ megathermic temperature regime. The average annual temperature is 23.2° C, annual rainfall 1305 mm, annual evapo- transpiration potential 1271 mm, real annual evapo-transpiration 1207 mm, annual water-deficit 64 mm and annual excess 98 mm (Fig. 2). On the other hand, the eastern slopes have a sub-humid humid climate (C2) n a B4 meso- bc apenas regime. is 23.2° C, annual rainfall 1207 mm, annual evapo-transpiration potential 1223 mm, real al evapo-transpiration 1163 mm, annual water-deficit 60 mm and annual excess 62 mm (Fig. 3). The greater rainfall and humidity encountered on the western slopes are due to the direct influence of humid on-shore ocean winds. Methods Las , e: GE to tl between 1980-85 and 1989-90. Plant collecti A LLO LLa 5 P. D e Andreata etal., Fl ictic di ity the M Alto Mouráo, Brazil 577 o E cd SC? de? — CUM. s Xs E Sei CS ms Veto Q2 Ao. 7 V lr E é DAE A A Gen dc NES. et E w b, pa f. KS br AL a de x r x Ae sod a CR iy b — A j e? E Les” " af "i — EU 4 *ROPATLA wë Gët r5 x "Era OM tae Pc ez a " No Tio EA E d D ve E SE e. Sa dÉ e Á Fa Le f 7 E e A Ju M EF. n WT ; Ea a a a 1 us 5 € x e "si à L i E Dv RN Convenções Cartográficas F ee E E ger — Me CUM a mem me = Limite Estadual stroda d Kl Zéi A ES mile Municipal — Estrada Prvimenladn bmi, | UM fr: : : r — Tráfego Patt Ze "P z C L3 Area Urbana m ausi emis Cu = Prefixo Fudtral n Tax. ets . e Side de Municipio bd 8t. E Di al ka ais a = = E if E "ëm V) Localidade MS 1 E a à — Bt Estadi te Pera d air E [J] Parque Estaria m zz EF. dp Ea iL [E pe-nea = Curva ce Hiel HR Ge 7/7 SC 4000 0 div. 800) m MT Hidrografia i EX, a CAR i VA ff fy PU Lë 7, M H ogi ——————————————————Ó——— Fic. 1 | ps Ab E ok Rã Ala RA A DI J I M Chad Collections were undertaken on the Morro Alto Mourão, as well as on the Morro do Elefante and Pedra de Itacoatiara, which have a similar vegetation cover. During the period from 1986 to 1988, collections were less frequent. However, these were generally accompanied by certain plant group specialists or undertaken especially to prepare schematic diagrams representing the phytophysiognomy of the vegetation. All collected material was identified and registered at the herbaria of the Instituto de Pesquisas Jardim Botânico do Rio de Janeiro (RB), the Universidade Santa Úrsula (RUSU) and GUA, HB, FFP (Orchidaceae). The plant lists are presented in an appendix, with the family names in alphabetical order. The Angio- sperm families are named according to Cronquist (1988). Those of the Pteridophytes are named according to Kramer and Green (1990). The flowering and fruiting periods are derived from the specimens collected in the area. A map of the area, g the water tic vegetatio fil ided e The different phytophysi j deia as either arboreal, shrub- My or Del ee in order P to acen the Econ cover. Climatic data for the Serra da Tiririca was obtained from the Niterói Weather Station (representing the climatic conditions prevalent on the western slopes, directly exposed to humid winds and intense rainfall) and from the Maricá Station (representing the climatic conditions prevalent on the eastern slopes that are shielded from the humid winds and intense rainfall characteristic of the western slopes). The techniques of Thornthwaite and Mather (1955) were employed to prepare local water balance diagrams, quantify the available soil humidity (evapo-transpiration) and determine the degree (either deficiency or surplus) of the water balance during the year based on the monthly rainfall data obtained from these stations. 578 250 ——— een E J— M E : VP d 200 NS A A EA A + À ded | LE DT —— Le ER X - ` r d à EA | 100 e=* - ; 3 — " Pn — maeme X ula m 50 = 7 0 1 D L| t L D 3 1 D i D i j f m a m j j a s o n d Fis. 2. Diag tl balance of Niterói (1958-67), Water Bal g to Thornthwaite & Matl (1955). Latitude: 22° 54'S- Longitude 43º 07'W. Annual Precipitation: 1305 P. Annual Evapo-t wes ; Two soil samples were taken (0-20cm deep) and analyzed at The Soil Fertility Division of the National Service of Research and Conservation of Soils at EMBRAPA, in the city of Rio de Janeiro RESULTS AND DISCUSSION Taxonomic diversity A total of 369 taxa were sampled within the study area. Seven of those were Pteridophytes (3 families), 295 were Dicotyledoneae (63 families) and 67 Monocotyledoneae (18 families) (Appendix 1). Leguminosae (40 taxa), Euphorbiaceae (23), Rubiaceae (20), Myrtaceae (15), Solanaceae (16), Brome- liaceae (12), Orchidaceae (12), Araceae (11) and Sapindaceae (11) represented the greatest taxa richness These totaled 43% of the Angiosperms encountered. Solanum (10 taxa), Rhipsalis (6), Machaerium, Philodendron, Anthurium and Trichilia (5 taxa each) were the most representative genera. Classifying these plants by habit, the largest diversity was observed among the herbs and sub-shrubs which were regionally represented by 115 taxa. The trees comprised 96 taxa, followed by the shrubs, climb- ers and epiphytes, with 92, 52, and 18 taxa, respectively (Appendix 1) Comparisons of the plant species richness of Morro do Alto Mouráo with other forest areas within the Serra do Mar Range in Rio de Janeiro State are shown in Table 1. A tendency towards reduced species rich- ness can be seen within coastal forest areas near the city of Rio de Janeiro (Morro do Alto Mouráo, Horto/ JBRJ, and Morro das Andorinhas) in relation to humid forests in the Serra do Mar Range (Macaé de Cima, Cairucu, and Tinguá). However, the high degree of local endemism of the coastal forests is very notable, with 20 species restricted to only Rio de Janeiro State (Appendix 1). This same pattern of endemism was reported by Lima (2000) and Sá (2006) who noted that most of the endemic species occurred in seasonal forests. Andreata et al ' Flaristic di it the M Alto Mourao, Brazil 579 180 + ———— 160 +p — 140 120 e Epa E71 [Pas 68 tup — 100 EN |-— 80 SX E — 60 40 ea 20 0 mmm mmm. mmm mmm Eee j f mamj jasond Fic. 3. D ftl bal f Maricá (1987—99).1 Bal ling to Thornthwaite & Mather (1955). Latitude: onion Gage 42º 49'W. Average Annual Temperature: ex PCA A | Precipitation: 1207 P. Annual Evapo-t pirat I , True Annual Evapo E DE i | E ou Fa A (TT Bn (e F^" nar A (TT UO! E o” LAI A nm | | A at ram rie "LA ree. ui IT LE LP" [| L kirma’ dor fJ E TL | 4 DA esoti e C The floristic diversity of the Morro Alto Mouráo flora was high in comparison with these other areas of the Atlantic Coastal Forest (Horto/JBRJ and Morro das Andorinhas), even when allowing for the differ- ent methodologies and sample sizes used in the different surveys. The significant floristic diversity of this coastal mountainous region within the Serra "e Mar Range is Pea due to numerous factors, the most important likely þei ificant local climat ity (due to t ition of the Serra da Tiririca Range). Physiognomic-floristic analysis The vegetation on the Morro Alto Mourão is typical of the Serra do Mar Range. It can be classified as low- montane rain forest (Rizzini 1979). E Sd ONA ane s eastern bis S the be E ino Mi EE the coastline, the forest | n stature Tara de CA 1 f 11 trees and acquires many elements of the near-shore (retina) vegetation, Th and herbaceous vegetation near the rock outcrops. The different physiognomies observed on the Morro do Alto Mouráo are illustrated in Figure 4. In addition to the secondary forest formations in the study area (resulting from human intervention 580 Jou t tani itute of Texas 2(1) Taste 1, Numbers of families, gen | ifi | infra-specific taxa among g the vascular flora in areas of Atlantic Coastal Forest within the state of Rio de e Brazil. Alto JBRJ/Horto1 Andorinhas2 Reserva Ecológica APA- Reserva Biológica Mouráo MacaédeCima3 ^ Cairucuá do Tinguá5 Families 84 74 60 122 133 124 Genera 244 168 103 413 483 421 Total taxa 369 277 105 1103 1087 1037 !Marquete et al. (1992); 2Araujo and Vilaça (1981); ¿Lima and Guedes-Bruni (1997); “Marques (1997); »PMA/JBRJ (2002). 2 n^ 2 eg rite "e A AA PAY s yn Ai lj | MERO ^ 2 3 e. Ma a Rg -— i wd p | lu ALTE ne d y : Hr Í r 3 " (1) SDE yl 1 qu ol Oso Sé "i Se Fic. A E | 7 £i ZA Lt L H » E sl RA Als AA = , n: JJ | at Cont : Brazil 1 A VE LE + ; Lo | Shrub-arboreal formation; 3. Herbaceous formation. during the last tl turies) tl ] 11 pri imary forest f Iragments. According to Pontes (1987), the Atlantic Coastal polest once red the entire region around the SES da Tiririca, but was subsequently subjected to ext tation through logging and clearing. Only isolated sections of the original forest remained in relatively inaccessible areas, such as Morro do Telégrafo and Morro Alto Mouráo. Edaphic and climat t t have played important roles in the establishment of plant communi- ties or large populations in the area, resulting in the mosaic of successional stages found there today. Despite the often imprecise limits of distinction between them, three principal physiognomic forma- tions are recognized in this study: arboreal, shrub-arboreal, and herbaceous. Arboreal formation Arboreal formations dominate the study area and they presumably occupied an even greater proportion of the slopes in this region in earlier times. The topography is generally gentle and allows the accumulation of organic and inorganic material (the latter formed by the gradual breakdown of the rock substrate) that is necessary for soil formation and forest establishment. These arboreal formations now occupy the slopes and depressions, resulting in the formation of deep and fertile soils. There are two forest layers—the upper and lower canopies, determined principally by the height of the component individuals (Fig. 5). The upper canopy is closed and is composed of individuals that are generally between 12 and 25 m tall (remnants of the original fores). Some emergents are to be found, with average heights between 25 and 30 m, such as Cariniana legalis, Anadenanthera colubrina, Pterogyne nitens, Schinus terebenthifolius, Gallesia integrifolia and Coussapoa curranii. The lower canopy is composed of individuals that are between 5 and 12 m tall. This layer is more open, and the tree crowns being more widely spaced. Some representative species are: Piptadenia gonoacantha, Andreata et al., Floristic di it the M Alto Mouráo, Brazil 581 LEGEND " deere ale d 2- Metternichia princips var. princips 3- Alseis sp. 4, 6, 9- Machaerium firm 5- Piptadenia jain T- Didymopanax s 8- Randia e gien in 10- Actinostem munis 11, 13, 15, 16, 17, 21- Syagrus romanzoffiana 204 | 12, 20, 22- dune opposita 14, 19- "paa eer var, fluminensis 18- Aspidosperma parvifolium 41 1 1-1 fe ral £4. y DE : Sab : e IT) 1 ka : iis] M Vel AP LPPLL LPL P M , Guapira Ery I E r [e Ò opposita, p Coari NS var. e The shrub-arboreal formation A shrub-arboreal vegetation predominates on the ocean-facing mountain slopes. This vegetation is very dense, with a Ron a den However, it has a small number of n a d E) pedes 1 "n 1 common to the sandy near-s q ga) ) vegetation, such as C J ji are also present. This formation has reasonably a popu ations of pans that stand out physiognomically in the veg- etation, such as Marli laris, M. richardiana, and Psidium cattleyanum (all of the Myrtaceae family) har: añ in Beste [rom 5-7 m. Pachystroma longifolium can reach up to 10 m. There are also smaller populations of Esenbeckia febrifuga growing together with the palm tree Syagrus romanzoffiana (that can reach 10-15 m). Clavija spinosa, Jacaranda jasminoides, Inga congesta, Brunfelsia uniflora, Norantea brasiliensis, Cupania racemosa, Tibouchina corymbosa, and Faramea stipulacea are a apne ee among the SE This same shrub-arboreal vegetation bart 1b species (e.g., Justicia beyrichii, heic M aggregate, m Li Md Julocroton triqueter, Capparidastrum brasilianum, Cyperus diffusus, Cal Alstroemeria caryophyllea, and Neomarica sp.). These cannot be considered a distinct — in ens as they are very thinly distributed within the area. On the other hand, there are dense populations of Schaueria calycotricha in shaded areas at the forest edge, near rock outcrops with many weedy plants such as Petiveria alliacea, Sida rhombifolia, Oxalis barrelieri, Solanum alternopinnatum, and Stachytarpheta polyura, as well as epiphytes and vines. oe ane hemi- KG bk are pne uncommon in the area. They include Anthurium pentaphyl- lum, A p. scandens, Monstera adansonii var. EE Philodendron bipennifolium, P. cordatum, Aechmea nudicaulis var. nudicaulis, Neoregelia ampullacea, Bill idalis var. pyramidalis, Rhipsalis teres f. capilliformis, R. mesembryanthemoides, R neves- EE and R. Pe among other taxa of Gesneriaceae, Orchidaceae, Piperaceae, and Pteridophytes. Some epiphytes are encountered in the area with other life forms, such as Anthurium scandens subsp. Ta l- Rudgea interrupta 2- Ruprechtia laxiflora 3,4,5,6,9,11,13,15,17, 19 ,20,21, 35,39,41,44,45,47,48,49,53- Cusparia ovata Ka kA + Z "&g E , Kë QC Ki DO a 10,32,33- SM brasiliensis 5,30- Syagrus zoffiana 26,40,57- Myrciaria floribunda 27- a virgiltoides 28- eolens Ast 29,56- Ecos um SEN 12- Engenia sulca 3l- Cec — 14- Psidium catan 34- Cous us 16- Acac 36- E. brasiliensis 18,43, E GE comunnis var. spathulatus 38- aci a e 22,55- Kielmeyera membran 42- Myrta et ` 5m 46,50- SC involuta Qi a 50 s4 58 j^ 37 Ja G ES A 42% 48.) NAS cl. a AT VE) SS "UN ES E Es: NON, ER] P > RAR Mina d mg scandens, as rupicolous, and terrestrial individuals of Philodendron cordatum. Also Oncidium and Pleurothallis species are encountered both as epiphitic and rupicolous. Both woody and SUONI vines are found on the Morro Álto Mouráo, especially species belonging AA e to the genera , Bauhinia, Chaetocalyx, Centrosema, Serjania, Paullinia, Urvillea, Heteropterys, antes IT En Gage Passiflora, Ipomoea, Jacquemontia, Reissekia, Dioscorea, Smilax, and Herreria. A single species of the hemi-parasite Struthanthus marginatus was encountered. The herbaceous formation This vegetation type is typically found on rock outcrops where plants grow directly on the rock surface (rupicolous) or within thin sandy mips of Së near the rock outcrops (saxicolous). The plants at these sites are exposed to extreme nditions due to their proximity to the ocean, including high levels of solar radiation, strong, constant winds and high salinity (Fig. 7). Tillandsia dura, Aechmea ramosa var. ramosa, Neoregelia ampullacea, Peperomia incana, P. corcovadensis, Cyrtopodium polyphyllum, Anthurium coriaceum, A. olfersianum, Philodendron speciosum, Coleocephalocereus fluminensis, Rhipsalis cereoides, and Hylocereus setaceus are among the most frequent Angiosperm pioneer species. There are also small clumps of plants found in deeper soils areas, including individuals of Tibouchina corymbosa, Tabebuia chrysotricha, and Stillingia dichotoma, in addition to the pioneer species. Other communities are found on flat isi EE in Ma! clearings within the forest (240—260 m Gees urens, Furcraea gigantea, Galactia altitude) on very humid soils and inch 1 striata, and Stylosanthes scabra, as well , such as Cleome dendroides subsp. dendroides, Manihot palmata, iia thyrsiflora, and Tradescantia abria Conservation This area is significant because it contains a large number of species (20) endemic to the State of Rio de Janeiro, some of which are limited to the the Serra do Mar forests (Appendix 1). Among the most interest- ing of these are SINUM grandifolia and EH speciosum, both of which were considered practically Atlantic Coastal Forest areas (Pirani 1990; Mayo extinct and 1991). Wilbrandia a cn) had not been found growing in the wild since the 19th century from only t Andreata et al , Floristic di it the M Alto Moura 0, Brazil 583 d LEGEND OË mum Selaginella sellowii * Stillingia dichotoma ; f m . Ac Bilbergia pyramidalis var. pyramidalis | Ka Furcraea gigantea Q ibouchina corymbosa Y Vellosia candida y Bulbostylis capilaris Tillandsia arauiei Y Cyperus coriifolius H Coleocephalocereus fluminensis + Pitcairnia staminea XN ^ Y EY ta DE + E AN ae WEEE EVI S Lë 2m En 7 Sct Pr FT fal. | L £ Ai fai. MA Alen LA an Din da | iro State. Brazil (Klein 1990). Also relevant is the finding of species that are considered either rare and/or threatened, such as Pterogyne nitens, Pouteria subsessilifolia (UCN 2004) and Astronium glaziovii. The latter species has only a few collections registered in all of the Brazilian herbaria (Santin 1989). Among the plants encountered that are threatened with extinction in the municipality of Rio de Janeiro are Inga congesta, Inga cordistipula, and Lonchocarpus virgilioides (Lima 2000a). Additionally, Ficus enormis and Coleocephalocereus fluminensis are considered to be threatened due to destruction of their habitat in the State of Rio de Janeiro (Duarte et al. 2003; Calvente et al. 2005). Other rare species are Gomidesia sonderiana, which has rarely been found in the Atlantic Coastal Forest (Nic Lughadha, pers. comm.); Alstroemeria caryophyllaea, which had only previ- ously been encountered growing in the wild humid and shady areas in the States of Minas Gerais and Rio de Janeiro (Assis 2004); Pleurothallis pardipes, recollected recently in Parque Municipal da Prainha, Rio de Janeiro State (Pinheiro 1999). The occurrence of these rare and/or threatened species, together with numerous cases of endemism, illustrates the importance of conserving the Morro Alto Mouráo area. These (and other) species could po- tentially be utilized as germplasm sources for seedlings production used to replant other fragments with a similar floristic compositions in the Rio de Janeiro Atlantic Coastal sica It is important to note that the Morro Alto Mourão area contains a bl | f j| ful A ML op be > 584 t tani Institute of Texas 2( to humans or with significant direct economic potential (see Lopes 1992), including those with medicinal (Lopes & Andreata 1989, 1990, 1991, 1992) and ornamental (Lopes et al. 2004) uses. The economic potential of these plants, together with the proximity of the Morro Alto Mouráo area to a large urban centre, complicates its preservation. Barros and Seone (1999) have proposed strategies for the conservation of areas within the Serra da Tiririca State Park including monitoring to combat plant resources extraction and fires, as well as controlling tourism. CONCLUSIONS The best represented plant families observed in the Morro Alto Mouráo area followed the same patterns seen in other floristic inventories undertaken in the State of Rio de Janeiro (Marquete et al. 1992; Araujo & Vilaca 1981; Lima € Guedes-Bruni 1997; Marques 1997; PMA/JBRJ 2002). The phyto-physiognomic analysis of this area revealed a significant vegetation cover (predominantly arboreal) in spite of both the historical and recent environmental aggression. The eastern portion of the Guanabara Bay in the municipality of Niterói has only a few small forest fragments HAM Seene the idis da Tipo. (PEEMA um im SS and the other data presented J]: in this work ] p B | y Pi esent within the Atlantic Coastal Forest in the State of Rio de nero. APPENDIX 1 List of the vascular plants of the Morro Alto Mouráo, Rio de Janeiro State, Brazil by alphabetical order of the families, citing the specialists consulted, Latin name, habit, flowering (fl), fruiting (fr), and collector's name (*endemic), (**introduced). PTERIDOPHYTES (P. Windisch & C. Mynssen) Amaranthaceae Alternanthera sp. — herb, fl Jan), R. Andreata 405 Alternanthera brasiliana (L) Kuntze var. villosa (Moq.) Kuntze — herb, fl (May-Jul), R. Andreata 72, 704 Pteridaceae lil collina (Raddi) J. Sm. — herb, R. Andreata 330 Doryopteris lomariacaea Klotzsch — herb, R. Andreata 898 Doryopteris patula (Fée) Fée — herb, V. Ferreira 2107 Anacardiaceae (F. Gil) Doryopteris varians (Raddi) J. Sm. — herb, R. Andreata 34,530 Astronium glaziovii Mattick — tree, fl (sterile), H Andreata 359, 361, Schizaeaceae Anemia phyllitidis Sw. — herb, R. Andreata 35, 443 Astronium graveolens Jacq. — tree, fl Jul), R. Andreata 85, 360, 69, Selaginellace ius EE Raddi — tree, R. Andreata 349 Selaginella ven (Desv. ex Poir.) Spring ex Mart. — herb, V. Spondias mombin L. — tree, fl (Sep), fr (Jan), R. Andreata 357, i Selaginella Soe Hieron. — herb, R. Andreata 51, 879 Spondias venulosa Mart. ex Engl. — tree, R. Andreata 915; G.Lewis 3988; D.Santin s.n. DICOTYLEDONS A h Annonaceae (C.G.P. Quin camina stas Rolliniopsis parviflora (A. St. na small tree, fl (Nov), fr Jan), Aphelandra longiflora (Lindl.) Profice — shrub, fl (Nov), fr (Nov), R. Andreata 403, 615; C. Farney 363 R. Andreata 602 DN H i hii N herb, fl (Apr), R. Andreata P4 Apocynaceae Aspidosperma sp. — tree, fl Jan), V. Ferreira 2094 Aspidosperma parvifolium A.DC. — tree, fl (sterile), R. Andreata 696 426 Justicia beyrichii Nees — herb, fl (May, Sep, Nov), R. Andreata Justicia sp. — - sub-shrub, f (Apr-May), R. Andreata 415, S. Aspidosperma tomentosum Mart. — tree, fl (Sep), fr (Sep), R. 2 Profice 577 Ruellia solitaria Vell. — herb, fl (Aug), R. Andreata 92 Mandevilla ee ~ K. Schum. - climbing, fl (Jul), fr *Schaueria calycotricha (Link. & Otto) Nees — shrub, fl (Jul, Qul), R.A à Oct-Deo), fr Jul-Sep, Oct-Nov), R. Andreata 47, 91, 131, ros sp.— GH fl (Nov), R. Andreata 609 Tabernaemontana hystrix Steud. — tree, fl (Nov-Dec), R. An- ga SE n ex Sims. - climbing, fl (Nov), fr Jul), R drea Farney 368 Andreata Prestonia cf. calycina Müll. Arg. — climbing, sterile, R. Andreata 118a Andreata eta | ri HE ES Jė HN Ai RA Li Prestonia coaiita (Vell) Woodson - climbing, R. Andreata Araliacae Schefflera succinea Frodin € Fraschi — tree, fl (sterile), R. An- Aristolochiaceae Aristolochia rumicifolia Mart. € Zucc. — climbing, fl (Sep), R. Andreata 525 E cymbifera ae 8 Zucc) Klotzsch - climbing, fl erile), R. Andreata 48 mes raja (Mart. e Zucc) Klotzsch - climbing, fl (Oct- Dec), fr (Sep), R. Andreata 37, 147, 193, 513 Asclepiadacede (N. rt de ibi Dep banksii - climbing, E (Jul, Sep), R. Andreata 96, 544 Asteraceae (R. Esteves) trat tatum Ca herb, fl (Dec) R. Andreata 19 Cyrtocymura scorpioides (Lam. H. Rob. — sub-shrub, fl (Apr, Aug, Dec}, R. Andreata 33, 87, 674 Eupatorium organense Gardner — sub-shrub, fl (Nov-Dec), R. Andreata 57, 314 Eupatorium vitalbae DC. — herb, fl id ` es 105 Gochnatia sp. — tree, fl (Jan), R. Andreat Tilesia baccata (L) Pruski — shrub, fl Sen ái ai R. Andreata 123, 127 Begoniaceae Begonia sp. — herb, fl (Nov-Dec), R. Andreata 46, 194 Bignoniaceae Adenocalymma paulistarum Bureau ~ climbing, fl Jan, Dec), A a E Anemopaegma chamberlaynii (Sim) Bureau & K. Schum — climbing, fr (May), R. Andreata 55 Arrabidaea af. rego (Vell) DC. — tree, fl (Apr), fr (Apr), R. An- dreata 669 R. Andreata 318, 347, 666 Jacaranda jasminoides (Thunb.) Sandwith — shrub, fl (Aug, Oct-Dec), fr Jul, Oct-Nov), R. Andreata 55, 89, 122, 165, 2301, 318.223 Ree crucigerum e A.H. Gentry — climbing, fr (May), . Andreata163; C. Farney 740 zapen vernicosum cm & K. Schum. - tree, fl (steri n tran El EK Aas Klo zt fr (Jan) labe bu ia sp. - tree, fl Ges H Ancreata 988 Tabebuia chrysotricha (Mart. ex DC.) Standl. — tree, fl (Aug), fr (Aug), R. Andreata 104 mbac Gees Se (Pasq.) A. ido a fl E fr Jan- Feb, Apr), R. Andreata 346, 496, 665; T. Plowman 12858 n lec (Kunth) PE. Gibbs & e - tree, «f (Apr), R d 376, 897 lifl (Cav.) A. Robyns - tree, ff (Jul), R. Andreata 455, 713,985 585 Quararibea turbinata (Sw.) Poir. — shrub, fr (May, Jul), R. An- dreata 461, 581 Boraginaceae Cordia ochnacea DC. — shrub, fl (Nov), fr (Apr-May), R. Andreata , 287, 589, 610, 667 Cordia trichotoma (Vell.) Arrab. ex Steud. - tree, fl (Apr-May), ; ta 549, 663, 910, 912 *Tournefortia salicifolia A. DC. — sub-shrub, fl (Sep, Nov), R. Andreata 334, 546 Cactaceae ( L. Scheinvar and A.M. Calvente) Brasiliopuntia brasiliensis (Willd.) A. Berger — tree, fl (Oct), fr s D, H E 130 TRA" ND st Ckeb. — herb, fl (Apr), R. Andreata 889 Hylocereus setaceus (Salm-Dyck ex DC) Ralf Bauer — epiphytic, Andreata 304 Rhipsalis cereoides (Backeb. & Voll.) Backeb. - epiphytic, fl E ` (Sep), R. Andreata 326, 545, 650, 886; T. Plow- 3935 Rhipsals deen K, schum. - “epiphytic V. SE a a Andreata 301; E Martinelli 8154 Rhipsalis neves-armondii K. o — epiphytic, fl (May), fr ndreata 303,5 Rhipsalis GC Pfeiff. — inne fl (sterile), R. Andreata 302 Sp ES Rhipsalis teres Steud. f. capilliformis (F.A.C. Mice Barthlott & Taylor — epiphytic, fi (sterile), R. Andreata Capparaceae Capparidastrum brasilianum (DC) Hutch. — sub-shrub, fl (Nov), fr Jul), R. Andreata 467, 611, 950 Cleome dendroides Schult. subsp. dendroides — shrub, R. Andreata 42, 106, 185 Celastraceae (A. Joffily) Maytenus brasiliensis Mart. — shrub, fl (Apr), fr (Nov), R. An- dreata 425, 97 Maytenus communis Reissek — shrub, fr (Aug), R. Andreata Clusiaceae Clusia fluminensis Planch. & Triana — tree, fl (Nov), R. Andreata 306 Garcinia gardneriana (Planch. & Triana) Zappi — shrub, fl (Nov), 2 Kielmeyera membranacea Casar. — tree, fl (Jan—Feb), fr Jan- Feb, Jun-Jul), R. Andreata 370, 456, 714,725; T. Plowman 13929 Convolvulaceae Argyreia baronii Deroin — climbing, fl (Sep), R. Andreata 538 Ipomoea grandifolia (Dammer) O'Donell — climbing, fl (Jan), fr Jan), R. Andreata 367 Jacquemontia martii Choisy — climbing, fl (Apr, Dec), R. Andreata 411 Cucurbitaceae (V.L.G. Klein) *Wilbrandia glaziovii Cogn. — climbing, fl (Oct), fr (Sep), R. Andreata 129, 924 586 Elaeocarpaceae Sloanea garckeana K. Schum. - tree, fl (Jan), fr Jan), R. An- dreata 382 Erythroxylaceae (T. Plowman) Erythroxylum frangulifolium A. St.-Hil. — tree, fl (Feb, Jul, Aug- Sep, Nov), fr (Feb, Jul, Sep-Nov), R. Andreata 70, 169, 501, 641; C. Farney 362; T. Plowman 12860, 13922, 13926; V.Fereira 2239 Erythroxylum o Peyr, — shrub, fl Jan- PE fr Jan- Feb), ta 400; T. Plowman 13929, 13 Entram puchun A St.-Hil. — tree, fl (sterile), É ee Baken (L.S. Vale and A. Sousa) Acalypha brasiliensis MOL Arg. var brasiliensis — sub-shrub, fl (Aug, Dec), R. Andreata al, 73,919, 979 Actinost |. Arg.) Pax var. st Arg. — shrub, fl (Aug), fr (Aug-Sep), R. Andreata 77, 79, 686, 689, 966, 979, 982 Actinostemon concolor (Spreng.) Mull. Arg. var. obovatus Mul. Arg. — shrub, fl (Aug, Oct-Nov), fr Jul, Sep-Nov), H Andreata 162, 300, 536, 618 Cnidoscolus urens (L.) Arthur — herb, fl (Dec), R. Andreata 43 Croton compressus Lam. — shrub, fl (Dec-Jan), R. Andreata +l T4 A Ar Croton klotzschii (Diedr) Müll. Arg. — sub-shrub, fl (Dec), R. Andreata 58 Croton urticifolius Lam. — sub-shrub, fl (Dec), R. Andreata 26 ampia alata Müll. Arg. — climbing, fl (Dec-Jan), R. Andreata 61, 362 Dalechampia micromeria Bail. — ciimbing, R. Andreata Euphorbia comosa Vell. — sub-shrub, fl (Oct, Dec), fr (Oct), R Andreat fl (Nov), fr (Nov), Julocroton triqueter (Baill) Müll. Arg. — sub-shrub, fl (Dec), fr (Jan) ndreata 22 Joannesia princeps Vell. — tree, fl (sterile), R. Andreata 992 in inflata Müll. Arg. — shrub, fl (Jan), fr (Jan), V. Ferreira "moo Sc? Müll. Arg. — shrub, fl (Jan, Dec), R. Andreata Marta c Müll. Arg. subsp. dle SC? Arg.) rub, fl (Nov), fr (Nov), R. Andre cod eins (Nees) LM Johnst. — SS i a Nov), fi , Aug), R. Andreata 463, 957; T. Plowman 13927 Phyllanthus pa Müll. Arg. — sub-shrub, fl (Sep), fr (Sep), R. Andre Plukenetia serrata Ss M Gillespie — climbing, v. Romanoa tamnoides = Juss.) Radcl.-Sm. — Eege fl (Apr), fr (Apr), Andreata 8 e cd lid e Morong - tree, fl (sterile), R. An- dre sb Se Spreng. — shrub, fr (Apr), R. Andreata stilingia y Müll. Arg. - shrub, fl (Apr, Nov), fr (Apr), R Andreata 339, 675, 885 Tragia volubilis L. — climbing, fl (Jan), V. Ferreira 2117 [| [| £ al n.a H 1n LI FIR £T. TE A! 17 Flacourtiaceae (R. Marquete) Casearia leutzelburgii Sleumer — tree, fr (Aug), R. Andreata 222.127 o o Png tee 2n (sterile), R. Andreata 914 tree, fl (Aug-Sep), fr (Sep), R. Andreata 909, 925, 991; V. Ferreira 2238 Gesneriaceae (A. Chautems and T.C.C. Lopes) Codonanthe gracilis (Mart.) Hanst. — herb, fl (Nov), R. Andreata 340 tao prasinata (Ker Gawl.) Benth. — sub-shrub, fl (May, Jul), fr (Aug), R. E iin Son C. Farney 736 i r—herb, fl (Sep), fr (Sep), R. Andreata 62, 542: G. Martinelli 8520 Lauraceae (A. Quinet) Aniba brittonii Mez. — tree, fr (Apr, Jul), R. Andreata 442, 489; V. Ferreira 21 Ocotea aniboides Mez — shrub, fr (Nov), R. Andreata 941 Ocotea brachybotra (Meisn.) Mez - tree, fr (Nov), R. Andreata 943 Phyllostemonodaphne geminiflora (Mez) Kosterm. — shrub, fl (Jan), R. Andreata 389, 446 Lecythidaceae Cariniana legalis (Mart) Kuntze - tree, fl (sterile), H Andreata 381 Leguminosae (Caesalpinioideae) Bauhinia radiata Vell. — climbing, fl (sterile), R. Andreata 196 Bauhinia microstachya (Raddi) J.F. Macbr. - climbing, fl (sterile), R. An dre ata 164, 3 A R. Andreata 28, 112 Chamaecrista glandulosa (L.) Greene var. brasiliensis (L) H.S. Irwin € Barneby — sub-shrub, fl (Nov), fr (Apr, Nov), R. Andreata 651, 670; H.C. Lima 2069 Copaifera trapezifolia Hayne — tree, R. Andreata 164, 576 ib pens Tul. - tree, fl (Apr), fr (Apr), R. Andreata 394, rt hn wh H (Mer) lH Up, ALAS, Senna E (Benth) HS Irwin € Barneby — shrub, fr (Apr), R. Andreata 410 Senna macranthera (DC. ex Colladon) H.S. Irwin & Barneby — tree, fl (May), fr (May), C. Farney 729 tenuifolia (Vogel) H.S. Irwin & Barneby = shrub, fl (Apr), R. Andreata 410; H.C. Lima 3714 Leguminosae (Papilionoideae) Acosmium lentiscifolium Schott ex Spreng. - shrub, fi (sterile), R. Andreata 931; H.C. Lima 3734 Centrosema sagittatum (Willd.) Ridley — climbing, fl (Ma . Lima 2592, 2600 Chaerocalycandens ) var. pubescens (DC) Rudd - a fl (Oct-Dec), fr (Oct, Dec), R. Andreata 36, 128, 17 Dalbergia Ee (Vell) Britt. small tree, H.C. Lima ^ 5 Desmodium affine Schltdl. — herb, fl (Apr), R. Andreata 447 Desmodium incanum (Sw.) DC. — herb, fl (Dec), fr (Dec), R. oo May), fr Andreata 25 Galactia striata Jacq.) Urb. — herb, fl (Jan), fr Jan), R. Andreata 66 | Fi * 4t | EJ ta Al IT Andreata et a ; Lonchocarpus virgilioides (Vogel) Benth. — tree, fl (Sep), fr (Sep), R. Andreata 141, 529, 724, 726, 904, 916 d firmum (Vell. Benth. — tree, fl (sterile), R. Andreata , 685 E SH (Vell) C. Stellfeld — tree, fl (sterile), R. Andreata Machaerium om Vogel - climbing, fl (sterile), R. ndreata 891 Machaerium pedicellatum Vogel, small tree, fl (sterile), R Machaerium violaceum Vogel — climbing, fl (sterile), R. An- dreata 899; H.C. Lima 2596 Pterocarpus rohri Vahl — tree, fl (sterile), R. Andreata 895 oe scabra Vogel — herb, fl (Oct), R. Andreata 135 Vi sub-shrub, fl (May), C. Farney 733 Zornia dites Sm. — sub-shrub, fl (sterile), R. Andreata 887 Leguminosae (Mimosoideae) Acacia plumosa Lowe — shrub, fl de al ~ Apr, Nov), R. Andreata 600, 884, 964; H.C. Lim Acacia mikanii Benth. — shrub, fl gek 4 See 575,616 Albizia polycephala (Benth.) Killip — tree, fl (Jan), fr Jan, May), a ; 550 diio es colubrina (Vell) Brenan - tree, fr (Jan), R Andreata 34 Inga Pus Ee — tree, L.A.Rodrigue ee Penn. - shrub, fl Jul- us fr Jul-Oct), H.C. 26034, 3991; R. Andreata 87, 125, 449, 451, 509 Inga cada Mart. - tree, fl (Jan—Feb), fr (May), R. Andreata 397, 583, 639, 1003 ), R. Andreata 0 Mimosa bimucronata (DC.) Kuntze — shrub, fl (Apr), fr (AprJun), R. Andreata 409, 452, 902 Mimosa extensa Benth. — climbing, fl (Nov), H.C. Lima 3733 Mimosa velloziana Mart. — shrub, fl (Apr), fr (Apr, Jul), R. An- , 66 Piptadenia gonoacantha (Mart.) J.F. ibid — tree, fl (Feb), fr (May), R. Andreata 586; H.C. Lim o paniculata Benth. — tree, i ran fr (May), R. An- ata 2 393, ud gu e SCH a371 D. mao PR AA DAA | ima - tree, fr (May), R. Andreata 5 676 Loganiaceae Strychnos acuta Progel — sub-shrub, fl (Sep), dreata 421, 944 Strychnos trinervis (Vell) Mart. — shrub, fr (Jan), R. Andreata fr (Apr), R. An- Loranthaceae Struthanthus marginatus (Desr.) Blume, hemi-parasite, fl (Oct), fr (Oct), R. Andreata 115, 117 Lythraceae Cuphea carthagenensis (Jacq) J.F. Macbr. — herb, fl (Dec), R. Andreata 39 AA E | LR AT Tel A I J A Am Amonmiarigida (A. Juss) W.R. Anderson - climbing, f (Jan), fr un), R. Andreata 373, 708 587 Lag, **Bunchosia glandulifera (Jacq.) H.B.&K. — shrub, fr (Apr), R. Andreata 664 o maritima (Vell) J.F. Macbr. — shrub, fl (Oct-Dec), fr R. Andreata 41, 111, 166, 558, 8 0 een bicolor A. d - climbing, fr (Jan), V. Ferreira 2100; R. Dee pacos (A. Juss.) A. Juss. — shrub, fl (Sep), fr (Jul), R. Andr E H SC sericea Se A. Juss. — climbing, fl (Jan), R. An- dreata 390 *Heteropterys ternstroemiifolia A. Juss. — climbing, fr (Jul), R Niedenzuella acutifolia (Cav) W.R. Anderson — shrub, climb- ing, fl (Aug—Dec), fr Jan, Aug), R. Andreata 152,155, 322, 345, 510 ae Malvaceae (M.G. Bovini) Abutilon bedfordianum (Hook. A. St.-Hil. € Naudin — shrub, fr (May), R. Andreata 524, 590 Pavonia sepium A. St.-Hil. - sub-shrub, fi (Apn, fr (Apr), R Andreata 431 Sida rhombifolia L. — sub-shrub, fl (Dec), R. Andreata 15 Marcgraviaceae Norantea brasiliensis Choisy — shrub, fl (Nov), fr Jan), R. An- dreata 309, 401, 986 Melastomataceae Clidemia hirta (L) D. Don — shrub, fl (Nov), fr Jul, Sep), R. ta 188, 492 Miconia cinerascens Miq. — shrub, fl (Apr-May, Sep, Nov), fr (May, c Nov), R. Andreata 138, 184, 414, 493, 933; C. Farne Miconia staminea (Desr.) DC. — shrub, fl (Nov), fr (Nov), R. Andreata 186 Ossaea marginata (Desr.) Triana — shrub, fl (Sep-Nov), R. Andreata 144, 187, 515 Tibouchina corymbosa (Raddi) Cogn. - shrub, fl (Nov—Jan), R. Andreata 59, 317, 377, 901 Meliaceae Guarea quidonia (L.) Sleumer, fl (Nov), R. Andreata 189; V. Ferreira Trichilia alternans DC. — tree, fl (Oct-Nov), R. Andreata 143, 170 Trichilia casaretii DC. — tree, fr (Apr, Jul), R. Andreata 423, 450 Trichilia elegans A. Juss. subsp. ijs Penn. — shrub, fl (Nov), fr (Jul), R. Andreata 308, 4 Trichilia pseudostipularis (A. Juss.) E — shrub, fl (Jan, Nov), fr (May), R. Andreata 402, 594 Trichilia silvatica DC. — shrub, fl (Nov), fr (Aug), R. Andreata f Menispermaceae (J.M.A. Braga) Chondrodendron platyphyllum (A. St.-Hil.) Miers — climbing, fr Apr, R. Andreata 419 *Hyperbaena oblongifolia (Mart) Chodat — climbing, fl (Jul, Sep), fr (Sep), R. Andreata 502; V. Ferreira 2235 Moraceae (J.P.P. Carauta) Cecropia glaziovii Snethl, — tree, fl (sterile), R. Andreata 937, mmm, UO “i A 588 Coussapoa curranii J.M. Blake — tree, fl (Jul), R. Andreata 457 Dorstenia arifolia Lam. — herb, fl (an, Nov), R. Andreata 183, 95, 646; V. Ferreira 2124 Dorstenia cayapia Vell. — herb, V. Ferreira 2102 orstenia turnerifolia Fisch. & C.A. Mey. — herb, fl (Nov), R. Andreata 438; V. Ferreira 2111 Ficus arpazusa Casar. — tree, fr (Nov), R. Andreata 612 Ficus enormis (Mart. ex Miq.) Mig. — T. fl (May, Jul), fr (May, Jul), R. Andreata 478, 593; C. Farn Sorocea hilarii Gaudich. — tree, fl un fr (May), R. An- dreata 436, 555, 571; V. Ferreira 2112 i irn (G.M. Barroso, M.C. Souza and M. Sobral) mpomanesia e (O. Berg) Nied., small tree, A rile), R. Andre Eugenia jurujubensis e. - b fl (Oct-Nov), fr ns ~ Oct), R. Andreata 148, 149, 190, 324, 332, cl Eugenia sulcata Spring. ex Mart. — tree, R. Andre e Eugenia mE L. — tree, fr (Apr), R. Andreata a 495, 967, 9/4,9 m en Ma gestasiana (Cambess.) Legrand - shrub, fl (Nov), r dreata 434 = QR Gomidesia sonderiana O. Berg — shrub, fl (Nov), R. Andreata Marliera racemosa (Vell) Kiaersk. — tree, R. Andreata 4/3 Myrcia insularis Gardner — shrub, fl (Jan), R. Andreata 387, 940 Myrcia racemosa (O. Berg) Kiaersk. - tree, fl (sterile), R. An- dreata 926 Myrcia richardiana (O. Berg) Kiaersk. — tree, fl (Nov), fr Jan), R. Andreata 388 LAI |) E Berg => shrub, fr Uan), R. Andreata 606, 968, 976 Myrciaria glazioviana (Kiaersk) G. Barroso ex Sobral — shrub, r ), R. Andreata 499, 506 Psidium cattleyanum Sabine - tree, R. Andreata 622, 962 Indet.1 — tree, fl (sterile), R. Andreata 978 Myrsinaceae (M.F. Freitas) Myrsine coriacea (Sw.) R. Br. ex Roem. - tree, fl (sterile), R Andreata 939 Nyctaginaceae (C.F.C. Sa) Bougainvillea spectabilis Willd. — shrub, fl (May), R. Andreata G (Vell) Reitz — shrub, fl Jul-Sep), fr (May, Sep), R. Andresta 75, 80, 108, 486, 540, 559, 688, 698, 700, 950, 975, 989, 990; V. Ferreira 2240 chnaceae Ouratea parviflora (DC. Baill. — shrub, fl (Sep), fr (Sep), R. Olacaceae Heisteria perianthomega (Vell) Sleumer - tree, fl (sterile), R Andreata 720 Oxalidaceae Oxalis barrelieri (L) Smali. — herb, fl (Aug, Dec), R. Andreata 13,09 [EE PE D LA: L Ge TË IE AAA A sub-shrub, fl (Feb), R. Andreata 643: T. Plowman 13928 Oxalis sepium A. St.-Hil. — sub-shrub, fl (Nov), R. Andreata 305 Passifloracea *Passiflora racemosa Brot. — climbing, fl (May, Jul-Aug, Oct- Jan), fr (May, Jul), R. Andreata 31, 121, 198, 459, 588, 921 Phytolacaceae Gallesia integrifolia Goreng) Harms - tree, fi (sterile), R. Andreata 39 Microtea paniculata Moq. — herb, fl (Jan), V. Ferreira 2094 Petiveria alliacea L. — sub-shrub, fl (Aug, Dec), R. Andreata 20, 82; V. Ferreira 2096 Picramniaceae (J.R. Pirani) *Picramnia iibi il — shrub, fl (Jul), R. Andreata 458 Pi (E.F. G | D.M. Ferreira) Peperoni corcovadensis Gardner - epiphytic, fl (Oct), R. ta 157 Peperomia incana (Haw.) Hook. — herb, fl (Sep, Nov), fr (Feb), R ndreata 315, 649; T. Plowman 13934; G. Martinelli 8528 Peperomia tetraphylla (Forst) Hook. & Arn. — epiphytic, fl (Nov), Andreata 201 Piper amalago L. var. SH (Jacq) Yuncker - shrub, fl (Dec), R. Andreata 168, 9 e SE Kunth — im fl (Feb), fr (Feb), T. Plowman Plumbaginacea Plumbago enee L. - climbing, fl (Nov), R. Andreata 325 Polygonaceae Coccoloba confusa R.A. Howard - shrub, fl (Sep), R. Andreata 531 Ruprechtia laxiflora (Meisn.) Kuntze — tree, R. Andreata 952 Ruprechtia lundii (Meisn.) Kuntze — tree, fl (Sep), R. Andreata 718 Casar) Coccuci - tree, R. Andreata 918 a Triplaris scandens Portulacaceae Portulaca mucronata Link — herb, fl (Nov), R. Andreata 338, 239 Talinum GE X Willd. — herb, fl (Oct-Nov), fr (Oct-Nov), R. Andreata 134, 177 Talinum racemosum (L.) Rohrb. — herb, fl (Aug), R. Andreata 6 Rhamnaceae Reissekia smilacina Endl. - climbing, fl (Nov), R. Andreata Rubiaceae ( M. Gomes) Alseis sp. — tree, fl (sterile), R. Andreata 678 Alseis CHEN Schott - tree, fl (Sep), fr (Jan-Sep), R. An- dre Alseis miu SH —tree, fl (Sep), fr Jan-Sep, R. Andreata 358, 50 ee Ee Rich. — shrub, fl (Feb), fr (Feb), R. An- 40; T. Plowman 13925 dia area sp. — o fl (sterile), R. Andreata 972 *Coussarea capitata (Benth.) Benth. € Hook. — tree, fl (Oct), R. Andreata 137 Andreata ef al e Floristi Cout j Ira () IK Schum fi ek Schum - tree, fl (Apr), fr (Apr-May), R. Andreata 661, 691, 697 Faramea sitpulacea (Cham. & Schltdl.) DC. — shrub, fl (May, Sep), f = —Jul), R. Andreata 469,491, 557; C. Farney 737 Lem M" *Fara - tree, fl (Dec), R. Andreata 54; V. Ferreira 2110 enn Pozvan Cham. — herb, fl (Nov), R. Andreata 176 Posoqueria acutifolia Mart. — shrub, fr (Jan), R. Andreata 384 Psychotria brachyceras Müll. Arg. — shrub, fl (Oct-Dec), R. dreata 30, 116, 171 *Psychotria stenocalyx Müll. Arg. — shrub, fl (Oct-Nov), fr (Apr, , Nov), R. Andreata 153, 418, 604; C. Farney 369 Randia armata (Sw.) DC. — shrub, fr (Apr), R. Andreata 441 Richardia E Gomes - herb, fl Jul), R. Andreata 16 Rudgea minor (Cham.) Standl. subsp. o (Benth) Zappi a ps (Mar), H.C. Lima and FCP Garcia 5027 Rudgea ia Müll. Arg. — shrub, fl Er fr (Sep), R. A 491,5 iE item Benth. — shrub, fl Jan, Sep, Nov), fr (Jan, Ap , Jul), R. Andreata 396, SCH 471, 474, 512, 572, 951; T rney 61; V. Ferreira 210 Rudgea ta Müll. Arg. — eh fl (Oct), fr (Apr), R. Andreata 118 Simira sampaiona (Standly) Steyerm. - tree, fl (Oct), R. An- dreata 124, 683, 723 Calo | ^ oC H —tree, fr (May, Jul, Nov), R. Andreata 195, 460, 574 Rutaceae (M. Emmerich and J.R. Pirani) E rubra A. St.-Hil. - shrub, fl (May, Jul, Nov-Dec), , Nov), R. Andreata 53, 101, 467, 596; C. Farney 370 Concha ovatus A. St-Hil. — shrub, fl (Nov), C. Farney _ fr (May, "un ovata (A. St-Hil) Tul. small tree, fl (Nov-Dec), fr (Apr), : ta 197, 442, 158 Esenbeckia febrifuga (A. St.-Hil) A. Juss. ex Mart. — tree, fr (Jul), Andreata 48 Galipea jasminiflora (A. St.-Hil.) Engl. — shrub, fl (Apr), fr (Apr), 416 Neoraputia alba (Nees €: Mart.) Emmerich - tree, fr (Apr), R. Sapindaceae (G.V. Somner) Allophylus edulis (A. St.-Hil.) Radlk. — shrub, fl (Apr), R. Andreata 444 Allophylus heterophyllus Radlk. — shrub, fl (Oct), fr (Apr), R Andreata 130, 440 Allophylus laevigatus Radlk. — shrub, fr (Oct), R. Andreata 109 Allophylus semidentatus Radlk. — tree. fl (Sep, Nov), R. Andreata Cupania sp. — tree, fl (sterile), R. Andreata 934 Cupania racemosa Radlk. — tree, fr (Jul), R. Andreata 374, 472,728 Paullinia meliifolia A. Juss. — climbing, fr (Apr-May), R. Andreata 445, 598 Serjania corrupta Radlk. — climbing, fr (May), R. Andreata 589 Serjania cuspidata Cambess. — climbing, fr (Apr), R. Andreata 703 589 Serjania fuscifolia Radlk. — climbing, fl (Apr-May), fr (Apr), R Urvillea stipitata Radlk. f. stipitata — climbing, fr (Apr), R. Andreata 668 Sapotaceae (M.H.D. Monteiro) Chrysophyllum flexuosum Mart. — tree, fl Jan, Apr), fr (Apr), H 424: V. Ferreira 2 Pouteria subsessilifolia Cronquist — tree, fl (Aug, Oct), fr (Oct), ta 133 Pouteria psamophila (Mart.) Radlk. — tree, fl (Oct), R. Andreata 161 Solanaceae (L.F.d'A.Carvalho) Athenaea anonacea Sendtn. — shrub, fl (sterile), R. Andreata } PINO m fr (Oct), R. Andres 114 o uniflora (Pohl) D. Don - shrub, fl (Aug), R. Andreata 00 var. fasciculata, small tree, SR schottianum Sendtn. — shrub, fl (Sep), R. Andreata 522 Dyssochroma viridiflorum (Sims) Miers — shrub, fl (Aug-Sep), R. Andreata , 60 *Metternichia principis Mikan var. principis, small tree, fl (Feb, ¡fr (Jan, Jul, Nov), R. Andreata 352, 480, 508, 638, 677; T. Plowman 13923 Solanum alternatopinnatum Steud. — climbing, fl (Apr), fr (Apr), Andreata 41 olanum americanum Mill. var. americanum — sub-shrub, fl (Nov), fr (Nov), R. Andreata 180 Solanum arenarium Sendtn. — shrub, fr (Jan), R. Andreata 404 Solanum did Dunal — shrub, fl (Aug), fr (Aug-Nov), R. Andreata 71, 1 Solanum caavurana Vell. — shrub, fl (Nov), R. Andreata 328, 608 Solanum incarceratum Ruiz & Pav. — shrub, fl (Dec), R. An- dreata 40 Solanum megalochiton Mart. var. megalochiton — climbing, fr (May), R. Andreata 597 Solanum melissarum Bohs — shrub, fl (Sep), R. Andreata 936 Solanum swartzianum Roem. & Schult. var. swartzianum shrub, fl (Oct), R. Andreata 133 Solanum sp.1 — sub-shrub, fr (Jul), R. Andreata 490 Theophrastaceae Clavija spinosa (Vell) Mez — shrub, fr (Apr), R. Andreata 437 Trigoniaceae (E.F. Guimaráes) Trigonia eriosperma (Lam) Fromm & Santos — sub-shrub, fl (Oct), fr (Aug), R. Andreata 76, 110 Turneraceae bi edi Il. — sub-shrub, fl Jun, Jul, Sep), fr (Aug, Sep, 1, R. Andreata 84, 181, 497, 511,710 Ulmaceae (J.P.P. Caraut Celtis brasiliensis (Gardner) Planch. — shrub, fl (Sep), fr (Jan, , R. Andreata 516, 913; H.C. Lima and J.F.C.P. Garcia 5026 bk E 590 Urticaceae Urera mitis (Vell) Mig. — shrub, fl (Apr), R. Andreata 429 Verbenaceae Aegiphila mediterranea Vell. — tree, fl (Aug), R. Andreata 78 Lantana sp. — sub-shrub, fl (Aug), fr (Sep), R. Andreata 74, 498 Stachytarpheta cf. maximilianii Schauer — herb, fl (Dec), R Andreata 23 Stachytarpheta polyura Schauer — herb, fl (Oct), R. Andreata 46 1 Vitex polygama Cham. — shrub, fl (Sep, Nov), fr (Sep, Nov), R. Andreata 331, 543 Viplacsae 4 ZC | PoE hri fr (Aug, Nov), R. Andiésia 88, 532, 949 MONOCOTYLEDONS Agavaceae Furcraea gigantea Vent. — herb, fl (Feb), T. Plowman 12855 Alstromeriace Alstroemeria e Jacq. - herb, fl (May, Jul, Aug), fr (May), R. Andreata 102, 412, 414, 462, 570, 573; C. Farney 732 Amaryllidaceae Hyppeastrum reginae Herb. — herb, fl (Apr, Jul, Sep), R. Andreata 514, 946 Araceae (SJ. Mayo and M.A.C. Nadruz) Anthurium coriaceum (Graham) G.Don - herb, fl (Sep), R Andreata 343, 539 RE EH ~ herb, fl (Feb, Nov), S. Mayo 607; V. Ferreira Anthurium Ce ex — herb, (Feb, Nov), R. Andreata 206; S. Mayo 606; V. Ferreira 2119 Anthurium um (Aubl) G.Don - epiphytic, fl (Dec), R Andr rea a 0 (Aubl.) Engl. subsp. scandens — epiphytic, fl (Nov), R. Andreata 207 — herb, v.v. Philodendron bipennifolium Schott — herb, S. Mayo 603 Philodendron corcovadense Kunth — herb, v.v. Philodendron cordatum Kunth — herb, v.v. Philodendron crassinervium Lindl. — herb, S. Mayo 605 *Philodendron speciosum Schott ex Endl. - herb, S. Mayo 602 Arecaceae Attalea sp. — tree, fi (Jan), R. Andreata 391 Syagrus romanzoffiana (Cham) Glassman - tree, fl (May), fr May), R. Andreata 553, 687, 690, 692, 693, 695, 699 Bromeliaceae (G. Martinelli) Aechmea nudicaulis (L) Griseb. var. nudicaulis — epiphytic, fl (Nov), R. Andreata m | he Gerbe ramosa Mart é ex Schult var. ramosa — Sa? fl Jul), R. Andreata 465 Aechmea sphaerocephala Baker — herb, fl (Sep), G. Martinelli 8525 Alcant lazioviana (Lem.) Leme — herb, fl (S 1;G. Martinelli 8527 (cultivated) be, end (Sims) Lindl. var. pyramidalis — herb, fl ec), R. Andreata 65, 205 Be KE Bertol. — herb, fl (Sep), G. Martinelli p), R. Andreata 8523 Neoregelia ampullacea (E.Morren) L.B.Sm. — herb, fl Gan), V. Ferreira 2114: G. Martinelli 8515 Neoregelia sarmentosa (Regel) L.B.Sm. — herb, fl (Nov), R. Andreata 20 Pitcairnia staminea Lodd. — herb, fl (Apr), R. Andreata 878 Tillandsia araujei Mez var. araujei — R. Andreata 896; G. Martinelli 8521; T. Plowman 13933 Tillandsia dura Baker — herb, fl (Apr, Nov), R. Andreata 321; G. Martinelli 8519; T. Plowman 13924 Cannaceae (J.M.A. Braga) Canna glauca L. — herb, fl (Nov), fr (Nov), R. Andreata 517 melinaceae Dichorisandra thyrsiflora J.C. Mikan — herb, fl (Apr, Nov), R. 6 * Siderasis fuscata (Lodd.) H.E. Moore — herb, V. Ferreira 2104 Tradescantia zebrina Bosse — herb, v.v mn, Cyperaceae (C.S. Muniz) Bulbostylis capilaris und herb, fl (Nov), R. Andreata 313 Cyperus coriifolius Boeck — herb, fl (Sep), R. Andreata 533 Cyperus diffusus Vahl — herb, fl (Dec), R. Andreata 18 Trilepis eximia (Clarke) Pfeiff. — herb, fl (Sep), R. Andreata 534 Dioscoreaceae Dioscorea sp.1 — climbing, H (Nov), H Andreata 613 Dioscorea sp.2 - climbing, fl Jan), fr Jan), V. Ferreira 2120, 2122 Dioscorea sp.3 — climbing, fl (Nov), R. Andreata 150 Dioscorea glomerulata Hauman — climbing, fl (Jan), R. An- dreata 406 Heliconiaceae (J.M.A. Braga) Heliconia laneana Barreiros var. laneana — herb, fl (Oct), V. Ferreira 2116 Herreriaceae (R.C. Lopes Herreria salsaparrilha Mart. — climbing, fl (sterile), R. Andreata Iridaceae Neomarica sp. — herb, fl (Apn, R. Andreata 433 Marantaceae (J.M.A. Braga) Calathea eichleri R.H. Petersen — herb, fl (Nov), fr (Jan), R. Andreata 680 Calathea truncata Lodd. — herb, fl (Nov), R. Andreata 45, 621 Maranta foliosa Kàrn. — herb, fl (Nov), fr (Apr), R. Andreata 342, 578 Ei (L. Toscano and F.C. Pinheiro) olous, fl Jul), T.B. KE Ix € E E nica rupico Cro 3t 53728; n Mowe ho 01 ree ee polyphyllum (Vell) Pabst ex F. Barros — herb, fl (Sep), fr (Sep), R. Andreata 537 Andreata et al., Floristic diversit the M Alto Mouráo, Brazil 591 dinis Pi d Barb. Rodr. - herb, fl (Nov), R. Poaceae (A.G.Burman) Andre Bambusa sp. — herb, fl (Oct), R. Andreata 920 tendum cut Lindl. - epiphytic rupicolous,fl (Nov, Dec), Ichnanthus sp. — herb, fl (Oct), R. Andreata 126 7; F. Pinheiro 95 Olyra brasiliensis Bertol.) Spreng. — herb, fl (Oct), R. Andreata E Dese (Lindl) Fenzl — rupicolous, fl (Oct), F. 156 Pinheiro 181 Pennisetum setosum (Sw.) Rich. — herb, fl (Oct), R. Andreata F. Pinheiro am Octomeria alpina Barb. Rodr. — epiphytic, fl (Sep), 145 173 Stipa latifolia (L) Raspail - herb, fl (Oct-Nov), R. Andreata Oncidium fimbriatum Lindl. — epiphytic rupicolous, fl (Oct), 319 F Pin 0 ium pu n um Lindl. — epiphytic, fl (Apr), F. Pinheiro 202 et grobyi Lindl. - rupicolous, fl (Feb, Mar), F. Pinheiro Smilacaceae Smilax sp. — climbing, fr (Oct), R. Andreata 140 Smilax quinquenervia Vell. — climbing, sterile, R. Andreata "Pars pardipes Rchb. f. — epiphytic, fl (Mar), F. Pinheiro Smilax subsessiliflora Duham. - climbing, fl (Sep), fr (Apr), R. Andr eata 95, 159, 167, 200, 435, 518, 527 Pleurothallis iie Rchb. f. - epiphytic rupicolous, fl (Man), E Pinheiro 143 Velloziaceae | Sophronitis cernua ZU - epiphytic, VV. Vellozia candida J.C. Mikan — herb, fl (Apr), R. Andreata 881 ACKNOWLEDGMENTS The authors of this work acknowledge the contributions of the professors Geisa Lauro Ferreira and Valério Ferreira and students of the Laboratório de Angiospermas da Universidade Santa Úrsula for their help with field work, Lucy Pinto Hack of the Pontifica Universidade Católica do Rio de Janeiro for preparing the water balance data, Paulo Botelho de Macedo and Paulo Cesar Silvestre Onofre for the graphic layout, the special- ists for their identifications and/or revisions of the taxa listed, the Universidade Santa Úrsula and Instituto de Pesquisas Jardim Botánico do Rio de Janeiro for assistance in various stages during this work and to the Conselho Nacional de Desenvolvimento Científico e Tecnológico for the research and productivity grants provided to the authors Regina Helena Potsch Andreata and José Fernando A. Baumgratz. REFERENCES Araujo, D.S.D. and A.M.N. ViLaca. 1981. Avaliação da cobertura vegetal remanescente de Itaipu. In: LM. Kneip, L. 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Revisáo tax nero Astronium e revalidacao do ceae). Tese de Mestrado, vedado ae eens Campinas. Pp. 1- 178. THoRNTHWATIE, CW. and J.R. MATHER. 1955. The water balance in climatology. Ceterton 8:1-104, LA FAMILIA NYMPHAEACEAE EN EL ESTADO DE NUEVO LEÓN, MÉXICO Carlos Velazco-Macías Rahim Foroughbakhch Pournavab, Marco A. Alvarado Vázquez, y Glafiro J. Alanis Flores Parques y Vida Silvestre de Nuevo León Calle París 343, Col. Mirador, C.P. 64070 Facultad de Ciencias Biológicas, U.A.N.L. Monterrey, Nuevo León, MÉXICO an Nicolás de los Garza carlos.velazcoegmail.com Nuevo León, MÉXICO RESUMEN Tixevebtaen ] € 1 = = " fi er a =] PE | ] MT T A Ce A 1 de Aa EI 1 a tende a a llenar ea vacíos in vnde una revisión de | D de la familia Nvmph sta entidad. A través de una revisión bibliográfica y jemp i gi iderad históricos, además a t de visitas SE d hábit ] I d famil 1 1 à Jo oeta Pami] £ BAH OTA Nuevo León, l (Nuphar y ee y dos especies (Nuphar advena y Nymph ampla) en 2 ee p el a E y3 NEM para el | trabajo d tró d j N. elegans) o E E [ DN A la 1 ladad N 1 UA A A ] ] Aad " ] d t E SEL | 1; L i D Ed D aa E ] 1 1 1:4 ] s] A de S lE E 1 = ] 3 1 1 s n J el x WW AT 1 CH a a | E E 3 3 J ] T 1 LE? ] 1 J TY J 1 J J ] 1 1 os | AT 7 e I I I , Se estima f 1 | J Jn] J 1 10 anos L D E r + ABSTRACT There are EH in our existing l ledge of the floristics of the State of Nuevo Leon in ME Mexico. e pape pe the 1 1 1 1 E e) AT. 1 Dal: q NEG ] 1 1 2 d A e field work at 17 dno New lor ind herbari tudies d ttwog (Nuphar and dL i for Nuevo e Two new state records are documente . Nymph den splemnnc and N me cand, each t la | lit; am mhnen nt tt new localities while it is now absent at three historical pis ee on ci specimens. Aso. Nuphar advena was eradicated uae one of the SE sites. Given n rate se habitat destruction and loss of wetlands in Nuevo León, we estimate members of the +l AARAA qa ax O years or less. INTRODUCCIÓN El estado de Nuevo León no cuenta con un listado florístico actualizado, para el año de 1965, se regis- traban un total de 1,296 EDO entre plantas nativas, cultivadas y adventicias, distribuidas en 657 géneros y 148 ee de a vasculares (Rojas-Mendoza 1965); otras estimaciones proponen cerca e 2,400 ies d ı vascular (Alanis-Flores et al. 1996). De manera puntual, la vegetación acuática o semiseudtira ha recibido poca o ninguna atención en los estudios florísticos, se conoce solamente un estudio de la Presa Rodrigo Gómez y sus afluentes en el municipio de Santiago (Moreno 1984). Para la familia Nymphaeaceae en Nuevo Eom solamente existen dais en la literatura de Nymphaea odorata y N. mexicana, ambas son como especies Rojas Mendoza 1965), sin que existan registros de herbario de estas especies hasta di See La única pe de la familia Nymphaeaceae que ha sido publicada de manera formal con respaldo de un ejemplar herborizado es N. ampla (Lot et al. 1986; Novelo y Bonilla 1999). De estados vecinos como Coahuila, se reporta solo N. ampla (Villarreal-Quintanilla 2001; Villarreal-Quintanilla et al. 2006), mientras que para Tamaulipas se reportan Nuphar luteum, Nymphaea ampla, N. elegans N. jamesoniana y N. amazonum (Lot et al. 1986; Martínez y Novelo 1993; y Mora-Olivo y Villaseñor 2007), y en el estado de Texas, en Estados Unidos de Norteamérica se reportan 4 especies de Nymphaea: N. ampla, N. mexicana, N. odorata y N. elegans, además de Nuphar luteum subsp. macrophyllum (Correll y Johnston 1979; Wiersema y Hellquist 1997). Cabe mencionar que para el caso particular del género Nuphar, ha existido una gran controversia en J. Bot. Res. inst. Texas 2(1): 593 — 603. 2008 SM JO al of t tani ti Texas 2( lo referente a la aplicación del epíteto específico, ya que N. luteum es de origen europeo, mientras que a través de trabajos genéticos y moleculares las especies americanas han mostrado una separación de las especies del viejo mundo, se aplica el nombre N. advena a los ejemplares reportados en trabajos recientes (Wiersema y Hellquist 1997). El objetivo del presente trabajo es d tar los registros de la familia Nympl en el estado de Nuevo León, así como conocer el rango de distribución actual de las m presentes como una contribución al conocimiento florístico. METODOLOGÍA Área de estudio El estado de Nuevo León se localiza en la part te de México, entre los 98? 17' y 101? 07' de longitud oeste, y los 23? 06' y 27? 50' de latitud norte. Limita al norte con los Estados Unidos de América y los estados de Coahuila y Tamaulipas; hacia el poniente limita con los estados de Coahuila, San Luís Potosí y Zacatecas, mientras que hacia el sur colinda con San Luis Potosí y Tamaulipas, con este ultimo también comparte la totalidad de su límite oriental (Fig. 1).Su superficie es de 64, 081.94 km”, las elevaciones van desde los 200 a los 3,700 m sobre el nivel del mar. Se presentan porciones importantes de tres regiones fisiográficas de México, la Sierra Madre Oriental, la Gran Llanura de Norteamérica y la Planicie Costera del Golfo (Alanís-Flores et al. 1996). Predominan los climas semisecos extremosos, la precipitación pluvial es variable, el promedio general va desde los 300 a los 600 mm de precipitación anual. Los climas secos y semisecos se distribuyen principalmente de la región nororiental y en la región suroc- cidental, ambas áreas están separadas por la Sierra Madre Oriental, siento en esta última donde además se presentan climas, semicálidos, templados y semifrios en la zona centro y sur de la entidad (INEGI 1986). La vegetación se compone principalmente de matorrales desérticos y semidesérticos de diversa composición florística, en la región de la Sierra Madre se presentan bosques con distintas asociaciones de encinos, pinos, y coníferas. En menor grado se presentan sucesiones edafológicas como vegetación gypsófila o vegetación halófita; la vegetación acuática solamente se encuentra registrada como bosque de galería en algunas corrientes de agua permanentes del estado (Treviño-Garza et al. 2001). Si bien existen en Nuevo León áreas con hábitats de tipo lótico, como escurrimientos temporales en el área de la Sierra Madre Oriental y vegetación riparia, son pocos los ríos que actualmente mantienen el flujo de SEH a lo largo del año; por otra pue los ecosistemas lénticos que son el hábitat potencial para la familia Nymy , Son sumamente escasos , algunos han desaparecido en los municipios de Galeana y mene We por el uso desmedido del recuso hídrico en las actividades agropecuarias, otros como en el caso del Cañón de la Huasteca en el municipio de Santa Catarina, han sido canalizados para el abastecimiento de la zona metropolitana de Monterrey, dejando en la actualidad solo unos cuantos sitios restantes en el estado para el desarrollo de comunidades de vegetación acuática y semiacuática. Métodos Primeramente se realizó una revisión y consulta bibliográfi la básqueda de registros de la familia Nymphaeaceae en Nuevo León, además de conte en las bases de datos de ee E herbarios: TEX, LL, UNL, MO, NY y el herbario de George B. Hinton para la búsqueda de de la familia. Se realizó trabajo de campo en las diversas localidades q ]taron de la búsqueda de ejemplares de herbario para corroborar la presencia de las especies en la actualidad, en total se realizaron visitas a 17 localidades con hábitat potencial en di municipios de Nuevo León para documentar los sitos de distribución de las especies (Tabla 1). Las diagnosis de la familia, géneros y especies, así como la identi- ficación de los ejemplares colectados están basadas en Zomlefer 1994; Novelo y Bonilla 1999; Calderón y Rzedowski 2001; Correll y Johnston 1979; Wiersema y Hellquist 1997. El material colectado, fue herborizado y depositado en el herbario de la Facultad de Ciencias Velazco et al 595 249 T LEYENDA TT Presas Escala 1:2,300,000 Corriente de agua 9 $ ' __ | Limites municipales Curvas a niv Kilómetros Monterr E 29" ago 101° 100° 99° git Fic 1 ih 506 I [| fal Das H In LI rr f Texas 2(1) Biológicas de la Universidad Autónoma de Nuevo León (UNL) siguiendo las recomendaciones de Lot y Chiang, 1986 para el montaje y herborización de ejemplares botánicos. RESULTADOS En el estado la familia está representada por 2 géneros (Nymphaea y Nuphar) y 4 especies (Nymphaea ampla, N. elegans, N. mexicana y Nuphar advena), las cuales se distribuyen en 4 municipios en la Planicie Costera del Golfo y los límites con la Sierra Madre Oriental (Figs. 1-3). Los registros de herbario muestran que se han extirpado diversas poblaciones, Nuphar advena ha desparecido del municipio de Paras, mientas que Nymphaea ampla se ha eliminado de diversos hábitats en los municipios de Fa Lampazos de Naranjo y Apodaca (Tabla 1). Las 17 localidades visit tran en la Tabla 1, en donde se muestra también el estado actual de cada una de ellas, y la presencia o ausencia de la lia Familia Nymphaeaceae Salisb. Plantas herbáceas, acuáticas, perennes, rizomatosas, a veces acaules; hojas simples, usualmente enteras, alternas, con venación bai o pinnada, NN directamente del rizoma, pace pecioladas, + láminas flotantes del agua; s solitarias, axilares, acti ficas, perfectas, hipog- inias, periginias o ice veces epiginias, comunmente Eudes y vistosas, flotantes o sobresaliendo del espejo de agua, provistas de pedúnculos largos; cáliz usualmente de 4 a 6 sépalos, libres, algunas veces petaloides, imbricados; corola con pétalos de 3 a muchos, dispuestos numerosas series, los cuales a menudo no se diferencian claramente de los estambres, transformándose de forma gradual en esta- minodios o estambres, algunas veces con nectarios abaxiales, de colores vistosos como amarillo, rojo, azul o blanco, imbricados; androceo compuesto de 3 a numerosos gd ae y arreglados de manera espiral, filamentos lineares o petaloides, pobremente teras, libres o unidos a los pétalos, anteras con dehiscencia longitudinal intrusa o extrosa; gineceo apocárpico o sincarpico, con 3 o más carpelos, ] n cada lóculo, anátropos, placentación lateral o lamelar, estilos ausentes, estigma(s) unidos y formando un disco con el mismo nümero de rayos que los carpelos, área recpetacular en toda la super ficie discoidal o solamente restringida a las áreas con rayos; fruto parecido a una baya, esponjoso, con dehiscencia irregular (debido a la expansión producida por el mucílago que rodea a las semillas), semillas pequeñas, usualmente operculadas, a menudo con arilo, endospermo escaso, perispermo copioso. Familia con una amplia distribución mundial, desde las regiones frías y templadas en el hemisferio norte hasta el trópico, habitando cuerpos lénticos de agua dulce; se incluyen en Ja familia 6 géneros con alrededor de entre 50 y 66 especies. En México se presentan 2 géneros con entre 11 y 12 especies. En el pasado se incluían en esta familia a los miembros de Cabombaceae y Nelumbonaceae, las cuales ahora se encuentra segregadas gracias a estudios moleculares. CLAVES PARA LOS GÉNEROS DE LA FAMILIA NYMPHAEACEAE EN NUEVO LEÓN P 1 4- | Kach zx | | Se el um | abet pa al E x amarillos (a Vu VTA nu mer oe pétalos escuamiformes O estamiformes insertados c con nos SE en n cepacia p por Sek ; Can , estig y sésiles en u 1. Nuphar | Pe MAN |; Sc Seu Stalos (más de 10) muy vistosos, de colores blanco, merlo O azul; carpelos ERR en un receptáculo c camoso con js E GE o hipantio en la led externa en la sa al lo pelos) esta i a arriba en forma pros g y curvas (esti pelares); ¡llas con arilo 2. Nymphaea 1. Nuphar Sm. Nymphozanthus Rich. Plantas herbáceas, acuáticas, con rizomas, sin estolones; hojas flotant nergidas o emergentes, desde Velazco et al., Nympl lestado de N León, México 597 -101 -100 -99 N | : ST E LEYENDA al Género Nuphar S N. advena X Población extirpada |. ]Límites municipales ."N. Curvas de nivel a 20 m . Vías de comunicación Escalal 1:2, 200 090 ANS Cuatro carriles 20 0 20 40 /N/ Dos carriles Kilómetros ME Monterrey Sr eer te S E E A n E -101 -100 -99 598 « C NUS LEYENDA Género Nymphaea N. ampla SE EN BH N. dico x AN. mexican X Población pad [|] Limites municipales - Curvas de nivel a 20 m Escala 1:2,500,000 EPI Cuatro caries 20 O 20 40 Dé: VAN anle Kilómetros Esp A deni | | -101 -100 -99 í li Jas en la Tabla 1 Velazco et al do de N | on, México RI [| I d LI J ps ME E 599 TABLA 1. Localidades visitadas, su y pre ) Localidad Municipio Estado de conservación/ Uso actual Presencia de la familia Ojo de agua de Lampazos Lampazos de Naranjo o / Uso turístico Histórica, extirpada en recreati la actualidad Presa de Paras Paras mpactado/ Uso local riego Histórica, extirpada en y abrevader la actualidad Ojo de agua de Bustamante Bustamante Impactado j Uso turístico Histórica, extirpada en recreativo la actualidad Ojo de agua de Sabinas Hidalgo Sabinas Hidalgo Buen estado / Uso Histórica, presente en turístico recreativo la actualida Presa de Sombreretillo Sabinas Hidalgo Buen estado / Uso Sin registros, no presen- turístico recreativo te en la actualidad Pantano de China China Buen estado / Sin uso aparente Sin registros, presente en la actualidad Presa El Cuchillo China Buen estado / Sin uso aparente Sin registros, no presen- Cañón de San Ana Benito Juárez Impactado / Uso local riego y abrevadero te en la actualidad Baño de San Ignacio Linares Buen estado / Uso local riego Sin registros, present y abrevadero en la actualida Presa Cerro Prieto Linares Buen estado / Uso Sin o no presen- turístico recreativo te en la actualida Ojo de agua de Apodaca Apodaca Impactado / Uso Histórica, Ee en Ea recreativo la actualida Manantial del Potosí Galeana Desecado en su totalidad Sin registros Manantial de San Juan y Puentes Aramberri Ge en su totalidad Sin registros Ojo de agua de Pesquería Pesquería Impactado Sin registros, no presen- turístico recreativo te en la actualidad Ojo de agua de Anahuac Anahuac Impactado / Uso Sin registros, no presen- turístico recreativo te en la actualidad Presa Rodrigo Gómez (La Boca) Santiago Impactado / Uso Sin registros, no presen- Presa Los Mimbres Montemorelos turístico recreativo Impactado / Uso turístico recreativo te en la actualidad Sin registros, no presen- te en la actualidad orbiculares a lineares, lóbulo ] E E + traslapados, márgenes enteros, el ápice del lóbulo de delgadamente obtuso a ampliamente idade do. venación primaria mayormente pinnada; flores flotantes o sobresaliendo del espejo de agua, diurnas, de forma casi globosa en la antesis; sépalos de 5 a 9, amarillos a Rs en la pane externa a menudo con tintes rojizos, Mi bd u amo hasta algo 2, gl A wagpmemleseseciu ema a 1 am Avon: r estambres amarillos o rojizos, i insertos por BE del ovario, récurvados cuando dehiscentes, apendice y estambres, disco estigmático con margen entero orbic ulares; É conectivo dectal arcon crenado o dentado; Bute aciendo en vedunculbs rectos, semillas ovides, arilo ausente. Se distribuye en América, Europa y Asia; un genero complicado desde el punto de vista taxonómico, muchas veces se presentan híbridos y rasgos intermedios entre poblaciones; se reconocen entre 10 y 12 especies, las cuales a algunas veces se tratan como subespecies, en México ocurren l o 2 especies dependiendo del sistema de clasificación que se adopte. En Nuevo León ocurre solo una especie. 1. Nuphar advena (Aiton) W.T. Aiton Nymphozanthus advena ee Fernald Nymphaea advena Áito 600 t tani | Texas 2(1) Plantas herbáceas acuáticas, enraizadas, rizomas de entre 5 y 10 cm. de diámetro; hojas comúnmente emergiendo del agua, pecíolo terado, o adaxialmente algo aplanado, laminas adaxial y abaxialmente de color verde, ampliamente ovadas hasta casi orbiculares, de 1 a 2 veces más larga que ancha, de hasta 40 cm de largo por hasta 30 cm de ancho, seno de la lámina de 1/3 a Y de la longitud de la nervadura central, los lóbulos se sobreponen o son divergentes, a menudo formando un ángulo de 90? o mayor, la superficie abaxial de la lámina es glabra a escasamente pubescente; flores de hasta 5 cm de diámetro; sépalos comúnmente 6, abaxialmente de color verde y de color amarillo en la superficie adaxial, a veces teñidos de rojo hacia la base, pétalos oblongos y gruesos; anteras de 3 a 7 mm, más largas que los fila- mentos, disco estigmático verde, algunas veces rojizo, de 13 a 25 mm de diámetro, entero o crenado, rayos estigmáticos de 9 a 23, lineares a lanceolados, terminando de 1 a 3 mm desde el margen del disco; fruto verde, ovoide, con costillas poco prominentes algunas veces, algo constreñido por debajo del disco estigmático, semillas de 3 a 6 mm de largo. Se ha registrado la desaparición de la población del municipio de Paras, esto es atribuible a la dese- cación de los cuerpos de agua en algunas temporadas; las poblaciones del municipio de Sabinas Hidalgo son abundantes, sin embargo se le considera como una especie problemática por su rápido crecimiento y puede llegar a cubrir rápidamente grandes extensiones de cuerpos de agua someros Ejemplares examinados: MÉXICO. Nuevo León. Municipio de Sabinas Hidalgo: Río Sabinas, 1 May 1981, Garza M, N. s.n. (UNL); Parque La Turbina, 26°29'12"-100°12'54°, 320 msnm, 28 Jun 2007, Velazco-Macías 069 (UNL). Municipio de Paras: Presa Paras, 4 Oct 1971, Valdez González, A. s.n. (UNL). 2. Nymphaea L. Castalia Salisb. Es] ] bá , lamina flotante o Plantas] , acuáticas, ri { ; hojas | expuesta fuera del agua, ales suborbicular, peltada o benda dem manera Profunda en la base, borde de la lámina entero o dentado- ondulado; flores solitarias, con pedúnculo largo, flotantes o saliendo fuera del agua, muy PER Ge EE Ge algunas veces con un tenue olor agradable; sépalos 4; petalos Dunn numerosos, manera gradual en estambres o EE colocados en i 1 + s 1 NH + con anteras ind sandes carpelos hundidos en un receptáculo carnoso, linde con él, un BEE carnosos semi-ínfero, estigma discoideo con pocos o muchos rayos, óvulos numerosos, pegados a las paredes divisorias de los lóbulos; fruto madurando enu del agua, de consistencia esponjosa, globoso u mide a veces cubierto con las bases persistentes de p y estambres, semillas numerosas envueltas en un arilo en forma de saco. Se estima que existen unas 50 especies distribuidas de manera amplia en todas las regiones tropi- cales y subtropicales. Utilizadas como plantas ornamentales, algunas especies se han considerado como invasivas. En México existen alrededor de 11 especies, distribuidas prácticamente en todos los estados, de manera particular en aquellos de afinidad tropical; en Nuevo león se presentan 3 especies. CLAVES PARA LAS ESPECIES DEL GÉNERO NYMPHAEA EN NUEVO LEÓN 1. Pétalos de color amarillo; sépalos de color verde claro, sin marcas oscuras en la parte externa; flores con un leve olor agradable 1. Nymphaea mexicana 1. Pétalos de color blanco o azul ; sépalos de col de cl llent o líneas en pare externa; flores ai volor. La) aloe k | | | acorracdnac Pe Ix IV d ICI a D 2 Pétalos de color azul bl | | treo; borde de las hojas li 3. N. elegans 1. Nymphaea mexicana Zucc. Castalia flava (Lei ) Green Nymphaea flava Leitner Velazco et al., Nymph | estado de N León, México 601 Rizoma recto y alargado, de color café, verrucoso, estolonifero en el ápice; pecíolo de hasta 1 m de largo, láminas flotantes, glabras, ovadas, de hasta 20 cm de largo o ancho, ápice no escotado, borde liso o ligeramente ondulado, base escotada, venación radiada, sin patrón en forma de telaraña, haz de color verde, envés de color café o café —rojizo; flores diurnas, comünmente flotantes, algunas veces sobresa- liendo del agua, de hasta 13 cm de diámetro; sépalos cuatro, lanceolados y agudos en el ápice, de hasta 8 cm de largo, de color verde claro, sin manchas en la parte externa, amarillo claro en la parte interna; pétalos de 30 a 40, de color amarillo o amarillo claro, de hasta 6 cm de largo por 1 cm o menos de ancho; estambres de 40 a 60, de color amarillo, apéndice conectivo diminuto o ausente, filamentos más anchos por debajo de la mitad, más largo que la antera; estilos de 7 a 9; fruto ovoide de hasta 2.5 cm de largo; semillas globosas de hasta 5 mm de diámetro. Florecen principalmente en verano, pero se pueden encontrar en floración durante todo el año; se distribuye en Estados Unidos en los estados de Oklahoma, Alabama, Florida, Georgia, Louisiana, Mississipi, Carolina del Norte, Carolina del sur, Texas y Arizona; en México se registra para el Distrito Federal y los estados de Hidalgo, México y Michoacán, considerada en algunos lugares como una planta invasora. En Nuevo León solo se ha encontrado en una localidad en el municipio de Benito Juárez, en una represa artificial en el Cañón de Santa Ana, municipio de Benito Juárez. Ejemplares examinados: MÉXICO. Nuevo León. Municipio de Benito Juárez: Cañón de Santa Ana, 25%33'12" — 100°11'09", 590 msnm, 15 May 2007, Velazco-Macías 063 (UNL) 2. Nymphaea ampla (Salisb.) DC. Castalia ampla Salisb. Rizoma recto, no ramificado, sin estolones; pecíolo de hasta 1 m de largo glabro, láminas flotantes, haz de color verde, envés de color purpúreo frecuentemente manchado, de forma ovoide, y hasta 45 cm de largo por ancho, márgenes dentados a espinoso-dentados, venación radial, sin un patrón en forma de telaraña, superficie glabra; flores emergiendo del agua, de hasta 18 cm de diámetro, diurnas; sépalos cuatro, lado abaxial de color verde claro con manchas o líneas de color oscuro, lado adaxial de color blanco sin manchas; pétalos de 12 a 21 de color blanco; estambres de 50 a 190, amarillos, los más exteriores con un apéndice conectivo proyectándose de 3 a 10 mm más allá de la antera, filamentos ensanchándose por debajo de la mitad, comúnmente iguales o más cortos que las anteras; pistilo con 14 a 25 lóculos, apéndices al margen del disco estigmático cortamente triangulares hasta 3 mm; semillas globosas a elipsoides. Florece durante todo el año pero principalmente en verano; presenta una amplia distribución, en Estados Unidos se ha reportado para los estados de Florida y Texas, mientras que para México se ha presenta en los estados de Coahuila, Nuevo León, San Luís Potosí, Tamaulipas, Colima, Guerrero, Jalisco, Michoacán, Oaxaca, Puebla, Querétaro, Veracruz, Campeche, Chiapas, Quintana Roo, Tabasco y Yucatán, se ha reportado también para Belice, Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua, Panamá, Islas Caimán, Cuba, Republica Dominicana, Haití y Jamaica. En el estado de Nuevo León, es la especie con distribución más amplia, ya que se ha colectado para 5 diferentes municipios en el norte y centro del estado, aunque en la actualidad solamente se presenta en 2 de ellos, Sabinas Hidalgo y Linares. Ejemplares examinados: MÉXICO. Nuevo León. Municipio de Apodaca: Ojo de agua de Apodaca, 22 Oct 1971, Valdez González s.n. gars Municipio de Bustamante: 1 Ago 1938, H. editei s.n. (TEX). a de "cpm de sisi Río Candela, cerca de icipal, 290 msnm, 22 Jul 1980, Briones V. 233 (UNL). Munici 4°51'56"- 99°20'10", 240 msnm, 13 Jun | 2007, Velazco-Macías 068 (UNL). Municipio de Sabinas Hidalgo: Parque La Sege 26°29 12"-100°12'54", 320 msnm, 28 Jun 2007, Velazco-Macias 070 (UNL) 3. Nymphaea elegans Hook. Castalia elegans (Hook.) Greene 602 | tani titute of Texas 2(1) Rizoma recto, no ramificado, sin estolones; pecíolo de hasta 1 m de largo, glabro, láminas flotantes, haz de color verde, envés de color purpüreo, de forma ovoide, de hasta 25 x 30 cm, márgenes enteros o algo ondulados, venación radial y prominentemente central, sin un patrón en forma de telarafia, superficie glabra; flores diurnas, emergiendo del agua, de hasta 13 cm de diámetro; sépalos cuatro, lado abaxial de color verde claro con manchas o líneas de color oscuro, lado adaxial de color blanco sin manchas; pétalos hasta 27 de color púrpura claro hasta casi blancos; estambres de 55 a 145, amarillos, apéndice conectivo proyectándose 1 mm (raramente más), más allá de la antera, filamentos ensanchándose por debajo de la mitad, más cortos que las anteras; pistilo con 12 a 20 lóculos, apéndices al margen del disco estigmático triangulares de 1 mm o menos, semillas casi globosas. Florece de primavera a otoño; presenta una amplia distribución, en Estados Unidos se encuentra en Florida, Louisiana y Texas, en México sel tado en Sinaloa, Sonora, Tamaulipas, Guerrero, Jalisco, Morelos, Nayarit, Oaxaca y Veracruz, bien se ha reportada para las Bahamas y Argentina. En el estado de Nuevo León se ha encontrado solamente en una localidad en el municipio de China, sobre la carretera libre Monterrey — Reynosa, en áreas inundables dominadas por Acacia farnesiana y ciperáceas diversas. ] MEXICO Nvevo lean M oo to Chi al hl M 1 R ,km 93, 25°42'08"- 99092547", 323 m msnm, 7 Nov 2006, Velazco-Macías s.n. (UNL), DISCUSIÓN Se identifican dos situaciones preocupantes, en primera instancia, el presente estudio pone de mani- fiesto la reducción en la distribución geográfica de las especies de flora asociadas a cuerpos de agua, por mencionar un ejemplo, N. ampla, ha d ido de 3 localidades en las cuales estaba documentada con ejemplares colectados entre los años de 1938 y Ro esta situación es Ao en ud parte a la dese- ción intenrinnal] de q cuerpo sae agua ;€ uso ] volú C gua F superficial para riego y consumo humano (Recdowaki 1978), pe manera stet a estos factores, el uso recreativo en algunas zonas ha sido determinante en la modificación del hábitat y la desaparición de especies; así de un total de 17 localidades visitadas en el estado, solamente 4 albergan poblaciones de esta familia. Una EES situación de E es a poca representación de esta en los herbarios consultados, esto se pone te 4 ejemplares herborizados y 1 una especie reportada en la literatura, aún más se contabilizan un total de 11 especies de angiospermas acuáticas estrictas registradas para Nuevo León (Lot et al. 1998), siendo los miembros de esta familia, especies llamativas y conspicuas por sus características fisonómicas, es difícil oca el poco interés por feo e esta familia en las colecciones botánicas, aún asi, el pres arroja que l I (Nymphaea elegans y N. mexicana) se presentan como nuevos registros para Nuevo bein: En base a las colectas históricas, así como lo observado en las distintas localidades visitadas, en donde se ha observado un alto impacto por actividades del hombre e incluso la total desaparición del hábitat en algunas localidades, se estimaría que en un lapso de 10 años las especies aquí reportadas puedan desaparecer del hábitat restante en el estado de Nuevo León, esta EE ha sido denotada en otros estudios, tales como el de Novelo y Bonilla (1999) qui a Nymphaea ampla como "rara en la región de estudio y por lo tanto vulnerable a la extinción," esto a pesar de ser una de las especies con mayor distribución en México. AGRADECIMIENTOS 4 1 f :1: 1... | E. 14. y peser A | jemplares A la Dra. M la González AI y en el herbario institucional de la Facultad. de Ciencias Biológicas (UNL) de la Universidad «tonos de Nuevo León. A los Biols. Liliana Ramírez Freire, Héctor M. Villalón Moreno y al Ing. Jorge Alberto Martínez Pineda por su colaboración en el trabajo de campo del presente estudio, en especial a este ültimo por su colaboración en el Sistema de Información Geográfica. Velazco et al N h | estado de Nuevo León, México 603 P d LI F REFERENCIAS ALANIS-FLORES G.J., G. Cano y Canc y M. RovaLo Merino. 1996. Vegetación y Flora de Nuevo León, una guía botánico- ecológica. Impresora Monterrey, S.A. de C.V.. México. Pp. 1-9. CALDERÓN DE Rzepowski, G. y J. Rzepowski. 2001. Flora fanerogámica del Valle de México. 2° edición. Instituto de Ecología, A.C. y Comisión Nacional para el Conocimiento y Uso de la Biodiversidad. Michoacán, México. Pp. 168-169. CORRELL, D.S. y M.C. Jounston. 1979, Manual of the Vascular Plants of Texas. The University of Texas at Dallas. Estados Unidos de América. Pp. 630-632 INEGI. 1986. Síntesis geográfica del estado de Nuevo León. Secretaría de Programación y Presupuesto. México, Lor, HA. A. Novelo y P. RAMfREZ-GARCÍA. 1986. Angiospermas acuáticas mexicanas 1. Vol. V. Listados florísticos de México.Instituto de Biología, Universidad Nacional Autónoma de México. México, D.F Lor, HA. A. NoveLo y P. Ramirez-Garcia. 1998. Diversidad de la flora acuática Mexicana. En: T.R: Ramamoorthy, R. Bye, A. Lot y J. Fa, comps. Instituto Se SE Universidad Nacional DEENEN de México. Pp. 563-578. LOT, A y F. CHIANG. 1986. Manual de herbario, administración y manejo de colecciones, técnicas de recolección y preparación de elas botánicos. Consejo Nacional de la Flora de México. México. Pp. 87-92. Martinez, M. y A. NoveLo. 1993. La vegetación acuática del estado de Tamaulipas, México. Ann. Inst. Biol. UNAM, Ser. Bot. 64(1): 59-86. Mora-OLIVO, A. y J.L. VILLASEÑOR. 2007. Diversidad y distribución de la flora vascular acuática de Tamaulipas, México. J. Bot. Res. Inst. Texas 1:511—527. Moreno, N.P 1984. Contribución al conocimiento de la flora vascular acuática y las asociaciones más comunes de la presa Rodrigo Gómez y sus afluentes, Mpio. de Santiago, Nuevo León, México. Tesis de licneciatura. Facultad de Ciencias Biológicas, Universidad Autónoma de Nuevo León. NoveLo, A. y J. Bonita. 1999. Nymphaeaceae. In: Rzedowski, J. y G. Calderón de Rzedowski (eds.). Flora del Bajío y regiones adyacentes. Fascículo 77. Instituto de Ecología, A.C. Centro Regional del Bajío. México. RoJAs-MENDOZA, P. 1965. Generalidades sobre la vegetación del estado de Nuevo León y datos acerca de su flora. Tesis doctoral. Facultad de Ciencias, Universidad Nacional Autónoma de México. México. Rzepowsxı J. 1978. Vegetación de México. Editorial Limusa S.A. México. Pp. 59-74. TREVIRO-GARZA, E.J., C. Cavazos-CAMACHO y O.A. AGUIRRE-CALDERÓN. 2001. Distribución y estructura de los bosques de galería en dos ríos del centro sur de Nuevo León. Madera y BOSQUES 7(1)13-25. VILLARREAL-QuINTANILLA, J.Á. 2001. Flora de Coahuila. Listados fl México. Instituto de Biología. Universidad Nacional Autónoma de México. México D.F. VILLARREAL-QUINTANILLA, J.A., M.A. CARRANZA, E. ESTRADA-CASTRILLÓN y A. RODRÍGUEZ. 2006. Flora riparia de los ríos Sabinas y San Rodrigo, Coahuila, México. Acta Bot. Mexicana 75:1-20. Wiersema, J.H. y C.B. HeLLQUIST. 1997. Nymphaeaeceae. En: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. New York and Oxford. Vol. 3:66-77. ZOMLEFER, W.B.. 1994. Guide to flowering plant families. The University of North Carolina Press. E.U.A. Pp. 49-54, 604 1 Lofal Das an Lis ET. 2(1) BOOK NOTICE GORDON W. FRANKIE, ALFONSO MATA, AND S. BRADLEIGH VINSON (eds.). 2004. Biodiversity Conservation in Costa Rica: Learning the Lessons in a Seasonal Dry Forest. (ISBN 0-520-24103-7, pbk.). University of California Press, 2120 Berkeley Way, Berkeley, California 94704-1012, U.S.A. (Orders: California/ Princeton Fulfillment See 1445 Lower Ferry Road, Ewing, New Jersey 08618, U.S.A, 1-800-777- 4 edu @cpfsinc.com). $34.95, 341 pp., bie photographs, line drawings, maps, + 726, tables, 7 x 10". Contents: Preface (with Acknowledgments) Introduction—Alfonso Mata and Jaime Echeverría PART I. BIODIVERSITY AND ECOLOGICAL STUDIES A. COSTA RICAN DRY FOREST 2. Flowering Phenology and Pollination Systems Diversity in the Seasonal Dry Forest— Gordon W. Frankie, William A. Haber, S. Bradleigh Vinson, Kamal S. Bawa, Peter S. Ronchi, and Nelson Zamora 3. Breeding Structure of Neotropical Dry-Forest Tree Species in Fragmented Landscapes—James L. Hamrick and Victoria J. Apsit T 4 Torch 1 (71 D H nh of Trees in doe i Ge S. Bawa 5. Tropical Dry-Forest Mamas of Palo Verde: Ecology and Kathryn E. Stoner and Robert M. Timm 6. The Conservation Values of Bees and Ants in the Costa Rican Dry Forest—S. Bradleigh Vinson, Sean T. O'Keefe, and Gordon W. Frankie 7. | Ecology of Dry-Forest Wildland Insects in the Area de C ión G te—Daniel H. Janzen e D B. BIOTIC RELATIONSHIPS WITH OTHER iria RICAN FORESTS . . Diversity, Migration, and Conservation of Butter flies i rthern Costa Rd Haber and Robert D. Stevenson 9. — Watershed Ecology and C tion: Hydrological R f Costa Rica—Alfonso Mata 10. Where the Dry Forest Feeds the Sea: The Gulf of Nicoya Estuary—José A. Vargas and Alfonso Mata 11. Mangrove Forests under Dry Seasonal Climates in Costa Rica—Jorge A. Jiménez qoem C. BIOTIC RELATIONSHIPS OTHER À CAL AREAS 12. Geographical Distribution, Ecology, and Conservation Status of Costa Rican Dry-Forest Avifauna—Gilbert Barrantes and Julio E. Sánchez 13. An Ultrasonically Silent S ps PA Dry Forest w Dum pele ME K. ra 14. Biodiy ersity a d Cc American Dry-F c 1 ZE E DAloz 15 A T D 1 y H T f. "XT Td 1D 1 jr AK XE . 15 Years of Conservation of Sea Turtles in Costa Rica—James R. Spotila and Frank V. Paladin 16. Prospects for Circa Situm Tree Conservation in Mesoamerican Dry-Forest Agro-ecosystems— David H. Boshier, James E. Gordon, and Adrian J. Barrance PART II. TR O ACTION: THE RECORD 17. Biodiversity Inventories in Costa Rica and Their Application to conservation—Paul Hanson Conflict Reenhitian: R izi iM ing Di linR Protecti Gregory A. Giusti o [e] Lei a Kr © ud 19 Y^" +: IC 2 ¿o oA E G T 1 WT +1 4 on D: T E 1 [XT ial Organization— Gordon W. Frankie and S. Bradleigh Vinson 20. The Media and Biodiversity Conservation— Gilda Aburto 21. Threats to the Conservation of Tropical Dry Forest in Costa Rica—Mauricio Quesada and Kathryn E. Stoner 22. a Law of un Rica: id i as Enforcement— Roxana Salazar 23. Dispute over the P t ME pacta Bustos 24. The Policy Context for C tion in Costa Rica: Model or À Katrina Brandon 25. Conclusion and Recommendations— Gordon W. Frankie, Alfonso Mata, and Katrina Brandon Contributors Index J. Bot. Res. Inst, Texas 2(1): 604. 2008 REFUGIO DE FITODIVERSIDAD EN LA CIUDAD DE MÉXICO, FL CASO DE LA CUENCA DEL RÍO MAGDALENA V. Ávi ez y L. Almeida-Leñero eiis ae Ecosistemas de MON um cultad de one , UNAM, MÉXICO, D.F. Circuito exterior, Ciud | UNAM, C.P 04510, MÉXICO Emails Vicavakdanail com RESUMEN La cuenca del río Magdal t le los últi EE gi fitodi idad d 87 familias con 251 | géneros y 4 487 especies. Las famili E I de g i Asteraceae a Lamiaceae, yophy y : i Salvia (13) eet (10), Pi (8) C 1; ` c EE ] Iz 1 a NOM- 059- 9-SEMARNAT 2 2001, siendo tres — endémicas m r negundo v var. mexicanum, a wanes As plantas e E Lo 4 de esta área, abarca ca. del 2596 de la diversidad florística de la cuenca del valle de México, o el 2.2% del total de la Se? del NM Por esto, la cuenca del río Magdalena debe ser considerada un refugio para la conservación de la biodiversidad, ya que es la base para el » : : x 14. 1 sa E EE ARI " : 1.1 PALABRAS CLAVE: Floristica, fitodiversidad, lel rio Mag lal , México D.F ABSTRACT The Magdalena ES ing f the last living ri f Mexico City, | floristic diversity of 87 families, 251 genera, and 487 species TI E pl families i f speci de | Asteraceae, Poaceae, Lamiaceae, Caryophyl- laceae, and Rosaceae. TI ith tl peci Salvia (13), Ageratina (10), Pinus (8), and Quercus (7). T E ] ] NOM-059-SEMARNAT- 2001. the Mexican classification f langered plant sy : gtl | I Acer negun ndo var. mexicanum, F bedingl ii, Dahlia scapigera. T peci led li | indi ,Ácaena elongata a eee ni aiia ] les ricl f thi I | 2596 of tl l flora in the Valley of Mexico, and ca. 2.296 y e the M gd ] Ri hed should ! idered ti f rich biodi i watershed tl int ft] portant yst ji led to tł INTRODUCCIÓN México es un país megadiverso dada su ubicación dentro de las regiones Neártica y Neotropical (Rzedowski 1978). Su flora nativa oscila entre 75 órdenes, 304 familias, 2,804 géneros y 22,351 especies, en los 200 millones de ha de superficie que abarca el país (Villaseñor 2004). Este mismo autor, utilizó métodos no pa- ramétricos para estimar a EEN total del país, así se añadirían a estas cifras más de 6,500 especies, por lo q r en el orden de las 29,000 especies. Con estas cifras la República Mexicana ocupa el o lugar entre los países con mayor riqueza florística del mundo (Sarukhán & Dirzo 2001). Sin embargo, E conocimiento de su fitodiversidad es todavía incompleto aired i Lazonat ] subhümeda del t mexicano cubre la mayor parte d del país, con una — cie de ca. 33 millones de ha, que corresponde al 1596 d la superficie nacional (Toledo & Ordoñez 1998). Esta zona se caracteriza por contener elementos holárticos, predominantemente en el estrato arbóreo, mientras que los estratos arbustivo y herbáceo son neotropicales, formando un complejo mosaico con los elementos autóctonos (Rzedowski 1978). Los principales tipos de vegetación en esta región son los bosques de coníferas, bosque mixto y bosque de Quercus, mientras que el bosque mesófilo de mon- tafia, el cual pertenece a la zona templada hümeda, representa una transición entre las regiones tropicales y templadas (Rzedowski 1970). La flora de estos bosques de coníferas junto con la del bosque de Quercus representan cerca de 2596 del total de la flora vascular de México (ca. 7,000 especies), de las cuales ca. 4,900 especies son endémicas (Rzedowski 1991). J. Bot. Res. Inst. Texas 2(1): 605 — 619. 2008 606 t tani tit Texas 2( La cuenca del Valle de México (CVM) contiene bosques de filiación templada en las que se desarrollan cerca del 2% de las plantas del planeta (Calderón de Rzedowski & Rzedowski 2001). En esta cuenca endo- rreica de origen volcánico con una extensión ca. de 7,500 km?, se ubica el Distrito Federal que, a pesar de contener una de las ciudades más grandes del mundo, todavía cuenta con importantes áreas con vegetación natural, las cuales representan el 58% de su territorio (Corenader 2006). Dentro del Distrito Federal, se encuentra la delegación política Magdalena Contreras, la cual ocupa el 9? sitio en extensión territorial, y más del 70% de su superficie corresponde a la categoría de suelo de conservación. Sin embargo, el conocimiento sobre la fitodiversidad es limitado. En la Delegación Magdalena Contreras se localiza el río Magdalena, el cual representa uno de los cuer- pos de agua más importantes del Distrito Federal (Delegación la Magdalena Contreras, 2006). El tipo de suelo y la cobertura forestal que ésta presenta, promueve la infiltración del agua de lluvia hacia los mantos acuíferos de la CVM (Mazari 2000), por lo que un excedente hídrico de 10% podría ser aprovechado en la zona urbana (Jujnovsky 2006). Al río Magdalena se le reconoce como un río vivo del Distrito Federal y existe un interés reciente por parte de las autoridades de la ciudad en manejar integralmente la cuenca. Debido a que estos bosques enfrentan una gran presión antrópica, es importante conocer la fitodiversi- bd deg estas comunidades boscosas, con la finalidad de evaluar su estado de conservación y poder proponer Este trabajo presenta eli tario inicial de la fitodiversidad de plantas vasculares SS la cuenca del río Magdalena (CRM) y su zona de influencia. Antecedentes.—Algunos de los autores que han realizado estudios Pus de E y vegetación en la CRM y zonas adyacentes son: Rzedowski (1954) hizo un análisis de la li n el matorral xerófilo del pedregal de San Ángel, Madrigal (1967), estudió el bosque de Abies religiosa en » zonas montañosas de la CVM, Rzedowski (1970), realizó una descripción para el bosque mesófilo de montaña en la CVM, encontrando algunos fragmentos dentro de la CRM, Velázquez y Cleef (1993), definieron las asociaciones vegetales en los volcanes Tláloc y Pelado, y Nieto de Pascual (1995) describió la estructura y composición de los bosques de Abies religiosa en la CRM. En el 2001, Calderón de Rzedowski y Rzedowski actualizaron la información sobre la flora del valle de México, estimada en alrededor de 2500 especies, la cual ha sido base para estudios recientes como los de Ávila-Akerberg (2002) y Nava (2003), quienes presentan datos de cobertura de las especies registradas por comunidad vegetal, así como su relación con diversas variables ambientales en la CRM. Área de estudio.—Localización física.— La cuenca del rio Magdalena (CRM) se ubica dentro de la Cordil- lera Volcánica Transmexicana (CVT), en la provincia florística de las Serranías Meridionales, dentro de la región Mesoamericana de Montaña (Rzedowski 1978). Esta área forma parte de la CVM en la vertiente oc- cidental de la sierra de las Cruces con un intervalo altitudinal de los 2,530 alos 3,870 msnm y una extensión aproximada de 3,000 ha (Fig. 1). Relieve.—Los bosques de la CRM se desarrollan en un relieve montañoso, dada su ubicación dentro de la CVT. El modelado del pu se debe principalmente a la acción erosiva hídrica y a los procesos fisico- químicos, siendo el desgas a corriente del río Magdalena el que ha ido formando un valle intermontano longitudinal joven TOME 2000). Geología.—El área de estudio presenta diferentes etapas en su evolución relacionada directamente con el origen de la CVM (Ontiveros 1980). Esta zona está formada por material ígneo extrusivo, producto de manifestaciones volcánicas del Terciario y Cuaternario (Álvarez 2000). El basamento de la CRM, está con- stituido por macizos de la sierra de las Cruces, de principios del Terciario, alcanzando el Terciario Superior. Esta sierra es uno de los principales focos de actividad del CVT que se formó como consecuencia de una serie de emisiones pacíficas de lavas ácidas. Formaciones KL i dan lugar alos cerros de San Miguel y la Palma (Cervantes 1969). La CRM, es el resultado de un afallamiento en bloques que deja bancos abruptos de forma regular en donde el río se encajona en márgenes estrechos (Álvarez 2000). Hidrología.—El río Magdalena tiene un cauce de una longitud total de 21,600 m, de los cuales 15,200 m recorren los bosques de la cuenca del río Magdalena (Ávila-Akerberg 2004). Posteriormente, el río entra a Muta Akerberg at al $ Eitnrli idad I lal río M dal ; México D F 607 464000 466000 468000 470000 472000 474000 476000 478000 7 T H H Seed 17, g id 2136000 2134000 E ^am * rio. 2132000 Mexico LAE AE AES NS. mo ud 2130000 pim- 2128000 ][-—- (E Zona urbana CRM C] SCMC E 2126000 T T T L SEC MEL NE E E E E Jj Distrito Federal De 11 H tfaa | sf. RA Jal zona urbana hasta llegar a la presa Anzaldo en donde el río es entubado y dirigido hacia el río Churubusco. Las aguas continúan su recorrido para finalmente salir de la CVM a través de los túneles artificiales de Tequisquiac y llegar a la cuenca del río Tula (Álvarez 2000). Las características hidrológicas de la CRM se deben a factores como la altitud y la influencia de las masas marítimas, tanto del Golfo como del Pacífico, que favorecen las condiciones de humedad. Así mismo, la estructura geológica de la sierra de las Cruces y las precipitaciones han permitido que la zona mantenga una infiltración constante, generando una fuente de almacenamiento de agua (Ontiveros 1980). Suelo.—Fl suelo en la CRM es resultado de una formación de material ígneo extrusivo, producto de manifestaciones volcánicas del Terciario y Cuaternario, predominando las Andesitas y Dacitas (Ontiveros 1980). Los suelos son, en su mayoría, Andosoles del tipo húmico, mólico, ócrico con mezcla de Litosoles, con textura franco, migajón arcilloso y arenoso. Los suelos con textura franca se encuentran distribuidos en toda la región. En las partes altas, el pH es más ácido y el contenido de materia orgánica es mayor (Jujnovsky 2006). Clima.—En la parte baja de la CRM, entre los 2530 y 2800 msnm, se presenta el subtipo climático C (w;) (w) (b) i'g (Koppen modificado por Garcia 1987); templado subhúmedo, el más húmedo de los subhúmedos con régimen de lluvias de verano y porcentaje de lluvia invernal menor al 596. El verano es fresco y largo, la temperatura media anual oscila entre 12 y 18? C, la temperatura del mes más frío entre -3 y 18° C y la del mes más caliente entre 6.5 y 22? C, con poca oscilación térmica y marcha de la temperatura tipo Ganges. En la parte alta, de los 2800 a los 3500 msnm el clima es Cb’ (wj) (w) (b” i g, difiriendo del anterior por tener una temperatura media anual entre 5 y 12? C con oscilación térmica menor a 5? C, es decir, isotermal (García 1987). Los porcentajes de lluvia altos se registran para el período de mayo a octubre, siendo el mes de julio cuando se alcanza una precipitación que supera los 250 mm. Las mayores temperaturas acontecen desde marzo hasta octubre, siendo el mes de mayo el más caluroso y enero el más frío (Álvarez 2000). icos que estudio, tales como el relieve montañoso, el Tei Vegetación.—Los el | E sustrato rocoso -producto de evolución geológica-, el clima, la existencia de agua de manantiales y ríos, así 608 | tani titute of Texas 2( como los suelos, han permitido el desarrollo de la vegetación templada, donde se presentan gran variedad de especies de importancia biológica y económica. De Md a u Ge de EE ESCH pos Ra (1978), la vegetación presente en la zona Į níferas com y P twegii. También se tran mezclas I de estas especies d (feras con otras latifoliadas como Quercus spp a lo que se conoce como r bosque mixto. En pequeños parches es posible encontrar bosque mesoi de montaña, caracterizado por especies como: Quercus spp., Garrya laurifolia, Clethra mexicana y Viburnum stenocalyx, entre otras. MÉTODO La recolecta de ej ] f lizada durante las múltiples visitas a la zona de estudio entre 2004 y 2006, en 128 sitios de: muestreo, dentro de los diferentes tipos de vegetación de la CRM y zonas adyacentes. En estos sitios se registraron datos ambientales de altitud, pendiente, orientación y suelo desnudo. Los ejemplares fueron herborizados, etiquetados y determinados a nivel de especie con ayuda de claves taxonómicas especializadas para la zona (Beetle 1983 y 1987; Calderón de Rzedowski & Rzedowski 2001, Espinosa & Sarukhán 1997). Para las determinaciones botánicas se siguió la clasificación propuesta por KEE Ge pus E ? pas e la de eee et al. (1985). Las sinonimias n la base de datos del Missouri Botanical Garden Soen mobot3.org). El material colectado fue depositado en la colección del Herbario FCME (Facultad de Ciencias, UNAM). Se identificaron las especies arvenses y ruderales consultando los trabajos de Espinosa & Sarukhán (1997) y Calderón de Rzedowski & Rzedowski (2001). Se realizó un análisis de coeficiente de relación entre las especies y géneros de la familia Asteraceae contra el total de las especies para comprobar la intensidad de muestreo (Rzedowski 1991). RESULIADOS El estudio registró un total de 87 familias con 251 géneros y 487 especies en la lista florística (Apéndice 1). Las familias más importantes en cuanto al número de especies son: Asteraceae 21%, Poaceae 4.9%, Lami- aceae 4.7%, Caryophyllaceae 4.3, Rosaceae 3.7% y Brassicaceae 3.5% (Tabla 1), mientras que, las familias con mayor número de géneros son: Asteraceae 18.2%, Poaceae 5.7%, Brassicaceae 5.3%, Fabaceae 4.0%, Rosaceae 4.0% y Lamiaceae 2.8%. Los géneros con más especies son Salvia (13), Ageratina (10), Pinus (8), Quercus (7), y Roldana y Senecio con (6). En cuanto a las formas de crecimiento (por especie, Fig. 2) las herbáceas son las más abundantes (365 p seguido de las arbóreas (49), arbustivas GE rasantes (20), lea (5) y epífitas (4). imiento a nivel de familias | n hierbas Asteraceae con 79 espe- cies, Poaceae con 24, Lamiaceae con 23, Caryophylaceae 18, Brassicaceae con 15 y Fabaceae y Cyperaceae con 12. Para los arbustos, Asteraceae con 20 especies, y Ericaceae y Rosaceae con 3 especies cada una; en el arbóreo, Pinaceae con 9, Fagaceae con 7 y Betulaceae con 4 (Fig. 3 De la lista florística resultante (487 especies), se registraron 10 especies en alguna categoría de riesgo de acuerdo a la Norma Oficial Mexicana 059-SEMARNAT-2001 (DOF, 2002), de las cuales 3 son endémicas a México: Furcraea bedinghausii, Acer negundo var. mexicanum y Dahlia scapigera (Tabla 2). Otras especies, por el contrario, se ven favorecidas por condiciones de perturbación por lo que es comün encontrarlas en cultivos IN O Hein a a orilla de caminos (ruderales). Dentro de estas especies se encuentran: Acaena elongata, Achill iolaris, Cerastium nutans, C. vulcanicum, Drymaria laxiflora, Erigeron galeottii, Gerni seemannii, Tr ger Senecio cinerarioides y Sigesbeckia jorullensis, entre otras. El total de las especies arvenses y/o ruderales fue de 69. El valor de coeficiente especies-géneros (e/g) para la familia Asteraceae fue de 2.3, en tanto que el coeficiente e/g para el total de la flora registrada en este estudio fue de 1.94, dando una diferencia de 0.36. Áwila Albarhara at al Eitadi idar I lal ria M lal México DF 609 Y i f 7 tad la CRM, México, D. F. TABLA 1. Familias de plantas jor repre Familia No. 96 No. géneros géneros especies especies Asteraceae 45 182 102 210 Poaceae 14 3.7 24 49 Brassicaceae 13 5.3 17 3.5 Rosaceae 10 40 18 3.7 Fabaceae 10 40 17 3.5 Lamiaceae 7 2.8 23 4.7 Scrophulariaceae 6 24 10 23] lacea 6 24 9 1.9 Solanaceae 5 20 15 3.1 Crassulaceae 5 2.0 11 2.3 Rubiaceae 5 20 9 1.9 Caryophyllaceae 4 1.6 21 4.3 Cyperaceae 4 1.6 12 23 agraceae 4 1.6 9 1.9 Polygalaceae 4 1.6 9 1.9 Ranunculaceae 3 1.2 8 1.6 Pinaceae 2 0.8 9 1.9 Fagaceae 1 0.4 7 14 500 —— — ES TN 400 - 300 4 E No. especies a 96 del total 200 - 100 - 0 " ERAR ". E om — EM O CENE Herbáceo Arbustivo Arbóreo Rasante Epífita Trepadora Fic. 2. F | imiento domi tes, 1 yi t j | CRM, México, D.F DISCUSIÓN Los resultados de este di t que la CRM es una de las! lidad AS 1 tant refugio de fitodiversidad dentro del suelo de conservación del Distrito Federal. En el área "m estudio se encuentran todos los principales tipos de vegetación templados del país y la riqueza florística es mayor o similar a la encontrada en zonas cercanas (e.g., Cornejo-Tenorio et al. 2003; Castillo et al. 2004; Sánchez-González et al. 2005). En cuanto al número de especies, las del estrato herbáceo dominan, lo que corresponde a lo obser- 610 No. spp. Árboles A E Pinaceae Fagaceae Betulaceae Caprifoliaceae Ericaceae Arbustos ej L >= E Asteraceae Ericaceae Rosaceae Grossulariaceae Solanaceae Hierbas ff m Asteraceae Poaceae Lamiaceae Caryophyllaceae Brassicaceae Fic. 3 Principales familias por f le crecimiento en la CRM, México, D.F Ween am lel río Magdalena, México, D.F 611 Ávila Abarhara at al Fit M , vado por Rzedowski (1978) y Cornejo-Tenorio et al. (2003) para la generalidad de los bosques templados de México. Sin embargo, considerando la cobertura, el estrato arbustivo es el dominante, lo que puede ser un indicador del deterioro de la zona. Este se explica por la deforestación, lo que genera claros en el dosel del estrato arbóreo permitiendo o estimulando el crecimiento de los arbustos. El endemismo en las EE de la CRM, presenta proporciones similares a las evaluaciones previas de la flora de las regiones t de México, ya que Rzedowski (1991) estima que la proporción de especies endémicas del país para los bosques de coníferas y de encino equivale al 70% y para el bosque mesófilo de montaña, a 30%. La familia Asteraceae, con 102 especies, es la mejor representada en la cuenca, P wi: en menor proporción por Poaceae, Lamiaceae, Rosaceae, Fabaceae y Brassicaceae. Este patrón ha sido encontrado también en otros trabajos realizados en tipos de DECH similares a los de la cuenca, como es el caso del Parque Nacional Lagunas de Zempoala (Bonilla y Viana-Lases, 1997), en el volcán Po- pocatépetl (Almeida-Leñero et al. 1997), el área de San Juan Nuevo Paranguricutiro (Medina-García et al. 2000) y la Reserva de la Biosfera de la Mariposa Monarca (Cornejo-Tenorio et al. 2003). Se ha visto que en las latitudes donde la familia de las compuestas juega un papel importante, el cociente entre el número de especies y géneros de compuestas se aproxima al mismo valor para toda la flora fanerogámica de un área. Para la cuenca del río Magdalena se encontró que este cociente es de 2.3, y si se multiplica por el número total de géneros (250) nos da 579 especies, número relativamente mayor al total de especies en el área de estudio (487 spp). Esto indica que algunas de estas especies han desaparecido, que hay sinonimias, incon- sistencias en los listados o que aún es insuficiente el esfuerzo de colecta. Sin embargo, al revisar los trabajos realizados por autores como Reiche (1914), Madrigal (1967), Sánchez (1969), Rzedowski (1970), Nieto de Pascual (1995) y Calderón de Rzedowski y Rzedowski (2001), se cuantificó un mayor número de especies en la zona, que las registradas en este estudio. Por lo tanto, es fundamental continuar con el monitoreo de la flora en la zona, sirviendo éste como Aa paras medir el pato de conservación. Los resultados que se EE an en n el papel fund I que gesemipena la CRM como área de 1 de la j| , no sólo por la riqueza florística sino también por la cantidad de especies endémicas (ca. 85 especies) y en alguna categoría de EE La Delegación Magdalena Con- treras presenta más del 78% de su territorio como suelo de co 5n, por lo tanto la riqueza de especies en esa superficie es relevante si consideramos que por Seuples en el caso de plantas, se alberga poco mas del 25% de la diversidad floristica que existe en el valle de México o el 2.2% de la riqueza total del pais. Si ademas se toma en cuenta la presencia en la zona de uno de los ultimos rios vivos en el Distrito Federal y la presencia de elementos del bosque mesófilo de montaña casi desaparecidos de la CVM, es fundamental priorizar el cuidado y recuperación de esta cuenca. Se espera que la información florística proporcionada por la presente contribución pueda ser útil en la elaboración del pan de so EE ER la zona, que representa: un SE gp esta AA y que I a ala ciudad A partir de esta información, es importante fomentar estudios similares en zonas eer abordando juega un papel 1 además otros aspectos biológicos, sociales, económicos y políticos. APÉNDICE 1 PLANTAS NO VASCULARES PLANTAS VASCULARES Bryophyta Equisetophyta a ini Ei Vi Equisetaceae rimmia tolucensis Card.—Rastrera, VA-168 Equisetum hyemale L. ssp. affine (Engelm.) Calder & Roy L. Pottiaceae Taylor—Hierba, VA-147 Trichostomum cylindricum (Bruch.) C.M.—Rastrera, JB-8 Du Pteridopsida Thuidiaceae Thuidium del; (Herdw) Mitt.—Rastrera, BGH-2968 Adiantaceae Adiantum ampillus-veneris L.—Hierba, VA-2 612 Adiantum andicola Lieom.—Hierba, VA-3 Adiantum marginatum Bory—Hierba, MV-1 Aspleniaceae Asplenium monanthes L—Hierba, VA-16 Dryopteridaceae EE Magie (L J Benin SET JB-4 ze) Alston-—Hierba, JB-5 Plecosorus speciossisimus (A. AMA ex Kunze) T. Moore— A-145 Polystichum rachichlaena Fée—Hierba, VA-146 Polypodiac Cheilanthes m (Forssk.) Kaulf.—Hierba, JB-6 Cheilanthes hirsuta Link.— Cheilanthes marginata Kunth.—Hierba, VA-2 Cheilanthes sinuata (Lag. ex Sw.) Ces VA-267 Pteridaceae Pteridium aquilinum (L) Kuhn—Hierba, VA-27 1 Coniferophyta Cupressaceae Cupressus lusitanica Mill—Arbol, MR-22 Pinaceae Abies religiosa (Kunth) Schitdl. & Cham. —Arbol, VA-230 Pinus ayacahuite C. Ehrenb. ex Schltdl. var. veitchii (Roezl) Shaw—Arbol, VA-231 us hartwegii Lindl.—Árbol, V. Se EEN Schiede ex sod i VA-233 montezumae Lamb.—Árb Ce patula Schltdl. & Cham Aon VA-235 Pinus pseudostrobus eg |, VA-236 Pinus rudis Endl.—Árbol, V. Pinus teocote Schltdl. & Gs um VA-238 Magnoliophyta Aceraceae Acer negundo L.var. mexicanum (DC.) Standl. & Roy L. Taylor— Árbol, VA-1 AInaranenacene lresine diffusa Humb. & bcp UE GH- 297 Iresine elongata Humb. & Bompl. ex Willd.—Hierba, VA-7 a, BGH-2977 8 Amaryllidaceae Zephyranthes carinata (Lindl.) Bentn.—Hierba, VA-8 Zephyranthes fosteri Traub.—Hierba, VA-9 Apiaceae Arracacia atropurpurea (Lehm.) Benth. & Hook ex Hemsl.— Hierba, VA-10 Daucus montanus Humb. € Bonpl. ex Spreng.—Hierba, VA-11 Donnellsmithia ee (Humb. € Bonpi.) Mathias & Constan- ce—Hier -12 Eryngium Ge F. Delaroche—Hierba, VA-13 Eryngium proteaeflorum F. Delaroche—Hierba, MR-1 Eryngium serratum Cav.—Hierba, MV- Eryngium subacaule Cav.—Hierba, MR-2 £T mia A P Osmorhiza mexicana Griseb.—Hierba, VA- Tauschia alpina (J. C. Coult. & Rose) VN VA-15 Asclepiadaceae A ] x ta AA À Pp pol į ¡A fr» H NAA j Wi | Asteraceae Achillea millefolium L.—dHierba, ruderal, VA-17 Ageratina enixa (Rob.) R.M. King & H. Rob.—Hierba, VA-18 Ageratina glabrata (Kunth) R.M. King & H. Rob.—Arbusto, VA-19 Ageratina mairetiana (DC.)R.M. King & H. Rob.—Arbusto, VA-2 -21 Ageratina oligocephala (DC.)R.M. King & H. Rob.—Hierba, VA Ageratina pazcuarensis (Kunth) R.M. King & H. Rob.—Hierba, VA-22 Ageratina petiolaris King. & H. Rob.—Hierba, ruderal, MV-6 Ageratina prunellifolia (Kunth) R.M. King & H. Rob.—Hierba, MV-7 R.M. King €: H. Rob.— Arbusto, Ageratina rhomboidea (Kunth) Ageratina rivalis (Greenm.) R.M. King EH Rob.—Arbusto, MV-9 Ageratina vernicosa (Sch. Bip. ex Greenm.) R.M. King & H. —Arbusto, VA-23 Ageratum corymbosum Zucc. ex Pers— Arbusto, VA-24 Archibaccharis asperifolia (Benth.) S.F. Blake—Arbusto, VA-25 Archibaccharis hieracioides (S.F. Blake) S.F. Blake—Hierba, Z3 O eg MR-3 Archibaccharis hirtella (DC) Heering—Trepadora, MR-4 Archibaccharis schiedeana (Benth. in Oerst.) J.D. Jackson.— Hug MR-5 tifolia (Kunth) Blake —Hierba, VA-26 Artemisia ludoviciana utes ssp vont WS Ex Spreng.) D.D. Keck—Hierba, arvense y ruderal, M eco ichx.—Hierba, dup PEN Baccharis conferta Kunth—Arbusto, yes Baccharis multiflora Kunth—Hierba, V. Bidens anthemoides (DC.) Sh ef He arvense, MR-6 Bidens aurea (Aiton) Sherff—Hierba, Bidens odorata Cav.—Hierba, arvense, Ki 1 GE ostruthioides (DC) Sch. Bio.—Hierba, MV-12 Bidens serrulata (Poir) Desf.—Hierba, arvense, MV-13 Bidens triplinervia Kunth—Hierba, Brickellia nutanticeps S.F. Blake—Hierba, MR-8 Brickellia pendula (Schrad.) A. Gray—Hierba, MR-9 Brickellia scoparia (DC) A. Ge ee VA-31 Cirsium ehrenbergii Sch. Bip.—Hier -3 Cirsium jorullense (Kunth) Spreng. Ge jorullense—Hierba, — A-3 Cirsium nivale (Kunth) Sch. Bip.—Hierba, VA-34 Conyza schiedeana (Less) Cronquist—Hierba, VA-35 Cosmos bipinnatus Cav.—Hierba, arvense, VÀ-36 Dahlia coccinea NM are MV-14 Dahlia scapigera (A. Dietr. Knowles & Westc.—Hierba, VA-37 Erigeron galeottii (A. Gray) Greene—Hierba, ruderal, VA-38 Erigeron karvinskianus DC.—Hierba, VA-39 , México, D.F 613 Ávila Alrarharn at al Cit bh M F Erigeron longipes DC —hHierba, VA-40 Erigeron pubescens Kunth—Hierba, VA-41 Eupatorium lucida (Ortega) R.M. King €: H. Rob.—Arbusto, A-42 Eupatorium oreithales (Greenm.) B.L. Turner.—Hierba, VA-43 Fupatorium schaffneri Sch. Bip. ex Robinson—Arbusto, MV-15 Florestina pedata (Cav.) Cass.—Hierba, arvense, VA-44 Galinsoga parviflora Cav.—Hierba, arvense, MR-10 Gamochaeta americana (Mill) Wedd.—Hierba, arvense, ERA VA-45 Gnaphalium chartaceum Greenm.—Hierba, arvense y ruderal, VA-46 Gnaphalium inornatum DC.—Hierba, VA-47 e hal; lio! i Sch Bin ex Klat var gh) D.L. Nash.—Hierba, MR-11 Gnaphalium salicifolium (Bertol) Sch. Bip.—Hierba, MR-12 Gnaphalium s caule DC.—Hierba, arvense, VA-48 Haplopappus stoloniferus DC — Hierba, VA- Helenium id ii (DC) A. Sia Her MR-13 Helianthus lacinata A. Gray—Hierba, M Hieracium Pole S.F. Blake— e ma 7 E IRAR um Less. ma EES cC E DIAI D Maa 5 & Moc.—Hierba, VA-50 emsl.—Arbusto, Montana frutescens (Mairet. ex DC) H BEE adscendens (Sch. Bip. ex Hemsl.) R. L. Rob & Te o (Kunth) A. Gray—Hierba, VA-53 Packera sanguisorbae (DC) C. Jeffrey—Hierba, VA-54 Packera a eile (DC) WA Weber & A. Lóve—Hierba, -55 Parthenium hysterophorus L.—Hierba, MV-19 Perymenium berlandieri DC.—Arbusto, VA-56 Pinaropappus (Less.) Less. var. roseus—Hierba, VA-57 Piqueria pilosa Kunth—Hierba, VA-58 Piqueria trinervia Cav.—Hierba, MR-15 Pseudognaphalium arizonicum A. Gray—Hierba, MR-16 eo GIN UI del dl DC.—Hierba, MR-17 DC. var. nataliae EJ Espinosa— Hierba, VA-59 ide albonervia (Greenm.) H Rob. €: Brettell —Arbusto, A-60 mos angulifolia (DC.) H. Rob. & Brettell —Arbusto, VA-61 ee SC -johannis (DC.) H. Rob. & Brettell —Arbusto, otra plata (Benth.) H. Rob. €: Brettell —Hierba, ob reticulata (DC.) H. Rob. & e MV-20 Roldana sinuata B.L. Turner—Hierba, Sabazia humilis (Kunth) T arvense, VA-65 Sanvitalia procumbens Lam.—Rastrera, arvense, VA-66 Senecio callosus Sch. Bip.—Hierba, VA-67 Senecio cinerarioides Kunth—Arbusto, indicadora de incen- i 68 Senecio jacalensis Greenm.—Hierba, VA-69 Senecio multidentatus Sch. Bip. ex Hemls.—Hierba, VA-70 Senecio roseus Sch. Bip.—Hierba, MV-21 Senecio salignus DC.—Arbusto, VA-7 1 Sigesbeckia pis Kunth—Hier es arvense, MV-22 Sigesbeckia orientalis L—Hierba, VA- Sonchus oleraceus L.—Hierba, VA- "x Stevia glandulosa Hook. & Arn.—Hierba, VA-74 Stevia incognita Grashoff—Hierba, MV-23 Stevia monardifolia Kunth——Hierba, MV-24 Stevia myricoides McVaugh—Hierba, MV-25 Stevia ovata Willd. var. ovata.—Hierba, MV-26 Stevia purpusii B.L. Rob.—Hierba, VA-75 Tagetes erecta L.—Hier 7 Tagetes triradiata Greenm.—Hierba, VA-7 araxacum officin —Hierba, arvense y ruderalVA-78 Verbesina REUS B L. Rob. €: Seaton—Arbusto, MV-27 Verbesina virgata Cav.—Arbusto, VA-79 Vernonia alamanii DC.—Arbusto, VA-80 Villanova achillaeoides (Less.) Less.—Hierba, VA-81 Berberidaceae Berberis moranensis Schult. & Schult. f —Árbol, VA-82 Berberis schiedeana Schltdl.—Arbusto, VA-83 Betulaceae Alnus acuminata Kunth ssp. arguta (Schltdl) Furlow—Arbol, A-8 Alnus acuminata Kunth ssp. glabrata (Fernald.) Furlow— Árbol, VA-85 Alnus jorullensis Benth. ssp. jorullensis— Árbol, VA-86 Boraginaceae Hackelia mexicana (Schltdl. & Cham.) I.M. Johnst.—Hierba, MV-28 uo Lasiarrhenum trinervium (Lehm.) B.L. Turner—Hierba, VA-88 Lithospermum strictum Lehm.—Hierba, ruderal, VA-89 Lai sylvatica Hoffm.—Hierba, MV-29 Brass Se pes Ledeb.—Hierba, VA-90 Brassica campestris L. mire ~ 91 Brassica rapa L.—Hierba, M Capsella bursa-pastoris L. Med Herbs VA-92 Cardamine hirsuta L.—Hierba, C LONE EE EE e VA-93 (Cham. & Schitdl) O.E. Schultz—Hierba, arvense, VA- 94 Draba E Kunth—Hierba, VA-95 Draba nivicola Rose—Hierba, VA-96 Eruca sativa Mill.—Hierba, arvense, MV-31 Erysimum capitatum (Douglas ex Hook) Greene— Hierba, MV-32 eo Roo : var. pubescens (Greene) C.L. Hitchc.— ie N-33 me yide sn? Rollins—Hierba, VA-97 Raphanus raphanistrum a, arvense, VA-99 Rorippa mexicana ës & Sessé) Standl. & Steyermark— Hierba, VA-98 614 Rorippa nasturtium-aquaticum (L.) Hayek—Rastrera, MR-18 Sisymbrium officinalis (L) Scop—Rastrera, VA-100 Buddleja Buddleja ER Kunth ssp. cordata Kunth—Árbol, VA-102 Buddleja parviflora Kunth—Árbol, MV-34 Campanulaceae Diastatea micrantha (Kunth) McVaugh—Hierba, ruderal, MV-35 Diastatea tenera (A. Gray) McVaugh—Hierba, VA-103 Lobelia gruina Cav. var. gruina—Hierba, arvense y ruderal, VA-104 Lobelia laxiflora Kunth var. angustifolia A. DC.—Hierba, arvense eral, VA-105 Lobelia schmitzii E. Wimmer—Hierba, VA-106 Caprifoliaceae Sambucus nigra ver. canadensis L. (L) B.L. Turner—Árbol, VA-107 hori 'crophyll th—Arbusto, VA-108 Viburnum stenocalyx (Oerst) Hemsl.—Árbol, VA-109 Caryophyllaceae Analia Ge EE ER VA-110 Hierba, VA-11 Arenaria apo dies Wii ex x Schitdl —Hierba, A 112 Arenaria oresbia Greenm.—Hierba, VA-113 Arenaria paludicola RC VA-114 Arenaria reptans Hemsl.—Rastrera, VA-115 Cerastium molle Bartl. —Hierba, VA-116 Cerastium nutans Raf. —Hierba, arvense, VA-117 Cerastium orithales Schltdl.—Hierba, VA-118 Cerastium purpusii Greenm.—Hierba, VA- na Cerastium tolucense D.A. Good—Hierba, M Cerastium vulcanicum Schltdl. Mp WEG VA-120 Drymaria effusa A Sici ea Drymaria laxiflora Benth.—Hie ci o (Cham. ^ Soch Fenzl ex Rohrb.— MR-2 n mui (Ser) Ea 2 36 Drymaria is S. Watson—Hierba, M Drymaria alos Schltdl. & mision ruderal, VÀ-122 Stellaria cuspidata Willd. ex Schltdl.—Hierba, VA-123 Stellaria media L—Hierba, VA-124 Stellaria umbellata Turcz.—Hierba, MV-38 Cistaceae Helianthemum glomeratum (Lag.) Lag. ex DC.—Hierba, A-125 Clethraceae Clethra mexicana DC.—Arbol, VA-127 Cornaceae Cornus disciflora DC — Árbol, VA-131 Cornus excelsa Kunth—Árbol, VA-132 Crassulaceae Schitdl.) R Hierba, VA- Echeveria gibbiflora Mac & ae ex (DC. Sis s " e Echeveria secunda Booth ex Lindl.—Hierba, VA-135 Sedum batallae Barocio—Hierba, VA-136 —ÀÓ LT di £T. s/a ICAda £117 Sedum bourgaei Hemsl.—Hierba, VA-137 Sedum jaliscanum S. Watson—Hierba, MV-41 Sedum moranense Kunth—Hierba, MV-42 Sedum oxypetalum Kunth—Arbusto, VA-138 Sedum die A. DC. ssp. parviflorum (R.T. Clausen) RT. sen—Hierba, VA-139 Tillaea pm Ruiz & Pavón—Hierba, Villadia batessii Baehni & Macbride— a A 141 Cucurbitaceae Cucurbita o Naudin—Hierba, VA-142 Sicyos deppei G. Don—Hierba, arvense, VA-143 Sicyos parviflorus Willd.——Hierba, ruderal, VA-144 Ericaceae Arbutus xalapensis Kunth—Árbol, o 148 aphila umbellata L.—Arbusto, MV-4 Comarostaphylis discolor (Hook) ii var. discolor—Árbol, VA-149 E 3 Gaultheria lancifolia Smali —Arbusto, VA-150 Vaccinium caepitosum Michx.—Arbusto, VA-151 Euphorbiaceae Euphorbia furcillata Kunth var. furcillata—Hierba, VA-152 Euphorbia indivisa (Engelm.) Tidestr.—Hierba, MV-44 Euphorbia peplus L.—Hierba, VÀ-153 Euphorbia prostrata Ait.—Hierba, VA-154 Fabaceae Acacia melanoxylon R. Br.—Árbol, VA-193 iin guatemalensis Hemsl. var. brevidentatus neby—Hierba, VA-19 m acm (Ait.) Bullock—Hierba, MV-62 Desmodium aparines (Link) DC.—Hierba, MV-63 Desmodium molliculum (Kunth) DC.—Hierba, VA-195 Desmodium neomexicanum A. Gray—Hierba, VA-196 Erythrina coralloides DC.—Árbol, BGH-2983 Eysenhardtia polystachya (Ortega) Sarg.—Árbol, VA-197 Lathyrus odoratus L.—Hierba, VA-198 Lupinus elegans Kunth—Hierba, VA-199 Lupinus glabratus J. Agardh— "isi pua 2984 CUPIS E Kanthes GE Hernsl.) am n 2 Darme ab: Arkh ety » ihe LX BGH-298 Trifolium e Kunth—Hierba, VÀ-201 Trifolium repens L.—Hierba, VA-202 Vicia faba L.—Hierba, BGH-2986 Vicia pulchella Kunth (Hemsl) ssp. mexicana CR. Gunn— Trepadora, BGH-2987 Vicia sativa L—Trepadora, VA-203 agaceae Quercus candicans Neé—Arbol, VA-155 Quercus castanea Neé—Árbol, VA-156 Quercus crassipes Humb. & Bonpl.—Árbol, VA-157 Quercus dysophylla Benth.—Árbol, VA-158 Quercus laurina Bonpl.—Árbol, MV-4 Quercus obtusata (Willd) Pursh—Arbol, VA-159 Quercus rugosa Neé—Árbol, VA-160 um , México, D.F 615 Ávila Abarharn at al Est H A | J I eH F Garryaceae Garrya laurifolia Hartw. ex Benth. ssp. laurifolia—Árbol, Gentianaceae ouis Dau ud (G. an SE —Hierba, VA-161 enth. ) Gillett.—Hierba, VA-162 Halenia brevicornis (Kunth) E Don—Hierba, VA-163 Halenia pringlei B.L. Rob €: Seaton—Hierba, VA-164 Geraniaceae Erodium cicutarium (L) L'Hér. ex Aiton—Hierba, arvense, -165 Geranium latum EN an Geranium lilacinum Knuth—Hierba, Geranium die e DC. ens arvense, VA-166 Geranium seemanii Peyr.—Rastrera, arvense, VA-167 Grossulariaceae Ribes affine Kunth—Arbusto, VA-169 Ribes ciliatum Humb. €: Bonpl. ex Schult.—Arbusto, MR-33 Ribes coliosum Howell—Arbusto, MV-49 Guttiferae Hypericum silenoides Juss. var. mexicanum (Keller) Rodr.— Hierba, VÀ-170 Hydrophyllaceae Nama dichotomum Choisy var. oe (Ruiz & Pav.) ois oisy—Hierba, arvense, MV-5 Phacelia platycarpa (Cav.) Spreng. uM arvense, MV-51 Wigandia urens (Ruiz & Pav.) Kunth—Arbusto, ruderal, VA-171 Lamiac agostache mexicana (Kunth) Linton & Epling—Hierba, MV-5 Cunila o Benth.—Hierba, VA-177 Lamium purpureum L.—Hierba, VA-178 EE caulescens (Ortega) Epling—Hierba, arvense, -179 me vulgaris L.—Hierba, VA-18 Salvia amarissima e VA-181 Salvia concolor Lamb. —Hierba, VA-182 Salvia elegans Vahl ex Benth.—Hierba, VA-183 Salvia gesneraeflora Lindl. 8: Paxton—Hierba, VA-184 Salvia mexicana L. var. minor (L.) Benth.—Hierba, VA-185 Salvia mexicana L.—Hierba, VA-186 Salvia microphylla L.var. microphylla L.—Hierba, VA-187 alvia microphylla L.var. neurepia (Fern.) Epling—Hierba, VA-188 Salvia patens Cav.—Hierba, MV-57 Salvia polystachya Cav.—Hierba, MV-58 Salvia E Kunth.—Hierba, MV-59 Salvia reptans Jacq.—Hierba, VA-189 Salvia stricta Sessé & Moc. — Hierba, VA-190 Stachys agraria Schltdl. & Cham.—Hierba, arvense, VA-191 Stachys coccinea Ortega—Hierba, VA-192 Se EE SS ee n 2979 Stachys sanchezii Rzed. & À. Garcia Hierba, VA 333 Lauraceae Litsea glaucescens Kunth—Arbol, MR-34 ntibulariac fuse Gees (DC) E. Mey.—Rastrera, BGH-2988 Linaceae Linum orizabae Planch.—Hierba, VÀ-204 Linum usitatissinum L—Hierba, VA-2 Loranthacea EE globosum Hawksw. & Wiens—Epífita, VA-206 & Bonpl. ex Willd.) J. PresI— Epífita, TNAM CUm (DC) Oliv.—Epífita, VA-208 Lythraceae Cuphea aequipetala Cav.—Hierba, arvense y ruderal, VA-209 Malvaceae Kearnemalvastrum subtriflorum (Lag.) D.M. Bates—Hierba, Oleacea Fraxinus da (Wenz) Lingelsh.—Árbol, VA-212 Ligustrum japonicum Thunb.—Árbol, exótica, VA-213 Onagraceae ine ME Kunth ssp. microphylla—Arbusto, E RE Kunth ssp. thymifolia—Arbusto, VA-215 Gaura coccinea Pursh—Hierba, arvense, 16 Gaura mutabilis Cav.—Hierba, VÀ-217 Lopezia racemosa Cav. ssp. racemosa—Hierba, arvense, VA-21 a Oenotl ticola (Loes) Mu ierba, VA-219 Vihar lo PATOS RACE M E pubescens Willd, ex Soreng.—Hierba, arvense, cena pups uz Hei BGH-2951 era rosea L'Hér. ex Aiton—Hierba, arvense, BGH-2952 Orobanchaceae Conopholis alpina Liebm.—Hierba, VA-224 Oxalidaceae Oxalis alpina (Rose) R. Knuth—Rastrera, VA-225 Oxalis corniculata L.—Rastrera, arvense y o VA-226 Oxalis nelsonii (Small) R. Knuth—Rastrera, MR- Papaveraceae Argemone ochroleuca Sweet—Hierba, BGH-2953 Argemone platyceras Link & Otto—Hierba, arvense y ruderal, A-227 Passifloracea Passiflora Sum Zucc.—Trepadora, VA-228 Phytolaccaceae Phytolacca icosandra L.—Hierba, ruderal, VA-229 Piperaceae Peperomia campylotropa A.W. Hill—Hierba, VA-239 Peperomia hintonii Yunck.—Hierba, VA-240 616 Plantaginaceae mE australis Lam. ssp. hirtella (Kunth) Rahn—Hierba, uderal, MR-36 Td linearis Kunth—Hierba, MR-37 Plantago major L—Hierba, VA-241 Plantago nivea Kunth—Hierba, arvense y ruderal, VÀ-242 Polemoniaceae Loeselia mexicana (Lam.) Brand—Hierba, arvense, VA-302 Polemonium mexicanum Sessé € Moc. ex DC.—Hierba VA-261 a Polygalaceae Eriogonum jamesii Benth — Arbusto, BGH-2991 Monnina ciliolata (D. Dietr.) DC — Arbusto, BGH-2992 Polygonum aviculare L.—Hierba, arvense, BGH-2993 Polygonum hydropiperoides Michx.—Hierba, VA-262 Polygonum punctatum Elliott var. eciliatum Small —Hierba, rvense, VA-263 Rumex deal L.—Hierba, arvense, BGH-2994 Rumex us L.—Hierba, arvense, BGH-2954 Rumex Ee Rech. f—Hierba, VA- 264 Rumex obtusifolius L. ssp. agrestis (Fr) Danser—Hierba, ar- V VA-265 Portulacaceae Calandrinia megarhiza Hemsl.—Hierba, VA-268 CI j A E | tata f» VA ZIEL ssp = (Ry ]1 ) lohn M. Miller & K.L. Chambers—Hierba, BGH-2955 Montia chamissoi (Ledeb. ex Spreng.) Greene—Hierba, VA-269 Primulaceae Anagallis arvensis L.—Hierba, arvense, VA-270 Pyrolaceae Chimaphila umbellata (L) W.P-C. Barton—Hierba, VA-272 Monotropa uniflora L.—Hierba, VA-273 Ranunculaceae Clematis dioica L.—Arbusto, VA-274 Ranunculus dichotomus Moc. & Sessé ex DC.—Hierba, A-275 Ranunculus donianus Pritz—Hierba, VA-276 Ranunculus multicaulis D. Don ex G. Don—Hierba, BGH-2995 Ranunculus petiolaris Kunth ex DC.—Hierba, BGH-2956 Ranunculus praemorsus Kunth ex DC. var. amellus (Brig) T. Duncan—Hierba, VA-277 Ranunculus sibbaldioides Kunth ex DC.—Hierba, VA-278 Thalictrum pubigerum Benth.—Hierba, VA-27 ae Reseda luteola L.—Hierba, arvense, VÀ-280 Rhamnaceae Ceanothus coeruleus Lag — Árbol, VA-281 Rosaceae Acaena elongata L.—Arbusto, ruderal, VA-282 Alchemilla aphanoides L.—Hierba, BGH-2996 Alchemilla pectinata Kunth—Hierba, BGH-2997 Alchemilla procumbens Rose—Rastrera, BGH-2998 Alchemilla sibbaldiifolia Kunth var. bourgeaui (Rydb) Perry— Rastrera, VÀ-283 bclli vulcanica Schltdl. & Cham.—Rastrera, VA-284 melanchier denticulata (Kunth) K. Kock—Árbol, VA-285 Cenepa mexicana Moc. & Sessé ex DC.—Árbol, bes 286 Hierba, VA-28 Fragaria mexicana Schltdl. —Hierba, VA-288 Potentilla candicans Humb. €: Bonpl. ex Nestl.—Hierba, VA-289 EIA DN o laxiflora Drew—Hierba, BGH-2957 entilla ranunculoides Kunth—Hierba, VA-290 Ge rubra Willd. ex Schltd!—Hierba, VA-291 Prunus serotina Ehrh. ssp. capuli (Cav) McVaugh—Árbol, VA-292 Rosa canina L.— Arbusto, VA-293 Rubus liebmannii Focke—Arbusto, VA-294 Rubus pumilus Focke—Hierba, VA-295 Rubiaceae Bouvardia multiflora (Cav.) Schult. & Schult.—Arbusto, -2958 Bouvardia obovata Kunth—Hierba, BGH-2959 Bouvardia ternifolia (Cav.) Hierba, VA-296 Didymaea alsinoides ( Cham. & Schltdl.) Standl.—Hierba, VA-297 Galium aschembornii Nees & S. Schauer—Hierba, BGH-2960 Galium mexicanum Kunth—Hierba, BGH-2961 Galium praetermissum Greenm.—Hierba, BGH-2962 Hedyotis cervantesii Kunth—Hierba, VA-298 Sherardia arvensis L.—Hierba, VA-299 Sabiaceae Meliosma dentata (Liebm) Urb.—Árbol, VA-300 Salicaceae Salix oxylepis C. K. Schneid.— Arbusto, VA-301 Salix paradoxa Kunth—Árbol, VA-302 Saxifragace Heuchera mE Hemsl.—Hierba, VA-303 Scrophularia Castilleja arvensis cedi & Cham.—Hierba, arvense, VA-304 Castilleja moranensis Kunth—Hierba, VA-30 Castilleja tenuiflora Benth.—Hierba, arvense y ruderal, VA-306 Lamourouxia SE (Cham. € Schltdl.) W. R. Ernst.— Hierba, VA- Mimulus prium Kunth—Hierba, VA-308 Penstemon campanulatus (Cav.) Willd.—Hierba, ruderal, VA-309 Penstemon gentianoides (Kunth) Poir.—Hierba, indicadora de incendio, VA-31 Penstemon roseus (Cerv. ex Sweet) G. Don—Hierba, BGH-2963 Sibthorpia repens (L) Kuntze—Rastrera, VA-31 1 Veronica serpyllifolia L.—Hierba, VA-312 os Árbol, VA-314 Geen EH Kunth—Arbusto, VA-315 Agila AbaeRhavi atal Etadi idas ] Lal via AA laf a F d México, D.F 617 Lycianthes moziniana (Dunal) Bitter—Hierba, arvense, B E Lycianthes peduncularis (Schlecht.) 3itter—Hierba, VA-316 Nicotiana glauca Graham—Hierba, exótica, VA-317 Physalis chenopodifolia Lam.—Hierba, VA-318 Physalis coztomatl Dunal—Hierba, VA-319 Physalis orizabae Dunal—Hierba, bea ~ 2965 Physalis pringlei Greenm.—Hierba, Physalis stapelioides (Regel) Man VA-321 Solanum EE Mil mal ba VA 322 Solanum cervantesii Lag.—Arbusto, VA-324 Solanum demissum Lindl.—Hierba, BGH-2966 Solanum nigrescens M. Martens & Galeotti—Hierba, arvense,VÀ-325 Symplocaceae Symplocos citrea Lex. ex La Llave & Lex.—Árbol, BGH-2967 Urtic Urtica andas Pursh—Hierba, arvense, BGH-2969 Urtica urens L.—Hierba, ruderal, BGH-2999 Valerianaceae Valeriana clematitis Kunth—Hierba, BGH-3000 Valeriana sorbifolia Kunth—Hierba, BGH-3001 Verbenaceae Verbena bipinnatifida Nutt.—Hierba, BGH-2970 Violaceae Viola guatemalensis W. Becker—Hierba, BGH-2971 Viola hemsleyana Calderón—Hierba, BGH-2972 Viola hookeriana Kunth—Hierba, BGH-2973 Viola humilis Kunth—Hie A-4 Viola painteri Rose & dou ON VA-126 Liliopsida Agavaceae Agave atrovirens Karw. ex Salm- nm MV-2 Agave macroculmis Tod.—Hierba, MV- Agave salmiana Otto ex Salm- HOHER VA-5 Furcraea bedinghausii K. Koch—Árbol, VA-6 Manfreda pringlei Rose—Hierba, BGH-2976 Commelinaceae Commelina coelestis Wilkd.—Hierba, arvense, MV-39 Commelina diffusa Burm.—Hierba, VÀ-128 Commelina orchioides Booth.—Hierba, MV-40 Gibasis pulchella (Kunth) Raf—Hierba, VA-129 Tinantia erecta (Jaca) Standl.—Hierba, VA-130 Bromeliac Tillandsia no (Mez) L.B. Smith—Epífita, VA-101 Cyperaceae Carex anisostachys Liebm.—Hierba, MR-23 Carex boliviensis Van Heurck & Múell. Arg.—Hierba, MR-24 Carex chordalis Lieom.—Hierba, MR-25 Carex tuberculata Liebm.—Hierba, MR-26 Cyperus e (Jacq.) Standl.—Hierba, arvense y ruderal, MR- Cyperus manimae Kunth var. rae T MR-28 Cyperus niger Ruiz & Pav.—Hierba, Cyperus semiochraceus Boeck.— gert m 30 Cyperus seslerioides Kunth—Hierba, MR- "m Eleocharis dombeyana Kunth—Hierba, M = montevidensis Kunth var. Geer SE kunthii Nees—Hierba, JB-3 ypoxidaceae Hypoxis mexicana Schult—Hierba, VA-172 Iridaceae ene ee SST —Hierba, VA-173 Hierba, VA- 4 Sisyrinchium dia Caden 8 feedowski-—Hierba; m scabrum Schitdl, & Cham.—Hierba, MV-52 Sisyrinchium schaffneri S. Watson—Hierba, VA-176 Sisyrinchium tenuifolium Humb. & Bonpl. ex Willd.—Hierba, MV-53 Juncaceae Luzula denticulata Lieom.—Hierba, MV-54 Luzula racemosa Desv.—Hierba, MV-55 Liliaceae Echeandia mexicana Cruden—Hierba, BGH-2989 Stenanthium frigidum (Schltdl. & Cham.) Kunth—Hierba, BGH-2950 Nolinaceae Nolina parviflora (Kunth) Hemsl.—Árbol, VA-211 Orchidaceae Corallorhiza wisteriana Conrad.—Hierba, A 221 Malaxis myurus (Rchb. f) Kuntze—Hierba, 2 Spiranthes hyemalis A. Rich. & EE VA-223 Poaceae Agrostis subpatens Hitchc.—Hierba, MR-38 Agrostis tolucensis Kunth—Hierba, BGH-2974 Arena sativa L—Hierba, ee ) Nash—Hierba, BGH-2975 Brachypodium dll Seba ex Beal—Hierba, VA-243 Bromus dolichocarpus Wagnon—Hierba, VÀ-244 Calamagrostis Ge (Kunth) Trin. ex Steud.—Hierba, 5 formis (Kunth) Scribn. € Merr.—Hierba, VA-246 Festuca amplissima Rupr.—Hierba, VA-247 Festuca tolucensis Kunth—Hierba, VA-249 Muhlenbergia ciliata Kunth—Hierba, VA-2 Muhlenbergia macroura (Kunth) Hitchc.—Hierba, arvense, VA-251 Muhlenbergia nigra Hitchc.—Hierba, VA-252 Munien bergig quadriden Wd ( Ku ia Du ,—Hierba, VA-253 Hierha MR-40 + LJ: Piptochaetium seleri (Pilg.) Henrard— Hierba, VA-254 Schedonorus arundinaceus (Schreb. Dumort..—Hierba, VA-24 mii indicus (L.) R. Br.—Hierba, VA-255 Stipa ichu (Ruiz & Pav) Kunth—Hierba, VA-256 618 t tani itute of Texas 2( Stipa mexicana Hitchc.—Hierba, VA-257 Vulpia bromoides (L.) Gray—Hierba, MR-41 Trisetum kochianum Hern. Torres—Hierba, VA-258 Smilacaceae Trisetum spicatum (L) K. Richt.—Hierba, VA-259 Sm pem moranensis M. Martens & Galeotti—Trepadora, Trisetum virlettii E. Fourn—Hierba, VA-260 VA-313 AGRADECIMIENTOS A los especialistas Dra. Susana Valencia Ávalos (Fagaceae), Biol. Ramiro Cruz Durán (Fabaceae), Dra. Mar- tha Martínez (Euphorbiaceae) y M. en C. Rosa Ma. Fonseca (Pinaceae y Rosaceae). Los autores agradecen a Myriam Rubio Palacios, Lilia Alvarado García y Madai Velasco Vázquez por apoyar en la colecta, procesado y montaje del material botánico, así como en el manejo de la base de datos. REFERENCIAS ALMEIDA, L., A.M. CLeer v A. VELÁZQUEZ. 1997, Fitosociología del | | if lel volcán Popocatépetl, México - In: ALMEIDA-LENERO, L.: Vegetación, fitogeografía y ge del zacatonal alpino y b uc: montanos de la región central de México. Tesis doctoral, Universidad de Amsterdam Álvarez, KE 2000. Geografía de la educación ambiental: algunas propuestas de trabajo en el Bosque de los Dinamos, Área de Conservación Ecológica de la Delegación Magdalena Contreras. Tesis de licenciatura en Geografía, Facultad de Filosofía y Letras, UNAM, México. ÁviLa-AkerserG, V. 2002. La vegetación en la cuenca alta del río Magd f florístico, fitosociológico y estructural. Tesis de licenciatura en Biología, Facultad de Ciencias, UNAM, México. ÁviLA-AKERBERG, V. 2004. Autenticidad de los bosques en la Cuenca del rio Magdalena. Diagnóstico hacia la res- tauración. Tesis de Maestría, Facultad de Ciencias, UNAM, México. BASE DE DATOS en: http://mobot.mobot.org/wt3/s earch/vast.html. BEETLE, AA 1983, 1987. Las Gramíneas de México. Tomos | y Il. Comisión Técnico Consultiva para la Determi- nación de los Coeficientes de Agostadero (COTECOCA) y Secretaría de Agricultura y Recursos Hidráulicos (SARH), México. BONILLA-BARBOSA, J. R. v J.A. ViANA-LASES. 1997. Listados Florísticos de México XIV. Parque Nacional Lagunas de Zem- poala. Instituto de Biología, Universidad Nacional Autónoma de México, México CALDERÓN DE RzevowskI G. y J. Rzepowski (eds.) 2001. Flora Fanerogámica del Valle de T CONABIO-Instituto de Ecología, A.C. México, D.F. CASTILLO-ARGUERO, S., G. MONTES-CARTAS, M.A. RoMERO-RomERO, Y. MARTINEZ-OREA, P. GUADARRAMA-CHÁVEZ, |. SANCHEZ-GALLEN y O. CasniLo-NüREz. 2004. Dinámica y conservación de la flora del matorral xerófilo de la Reserva Ecológica del Pedregal de San Ángel (D.F, México). Bol. Soc. Bot. México. 74:51-75 CERVANTES, B.F. 1969. Algunas consideraciones geomorfológicas de la Cuenca del río Magdalena. Universidad Nacional Autónoma de México. Bol. Inst. Geogr. 2:89-107. CORENADER. 2006. El iii BE conservación del Distrito Federal. Secretaría de Medio Ambiente del Distrito Federal kl fl / / j V (http:// corenade CORNEJO-TENORIO, G., A. pee B. FARFÁN, J.L. wee Y G. lgarra-MansíquEZ. 2003. Flora y vegetación de las zonas núcleo de la Reserva de la Biosfera Mariposa Monarca, México. Bol. Soc. Bot. México 73:43-62. Cronauist, A. 1988. The evolution and classification of flowering plants. 2a. ed, Columbia University Press, New York. DELEGACIÓN LA MAGDALENA CONTRERAS. 2006. Información delegacional. ( mx). DaHLGREN, R.M.T., H.T. CurrorD y PF. Yeo. 1985. The families of the monocotyledons. Structure, evolution and tax- onomy. Springer-Verlag, New York. DIARIO OFICIAL DE LA FEDERACIÓN. 2002. Norma Oficial Mexicana-059-SEMARNAT-2001, México Espinosa, G.F.). y J. SARUKHAN. 1997. Manual de malezas del Valle de México. Universidad Nacional Autónoma de México y Fondo de Cultura Económica. |. flor if |=| a Ys LI ~E d Avila Akerberg at al i Erto di id ar | lal rio M dal i México, DF 619 fe, García, E. 1987. Modificaciones al sistema de clasificación climática de Kóppen. Instituto de Geografia. UNAM. México, D.F. JuJNOvskv, J. 2006. Servicios ecosistémicos relacionados con el recurso agua en la Cuenca del Río Magdalena, Distrito Federal, México. Tesis Maestría, Facultad de Ciencias, UNAM. México. MADRIGAL, S. 1967. Contribución al conocimiento de la ecología de los bosques de Oyamel (Abies religiosa H.B.K. Schl. Et Cham.) en el Valle de México. Boletín Técnico 18, Instituto Nacional de Investigaciones Forestales, México. Mazarl, M. (Comp). 2000. Dualidad población-agua: Inicio del Tercer Milenio. Ed. El Colegio Nacional. México, D.F Mepina-Garcia, C., F. Guevara-Férer, MA Martinez-Ropricuez, P. SiLva-SAENZ Y M.A. CHAvez-CARBAJAL. 2000. Estudio flo- rístico en el área de la comunidad indígena de Nuevo San Juan Parangaricutiro, Michoacán, México. Acta Bot. Mex. 52:5-41. MIRANDA, F. y E. HERNÁNDEZ-XOLOCOTZIN. 1963. Los tipos de vegetación de México y su clasificación. Bol. Soc. Bot. México 28:29-1 79. Missouri BOTANICAL GARDEN. 2006. Plant species database. (http:// Nava, M. 2003. 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Los géneros de plantas vasculares de la flora de México. Bol. Soc. Bot. México 75:105-135. org) j= sud | RAL — PA | lh La Lerdo £T. BILE 620 BOOK NOTICE PAMELA A. MATSON AND ÁSHOK GADGIL (eds). 2007. Annual Review of Environment and Resources: Volume 32. 2007. (ISBN 978-0-8243-2332-7, hbk; ISSN 1543-5938). Annual Reviews Inc., 4139 El Camino Way, PO. Box 10139, Palo Alto, California 94303-0139, U.S.A. (Orders: www.annualreviews.org, science@annualreviews.org, 1-800-523-8635, 1-650-493-4400, 1-650-424-0910 fax). $80.00 indiv., $197.00 inst. (USA), 449 pp., indexes, 7 5/8" x 9 3/8". Contents of Volume 32: Who Should Read This Series? Preface Earth’s Life Support System Feedbacks GE Terrestrial Ecosystems t ae n ena dedi Hes die lie Halton A. Peters, Nona R. Chiariello tem C n g ocene— Scott C. Doney, David S. Schimel The Nature and V. FE Servi An O iew Highlighting Hydrologic Servi Kate A. Brauman, Gretchen C. Daily, T. Ka'eo Duarte, See Moon ney Perspective—Cheryl Palm, Pedro Sanchez, Sonya Ahamed, Alex Awiti Soils: A Human Use of Environment RUE Resources Ambuj D Sagar, Sivan Kartha Models of Decision Making and gesidentil Energy Use— Charlie Wilson, Hadi Dowlatabadi Renewable Energy Futures: Targets, Scenarios, and Pathways— Eric Martinot, Carmen Dienst, Liu Weiliang, Chai Qimin ie ae a m a a RM T. Wolf d Nitrogen Cycles—Henning Steinfeld, Tom Wassenaar Global Envi lards for Industry— David P. indi Trina Hamilton, Matthew T. Huber Industry, Environmental Policy, and E Population and Environment—Alex de Sherbinin, David Carr, Susan Cassels, Leiyywen Jiang Management, EE and Governance A Resources and DERE E 10 Daniel P Prece £ Fla 1 oe pend A Review of | k—Donald R. Nelson, W. Neil Adger, Katrina Brown ee Themes omen, Water, and SE T 3 Cr FEMA! 42 Isha Ray hors, Volumes 23-32 o Chapter Titles, Volumes 23-32 J. Bot. Res. Inst. Texas 2(1): 620. 2008 PRAXELIS CLEMATIDEA (ASTERACEAE), A GENUS AND SPECIES NEW FOR THE FLORA OF NORTH AMERICA J. Richard Abbott C. LeAnn White pi rtment of Bot Yid Department of Infectious Diseases sity of Flori isis thology mess Gates Er USA. rsity of Florida Gainesville, Florida 32611, U.S.A. a Florida 32611, U.S.A irabbottabotany.ufl.edu WhiteC@vetmed.ufledu and sd of Horida ke IE son Hall, PO. Box os G nesville, Florida 32611, U.S S. Barry Davis University of Florida Herbarium (FLAS) Florida Museum of Natural jd 379 Dickinson Hall, PO. Box 11057 Gainesville, Florida 3261 1, SECH taxongfimnh.ufi.edu ABSTRACT Praxelis clematidea (Asteraceae) is a new genus and species for North America. Praxelis is compared with similar species, and their diagnostic characters are discussed. RESUMEN P lis cl tidea (A ) es un gé i América del Norte. Se compara Pra» elis con i i , Y se discuten sus caracteres diagnósticos. Field work in central Florida by the second author has led to the discovery ER a plant species new to North America, Praxelis clematidea (Griseb.) R.M. King & H. Rob. (Fig. 1). The sp to Árgentina, Bolivia, Paraguay, and Brazil (King & Robinson 1987). It is also erroneously reported from Venezia Waterhotise et al. 2003). It has been naturalized in Australia, China, and Hong Kong (Corlett & Shaw 1995; Veldkamp 1999; Waterhouse 2003). There are 13 additional species of Praxelis Cass., all restricted to South America, none of which are known as exotics elsewhere (King & Robinson 1987). A more complete description of P. clematidea is available on-line (U.S. Forest Service 2007), as are photos (Waterhouse et al. 2003), including comparison with the similar Ageratum houstonianum Mill. (Pollock et al. 2004). Voucher specimen data: UNITED STATES. Florida. Orange eh 28?24'80"N 81°37'13"W bibe N'W of Kissimmee, on private property near the junction o: M Road with FL 429, aband ( ), herb ca. 2' tall, flowers ini purple, rare, 26 Jul 2006, LeAnn White 1 [FLAS], det: S Barry Davis (FLAS), Oct 2006, v John Pruski (MO), Mar 2007. Orange Co.: 28?24'9.7"N 81°37'11.5"W (WGS84), WSW E Orlando, S f Reed | id Infil B Reedy C District along Old Hartzog Road EE E on FL 429 and NM Bay, W ep Reedy GEER, accessed S a Steeg Road. Roadside and edge of pine plantation with Bidens alba, H baxillaris, Lantana camara, P. tatum, and Rhynchelytrum repens, suffrutescent herb to ca. 1.3 m tall, usually with several branches from the base, with very n smell ion cat-spray like), flowers pale pinkish purple, locally abundant, forming a large patch with hundreds of stems, 17 Jul 2007, J. Richard Abbott 22887, with Tim Burns & Melissa Clark [B, FLAS, G, GH, HBG, IJ, K, L, LD, M, MAD, MAPR, MEXU, MICH, MIN, MO, MS TEX, U, US, boit WU, e E Co.: 28º26'56.9"N ad 46.6"W (WGS84), SW of Orlando, SE of Lake Hancock, NW corner of j ith | ] to N, accessed E of FL 429 on eun Road, Mn get turns N CoL J 1 T 4 LI L D 1 Roadside +. Did all a, Cyt llifoliur Paspal tatum, Sida rhombifolia, and Urena lobata, forming a large patch witl 50 WEM here, 17 Jul 2007, J. Richard Abbott 22888, I Ret Rac Inct Tavas 2(1): A — 626. 2008 D | Fal Dos H In LI PE ET TE Al VP 622 with Melissa Clark [BM, BRIT, C, CAS, CICY, CONN, DAV, DUKE, E, ETU, F, FLAS, FSU, FTG, GOET, JBSD, LSU, MO, PIHG, SEL, UC, GÁ RP, USF, UWFP, W] and 8 Mar 2008, J. Richard Abbott 24194 [FLAS].. Praxelis clematidea: herb to suffrut t shrub to ca. 1.3 m tall; stems soft pubescent (densely villous to hirsute, especially above); petioles usually very short, well less than 1/4 the blade length (except on the lowest leaves which may have petioles up to ca. 1/3 or more the blade length; these leaves are often gone by the time the plant is in fruit); blades ovate, rounded to cuneately narrowing at the base, generally about half (or more) as wide as long; margins irregularly toothed, i.e., some teeth nearly dentate, others more sharply serrate, with size variation even on same leaf; heads cylindrical-campanulate, clearly longer than wide (ca. 3-4 mm wide, 6—8 mm tall), in lax to dense clusters; phyllaries about 1 mm wide, flat, densely imbricate, although deciduous in fruit, with conspicuous dark striations from the veins. A visit to the region on 17 July 2007 discovered hundreds of individuals of Praxelis at several scattered localities, all disturbed roadsides. This local delen in so many different spots, in conjunction with the original collection in a relatively remote and i area, suggests that Praxelis has been in Florida for some time and has been spreading. The site where the species was first bcn in July oe is an open old field dominated by exotics [Rhynchelytrum repens (Willd.) C.E. Hubb.; Ricl } es; on edges, Urena lobata L.] and ‘weedy’ natives [Cenchrus spinifex Cav., Heterbtheca subaxillaris (Lam.) Britton & Rusby, Bidens alba (L.) DC.]. The site is a dry, well-drained sandy upland that was once a citrus grove. Although there are no indisputable indicators of the original vegetation, t that it may have been sandhill o e.g., Soma erecta L., Croton EEN L., Galactia elliotti Nutt., Opuntia humifusa (Raf.) R flora i I yrtifolia Willd. (only 1 simt seen), and, along the edges, Euthamia Moni (L. Greene ex Parier & Britton and Verbesina virginica L. The area is now mostly surrounded by dry pine woods that appear secondary, with a remnant swampy baygall nearby to the southwest. The site has about 200 Florida gopher tortoises (Gopherus polyphemus) and a density of about 29.1 tortoises/ha, which is very dense for gopher tortoise populations. Praxelis was completely absent on a follow-up trip on 5 November 2006, at which time the site was also seen to be heavily disturbed by hog rooting (Sus scrofa). One small patch was rediscovered in the same area during a follow-up visit on 23 August 2007, making it clear that the species is established at the site. A follow-up trip on 8 March 2008 to the Lake Hancock Road site found that, as part of a road construction project, the population had been nearly eradicated (although achenes are doubtless still persisting in the area); only one plant was seen (and it was in flower), with the remaining area covered in fresh sod. This did demonstrate that at least some individuals of Praxelis over- winter as above-ground perennials, flowering year-round (or nearly). Eupatorieae are distinguished by the combination of discoid heads, anther bases non-tailed (obtuse, rounded, or truncate), styles usually more or less filiform, style-branch appendages usually terete to clavate, and leaves usually opposite (FNAEC 2006a). Praxelis keys to either mea ue imd or Chromo- laena DC. in FNAEC (2006b), matching key features for both—ca. 20 unequal phyll (the outer ones are generally smaller), 25—30 florets per head, a 3—4 ribbed achene, a pappus of ca. 40 capillary non-plumose bristles, and a glabrous style base. Praxelis has been reported as an annual and a perennial (U.S. Forest Service 2007, Waterhouse et al. 2003) and may behave as either, depending on local climate. A modified SSH is ee here, based on species in North America that OK most similar to E clematidea, i.e., | Eupatorium L. s.l. with petiolate, opposit tri | from base and EM r purple corollas. For certainty al identification in a broader taxonomic context in the absence of compara- tive herbarium material or consultation with a specialist, the best references are the keys and descriptions in Cronquist (1980), King and Robinson (1987), FNAEC (2006a, 2006b), or, for Florida, Wunderlin and Hansen (2003). Abbott et al., Praxelis clematidea, new for North America C Fic. 1. Praxelis clematidea. A. Inflorescence structure with inset of leaf. B. Close-up of flowers. C. Close-up of heads showing conical receptacle. 624 t tani i Texas 2( ARTIFICIAL KEY TO DISTINGUISH PRAXELIS CLEMATIDEA FROM SIMILAR GENERA IN NORTH AMERICA NORTH OF MEXICO ke de Receptacle conical (usually strongly $0 | like structure), or of 5-6 flattened scales (some- times 5-6 tapering setae in A. conyzoides) Rgeratum [20-125 florets; 30-40 phyllaries, (sub-)equal, persistent t decumbent; only 4 spp. in North America, 3 are discussed i the fourth, A. corymbosum Zucc, occurs in AZ and N 2. Pappus of capillary bristles; bristl than 20. 3. Bristles ca. 30; More 35. 70+; phyllaries ca. 25, subequal, persistent Conoclinium [3 spp; 1 discussed Ge me SE 2 are C. dissectum A. Gray of TX, AZ, and n Mille ais King €: H. Rob. of T 3.Bristies ca. 40; florets 25-30; phyllaries 15-25, unequal, decid (al n fru Praxelis [13 Spp. Ro P clematidea all e E n South America] |. Receptacle flat to slightl vex (rarely very shallowly conical in Fleischmannia 4. Phyllaries cdo, 18- -65, unequal; b E ca. 40; florets 6-75 Chromolaena [only 4 spp. in North America, 3 discussed below, us C. bigelovii (A. Gray) R.M. King & H. Rob, a shrub of TX] 4. Phyllaries persistent, 20-30, unequal (to rarely subequal); bristles 20-40 (rarely O or 5), florets (1035-2550) Fleischmannia [only 2 spp. in North America, 1 discussed below, the other is F. sonorae (A. Gray) R.M. King & H. Rob. of AZ & NM] Despite superficial macro-morphological similarity, the genera in the above key are from several different subtribes and do not form a clade (Schilling et al. 1999; Schmidt & Schilling 2000): Ageratinae (Ageratum), Fleischmanniinae (Fleischmannia), Gyptidinae (Conoclinium), Praxelinae (Chromolaena & Praxelis). Most North American regions don’t have more than one or two of these Praxelis “look-alikes,” but a total of eight spe- cies in all four genera is present in Florida: Ageratum conyzoides L. (native to tropical America; rare exotic in southern Florida), A. houstoni Mill. (native to Mexico and northern Central America; occasional exotic in central and southern Florida), A. littorale A. Gray (rare in the keys; vegetatively glabrous, unlike most individuals of the other taxa here), Chromolaena frustrata (B.L. Rob.) R.M. King & H. Rob. (rare in southern Florida), C. ivifolia (L.) R.M. King & H Rob. (native to tropical America; rare exotic in central Florida), C. odorata (L.) R.M. King Ge H. Rob; cia in southern Florida), Conoclinium coelestinum (L.) DC. (common throughout), and Flei ta (Walter) R.M. King & H. Rob. (occasional in northern and central Florida). In central Florida where Praxelis is now known, the most similar species to Praxelis that are most likely to be one are id houstonianum, Een m EORUM and Geiger incarnata, Aserpirit ARO although Chromolaena ] possibility See tl pI ore of taxa similar to Praxelis. APPENDIX SUPPLEMENTAL DESCRIPTIONS The following descriptions are based on field and herbarium observations. This information supplements descriptions in Cronquist (1980) and FNAEC (2006b). All of these species are very different looking when placed side by side. These gen- O are gece to ao the E macroscopic differences and are offered here to facilitate preliminary field diagnoses. Agera , A. houstonianum, Praxelis clematidea, and some individuals of Chromolaena odorata and o HE are GER hairy to the naked eye (primarily just on the uppermost stem in the latter two). The hairs in Fleischmannia incarnata and many individuals of Chromolaena odorata and Conoclinium coelestinum are small enouch and sparse enough to escape detection by the untrained naked eye. Chromolaena flowers are often a very pale bluish with the overall head having a whitish appearance due to the whitish phyllaries (except for the conspicuous green striations from the veins), E IONES are often whitish ii Only: a pee or bluish tinge, and the others (Ageratum, Conoclinium, a strongel bluish purple, g All of these taxa have a resinous odor (probably M o e to CC UI idary | Is i the ganas, which a are often visible with magnification, on the lower leaf mee but apy tly, P at-urine-like odor (Waterhouse et al. 2003). Abbott et al., Praxelis clematid for North America 625 (A t ( lly id li | , at least to the naked eye).—Petiole very short, less than 1/4 plage iic. to nearly as long as the blade; blade mostly broadly ovate, rounded to truncate or date at the base (som es a small cu narrowing wedge of tissue at the apex of the petiole, but the blade rarely with a cuneately Sem DUE Except hen the blade is more narrowly ovate); blade often nearly as wide as long; margin mostly very regularly, evenly, and shallowly crenate-serrate; heads more or less subglobose, often wider than tall (ca. 8-9 mm wide, 6-7 mm tall, in asia: clusters; puteos not quite 1 mm aod more or less io (apex iiL EH somewhat involute), fairly in g them in fruit | ly striate (alt E NU veins ER stand c DUM against t green | hyll n ) hromola ly short, generally only 1/4-1/3 sometimes iB nearly 1/2) the blade length; blade bd to Geer ovate, sometimes lanceolate, mostly cuneately narrowed at base (someti this is present as a large manos wedge BEW s rounded to truncate broader portion); blade irt variable, (fomes Ca. 13 to more than half as g; oarsely, irregularly toothed, but the teeth can be e and fairly regular, orthe blade may be ti | ly cylindrical, much longer than wide (ca. 3-5 SEN 8- 12 mm long), not in dense clusters; phyllaries about 1 mm wide, flat, densely imbricate, with conspicuous gre Conoeinium E EXE general ee the blade lengi de ovate n bluntly triangular, cordate to truncate at the base (sometimes a small cuneately narrowing v euge of of tissue at the apex of the petiole, but the SES rarely SE 4 ly L J a Lf J 4l l I£ long); A | | | f +! hon vt | SH | 1] | I M, SV UI | VCI E SO Sat late); heads more or less subdflobose re 4- 5 mm Wide and 3-4 mm tall), in | dense clust hyllaries very narrow (much less than 1 mm wide), almost acicular in appearance, very loosely imbricate, with conspicuous gaps between Den: even in BUE not striated. ta.—Petiole often as | the blade or longer (sometimes only 1/2 or 1/3 as long); blade mostly usi SENSO sometimes eunt or ovate, mostly truncate a mE Dose Sometimes rounded or WER SU EE nar- Ini. BI PAV Vy wI ) J; y 2 id ld ed near base, ti ith shall led tions, ti benti pecially on upper a.6-7 mm wide and tall), ly lax (f ot de ensely packe d); phyllaries ca. 0.8-0.9 mm wide, flat, loosely imbricate, with conspicuous gaps, at least in ër not EEN striated (although veins quite clear with 10x magnification). i | L f4 ACKNOWLEDGMENTS We would like to thank John Pruski for confirming the identity of our specimen. We would also like to thank Kent Perkins, Collections Manager of the University of Florida Herbarium, for his assistance. We are grateful for the helpful suggestions Walter Judd, Guy Nesom, and Barney Lipscomb provided in reviewing the manuscript. We also thank John Wooding for helpful discussions. This research was supported in part by a grant (no. DEB0224953) to M.B. Brown, University of Florida, from the Ecology of Infectious Disease Program. REFERENCES Cortett, R.T. and J.C. Suaw. 1995. Praxelis clematidea: yesterday South CERS togay be Kong EE ue world? Mem. Hong Kong Nat. Hist. Soc. 20:235-236. Available at http://wwy gy | corlett-pdf/RTC-praxelis-1995.pdf CRONQUIST, A. 1980. Asteraceae. In: A.E. Radford et al., eds. Vascular flora of the Southeastern United States, vol. 1. University of North Carolina Press, Chapel Hill. [FNAEC] Flora or NORTH AMERICA EpiroriaL COMMITTEE, EDS. 2006a. Flora of North America, North of Mexico, vol. 19: Magnoliophyta: Asteridae, part 6: Asteraceae, part 1. Oxford University Press, New York. [FNAEC] FLora or NORTH America EDITORIAL COMMITTEE, EDS. 2006b. Flora of North America, North of Mexico, vol. 21: Magnoliophyta: Asteridae, part 8: Asteraceae, part 3. Oxford University Press, New York. King, RM. and H. Rosinson. 1987. The genera of the Eupatorieae (Asteraceae). Monogr. Syst. Bot. Missouri Bot. Gard. 22. PouLock, S. A. HOLLAND, W. SMITH, and s Price, 2004. RH SEN weed for Queensland: Praxelis. Queensland Herbarium Alert Sheet 1/2004. Q | tal Protection Agency. Available at http://www.epa. a LAA Nm Nue qld.gov.au/publications/ 0012483. pdf/Praxelis new alien weed, for Queensland.pdf 626 | a | A - | | L I BA A Texas 2( SCHILLING, EE. J.L. Panero, and PB. Cox. 1999. Chloroplast DNA restriction site data support a narrowed interpreta- tion of Eupatorium (Asteraceae). do Syst. Evol. 219:209- mii ScHMIDT, G.J. and E.E. ScHILLING. 2000. Phylogen | phy of Eu ium (, on nuclear ITS sequence data. imei J Bot 87:716- -726. U.S. Forest Service. 2007, Pacific Island Ecosystems at Risk (PIER). Online resource at http://www.hear.org/pier/ Accessed 30 March 2007. VELDKAMP, J.F. 1999. Eupatorium catarium, a new name for Eupatorium clematideum Griseb., non Sch. Bip. (Com- positae), a South American species naturalized and spreading in S.E. Asia and Queensland, Australia. Gard. Bull. Singapore 51:119-124. WATERHOUSE, B. M. 2003. Know your enemy: recent records of potentially serious weeds in northern Australia, Papua New Guinea and Papua (Indonesia). Telopea E WATERHOUSE, B., R. MCFADYEN, A. HOLLAND, and J. THore. 2003. ' | g t guide: Praxelis - Praxelis clematidea. CRC Weed Management. Available at http:// t.gov.au/ biodiversity/i ive/weeds/weeds alert/pubs/p-clematidea.odf WUNDERLIN, R.P. and BE Hansen. 2003. Guide to the vascular plants of Florida, 2"4 ed. University Press of Florida, Gainesville, Florida. EXOTIC SPECIES OF CELTIS (CANNABACEAE) IN THE FLORA OF NORTH AMERICA Alan T. Whittemore US National Arboretum 35 ork Ave NE Washington, DC 20002-1958 U.S.A. ABSTRACT "TA p : Sr Te 1: Jd: KT aT America ac bo 4 T: I 1 ll; in ipa 11 1 1 C Valley of northern California, while C. si is ( to tl lventive fl f North A ica) pesi iety of hal f idely sep | ites in Californi 1 in the District of Columbia, with seedlings found up to 300 m f dult plants. D ipti 1 illustrati of both species are provided. RESUMEN TY E 14/7514: n " 15 a "AT + A o FF " da 1 1 1 a : 117511 dax I I L L del norte de California, mientras que C. si is ( la flora adventicia de Norte América) escapa en una ariedad de habitats 1 B ) en sitios ia il Separados en ime yen nel Distrito de Columbia, plantas q tran hasta a 300 m de las plant 1 u D r D The genus Celtis L. (Cannabaceae, traditional Ulmaceae; Sytsma et al. 2002) is a widespread group of ca. 60 or 70 species of trees and shrubs, about half growing in the north temperate zone. Several Celtis species are cultivated in North America. Most commonly cultivated are two native species, Celtis occidentalis L. (hack- berry) and C. laevigata Willd. (sugarberry). Two exotic species, C. australis L. (native to the Mediterranean basin and southwest Asia; Browicz and Zielinski 1982; Tutin 1991; Zielinski 1979) and C. sinensis Pers. (native to China and Japan; Fu et al. 2004), are available in the nursery trade. Both are planted, primarily as street trees, mainly in warm interior valleys of California (Brenzel 2001). Celtis sinensis is occasionally planted in the eastern United States (Meyer et al. 1994; Dirr 1998), and is sometimes considered a species with potential for more widespread use as a landscape tree (Dirr 1998). Native Celtis species are well known to be weedy in gardens and disturbed urban areas. The single report of C. australis escaping in Butte Co., California by Oswald (1994) is the only report of a non-native Celtis sp. reproducing outside of cultivation in North America. No exotic species of Celtis is treated in any continental or regional flora for North America (Barker 1997; Wilken 1993). However, Batianoff and Butler (2002) considered Celtis sinensis one of the worst invasive plants in south-east Queensland, Australia, where they found it capable of crowding out native vegetation and forming ltures, suggesting that the spe- cies has the potential to become invasive in suitable conditions. Herbarium specimens and field sites for Celtis were examined. Several Hen sites in California were confirmed for C. australis and C. sinensis. The latter was also found to escape in g tion in the District of Columbia. Nesom (2000) suggests a simple, uniform terminology for describing the "reproductive status and la success’ "BE non-native EE Uniform treatment of invasives is Se eg to allow T rv difficult to em to newly escaping woody plants or other pensis with long P times. Distin- guishing between Nesom’s categories of “waif” and “naturalized” depends on the scientists’ estimate of the long-term reproductive behavior of the species in the habitat. This is often suitable for annuals or perenni- als that are short-lived or reproduce when young, but not suitable for woody plants, where it may be many years before the long-term ability of a population to persist and reproduce in the habitat can be estimated accurately—thus, before it can be known whether the population is naturalized. Unfortunately, the early J. Bot. Res. Inst. Texas 2(1): 627 — 632. 2008 628 tani titute of Texas 2( xs 1 v5 : + A : £ +1 tablisl I | | ly t] the period when inf needed. it becomes important to document the habitats these species colonize and record their dispersal distances from seed sources. Invasive species have often gone unrecognized for long periods because they are not included in regional herbaria, floras, or other literature (Luken & Thieret 1995; Whittemore 2004). These two species are not described or illustrated in any North American flora (except McMinn & Maino 1947, who included C. australis as a cultivated species). A key, descriptions and illustrations are provided below to ensure that their populations can be recognized by local botanists. TAXONOMIC TREATMENT Identification of Celtis in North America is greatly complicated by unresolved taxonomic problems in a variable complex of shrubs and small trees, distributed over most of the United States, that are probably apomictic (Whittemore 2005a). These plants (referred to as the "small tree complex" below) are treated in different fl under a variety of names (tl t commonly used, in the order they were first published, are Celtis pumila Pursh, C. tenuifolia Nutt. € xélitulatà Torr., C. lindheimeri Engelm. ex K. Koch, and C. georgiana Small). None of the published treatments presents the variation in these plants adequately. Furthermore, of the five names listed above, only C. pumila and C. reticulata have been typified, and the correct application of the other names is in question. Until an adequate taxonomic treatment of this complex is available, it is not possible to write a full key to the North American Tes of EE Furthermore, Celtis shows a very high level of within-plant leaf polymorphism, which greatly ification. Leaves of juvenile or highly stressed plants are often very different from leaves of — adults with the same genotype, and leaves of vigorous leaders may be much larger than Sis of fruiting laterals, and these two = ee a ie Sie? Al anann an af the mall Trasnonmnle strongly in shape, ceperant Inc. Celtis spp. may req ial, and « ing leaves that 1950, Whittemore 2005b). The key below includes all Celtis species that grow outside Bi wo in EN United States and Canada, but does not attempt to divide the small tree complex into natural taxa. For the taxonomy of Celtis gt Mertensia M Planch., see ui and E (2001). UE" 1: à 1. fT 1.] 1. Stems with | t nodal spine itt y male fl ers lower on the same cyme e scars visible on pedicel) Styles bisbifid (i.e. Each lobeis is again deo divided) Celtis da Mertensia 2. Nodal spines straight. Leaves 1-35) x 0.4-2 cm. Axils of basal veins with deeply invaginated dom Shrubs 2-6 m tall. Florida, Texas, New Mexico, Arizona Celtis oe renbergiana (Klotzsch) TE =Celtis pallida Torr.) 2. Nodal spines curved. Leaves 5.5-10 x 2.5-4 cm (to 13 x 6 cm in the tropics). Axils of basal veins without a domatia. Shrubs (often scandent) 5-12 m tall. Florida eeu (qs (Jacq) Sarg. l. St LH hroditic fl litary on axillary pedicels (without ! | | bifid (i.e. with two undivided lobes) P Se subg. Celtis 3. Abaxial surface of leaf uniformly pubescent over its whole surface, felty to the touch; leaf abruptly con- an ed B a sender acumen, each margin THER 20- 22 teeth _ Celtis australis veins and vein ane meow to the touch, the leaf blad | | | to tl | | with no more e tapered or gradually n 16 teeth on each side. 4. Large shrubs or small trees to 12 m tall, with leaning trunks and horizontal or arching leaders or (with age) TE e e H EE or penury aborted, Secon more GEN 10% of the grains sone acetocarmine n Q f pres sent) very number of pores, OT | i Il and indehi | “Small tree complex" 4. EXeUrrent trees to 30r m n tal EE with g ionall de Or 3 parallel upright trunks), i l| tl Anti Iways dehi t, pollen grai , uniformly 3- poets Gm TG praia wine acetocarmine. 5 te-elliptic, ariitararchart_ari iminate, | | J y veins EIN (0. 5 04 a O. ie PL of plage bark without corky outgrowths Celtis sinensis g | obliquely triangular or triangular-ovate, acuminate (usually -slenderly so), basal lary vei t 0.3-0.55 length of blade; bark with prominent corky warts and ridges. Whittemore, E a? DH Ec atas al Al al. À H y. ES 629 6. Leaves generally ! bright green on both EH es SEHR pedicels evenly tapering or gradually ger but similar in shape; mature drupes 6-8 mm in diameter | brownish- -orange or red Celtis laevigata Willd. 6. Leav lly paler al lly; | btending pedicels abruptly narrowed to the acumen; leaves of vigorous leaders larger and different in shape (often lanceolate or oblong, apex acute or short-acuminate, base often shallowly cordate); mature drupes 8-11 mm in diameter, dark brownish purple or purplish red (but immature drupes may be red or orange) Celtis occidentalis |. Celtis australis L., Sp. Pl. 1043. 1753. (Fig. la). Deciduous trees to 20-25) m tall. Bark gray, smooth or finely roughened. Branchlets brown or purplish, pubescent with spreading hairs to 0.5 mm long. Winter buds brown or purplish, 3-5 mm long, puberulent. btending pedicels with petiole 9—17 Stipules linear or lanceolate, 3-6 mm, pubescent, fugacious. Leaves g mm, tan, pubescent, dorsal sulcus usually a narrow sharp groove, seldom broad and shallow; leaf blade narrowly elliptical to lanceolate or lance-ovate, 8-11 x 3—5 cm, rather thick but flexible, adaxial surface dull dark green, abaxial surface light green or glaucous, pubescent with spreading hairs over veins and whole blade, felty to the touch; secondary veins 2-4 on each side, basal secondary veins extending (0.45-)0.5-0.7 length of blade; base cuneate to rounded, usually strongly -o margin serrate from base to apex with sharp teeth 2-3 mm long, teeth 20-32 on each side; leaf ap ptly contracted to a slender acumen; leaves of sterile leaders similar but often larger, to 16 cm ong Male flowers fascicled in axils of bud-scales and lower leaves, bisexual flowers solitary in lower leaf axils, reaching anthesis as leaves emerge in the spring. Pollen not seen. Drupes 1 per leaf axil, pedicel pubescent (at least proximally), 19-31 mm long, 1.6-2.7 times as long as subtending petiole. Drupe subglobose to pyriform, usually tapering to style-base, 10-12 mm in diameter, very dark brown (almost black) at maturity. Distribution and Flowering —Found in North America in riparian woodland, elev. 15-60 m. Flowering as leaves emerge in the spring: fruiting mid- to late fall. Feral North American psum: examined. U. > A. —— Butte Co.: spontaneous in riparian Quercus lobata - Platanus forest, Lower Bidwell Park bet Creek Geet a N, 121° 49! W, A T. bere 06-004 (NA, UC). Tree ca. 8 m tall, edg: leveloped lland 10m $ of footbridge at W edg Park, Chico, V. Oswald 417 (CHSC). Yolo S oods on SH nks of SE Ree a tributary of Willow Slough but now an artificial daniel that drains north into the Willow EE Bypass) just east of F street near jct. Faro Ave., 38? 34' N, 121? 44' W, A.T. Whittemore 06-010 (NA, UC). Celtis australis can be recognized by its smooth or finely roughened bark (not developing corky warts or ridges like the native North American species), its leaf blades that are sharply serrate and very slenderly long-acuminate from a narrowly elliptical to lanceolate or lance-ovate body and uniformly felty-pubescent beneath, and its large, dark brown or black drupes (the drupes ripen orange-brown, red or purplish in other American species). Celtis australis was reported by Oswald (1994) as an escape in Bidwell Park, Chico, California. This site was visited in August 2006. The road between the site of the former Forestry Station and the current Nature Center is lined with large, well-established trees, evidently planted many years ago and now fruiting heav- ily. Seedlings and saplings are common in the vicinity of these trees. The Duden d are found in disturbed roadside habitats, but well-established small t Iso found in open Q Platanus woodland in the vicinity of the planted trees. Some of these trees are fruiting in this habitat, so C. australis is definitely established in native habitats and should be considered locally naturalized. Feral plants of Celtis australis were also found along a Rate on the ee Se of Davis, in ie A secondary forest that grew up on disturbed soil exp when the waterway thirty years ago. No cultivated trees were seen nearby, but the storm-drain stem from much ob Davis Ense ties into this waterway, so seeds could easily have washed in from elsewhere in Davis (where C. australis is occasionally planted as a street did The RE is not ee a SO e£ a strict eee e the 2000 +] a ity to definitions of Nesom (2000) their reproduce themselves locally), but the site was heavily disturbed dünne the handed. and it receives 630 | H tanical h Insti Texas 2( Fic. 1. Foli id f introd | Celt; ies. A. C australis. B. C. sinensis extra water from urban runoff, so it might be more realistic to regard them as weedy rather than invasive. The site is worth noting, however, as it does suggest that C. australis is capable of dispersing over significant distances. Celtis sinensis Pers., Syn. Pl. 1:292. 1805. (Fig. 1b). See trees to id m Ln BAS Bray, smooth or finely roughened. Branchlets brown, brown-pubescent, ason. Winter buds dark brown, 1-3 mm long, glabrous or puberu- lent. Stipules [ineat or E lanceolate, 3—5 mm, pubescent, fugacious. Leaves subtending pedicels with petiole 3-10 mm, brown, pubescent, dorsal sulcus broad and shallow; leaf blade ovate to ovate-elliptic, 3-10 x 3.5—6 cm, rather thick but flexible, adaxial surface glossy dark green, abaxial surface light green, usually inconspicuously yellowish-brown puberulent when young, at maturity with hairs abaxially scattered on major veins and sometimes tufted in vein axils, smooth to touch, secondary veins 3—4 on each side, basal secondary veins extending (0.5-)0.6-0.75 length of blade; base rounded, obtuse, or obliquely truncate, subsymmetrical to moderately oblique, margin subentire to crenate on apical half, teeth 0—16 on each side, apex acute or short-acuminate; leaves of sterile leaders similar but often larger, to 9 cm long. Male flowers fascicled in axils of bud-scales and lower leaves, bisexual flowers solitary in lower leaf axils, reaching an- thesis as leaves emerge in the spring. Anthers dehiscent, pollen grains uniform in size, uniformly 3-porate, 97-99% stainable with acetocarmine. Drupes 1(3) per leaf axil, pedicel rather stout, pubescent (at least proximally), 4-10 mm long, 1-1.5 times as long as subtending petiole. Drupe subglobose, 5-78) mm in diameter, brownish orange at maturity. Whittemore, E DI è cre! Sos Al Al al. À AA Plus 631 Distribution and Flowering —Found in North America in riparian woodland, dry grassland, open wood- land margins, elev. 15-580 m. Flowering as leaves emerge in the spring; fruits ripe mid- to late fall. Feral North Ameri j ined. U.S.A. California. Butte Co.: O dge of Dry C dland SS y camores, alders M a ca. 0.5 km N Ne jct. Pentz and Mesilla ae be elev. ca. 120 m, B. Castro 436 (CHSC). Orange Co. g t Blvd. and Campus Dr., elev. 15 m, A.C. Sanders 27326 (UCR). San B dino Co.: S in alder forest i an Hot = ing li liate v ty; Cold Canyon and ee e Canyon ca. 0.5 mi E of the old — Hot Springs Hotel, San B lino Mtns., elev 580 m, Á.C. Sanders 13812 (UCR). Yolo telis d 1 ] banks of waterway (formerly a tributary of Willow Slough but now an artificial channel s drains north into the Willow v Slough ID Just east tol F street near jct. Faro Ave., 38? 34' N, 121? 45' W, A.T. Whittemore 06-011 (NA, UC). Spont | 101A and the railroad tracks, 0.6 km south of Road 29, th side of Davis, 38? 35' N, 121? 45' W, A T. Whittemore 06-012 (NA, UC). District of Columbia: Disturbed woodland margins, service road to Administration a Bonsai Pavilion HE docks, e States a Arboretum, 38? 55' N, 76? 58' N, A.T. Whittemore 04- 012 (NA). D g gI United States National Arboretum, 38? 55! N, 76? 58' N, A.T. Whittemore 04- 016 (NA). Celtis sinensis can be recognized by its smooth bark, its rather leathery, glossy leaves, with the margins usu- ally crenate or bluntly serrate and the apex often rounded-acute. Celtis sinensis has never been reported to escape in North America, but field sites and herbarium specimens were found documenting feral populations over a surprisingly wide geographic range and in a variety of habitats. It is not common at any of the sites and there is currently no evidence that it can be an aggressive competitor in our flora (as reported for Australia; Batianoff and Butler 2002), but saplings are often found several hundred meters from possible seed sources, suggesting that effective dispersal agents for this species exist in North America. On the grounds of the U. S. National Arboretum in Washington DC, young trees are occasionally found in open margins of native deciduous forest, often 100—300 m from the nearest possible seed source, but feral plants have not been observed in closed forest (except as a weed in cultivated beds). The Butte County site (Castro 436) is riparian woodland in dry, grassy low foothills of the northern Sierra Nevada, and according to the herbarium label, the feral plants are ca. 200 m from planted trees of C. sinensis. The tree near Road 101A in Yolo County (Whittemore 06-012) is in native alkali grassland that has never been cultivated but has probably been subject to some disturbance because of its proxim- ity to the county road and railroad. This tree is at least 300 m from the nearest possible seed source. The long dispersal distances seen at most of the sites are not surprising, since the sweet, fleshy drupes are well suited to bird dispersal. Fleshy bird-dispersed fruit has been shown to be a significant predictor of invasive behavior in woody plants (Widrlechner et al. 2004), so further tracking of this species is desirable. ACKNOWLEDGMENTS I would like to thank Lawrence Janeway and Warren Roberts for directions to field sites and help in the field, Andrew Sanders for providing comments on habitats of his Celtis collections, and Guy Nesom and Barbara Ertter for helpful comments on the manuscript. REFERENCES Barker, W.T. 1997. Celtis. In: Flora of North America North of Mexico volume 3. Flora of North America Editorial Committee, eds. Oxford University Press, New York. Pp. 376-379 Batianorr, G.N. and DW. Buter. 2002. Assessment of invasive naturalized plants in south-east Queensland. PI. Protect. Quart. 17:27-35. Bere, C.C. and S.V. DaHLBERG, 2001. A revision of Celtis subg. Mertensia (Ulmaceae). Brittonia 53:66-81. BRENZEL, K.N. (ed) 2001. Sunset western garden book. Ed. 7. Sunset Publishing Corp. Menlo Park. Browicz, K. and J. Zim ski. 1982. Celtis. In: Flora of Turkey, vol. 7. PH. Davis, ed. Edinburgh University Press, Edin- burgh. Pp. 649—652. Den M.A. 1998. Manual of woody landscape plants. ed. 5. Stipes Publishing, Champaign, IL. FERNALD, M.L. 1950. Manual of botany. Ed. 8. Van Nostrand, New York. | fal Das H In LE PA £T >» * F 632 Fu L-G, Xin Y-.Q,, and AT. Wurrrevone. 2004. Ulmaceae. In: Flora of China, volume 5. Z.-Y. Wu and PH. Raven, eds. Missouri Botanical Garden, St. Louis. Pp. 1—19. Luken, J.O. and JW. Thierer. 1995. Amur honeysuckle (Lonicera maackii, Caprifoliaceae): its ascent, decline, and fall. Sida 16:479-503. McMinn, H.E. and E. Maino, 1947. An illustrated manual of Pacific Coast trees. Ed. 2. University of California Press, Berkeley. Mever, FG, PM. Mazzeo, and D.H. Voss. 1994. A catalog of cultivated woody plants of the southeast United States U.S. Natl. Arbor. Contr. no. 7: i-vii, 1-330. U.S. Dept. of Agric., Washington, DC. Nesom, G.L. 2000. Which non-native plants are included in floristic accounts? Sida 19:189-193. OswaLD, V.H. 1994. Manual of the vascular plants of Butte County, California. California Native Plant Society, Sacramento. Sytsma, K.J., J. MoRAwtrz, J.C. Pires, M. NEPOKROEFF, E. Conti, M. ZuHRA, J.C. Hat, and MW. CHASE. 2002. Urticalean rosids: Circumscription, rosid ancestry, and phylogenetics based on rbcL, trnL-F, and ndhF sequences. Amer. J. Bot. 89:1531-1546 Tunn, T.G. 1991. Celtis. In: Flora Europaea vol. 1, ed. 2. T.G. Tutin, N.A. Burges, A.O. Chater, J. R. Edmonson, V.H. Heywood, D.M. Moore, D.H. Valentine, S.M. Walters, and D.A. Webb, eds. Cambridge University Press, Cam- bridge. P. 77. Whittemore, AT. 2004. Sawtooth oak (Quercus acutissima, Fagaceae) in North America. Sida 21:447—454. Wuitremore, A.T. 2005a. Sexual sterility and E in eastern Po ll SOUS Abstract 74, Botany 2005, Austin, Texas [http //www.2005.1 detail&aid=74] WHrTTEMORE, A.T. 2005b. Introgression, Genetic structure, and (S ONG status in the Celtis laevigata - C. reticulata complex (Cannabaceae). Syst. Bot. 30: 809-817. WibRLECHNER, M.P, J.R. THomeson, J.K. ILes, and PM. Dixon. 2004. Models for predicting the risk of naturalization of nonnative woody plants in lowa. J. Environm. Hort. 22:23-31. Wiken, D.H. 1993. Ulmaceae. In: The Jepson manual: higher plants of California. J.C. Hickman, ed. University of California Press, Berkeley. Pp. 1081-1082. Ze, J. 1979. Ulmaceae. [in Latin] Flora Iranica fasc. 142. KH. Rechinger, ed. Graz, Austria. LUZIOLA SUBINTEGRA (POACEAE: ORYZEAE), NEW TO FLORIDA AND THE UNITED STATES John M. Kunzer Michael J. Bodle Department of Biology South Florida Water Management District University of South Florida 3301 Gun Club Roa 4202 E. jo ie. SCA 110 West Palm Beach, Florida 33406-3007, U.S.A. Tampa, Florida 0-5200, U.S.A. mbodlegsfwmd.gov al ustedu ABSTRACT The MA Ls taxon ed subintegra e has iin d in See id e = o E ERE Florida, U.S i key to the genus Fus are given. RESUMEN El ] inoide Luziola subint Swall ] trad la bahía Fisheati lel Lago Okeechobee, en e] condado Glades, Florida, EE UU.E ta el pri ist ido de este taxón en la Florid yos Estados Unidos. Se presentan una 4 1 ] 1 ganern Liueinln Md Mr tl ee do ipci o EA i F o In early December 2007, the USF herbarium received a specimen of an unknown aquatic grass collected along ER northwestern shore of Lake Okeechobee, Glades County, Florida. The specimen was not satisfac- torily identifiable as anything currently known in the state's flora (Wunderlin & Hansen 2003; Wunderlin & Hansen 2008) and its morphological characteristics led to investigation of the genus Luziola. Examination of the revision of Luziola (Swallen 1965) revealed that there are two named species of very similar morphology, L. spruceana Benth. ex Dóll and L. subintegra Swallen. The material in question was determined to be closer to L. subintegra, and this was confirmed by Gerrit Davidse (MO; determination in duplicate). Consultation of the Flora of North America treatment of Luziola (Terrell 2007) and the USDA PLANTS database un NRCS 2008) revealed that a taxon is ipe new to the United States flora. +! bint known distribution from Mexi B through Central America and South nea a to Argentina, and the Caribbean basin (Judziewicz et al. 2000). It is apparently rare in Central America, at least in Costa Rica and Nicaragua (Morales 2003; Pohl & Davidse 2001). Description.—Stoloniferous perennial, emergent, floating, mat-forming aquatic or prostrate-creeping terrestrial. Culms nde to ca. d m long or more, 3-10 mm diameter, rooting from the nodes. Leaf sheaths inflated, spongy; ligule an , acuminate, 1— 4 cm long, eee 1-7.5 dm long, to 7-20 H T r | H E n a 11 3 f. wide I rarely with a few pistillate spikelets within the staminate panicle. Staminate panicle terminal! M spikelets 5—7 mm long, with one fertile floret per spikelet; glumes absent or obsolete; palea 3-nerved; lemma 4 or 5-nerved, both palea and lemma hyaline, similar; anthers 6(5—7) per floret. Pistillate panicle axillary, congested, 1—several per culm, barely exserted from the sheath, the primary branches sharply recurved when mature, with evi- dent pulvini, spikelets 3—5.5 mm long, with one fertile floret per spikelet; glumes absent or obsolete; palea 5-nerved; lemma 5-7-nerved, palea and lemma similar. Caryopses (achenes) with a hard, brittle pericarp, minutely longitudinally striate, asymmetrically ovoid, to 2 mm long, with a persistent style base. Luziola subintegra is unlike any other Luziola currently known to the United States. It is robust, with large, thick culms, and has leaves with inflated, spongy sheaths and long, broad blades. All of our other Luziola taxa can be described as delicate, slender plants with narrow culms, and shorter, narrower leaves. Luziola subintegra gives the overall impression of being quite similar in habit to Paspalum repens P.J. Berguis or to a lesser degree Hymenachne amplexicaulis (Rudge) Nees (Figs. 1-3). J, Bot. Res, Inst. Texas 2(1): 633 — 636. 2008 634 t tani i Texas 2( Fic. 1. Luziola subintegra staminate panicle. Fic. 2. Luziola subintegra pistillate panicles (upper—flowering, lower- mature). The following key modified from Wunderlin and Hansen (2003) serves to differentiate L. subintegra from the rest of the genus in Florida and the United States. MODIFIED KEY TO LUZIOLA . Culms robust, 3 mm in Ge or SES, eat pass 7 mm Weg or MD SE bad Gi ide L. subintegra : SE slender, 2 mm or les m long 2. l : pistillate spikelet tly included in tl btending sheatl L. fluitans 2. Infl le; pistillate spikelets all or mostly exserted from ti btending sheat! 3. Pistillate spikelet 2- E mm long; fru it smoo ei L. peruviana 3. Pistillate spikelet 3-5 | o frui t striate L. bahiensis In late December 2007, we investigated the extent of the population within Fisheating Bay, and found it to be nearly ubiquitous there. The population consisted of two large, near-monoculture mats, one mat of ca. 2 ha, to the east of Harney Pond Canal, and the other of ca. 80 ha, to the west of the canal (Fig. 3). The locality of these two mats was within a portion of the lake bed that during the record low-water level of early June 2007 was completely exposed and bare of all vegetation. Surrounding each of the large mats and scattered throughout the open bay were sporadic, small clumps of one to several mature individuals. It was also found as both an emergent aquatic and a terrestrial inside the mouth of Fisheating Creek. At this location, the terrestrial individuals of a mature state were evident only in small, isolated clumps, and they appeared to be less competitive against the other terrestrial flora. Fisheating Creek is the only remaining unregulated inflow to Lake Okeechobee. Based upon this infor- mation and the fact that L. subintegra is present in the creek, we speculate that Fisheating Creek is its point of introduction into Fisheating Bay and Lake Okeechobee. Given the extent of the current population and K | Bodle, Luziola subintegra, new to Florida 635 V = A aii TER , A OSI 8 d at, A i ERAN om "a TAS , ‘ral Tult ds fa MS H + d DA UM AA AN ! d P : 4 i e. ZER à ar Si = j ns pr erp Te DR ad y e El st E wt Pas mr E A Rr AER rm Pir Pr Leer A FT |! PARRA pR =. (oe zt d Dës Ee ice e Fic 3 i Salt | tas la nar £g y nm if Li "1 E bat , facing SW the plant's apparent fecundity, we suggest that this taxon be considered for listing by the Florida Exotic Pest Plant Council (FLEPPC). We also suggest that ecological research be immediately undertaken to determine its potential impact on Florida's natural systems. Voucher specimens: U.S.A. Fiorina. Glades Co.: Lake Okeechobee, Fisheating Bay, ca. 3.75 km S of the junction of C-721 and FL 78. 26? 58' 0.3" N, 81? 04' 9.5" W, 6 Dec 2007, Mike J. Bodle s.n. (MO, USF 244612, others to be distributed); Lake Okeechobee, Fisheating Bay, ca. 3.25 km SE of the junction of C-721 and FL 78. 26? 58' 13.8" N, 81° 03' 37.4" W, 20 Dec 2007, John M. Kunzer, Mike J. Bodle, Kurt Ramsey 2253 (USF 244608, others to be distributed). ACKNOWLEDGMENTS The authors would like to thank Bruce Hansen (USF) and Richard Wunderlin (USF) for their generous advice in research and editing, and Gerrit Davidse (MO) for confirming our determination of the initial specimen. We would also like to thank Rufino Osorio for the original Spanish language version of the abstract, and Annie C. Schmidt for putting the authors in touch with each other. REFERENCES Jupziewicz, E.J., RJ. Soreng, G. Davipse, PM. Peterson, T.S. FiLGUEIRAS, and FO. ZuLOAGA. 2000. Catalogue of New World grasses (Poaceae): 1. Subfamilies Anomochlooideae, Bambusoideae, Ehrhartoideae, and Pharoideae. Contr. U.S. Natl. Herb. 39:1-128, MORALES, J.F. 2003. Poaceae. In: B.E. Hammel, M.H. Grayum, C. Herrera, and N. Zamora, eds. Manual de plantas de Costa Rica Vol. 3. Monogr. Syst. Bot. Missouri Bot. Gard. 93:598-821. 636 Pou, RW. and G. Davwpst. 2001. Luziola. In: W.D. STEVENS C.U. Ulloa, A. Pool, and O.M. Montiel, eds. Flora de Nicara- gua: Angiospermas (Pandanaceae-Zygo| ). Monogr. Syst. Bot. Missouri Bot. Gard. 85:2072-2073. SWwALLEN, J.R. 1965. The grass genus Luziola. Ann. Missouri Bot. Gard. 52:472-475. Terre, EE 2007. Luziola. In: Flora of North America Editorial Committee, eds. 2007. Flora of North America north of Mexico. Vol 24. Magnoliophyta: Commelinidae (in part): Poaceae, part 1. Oxford Univ. Press, New York, New York. USDA, NRCS. 2008. The PLANTS Database (http://plants.usda.gov, 15 Feb 2008). National Plant Data Center, Baton Rouge, LA 70874-4490 USA WUNDERLIN, R.P, and B.F. Hansen. 2003. Guide to the vascular plants of Florida: S | edition. Gainesville: University Press of Florida. WUNDERLIN, R.P. and B.F. Hansen. 2008. Atlas of Florida ts (http:// lantat! fedu/).[S. M. LANDRY and K. N. CameseLL (application development), Haidi Centerfor SOEN Design and een Institute for Systematic Botany, University of South Florida, Tampa. NEW AND NOTEWORTHY RECORDS OF SEVERAL NON-NATIVE VASCULAR PLANT SPECIES IN ARKANSAS Brett E. Serviss James H. Peck Department of Biology Department of Biology Henderson State University peau of Arkansas at Little Rock Arkadelphia, Arkansas 71999-0001, U.S.A. Little Rock, Arkansas 72204, U.S.A. servisb@hsu.edu jhpeckeualr.edu ABSTRACT Collections of nine es a Mas pot all non-native to the United States, are hi Jin as new or noteworthy for Arkansas. Three species, Ma , and Thunbergia alata, I to Arkansas. gece ia pos Ch des} yl thalmi ted for their second occurrence in Arkansas. Án additional i 4 EST WEE a E : E de od EI. dA z DK? and A d H 1 1} de A I E Ech ph; 1 The current ROWE distribution fr ps H : A wl E e | 1 Finally, e Fu E] Te J ] PEL Pares | planis are M GM These d p pem aes male diia e (common —À ‘Camellia sasanqua (fall- wthorn), and Ternstroemia gymnanthera (cleyera); 11 f 1 1 ] " ]: Al - alii Ases all OVIL vvlIIDCLIL WCLO previously C 1t = mL] 1 1 H A : J ] 1 , todas ellas no-nativas - es Estados Unidos, AA 1 7 rg T3 rd 1 1 A para Arkansas. Tres especies, ilea q ifolia, iset i ta, se citan como nuevas para Arkansas maranthus Li. el Tolg. Lë ] Alae c ] DÉI SES, 1 dad J ESLLLRA TIL + L L [o] 1 I £4. ] 4 E H Ti á a EZ JI IM Aa A 1-4. € " c T dict DA 4 1 EH d _ acutissima en GE EES Finalmente, cit id t4 d lant ltivad Estas incluyen: Acer + 151 1. £1 H + x a : f. IE fam wt Tis f 14 pa] tafani T Suc Ed E dE ? Gë É Raphiolegis india a indio), yT t ia g) thera (cleyera); q taban d tad Arl t dició INTRODUCTION Non-nati j ti lly introduced into the United States tl gh bot! idental and intentional human acuvities. d introducti times lead to the f tion of | lations of these plants. Spontaneous is here defined as the autonomous occurrence drou sexual or qM MM tion of a non-native plant species in a region or flora to EHS? it is not native. e is SE to the term “escaped” as defined by Nesom (2000). Most non-nati duction do not success- fully establish, or if establishment occurs, ER not pores a serious nea to native pen pestes or iri Lini 4 (Williamson 1996). However, some of so to reduce native biodiversity. It is therefore garten to record first encounters with escaped papúla- tions and monitor these new spontaneous occurrences of non-native species to evaluate their potential for becoming the next wave of invasive species (Yatskievych & Raveill 2001). At present, non-native species comprise approximately 21% of the Arkansas flora (Arkansas Vascular Flora Committee 2006), with 3% annotated as invasive in Arkansas. Our field work focusing on ruderals and recent escapes in urban has led to tl of many non-native plant species being documented as new to Árkansas (Peck 2003; Peck & Service 2006; Serviss et al. 2006, 2007a, 2007b), with many of the species locally well established when discovered. At present, more Species are being documented as entering the Arkansas s than are pene lost from habitat loss, fragment , or human disturbance. Additionally, we have ol flocalized ` spontaneity in several species of non-native ornamental shrubs, and while the parent plants were cultivated, substantial reproduction of offspring was documented in the local vicinity of these plants (see end of paper for specimen data). J. Bot. Res. Inst. Texas 2(1): 637 — 641. 2008 638 t tani i Texas 2( The national occurrence and distribution of the below listed taxa was determined from the national flora database kept by NRCS (1999). SPECIES NEW OR NOTEWORTHY POR ARKANSAS Amaranthus blitum L. (Amaranthaceae). Purple th is a ruderal species that is native to the Mediter- D ranean region. It ross the coastal states from M husett th to Florida, and west to Louisiana and Texas. This is only the second record for A. blitum in Arkansas; it was previously documented in Pulaski NM (Peck & Serviss 2006). Clark C ivel plant ls in fl bed, Lake DeGray Visitor Center, Lake DeGray State Park, 28 Oct 2007, Serviss 7344 (HEND), url deral habitat, Arkadelphia, Hend State University campus, 25 Aug 2004, Serviss 6124 (HEND). Chamaesyce ophthalmica (Pers.) D.G. Burch (Euphorbiaceae). Florida hammock sandmat is an annual ruderal species that is native to the Gulf coast from Florida and Georgia west to Louisiana. This is only the second record for C. ophthalmica in Arkansas; it was previously documented in Pulaski County (Peck & Serviss 2006). Clark C ivel lant 1s in f] bed, Lake DeGray Visitor Center, Lake DeGray State Park, 28 Oct 2007, Serviss 7345 (HEND). Eichhornia crassipes (Mart.) Solms (Pontederiaceae). Water hyacinth, which is native to Brazil, is an aggressive and invasive non-indigenous aquatic species in North America. It was reported as present and then eradicated twice in Arkansas, once as an escape in EE Arkansas and once in southwestern Arkansas (Tumlison & Serviss 2006). It is ] ted as a persi d spreading population at its most northerly location yet in Arkansas, at the deeg of the Little Mianraelle River and the Arkansas River in Pulaski County, Arkansas. Here it co-occurred with Egeria densa, Hydrilla verticillata, Marsilea quadrifolia, and Myriophyllum spicatum. It was first discovered at this location in 2006 by Bob Spraggins, an accoun- tant for the Arkansas Natural Heritage Commission, who brought it to the attention of Theo Witsell, staff botanist of the same agency. In 2007 the locality was searched, and all five invasive species were noted as unchecked and a Da l Cr” rl ande River, 16 Jul 2007, Peck 07- 1352 (LRU). J]: 1 A 1 Ii J 1 1 Le: ^ [41 1 Y fe t km of swamp slough à a [e] H Hydrilla verticillata (L.f.) Royle (Hydrocharitaceae). Water-thyme is an aggressive and invasive non- indigenous aquatic species that is native to Asia. It was reported as present in several impoundments of tributary rivers to the Arkansas River (Peck 2003; Peck & Serviss 2006), and here for the first time in Pulaski County, at its most northerly location yet in Arkansas, at the junction of the Little Maumelle River and the Arkansas River. Here it co-occurred with Egeria densa, Eichhornia crassipes, Marsilea quadrifolia, and Myriophyllum spicatum. It was discovered at this location in 2007 with four other invasive species and was noted as unchecked and spreading. As it is known to survive multiple winters as far north as lowa (Peck & Smart 1986), it might al also be found in northern Arkansas impoundments. 1 1 4 d 1 r 1 ei f swamp, slough, d s] habitats of the ] Little Memes River, 16 Jul 2007, Pech 07-1358 (LRU). Marsilea quadrifolia L. (Marsileaceae). European Waterclover is native to Europe and Asia, and is an ag- gressive and invasive non-indigenous aquatic species. It was introduced into eastern North America as an aquarium plant, and has since escaped and spread westward as far as southwestern Iowa. lt is reported for the first time from Arkansas, where it was found along a muddy shore of the Little Maumelle River, Pulaski County, Arkansas. The patch was relatively small, suggestive farecent introduction, most likely by transient waterfowl. It was associated with extensive growths of other non-native and invasive species: Fichhornia crassipes, Egeria densa, Hydrilla verticillata, and Myriophyllum spicatum. Serviss and Peck, Records of non-native vascular plants in Arkansas 639 a oo Pulaski Co.: a single patch of floating-leaf shoreline plant spreading along p shore habitat of the 47.1] D " 1 ei £ +1 +) [+] Tittle M 1] +1 Arlrancac D Y mue Li Lr Jul 2007, Pech 07-1514 (LRU). repart Spica L. EE Eurasian water milfoil is an aggressive and invasive non- d Asia. It is reported for the first time in Pulaski County, at Gs cion of the Little Maumelle River set the Arkansas River. Here it co-occurred with Eichhornia crassipes, Egeria densa, Hydrilla verticillata, and Marsilea quadrifolia. TQ 1.5 es 1 J 1 J: 1 1 Fiel T 144] RA 17 D: ] Eu Eu CA e HEC "uum PTS CM TS River, 9 Jul 2007, Peck 07-1635 (LRU). i en Twin R a y Pennisetum setaceum (Forsk.) Chiov. (Poaceae). Fountain grass is a caespitose, perennial grass that is na- tive to Africa. This is the first documentation of this species in Arkansas outside of cultivation. It has been previously documented outside of cultivation in Arizona, California, Colorado, Florida, Hawaii, Louisiana, New Mexico, Oregon, and Tennessee. Voucher opem ben Co.: a Jw ee Gees Towne as epus in a B Wess on the Henderson State University the origin , and i couple of years, Arkadelphia, 24 Sep 2004, mpus I Serviss 6636A (HEND). Quercus acutissima Carruthers (Fagaceae). Sawtooth oak is native to Korea, China, Japan, and Vietnam. It has been documented outside of cultivation in several states, including Alabama, ud BAUER, Maryland, North Carolina, Mississippi, Pennsylvania, and Virginia. Though previously documented as pres ent in the Arkansas flora (Arkansas Vascular Ve Committee 2006); specifically, it has been enel outside of cultivation in 10 Arl ties (Cross, Drew, Hempstead, Jackson, Jefferson, Lincoln, Nevada, Pike, Union, and most recently, Clark). Sawtooth oak has demonstrated the ability to escape cultivation and subsequently naturalize in many areas of the eastern U.S., especially when cultivated plants are in the vicinity of open areas, such as grasslands and disturbed woods (Whittemore 2004). Hence, it appears to have the potential to become invasive in Arkansas, and its current known distribution in the state seems worthy of note. ile plant i d and under cultivated shrul ltivated Q. acutis- i Clark C , Spontaneous, j E sd L [e] [eJ Se : n jean ibly the parent, Arkadelphia, Ouachita r 4 Baptist University campus, 31 Oct 2007, Serviss 7342 2 (HEND). Thunbergia alata Bojer ex Sims (Acanthaceae). Blackeyed susan vine is native to Tropical Africa, but has become a pantropical weed. Thunbergia alata is currently only known outside of cultivation in the U.S. from Florida, Hawaii, and Texas. One spontaneous plant has been documented in Arkansas. This is the first documentation of this species in Arkansas outside of cultivation. gw Clark € ZE ET Toce di hatuni | of a home site, Arkadelphia, 420 made | , Ter I Cedar Grove Mg 5 i 2004, Serviss 6628 (HEND): one of th ial nf T Latent 1 4 des kad +l H 1 1 ADDITIONAL NOTEWORTHY SPECIES The following six non-native, woody, angiosperms were documented as spontaneous offspring in the im- mediate vicinity, or within a short (several meters) distance from the vicinity of cultivated and reproductive individuals of these species. Documentation of these initial reproduct by cultivated non-native plants is essential for dating their initial entrance into the flora by other than the dent assistance of human activities. Additionally, the ability of these species to successfully reproduce offspring is significant in that probably with regard to most of the non-native and invasive ornamentals in the U.S., their initial “entrance” into the flora was accomplished in similar fashion to the following examples. It is therefore crucial to rec- ognize and sul tly document these initial occurrences of spontaneity in non-native plant species. All six of the lonas species have displayed successful reproduction in Arkansas, evidenced by the presence 640 J ] tanical Insti Texas 2( of few to many spontaneous juveniles being present within a short distance of cultivated reproductive age plants of their respective species. Specimen data: Acer palmatum Thunb. (Sapindaceae). Japanese maple is native to Japan, and has been previously recorded outside of cultivation in Ohio, Pennsylvania, and New York. Voucher specimen: Clark Co.: dozens of seedlings scattered across a few disturbed flower beds in proximity to cultivated plants of A. palmatum, Arkadelphia, Ouachita Baptist University campus, 27 Nov 2007, Serviss 7352 (HEND). Camellia japonica L. (Theaceae). Japanese camellia is native to Japan, but has been documented outside of cultivation in Florida, Georgia, and the Carolinas. Voucher specimen: Clark Co.: a few spontaneous plants present be- neath a large, cultivated shrub of C. japonica, Arkadelphia, Henderson State University campus, 1 Nov 2007, Serviss 7344 (HEND). Camellia sasanqua Thunb. (Theaceae). Fall-flowering llia is native to Japan. It has been documented outside of cultivation in Florida, Georgia, and the Carolinas. It has been documented in several locations of Clark County by the presence of numerous (more than 100) spontaneous seedlings and juveniles in the vicinity of cultivated C. sasanqua plants. Voucher specimens: Clark Co.: several small seedlings and juvenile plants in the vicinity of two reproductive age plants of the species, Arkadelphia, edge of disturbed wooded area off of 12‘ Street, one block west of the Henderson State University campus, 18 Mar 2005, Serviss 6861 (HEND); Arkadelphia, Henderson State University campus, 29 May 2005, Serviss 6985 (HEND). Loropetalum chinense (R. Br.) Oliv. (Hamamelidaceae). Fringe flower is native to China. Voucher specimen (two locations located for Clark Co.; one cited below): Clark Co.: several spontaneous seedling and juvenile plants growing in a disturbed area in the vicinity of several cultivated, reproductively mature plants of L. chinense, Arkadelphia, Henderson State University campus, Oct 2006, Serviss 7094 (HEND). Raphiolepis indica (L.) Lindl. (Rosaceae). Indian hawthorn is native to China. Arkansas material luctive age plants of this species is represented by numerous spontaneous seedlings in the vicinity of rep of the bee present at a EE of locations in Clark RA Keiers pun SSC Eos numerous ightly larger juveniles in the vicinity of several cultivat tively mature plants of R. indicd, Arkadelphia, Henderson State Senay campus, 28 Feb 2007, B nds 7115 (HEND). (Wight & Arn.) Sprague (Theaceae). Cleyera is native to India, the Malaysian Peninsula, and jm This species is ena from Gees Si only a few spontaneous juvenile plants (one specimen cited below). Spont ] | meters from the vicinity of cultivated GE age plants of T. —— Voucher specimen: Clark Co.: one juvenile plant growing with a variety of other weeds under cultivated shrubbery; cultivated, reproductive age plants of T. gymnanthera list f the location of t ile, Arkadelphia, residence adjacent were present a short to Henderson Street and 13' Street, 28 Feb 2007, AM /113 (HEND); circumistamital evidence es dispersal by birds (the seeds of T. gymnanthera are bird-dispersed). ACKNOWLEDGMENTS We would like to thank Allen Leible, Will Seymour, and Jane Dunn (Henderson State University) for assis- tance in acquisition of some of the above cited specimens. Additionally, we thank Brent Baker (University of Arkansas) for providing distribution data for some of the above listed taxa. We also thank R. Dale Thomas (University of Louisiana at Monroe) and George P. Johnson (Arkansas Tech University) for their comments and suggestions regarding this paper. REFERENCES ARKANSAS VASCULAR FLORA Committee. 2006. Checklist of the vascular plants of Arkansas. University Herbarium, Depart- ment of Biology, University of Arkansas, Fayetteville. Nesom, G. 2000. Which non-native plants are included in floristic accounts. Sida 19:189-193. Peck, J.H. 2003. Additions, re-instatements, exclusions, and re-exclusions to the Arkansas flora. Sida 212737214757. Peck, J.H. and B.E. Serviss. 2006. New and noteworthy collections for Arkansas. Sida 22:817—820. Serviss and Peck, Records of non-native vascular plants in Arkansas 641 Peck, J.H. and M.M. Smarr. 1986. Ecological perspectives of the Upper Mississippi River: an assessment of aquatic and wetland vegetation of the Upper Mississippi River. Hydrobiologia 136:57-76. Serviss, BE N. FREEMAN, J. HERNANDEZ, A. Leige, and C. TALLey. 2007a. Tungoil tree (Aleurites fordii Hemsl.) (Euphorbi- aceae) new to the Arkansas flora. J. Arkansas Acad. Sci. 61:128-130. Serviss, BE N. Freeman, J. HERNANDEZ, A. LEiBLE, C. TALLEY, and B. Baker. 2007b. Negundo chaste tree (Vitex negundo L. (Verbenaceae) new to the Arkansas flora. J. Arkansas Acad. Sci. 61:131-133. Serviss, B.E., N. Freeman, and S. MeLancen. 2006. Chinese flame tree (Koelreuteria bipinnata Franch.) (Sapindaceae) new to the Arkansas flora. J. e Ge Sci. 60:197-199. TUMLISON, R. and B.E. Serviss. 2006. Naturali and extirpation of water hyacinth (Eichhornia crassipes, Ponted- eriaceae) in southwestern Arkansas, USA. Sida 22:749-754. USDA-NRCS. 1999. The PLANTS database. National Plant Data Center, Baton Rouge, LA 70874-4490 USA. Avail- able at http://plants.usda.gov. Accessed on 27 November 2007. WHITTEMORE, A.T. 2004. Sawtooth Oak (Quercus acutissima, Fagaceae) in North America. Sida 21:447-454, WILLIAMSON, M.H. 1996. Biological invasions. London: Chapman and Hall. YATSKIEVYCH, G. and J.A. Raven. 2001. Notes on the increasing proportion of non-native angiosperms in the Mis- souri flora, with reports of three new genera for the state. Sida 19:701—709. A" 642 BOOK NOTICE Doucras J. Furuyma, H. BRADLEY SHAFFER, AND DANIEL SIMBERLOFF (eds). 2005. Annual Review of Ecology, Evolution, and Systematics: Volume 36, 2005. (ISBN 978-0-8243-1436-0, hbk; ISSN 1543-592X). Annual Reviews Inc., 4139 El Camino Way, PO. Box 10139, Palo Alto, California 94303-0139, U.S.A. (Orders: www.annualreviews.org, science@annualreviews.org, 1-800-523-8635, 1-650-493-4400, 1-650-424-0910 fax). $79.00 indiv., 689 pp., indexes, 7 5/8" x 9 3/8". Contents of Volume 36: 1. Tl ti d evolut F f] gas; try—Larry J. Leamy, Christian Peter Klingenberg 2. Estan y evoluti ptil Richard Shine 3 TL f J + 1 + ence, +} m ] I lapa A Į le denne Carol Goodwillie, Susan Kalisz, i: dpi Bcher 4. Indirect interaction webs: h trait cl lants—Takayuki Ohgusl 5. SE See of sm Peter Linder, Paula J. Ruda Il 6. I f perm selection, and offspring viability—Leigh W. Simmons 7. Individual-based modeling of ecological and evolut y processes—Donald L nee vee Mooij 8. The influence tabolit I ti logy of! l vertebrates—M. Denise Dearing, William J. Foley, Stuart inii 9. MM and li position in t trial ecosystems Stephan a ie V. Tiunov, Stefan Scheu 10. T ifi fril 1 (DNA ion: i ecology g Lawrence J. Weider, James J. Elser Teresa J. Cera le Mateos mus B. Cotner, Therese A. Markow Kristy U. Brady, Arthur R. e H.D. Bradshaw Jr. llend 11. E Ly olutionary ELU WED H E E Ary a 12. Biodiversity € L f i I i it ] tt Diane S Sri astava, Mark Ve 13 co L +1 SE {Dal S t1 r 1 t Stev en D'Hondt H L e J Lev Li Ibid 14. Land l hat is the state of the science?—Monica G. Turne I Oy 15. Ecology and evolution of aphid-ant interactions—Bernhard Stadler, Anthony F.G. Dixon 16. Evolutionary | fi thology—Ándrea L. Graham, Judith E. Allen, Andrew F. Read 17 Th ] 1 ] f 1 Inon Schlupp D 4 dd Y VL 4 NON IT eO wd 18. M tofi t Timothy Wootton, Mark Emmerson 19. pius NR in phylogeneties—Jack Sullivan, Pal Joyce 0. P Tiffany M. Knight, Janette A. Steets, Jana C. Vamosi, Susan J. Mazer, Martin cua Diane R. Campbell, Michele R Dudas Mark O. Johnston, Randal) Mitchell, Tia-Lynn Ashman Jeffrey R. Stevens, Fiery A. Cushman, Marc D. Hauser 21 p 22. Nic! ism: i j lution, ecology, and j ] John J. Wiens, Catherine H. Graham 23. id M m Philippe, Frédéric ai Henner Brinkmann, Nicolas Lartillot 24. T g Ulrich G. Mueller, Nicole M. Gerardo, Duur K. Aanen, Diana L. Six, Ted R. Schultz 25.1 t plants: diversity of herbi blag d—Thomas M. POM Vojtech Novotny, Yves Basset 26. T} pulation biology of mitochondrial DNA and i ua y genetic imp J. William O. Ballard, David M. Rand 27. A sl oyes eco ff ications—Jennifer L. Ruesink, Hunter S. H Alan C. Trimble, Kimberly W. Heiman, lorena Micheli, James E. Byers, Matthew C. Kay Subject Index Cumulative Index of Contributing Authors, Volumes 32-36 Cumulative Index of Chapter Titles, Volumes 32- J. Bot. Res. Inst. Texas 2(1): 642. 2008 NOTEWORTHY VASCULAR PLANT COLLECTIONS FROM NORTHWEST LOUISIANA Christopher S. Reid and Barbara R. MacRoberts and Patricia L. Faulkner Michael H. MacRoberts Louisiana Natural Heritage Program Bog Research, 740 Columbia Louisiana Depart t of Wildlife and Fisheri ee Louisiana 71104, U.S.A. Baton Rouge, Louisiana 70898, U.S.A. and Herbarium, Museum of Life Sciences Louisiana St tate University in Shreveport Shreveport, Louisiana 71115, U.S.A. ABSTRACT TI LA! f. ] t ] t T ici g T i 1 lat Ç j Hi , Lechea can_ecoheangd , and Panicum geen Significant new aud are nes for ind b an Grindelia i lanceolata, s cardenas Geocarpon minimum, and Carex RESUMEN 7 : ` »" " : E. + I cot Cc TZ i halat Ç in A Na Terhen con-caheman H H BOX So M eu sg T. AI i 2. F^ E INTRODUCTION Recent floristic work in nort] tern Louisiana has resulted in the di of plants new to Louisiana and other significant records (Reid & Faulkner 2006; MacRoberts et al. 2007; Reid et al. 2007). In this paper we report four species new to Louisiana and four additional significant collections. ALISMATACEAE Sagittaria subulata (L.) Buchenau—Sagittaria subulata is not reported for Louisiana by USDA, NRCS (2007), Haynes and Hellquist (2000), Kartesz and Meacham (2005), MacRoberts (1984), or Thomas and Allen (1993). On 27 June 2006, we discovered hundreds of plants on a stretch of shoreline on Caddo Lake at Oil City, northern Caddo Parish. According to Haynes and Hellquist (2000), S. subulata ranges from Mas- sachusetts south to Florida and west to Mississippi. Our record extends the range of S. subulata over 500 km to the northwest of its previously known western range limit in Mississippi. I Caddo Parish: a! ] f oe Lake in Oil City, E1/2 SE1/4 $13 T20N = Aa E ee open "a lakeshore in saturated muc ln sand, with Eleoch ?), Nelumbo lutea, P Luziola tes irre cd a: os ge on da 100s of submerged plants, 27 Jun 2006, Reid 5809 (FSU, LSU); along shore oat ramp in Earl G. Williamson Park, Oil City, flowers white, 11 Jul 2006, L.R. Raymond 2166 (LSUS). ASTERACEAE Grindelia lanceolata Nutt.— Grindelia l latai ted for Louisiana by Riddell (1852) and Steyermark (1934). Steyermark (1934) cites Leavenworth specimens at G (= GH) and NY as his authority. MacRoberts (1984 and Thomas and Allen (1996) include G. lanceolata in the Louisiana dee on He basis of these older literature reports. We discovered G. lanceolata on 20 June 2007 int ine Parish. The plants occurred on open slicks with overall light vegetative cover. Daca a return visit on 7 August 2007, we observed an estimated 500 plants, many of which were beginning to flower. Our record of G. lanceolata is apparently the first in Louisiana since Leavenworth’s time, ca. 150 years ago (Ewan 1968). "— | Voucher specimens: Sabine Parish: Oakley's Pasture (a.k.a. Mary Lee's Pasture), N of LA 174 ca. 5 mi W of Converse, along E side of McDonald Bayou, NW1/4 NW1/4 512 & SW1/4 SW1/4 S1, TON R14W. lat/long: 3147 11N, 934544W, common in small area of remnant J. Bot. Res. Inst. Texas 2(1): 643 — 647. 2008 I L_ £al Das H In LI d. den ade fT TÁ E! A ,F 644 p tl t d within a large area of f li irie that ted to Bahia grass long ago, Talinum parviflorum, Cynodon da nis SE bolus pyramidatus, Evolvul i tc., remnant ca. 0.25 acres, 20 Jun 2007, Reid 6164, Faulkner et al. (LSU); N edge of pi i lj Oak] ture (a.k.a. Mary Lee's Pasture), N of LA id ca. 5 air mi W iod Converse, E side of McDonald avail SEI/4 NE1/4 S11 TON RIAW. lat/long: 314658N, 934554W, several | remnant with abundant Croton willdenowii, also w/ C. capitatus, Neptunia lutea, Talinum parviflorum, Gesten drummondii, & T amarum, 7 Aug 2007, Reid 6211, Faulkner & Gregory (LSU, LSUS, NO Senecio ampullacens Hook.—Senecio ampullaceus is not reported for Louisiana by MacRoberts (1984), Thomas and Allen (1996), Kartesz and Meacham (2005), USDA, NRCS (2007), or Barkley (2006). On 28 March 2007, we discovered a population of S. ampullaceus in the xeric sandhills (M.H. MacRoberts & B.R. MacRoberts 2006) of northern Caddo Parish. It was recently reported new to Arkansas (Singhurst & Hol- mes 1998) and Oklahoma (Holmes et al. 2006). Our record is apparently the first for 5. ampullaceus from Louisiana. Voucher specimen: Caddo Parish: 5 side of Kendrick Road near its W terminus at Vivian-Atlanta Rd., near NE corner of S15 & NW corner of 514, T22N R16W. lat/long: 325420N, 935957 W, sandy field with conspicuous Opuntia humifusa, 28 Mar 2007, Reid 6020 & Faulkner (LSU). BRASSICACEAE Arabis canadensis L.—On 13 April 2006, we discovered a small population of Arabis canadensis in De Soto Parish growing on a low ridge embedded in a matrix of poorly drained Quercus similis — Crataegus spp. flatwoods. MacRoberts (1989) and Thomas and Allen (1996) include A. canadensis in the Louisiana flora on the basis of a specimen collected by Leggett (1578 — NLU) from Bossier Parish. Leggett (1976) reported it from one station in Bossier Parish where it was "localized and uncommon on sandy soil in mixed woods. During recent visits to NLU, we have failed to locate this specimen. It is apparently not on loan and may have been misfiled or annotated to another taxon. Providing that the identification and locality data are correct, our record is the second from Louisiana. If the Leggett specimen was misidentified, then our re- cord is the first for the state. In either case, our record represents the only confirmed extant A. canadensis population in Louisiana. De Soto Parish: Dicl Prairie, S side of C B .2 air mi NE of Stonewall, NW1/4 S9 TI5N R14W, mixed ZE r hard 1-loblolly f l id iated with G d , Pinus M Aesculus pavia, Senecio obovatus, Celtis laevigata, Dioscorea villosa, and Sabal minor, 13 Apr 2006, Reid 5713 and Faulkner (LSU); same locality, 2 May 2006, Reid 5751, Faulkner et al. (FSU, LSU). CARYOPHYLLACEAE Geocarpon minimum Mackenzie Arkansas, Louisiana, Missouri, and Texas (Niendlier 2005). This Peel was initially discovered in Louisiana in Winn Parish in 1989 with an additional population discovered nearby in 1992 (McInnis et al. 1993). These were the only two Louisiana populations known until 2006 when a third occurrence was discovered in a saline prairie in Caddo Parish (B.R. MacRoberts & M.H. MASON 2006). A survey for G. minimum in saline prairies in Caddo, De Soto, Red River, and Winn parishes was ted in March 2007 (MacRoberts & MacRoberts 2007) that yielded three new sites representing two new populations, all in De Soto Parish. Because two of the collection sites, Dickson Prairie ane Daten ue are bi co by a narrow drainage, a Geet is federally listed as threatened and is known from we regard G. minimum from these two sit lation. The third collection was from another saline prairie, Rambin Bayou Prairie, which is well removed from the other sites. V I D h: Dickson Prairie, ca. 2 air mi NE of St 11 LAS all lat/long: 321827N, 934824W, : — d ER , Plantago pusilla, Gusti rosea, H. micrantha, Crassula pica e ai minima, 7 Mar 2007, Reid 5995, Faulkner, MacRoberts e EA (LSU); Dalton Prairie, ca. 2.5 air mi SE of Keithville, SE1/4 54 T15N R14W, lat/long; 321841N, 934807 W, saline prairie, in and around slicks and sparsely vegetated areas associated with Opuntia aide: Plantago pusilla, Talinum panic; Houstonia rosea and H. micrantha, etc., 7 Mar 2007, Reid 6003, Faulkner, MacRoberts & À ts (LSU); Rambin Bayou Prairie, N of Rambin Bayou, W of 1-49 via gas field e NELA S? TIAN RISW. lat/long en 93435 IW, ca. 250 plants counted, in and around slicks with Anagallis minima, lothoscordum bivalve, Krig talis, Plantago pusilla, Houstonia spp., and Talinum parviflorum, 19 Mar 2007, Rei 6010, Pauline & Smith (LSU). Reid et al., N A Ll 1 A il a? £ all A) - m 645 e Lechea san (Buckley) Hodgd Lechea san-sabeana was reported for Louisiana by Riddell (1852) as Lechea drummondii Torr. SA. Cay and luded from the flora by Thomas and Allen (1996). MacRoberts (1984) lists L. san-sabeana as questionable in Louisiana and cites the Riddell report. Correll and Johnston (1970), Carr (2002), Kartesz and Meacham (2005), and Diggs et al. (2006) regard L. san-sabeana as a Texas endemic. In May and June 2007, we collected L. san-sabeana at three saline prairies in De Soto and Caddo parishes. It was common at these sites and was associated with pimple (mima) mounds. Voucl i Caddo Parish: B Road Saline P T16NR14WS27, N of Barron Road, S of Boggy Bayou. Mima mounds, 13 2007, BR _& MH. o 7830 bnnc De Soto Parish: ee Prairie, ca. 2 air mi NE of Stonewall, NW1/4 S9 T15N R14W. e 321827N, 934824W May 2007, Reid 6064, Faulkner & Smith (TEX-LL); same e locality, common on n lower edges e pimple nen associated w/ m d Chamaecrista fasciculata, Rhynchospora harvey aciculare, 21 Jun 2007, Reid 6179, Faulkner, & U). Jones (LSU, NO); Eton Saline puiltie/Bstteti, TISNRIANSA, mima monid, 12 Jun 2007, B.R. & M. H. MacRoberts 7825 (LS CYPERACEAE Carex arkansana (L.H. Bailey) L.H. Bailey—Louisiana is not included in the range of Carex arkansana by Ball (2002) but is so included by Kartesz and Meacham (2005), USDA, NRCS (2007), NatureServe (2007), and Thomas and Allen (1993). MacRoberts (1989) and Thomas and Allen (1993) cite a specimen from Caddo Parish (Thieret 25936 — LAF) collected in 1967 as the only record for Louisiana. B.R. MacRoberts and M.H. MacRoberts (2006) examined this specimen and confirmed its identification. On 2 May 2006, the authors discovered a small population of C. arkansana growing in a depression in a saline prairie in De Soto Parish. On 4 May 2007, we found a much larger population in a wet swale in a different part of the same prairie complex, adjacent to Quercus similis-Crataegus brachyacantha flatwoods. Our record is only the second record for this species in Louisiana and came 40 years after Thieret first collected it from the state. V. De Soto Parish: Dickson Prairie, S side , 2 air mi NE of Stonewall, NW1/4 S9 TIS5N R14W, rare, tiny depression in saline prairie, associated with Carex Dd a sun, 2 Ms 2006, Reid 5748, Faulkner et al. (LSU, SWSL) POACEAE Panicum diffusum Sw.—On 20 June 2007 we found Panicum diffusum to be locally common in a saline prairie remnant in Sabine Parish. It was iated with open slicks with much bare soil and light vegetative cover. This record is apparently the first for Louisiana as the species is not reported for the state by Nature- s (2007), Mid NRCS (2007), Thomas and Allen (1993), Allen et al. (2004), nor Freckman and Lelong icum iously known in the United States only from Texas; its range extends south ( to the Caribbean and Latin p (Diggs et al. 2006; Freckman & Lelong 2002). Voucher specimen: Sabine Parish: Oakley's Pasture (a.k.a. Mary Lee's Pasture), N of LA 174 ca. 5 mi W of Converse, along E side of McDonald pij NWI/4 we SC & SW1/4 SW1/4 e TON R14W. lat/long: 314711N, 934544 W, common in slicks in saline prai- rie, , Talinum parviflorum & Sporobolus pyramidatus common and characteristic, 20 Jun 2007, Reid 6173, Faulkner et al. (Ft. Polk Herbarium, LSU) ACKNOWLEDGMENTS We thank George Dickson and Ralph and Becky Dalton for allowing our surveys on Dickson and Dalton saline prairies, respectively. Sidney Evans and James Taylor allowed us to survey Barron Road Prairie in Caddo Parish. Judy Oakley allowed our survey of Oakley's Pasture. We thank Tommy Smith of Red River Forest Management for taking us to rin Bayou Prairie in De Soto Parish. Red River Forest Management also owns part of the Grindelia | lation in Sabine Parish and we appreciate their kind permis- sion to botanize on their lands. Robert love of Louisiana Department of Wildlife and Fisheries informed us about Oakley's Pasture and an additional saline prairie site in Sabine Parish and led us to these sites. We thank DW. and Patricia Little for allowing access to their property in order to collect Senecio ampullaceus. Ryan Hook allowed access to the Senecio ampullaceus location through his family's property and we appreci- ate his kindness and interest. Jason Singhurst verified our first collection of Lechea san-sabeana. We thank 646 J tl tani h Institute of Texas 2( Charles Bryson for verifying a duplicate of Carex arkansana and Robert Haynes for verifying a duplicate of Sagittaria subulata. Charles Allen verified a duplicate of Panicum diffusum and we appreciate his help. Dennis Bell made one last search at NLU for the Bossier Parish specimen of Arabis canadensis and we appreciate his help with trying to track down that specimen. We thank J. Scott Peterson, R. Dale Thomas, and an anony- mous reviewer for providing helpful comments on this manuscript. REFERENCES ALLEN, CM, DA Newman, and H.W. Winters. Grasses of Louisiana, 3 edition. Allen's Native Ventures, LLC. Pitkin, LA. 374 pp. BALL, PW. 2002. Carex sect. Phaestoglochin. In: Flora of North America editorial committee, eds. Flora of North America. Volume 23. Magnoliophyta: Commelinidae (in part): Cyperaceae: 285-297. Oxford University Press, New York and Oxford. BAnkLEY, TM. 2006. Senecio In: Flora of North America editorial committee, eds. Flora of North America. Volume 20. Magnoliophyta: Asteraceae, part 2:544—570. Oxford University Press, New York and Oxford. CARR, W.R. 2002. No place but Texas: An annotated list of the plants endemic to the Lone Star State. Unpublished manuscript. CorreLt, D.S. and M.C. JoHNsTon. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner, Texas. Dicas, G.M., B.L. Lipscoms, M.D. Reep, and RJ. O'Kennon. 2006. Illustrated flora of east Texas. Sida, Bot. Misc. 26:1-1594. Ewan, J. 1968. A bibliography of Louisiana botany. SouthW. Louisiana J. 7:1-83. FRECKMAN, R.W. and M.G. LELONG. 2003. Panicum. In: Flora of North America editorial committee, eds. Flora of North America. Volume 25. Magnoliophyta: Commelinidae (in part): Poaceae, Part 2:450-488. Oxford University Press, New York and Oxford. Haynes, R.R. and C.B. HeLLquIsT. 2000 Alias In: Flora of North America SOLUM committee, eds. Flora of North America. Volume 22. Magnoli ta: Alismatidae, Arecidae, C li (in } nd Zingiberidae: 7—25. Oxford University Press, New York an Oxford. Homes, W.C., J. SINGHURST, and S.G. Powers. 2006. Senecio ampullaceus (Asteraceae): A West Gulf Coastal Plain en- demic new to Oklahoma. Phytologia 88:193-196. Kartesz, J.A. and CA MEACHAM 2005. Synthesis of North American flora. Version 2.0. North Carolina Botanical Garden. Chapel Hill. Leccett, V.S. 1976. A preliminary survey of the vascular flora of Bossier Parish, Louisiana. M.S. Thesis. Northeast Louisiana University. MacRoseats, B.R. and M.H. MacRoserts. 2006. An updated, annotated vascular flora of Caddo Parish, Louisiana, with notes on regional phytogeography and ecology. Sida 22:1191-1219. MacRostnrs, 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. 6:1-165. MacRoserts, D.T. 1989. A documented checklist and atlas of the vascular flora of Louisiana. Bull. Mus. Life Sci. 791-7595 MaAcRosenrs, M.H. and B.R. MAcRoBERTS. 2006. Xeric lylands (west ic sandhill woodlands) of the West Gulf Coastal Plain. Unpub. Manuscript. Louisiana Natural as Program, E SEN EE Louisiana. MacRoserts, M.H. and B.R. MacRoserts. 2007. Survey for G rthwest Louisiana. Unpublished survey report submitted to Louisiana Natural Heritage Biogr MacRosenrs, M.H., B.R. MacRoserts, C.S. Ben. PL. FAULKNER, and D. Estes. 2007. Minuartia drummondii (Caryophyllaceae) and Gratiola flava (Plantaginaceae) rediscovered in Louisiana and Gratiola flava historically in Arkansas. J. Bot. Res. Inst. Texas 1:763-767 McInnis, NM. L.M. Smith, and A.B. Pitman. 1993. Geocarpon minimum (Caryophyllaceae), new to Louisiana. Phy- tologia 75:159-162. Reid et al., Not thv plant collections f ti t Louisi 647 NaTURESERVE. 2006. Nat S Expl An onli lopedia of life [wel lication]. Version 6.1. NatureServe, Vu Y A ` Pi rr Ela “q / + rra Fr explorer. nf Arlington, Virginia. Available http NiENABER, M.A. 2005. Geocarpon. In: Flora of North America editorial committee, eds. Flora of North America Vol 5: Magnoliophyta: Caryophyllidae, Part 2:148-149. Oxford Univ. Press, New York and Oxford. Rep, C.S. and P.L. Faulkner. 2006. Loeflingia squarrosa (Caryophyllaceae): new to Louisiana. Phytologia 88:98/—90. Rep, C.S., PL. FAULKNER, B.R. MacRoserts, and M.H. MacRoserts. 2007. Saxifraga texana (Saxifragaceae) new to Loui- siana. J. Bot. Res. Inst. Texas 1:1251-1252. RiDDELL, J.L. 1852. Catalogus florae Ludovicianae. New Orleans Med. Surg. J. 8:734—754. SINGHURST, J.R. and W.C. Homes. 1998. Four new additions to the Flora of Arkansas. Castanea 64:2/6-277. STEYERMARK, J.A. 1934. Studies in Grindelia Il. A monograph of the North American species of the genus Grindelia. Ann. Missouri Bot. Gard. 21:433-608. Thomas, R.D. and C.M. Auen. 1993. Atlas of the vascular flora of Louisiana. Vol. |: Ferns, fern allies, conifers, and monocotyiedons. Louisiana Department of Wildlife and Fisheries, Baton Rouge. THomas, R.D. and C.M. ALLEN, 1996. Atlas of the vascular flora of Louisiana. Vol. Il: Dicotyledons: Acanthaceae - Eu phorbiaceae. Louisiana Department of Wildlife and Fisheries, Baton Rouge. USDA, NRCS. 2007. The PLANTS Database (http://plants.usda.gov). National Plant Data Center, Baton Rouge, LA 70874-4490 USA. 648 Jo BOOK NOTICE DoucLas J. FurUYMA, H. BRADLEY SHAFFER, AND DANIEL SIMBERLOFF (eds). 2007. Annual Review of Ecology, Evolution, and Systematics: Volume 38, 2007. (ISBN 978-0-8243-1438-5, hbk; ISSN 1543-592X). Annual Reviews Inc., 4139 El Camino Way, PO. Box 10139, Palo Alto, California 94303-0139, U.S.A. (Orders: www.annualreviews.org, science@annualreviews.org, 1-800-523-8635, 1-650-493-4400, 1-650-424-0910 fax). $80.00 indiv., $197.00 inst. (USA), 876 pp., indexes, 7 5/8" x 9 3/8". Contents of Volume 38: 1. Evolution of Animal Photoperiodism—William E. Bradshaw, Christina M Holzapfel 2. Virus Evolutio ights f Experimental Approach—Santiago F. Elena, Rafael Sanjuán 3. The Social Lives of Missdbes-— Suas A. West, Stephen P. Diggle, Angus Buckling, Andy FÉ Ashleigh S. Griffin 4. Sexual Selection and Speciation—Michael G. Ritchie 5. Kin Selection and the Evolutionary Theory of Aging— Andrew F.G. Bourke 6. Climate Ch d Invasibility of the Antarctic Benthos—Richard B. Aronson, Sven Thatje, Andrew Clarke, Lloyd S. Peck, Daniel B. Blake, Cheryl D. Wilga, Brad A. Seibel f Visual Signals in Animal C ication—Gil G. Rosenthal f Flight: Bi hanical Novelt N ity? —Robert Dudley, Greg Byrnes, Stephen P. Yanoviak, VL LYALL IO LL i 7. Ee Dimensions of Brendan Borrell, "c M. edt Jimmy. A. McGuire 9. How Mutational ^ 10. How Does It Feel to Be Like a Rolling Stone? Ten Questions about Dispersal e Ronce 11. o ES anobacterial led m cl M. Usher, ap de id John A. t, Present, and Future— Dav ichardson, ende W. —— Stephen T. Jackson, Robert O. Teskey, from RNA Models— Matthew C. Cowperthwaite, Lauren Ancel Meyers Ll RM Piin uro Man] ving sera Proches 13. Chemical C Hu pow ee Linda Partridge 14. A Global Review of the a Ge and 1 t d € L. oe Ce Smith t fel H FO Anrectrar Arturo AT LiAl A 15. The V Becerra, Luis Delaye, Sara me Antonio Lazcano . Functional Versus Morphological d in Macroevolution— Peter C. Wainwrig of Continuous Traits—Brian mec SEN Brown p C 17. Evolutionary Game Theory and Adaptive Dvnamics 18. The Maintenance of Outcrossing in MR Selfing Species: Ideas and E g Į Christopher G. QUAN Ken S. Moriuchi, Alice A. W 19. S peciation: Models and Empirical Lam ade Bolnich, odd M. D AM 20. The Evolution of Col I ypticity, ing Images, and Alan B. Bond 21. Point, Counterpoint: The Evolution of Pathogenic Vi 1 their H Hosts-—Michael V orobey, Adam SH Joel O. Wertheim 22. The Evolution of Resistance and Tole Herbi Juan Nüfiez Farfán, Juan Fornoni, Pedro Luis Valver 23. Plant-Ànimal Mutualistic Networks: The Architect f Biodiversity—Jordi Bascompte, Pedro Jordano 24. Gene ge and hd AMEN in Trees—Outi Savolainen, Tanja Pyhäjärvi, Timo Knü The nor Major Transition?— Richard K. Grosberg, S Um Strathmann J.T. Streelman, C.L. Peichel, D.M. Parichy 26. Developmental Genetics of Adaptation in Fishes: The Case for Novelty 27. T trial Carl Cycle Feedback to Climate Warming—Yigi Luo 28. Shortcuts for Biodiversity Conservation Planning: The Effectiveness of Surrogates—Ana S.L. Rodrigues, Thomas M. Brooks 29. Understanding the Effects of Marine Biodiversity on Communities and Ecosystems—John J. Stachowicz, John F. Bruno, J. Emmett Duffy 0 Ei 1 y f Plant-Water Interacti Gabriel Katul, Amilcare Porporato, Ram Oren 31. Evoluti y Endocrinology: The Developing Synthesis] Endocrinology and Evoluti y Genetics — Anthony J. Zera, Lawrence G. Harshman, Tony D. Williams 32. The Role of Behavior in the Evolution of Spiders, Silks, and Webs—Fritz Vollrath, Paul Selden 33. Applicati f Flow Cytometry to Evalaif onar y and Population Biology— Paul Kron, Jan Suda, Brian C. Husband umulative Index: Contributing Authors, Volumes 34-38 Chapter Titles, Volumes 34-3 J. Bot. Res. Inst. Texas 2(1): 648. 2008 SILENE FLOS-CUCULI SSP. FLOS-CUCULI (CARYOPHYLLACEAE) AND EUPHORBIA PEPLUS (EUPHORBIACEAE) NEW TO NORTH CAROLINA Derick B. Poindexter Appalachian State University Biology Department Boone, North Carolina 28608, U.S.A poindexterdb@appstate.edu ABSTRACT T, : 1 ] ] f£. 1 1 D CNT de oi E 1 decl Tal 1 E counties located in extreme northwestern North MM. in p Blue EES E siographic Province of the Southern Appalachians Mel m Tus taxon was aD Silene flos-cuculi ssp. eege is a Euroy from a ruderal, old-fi it theast Alleghany Coma The E tic Eupl x. at 1: 1 E GE d 1 NT 1 : TL: E : ` : letnirl 1 America y disturbance 1n FO Watauga County. RESUMEN T+» t J 4 1 7 [1 1 ] Le 1; dal KT + - "n A 1 P 1 L L Eu A ] ] EN 1 TE J 1 ees E Lk Jal NT de 1 [- T. DI D: (n A 4 Ty H E e [9] hi flag h 1 A F1 11 1 1 ] T da | de los are del sur. Sine as Rom ssp. flos p p | I I Estados Uni E iej leral l te del dado Alleghany. I E J i [e] Pi E 1 1 n | A 1 J = zt 1 A 4 E ARE a à ES "m E ZE 4 4 4 X Eu I aheeryvada | ] ] T "P mriarnrin de éi I Watauga. Silene flos-cuculi (L.) Clairv. ssp. flos-cuculi (syn. Lychnis flos-cuculi L.), Ragged Robin, is a showy intro- duced taxon in the Caryophyllaceae with deeply dissected, pink petals. This perennial European native is presumed to be established in the northeastern United States (Weakley 2007), but has also been reported in several other U.S. localities, including fifteen states and the District of Columbia. States with records of this plant are Connecticut, Maine, Maryland, Massachusetts, Montana, New Hampshire, New Jersey, New York, Ohio, Pennsylvania, Rhode Island, Vermont, Virginia, Washington, and Wisconsin (USDA, NRCS 2007). Silene flos-cuculi ssp. flos-cuculi in abundance, with approximately four individuals observed in an old-field across from Rich Hill Church in the rural Glade Valley community. Associated taxa include Dactylis glomerata L., Pinus virginiana Mill., Pteridium aquilinum (L.) Kuhn var. latiusculum (Desv.) Underw. ex A. Heller, Rubus argutus Link, and Sassafras albidum (Nutt.) Nees. The four-lobed, lacerate petal morphology and pink coloration differs substantially from other Silene spp. in North Carolina. Voucher specimens: NORTH CAROLINA. pen pen Co.: Glade Valley, Rich Hill Church, at the jct. of Shawtown Rd. and Rich Hill Rd., at latitude 36°28'20" N and longitude 80'59'31" W, 18 May 2007, Poindexter 07-136 (BOON, NCU A pepius- L. o Spurge, is an exotic annual in the Euphorbiaceae. This Eurasian introduced taxon ] l distribution in the United States, E in Ger of thirty-five states, includ- ing Hawaiiand deen Ge Pos SE This her! es its short fleshy stems, primarily alternate, | | nd a 1 mor phology: of outer pitted faces andi mner furrows. This plant was discovered to be frequent in ruderal communities created by regular human perturbation on the campus of Appalachian State University. An estimated 50-100 individuals were observed. Associated taxa include Chamaesyce maculata (L.) Small, Ranunculus repens L., Cyperus strigosus L., Sonchus asper (L.) Hill, Senecio vulgaris L., Portulaca oleracea L., and Galinsoga ani dni Ruiz & Pav. A C T Voucher specimens: NORTH CAROLINA. Watauga Co.: Boone; App iversity, growing pus g j J. Bot. Res. Inst. Texas 2(1): 649 — 650. 2008 650 I | £al Dos H In LI d. dien adis LT [41 to Rankin Science Bldg., parallel to Rivers St., in disturbed areas and permanent flower beds, at latitude 36°12'49" N and longitude 81°40'49" W, 15 Jun 2007, Poindexter 07-205 (BEREA, BOON, NCU). ACKNOWLEDGMENTS I would like to thank Zack E. Murrell, Justin Wynns, and two anonymous reviewers for their critical evalu- ation of this manuscript. 1 would also like to thank Jorge Esquivel for his translation of the abstract. REFERENCES USDA, NRCS. 2007. The PLANTS database. National Plant Data Center, Baton Rouge, LA. (http.://plants.usda. gov/). Accessed 16 July 2007. WEAKLEY, AS. 2007. Flora of the Carolinas, Virginia, and Georgia, working draft of 11 January 2007. University of North Carolina Herbarium, N.C. Botanical Garden, Chapel Hill, NC. (http//www.herbarium.unc.edu/). Ac- cessed 16 Jul 2007. ANNOTATED CHECKLIST OF THE VASCULAR FLORA OF THE BEECH CREEK UNIT OF THE BIG THICKET NATIONAL PRESERVE, TYLER COUNTY, TEXAS Larry E. Brown Herbariu é Ru Branch Science Center estview Drive custon, s 77055, USA, pu theplantman@att.net 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, 71115, U.S.A. Paul A. Harcombe, Warren W. Pruess, and |. Sandra Elsik ny A A ETT Ê (fe I r: Dic-J Suzanne Birmingham Walker Azimuth Forestry Services, Inc. 14671 State Highway 87 South Shelbyville, Texas 75973, U.S.A Rice University Houston, Texas 77005, U.S.A ABSTRACT Án annotated, hered checklist i ji ided for the vascular pl of the 2,059 1 Beech Creek Unit, Big Thicket National Preserve, Tyler County, southeastern Texas. D ted in t] it 470 taxa of which 52t 196) ti RESUMEN Se aporta un catálogo con anotaciones y testigos de los taxa de plantas vasculares de las 2,059 hectáreas de la unidad Beech Creek, Big Thicket National Preserve, Tyler County, sureste de Texas. Los taxa documentados en la unidad son 470 de los que 52 (11%) son exóticos. INTRODUCTION This manuscript is the 5th part of an intended inventory, based on voucher specimens, of the vascular flora of the Big Thicket National Preserve, southeastern Texas. In 4 previous papers, we provided an annotated checklist of the Hickory Creek, Turkey Creek, Big Sandy Creek, and Lance Rosier units (MacRoberts et al. 2002; Brown et al. 2005, 2006a, 2006b). In this paper, we provide an annotated checklist of the Beech Creek Unit. SITE AND METHODS The Big Thicket is located mostly within the longleaf pine region of the West Gulf Coastal Plain in south- eastern Texas (Ajilvsgi 1979; Harcombe & Marks 1979; Watson 1979; Marks & Harcombe 1981; Harcombe et al. 1993; Diggs et al. 2006 and references therein). The Big Thicket National Preserve consists of 12 units scattered over 7 Texas counties. The units range from 222 to 10,100 hectares and total about 34,000 hectares (Peacock 1994). The Beech Creek Unit consists of 2,059 hectares and is located in western Tyler County (Fig. 1). It is relatively flat with poor drainage over most of the area. Elevation ranges from about 43 meters to 62 meters. Little Beech Creek and Beech Creek run through the unit. The Beech Creek Unit was chosen for inclusion in the Big Thicket National Preserve because of its extensive stands of beech forest (Ajilvsgi 1979; Harcom be & Marks 1979; Watson 1979; Marks & Harcombe 1981). It is dominated by lower slope hardwood-pine forest (mesic lower slopes and terraces), with lesser amounts of baygall, mid-slope oak-pine forest, floodplain hardwood-pine forest (mesic stream bottoms), and floodplain hardwood forest (seasonally flooded river floodplain) (Harcombe & Marks 1979; Marks & Harcombe 1981; Van Kley 2006). A map of community types can be found in Harcombe and Marks (1979). Fire suppression over the past 30 years has resulted in extensive midstory development and thus reduction of herbaceous vegetation that | Rat Dee Inet Tavac 2(1): 451 _ 660. 2008 652 | t tanical h Insti Texas No 4 S | “XA Y Pd S: ALABAMA - COUSHATTA a Y \ WOODVILLE myn ^ INDIAN RESERVATION ARTT UPPER NECHES RIVER CORRIDOR UNIT Beech "E \Creek Unit' Sé 7 judi E HILLISTERGS o mo SS BIG SANDY / WARREN (E NECHES BOTTOM 4 JACK GORE BAYGALL UNIT KOUNTZE SN SE V Con Ey” Ji Ze H d EVADALE ' | LOWER be RIVER ROSIER UNIT 2 | LITTLE PINE ISLA ISLAND ORANGE CO. BAYOU CORRIDOR UNIT i BEAUMONT UNIT o, ^ unm i fmm HARDIN SO. ; ei (10 ) LIBERTY CO. s JSOUR LAKE J E EN E ae a! ` P. j Bn S BEAUMONT gl al Ar Ze $ n sana e Jä | i — e: a9 P ` m ue, fé ul 5 Í Fic 1 1 at E al nm L fe it, Big kd WI sz AT PR! IR j Tayac is adapted to frequent fires. Hurricane Rita, a category 3 hurricane, passed over the unit on 24 September 2005 damaging and toppling thousands of trees. It will probably take decades for this litter to decompose. Details about soils, topography, and climate can be found in Deshotels (1978); Watson (1979, 1982, 2006), Harcombe and Marks (1979); Marks and Harcombe (1981); and Diggs et al. 2006). Like most units of the Big Thicket National Preserve, the Beech Creek Unit has not been the subject of a detailed floristic inventory. Larry Brown collected 290 specimens during 9 field days beginning 9 November 2006 and ending 17 October 2007. The MacRoberises collected 154 specimens during 4 field days in 2004 and 1 day in 2007. Suzanne Walker collected 101 specimens during 2 days in 2004 and 1 additional day in 2007. Sandi Elsik and Warren P together gathered 82 specimens during 2005. Geraldine Watson collected 25 specimens Brown et al., Vascular fl it, Big Thicket 653 over several years in the late 1970s and early 1980s. Warren Pruess collected another 11 specimens over a 2 year period. Other collectors totaled 12 specimens. We c (TAMU) herbarium consortium data base (http://www.csdl tamu.edu/FLORA/t onducted a search of the Texas A&M University 1) for ad- f. AE Pi ditional specimens but found no additional taxa not reported here. Because our purpose was to produce a list of taxa known to occur on the Beech Creek Unit, a vouchered specimen was considered to be the only evidence acceptable for inclusion in the list. In all, 675 herbarium specimens form the data for this report. All specimens have been annotated by Larry Brown. In most cases, nomenclature follows Jones et al. (1997), Kartesz (1999), Turner et al. (2003), and Diggs et al. (2006). In cases of multiple collections, no more than 4 are listed for each taxon. RESULTS The following is an annotated list of the Beech Creek Unit taxa. Abbreviations used are given below. GW = Geraldine Watson. Her collections are at TÃES. JL = Joe Liggio. His collections are at TAES LB = Larry Brown. His specimens are at TÃES. MM = Barbara and Michael MacRoberts. Their specimens are at LABS PH = Paul Harcombe. His collections are at TÃES. PM = Phil Malnassey. His collections are at TÃES. RH = Roland Harper (grandson of LB). Acanthaceae Justicia ovata (Walter) Lindau var. lanceolata (Chapm.) RW. F E F 6 Ruellia caroliniensis UE Gmel) Steud., SEWPLB 488; LBSH 3215] Ruellia hurnilis Nutt., MM 7150 Aceraceae Acer rubrum L, MM 7078 Agav dila eer L, LB 31939; 2 272 Yucca louisianensis Trel., MM 714 Anacardia Rhus m L., SEWP 249 Toxicodendron radicans (L) Kuntze, SEWP 337; LB 32783 Toxicodendron vernix (L.) Kuntze, SEWP 341 Annonaceae Asimina parviflora (Michx.) Dunal, SW 03; LB 31996, 32178; MM 7097 Apiaceae entella erecta (Lf) Fernald, SW 03 Chae vophyllur tainturieri Hook. var. tainturieri, WP 113 E. P41 *Cyclospermum leptophyllum (Pers.) Sprague ex Britton & P. Wilson, SW 58 Eryngium prostratum Nutt. ex DC., SW 68 Hydrocotyle verticillata Thunb., SEWP 262; L Ptilimnium capillaceum o Raf, LB o es 32122 Sanicula canadensis L Sanicula smallii E.P. AT "T 111 Apocynaceae Trachelospermum difforme (Walter) A. Gray, LB 31998 Aquifoliac ilex pine VN ) Torr, PH 001 SEWP - Sandra Elsik and Warren Pruess. Their specimens are at TAE SH - Shawn Harper (grandson of LB). SW - Suzanne Walker. Her specimens are at TAES. WP - Warren Pruess. His specimens are at TAES. WRC = Bill Carr. His collections are at TEX/LL. * z exotic + = see note at end of list. llex coriacea (Pursh) Chapm., SW 012 llex longipes Chapm. ex Trel., PH 002 Hex opaca Aiton, MM 705 ilex vomitoria Sol. in Aiton, MM 7058 Araceae Arisaema triphyllum (L) Schott, SW 10, 31 Araliaceae Aralia spinosa L., SEWP 245 Arecaceae Sabal minor (Jacq.) Pers, MM 7131 Aristolochiaceae Aristolochia serpentaria L., SW 05; GW 2261 Asclepiadacea Asclepias E Walter, SEWP 230 Asclepias variegata L., LBRH 32061 Aspleniaceae Asplenium platyneuron (L) Britton, Sterns, & Poggenb., WP 116 Asteraceae Ambrosia artemisiifolia L., MM 7198 Ambrosia trifida L., SEWP 265 Baccharis halimifolia L, LBRH 32071; SW 105 Bidens aristosa (Michx.) Briton, LB 33034 Bidens bipinnata L., SEWP 252; LB 33030 Bidens frondosa L., LB 33003 Boltonia diffusa Ell, MM 7199 +Bradburia pilosa (Nutt.) Semple, SEWPLB 491, 501; MM 7197 Cirsium horridulum Michx., SEWP 240; SW 111 Conoclinium coelestinum (L.) DC., SW is LB 33039 Conyza bonariensis (L) Cronquist, LB 32466 Conyza canadensis (L) Cronquist, 335; LBJL 32616a Coreopsis lanceolata L., SW 063 Coreopsis tinctoria Nutt., SEWP 253 654 Elephantopus carolinianus Raeusch. SW 74; LBJL 32959 Elephantopus nudatus A. Gray, LB 33002 Elephantopus tomentosus L., LB 28598a, 28353; SEWP 327; Erechtites See (L) Raf. ex DC., SEWP Erigeron s Muhl. ex Willd., MM 7139; n 32060 Bm IER (Small) Lam., PM 03; LB 32457 Fupatorium lancifolium (Torr. & A. Gray) Small, LBSEWP 31899; Eupatorium mohrii Greene, SEWPLB 496 Eupatorium perfoliatum L., SW 83; MM 7147 Eupatorium serotinum Michx., SW 43; PM 02; LB 33016 a ER Jor & A. Gray) Greene ex Porter & n, LBSEWP 3 E. retusa (Lam) n on LBRH 32048; MM 7683 Camarda See Foug.. \ var. pulchella, SEWP 255 M 7106 Gamochaeta purpurea (L J L) Cabrera, SW gn Helenium amarum (Raf) H. Rock var. amarum, MM 7135 Helianthus angustifolium L., ~ 403b; LB 33023 Hieracium gronovii L, LB 32 ES E DC. var. artemisiifolius, MM 7148 *Hypochaeris microcephala (Sch. Bip.) n var. aibiflora (Kuntze) Cabrera, LBRH 32075; MM 7 Krigia cespitosa (Raf) Chambers, LB SCH Ce 32051 Krigia virginica (L.) Willd., MM 7678 Mikania scandens (L.) Willd., SW 33, 86 Lactuca canadensis L., LB 32166 Pluchea camphorata (L) DC., MM 7211, 7212; LB 32804 Pluchea foetida (L.) DC., MM 7236; SW 92 Pyrrhopappus carolinianus (Nutt.) Wooten & Standl., LB Pyrrhopappus pauciflorus (D. Don) DC., WP 119; SW 125 Rudbeckia hirta L., MM 7112; LBRH 32074 Solidago altissima L., SW 100; LB 33015 Solidago ludoviciana (A. Gray) Small, SEWPLB 502 Solidago rugosa Mill. var. aspera (Aiton) Fernald, LBSEWP 31887; SEWP 331; LB 33024 *Sonchus asper (L.) Hill, LB 32006; SEWP 544; SW 113 Symphyotrichum dumosum (L.) G.L. Nesom, MM 7206 Symphyotrichum lateriflorum (L) A. SD Love, WP 120; MM 7668; GW 2248; SEWPLB 481 Symphyotrichum racemosum (Elliott) G.L. Nesom, LBSEWP 31886; LB 33001 Berberidac idi. pe L, MM 7109 Betulaceae Alnus serrulata (Aiton) Willd., GW 2250 Betula nigra L., SEWP 360 Carpinus caroliniana Walter, MM 7083 Ostrya virginiana (Mill) K. Koch, MM 7126 Bignoniaceae Bignonia capreolata L, MM 7065 Campsis radicans (L. Seem. ex Bureau, SW 02, 49 Blechna jdn areolata (L) T. Moore, SW 47; MM 7659 Woodwardia virginica (L.) Small, MM 7238; LB 33036 Brassicaceae *Cardamine hirsuta L., LB 31943 Cardamine parviflora L. var. arenicola (Britt.) Schultz, LB Bromeliaceae Tillandsia usneoides (L) L, SEWP 395 Buddlejaceae Polypremum procumbens L., PM 04; LBJL 32601 Burmanniaceae Apteria aphylla Barnhart., MM 7224; SW 88; LB 33009 Burmannia biflora L., SEWP 422 Callitrichac e Ge? Nutt., LB 32017 Campanulaceae Lobelia cardinalis L., SW 84; MM 7 Lobelia appendiculata A DC. Kë a LBSH 32149 Lobelia puberula À SEN SW 34; GW 2249, 2336; LB 31934 ne | R220090 I BDLI 359^&oO H mio, SCH (L) Nieuwl. LBRH 32059 Wahle o marginata (Thunb.) A. DC., WP 121; LB 32487; W 1 Caprifoliaceae *Lonicera japonica Thunb., MM 7162 Sambucus nigra L. subsp. canadensis (L.) R. Bolli, SW 60, Viburnum acerifolium L., oe : Viburnum dentatum L., SW Viburnum nudum L. var. eum GW 2260b; LB 32170 Celastraceae uonymus americanus L., SEWP 418 Cistaceae Lechea mucronata Raf., GW 06b; PM 06 Lechea tenuifolia Michx., LBRH 32058 Clethra Clethra GE L. SW 20; MM 7119 Commelin E Bain L., LBSH 321 Commelina virginica L, MM 71 e LBSH 32120 Convolvulaceae Dichondra carolinensis Michx., MM 7161 Ipomoea cordatotriloba Dennst var. cordatotriloba, SEWP 350; LB 33031 Ipomoea pandurata (L.) G. Mey., LB 324 Jacquemontia tamnifolia (L.) Griseb., F 500 Cornaceae Cornus florida L, MM 7087 Nyssa aquatica L., SEWP 362 Nyssa biflora Walter, SEWP 397 Nyssa sylvatica Marsh., SEWP 275; MM 7081 | JE £ al n Le Brown et al a it, Big Thicket Crassulac LS ee L, LB 32805 Cupressac Juniperus ef? L. var. virginiana, Hd 7076; LB 33032 Taxodium: distichum (L) L.C. Rich., SW 4 Cucurbitaceae *Citrullus lanatus (Thunb) Matsum & Nakai LB 33011 Cuscutaceae Cuscuta compacta Juss. ex Choisy, MM 7231; LB 33007 Cyperaceae Carex albicans Willd. ex Spreng. var. australis (L.H. Bailey) Rettig, LB 31936 Carex brevior (Dewey) Mack. ex Lunell, SW 127 arex cherokeensis Schwein., LB 32004 Carex complanata Torr. & Ron | E 31953, 31989; MM 7677 Carex corrugata Fernald, LB 320 Gs debi lis Mi de var. EPA o 723] S | illd. var. foridana (L.H. Bailey) Naczi & 8n ryson, LB 31947, 32007; i a 32133; MM 7672 Carex flaccosperma Dewey, LB 32014, 32015 les frankii Kunth, LB 32175 tt, MM 7210; LBJL 32600 e intumescens Rudge, LBRH 32046; MM 7663; SW 129; 32065 Carex joor? L.H. Bailey, SEWP 351 Carex leptalea Wahlenb subsp. harperi (Fernald) W. Stone, LBSEWP 31889; LB 32023 Carex lurida Wahlenb., LBSH 32148; LBJL 32618; SW 119 +Carex section ovales, LB 31 L07 Cyperus croceus Vahl, LBRH 32067 *Cy, entrerianus Boeck., SEWPLB 503 Ge entra Muhi., LB 32794 Ly SEWP 3 a zt 32481 Cyperus odoratus L SEWP 4 Cyperus pseudovegetus nien LB 32165 Cyperus virens Michx., MM 7666 SEET microcap Tort, LBSH 32134a & Schult., LBSEWP 31895; MM 7233; LBRH 22047 E 7662 Kyllinga odorata Vahl, LB 3 Kyllinga pumila Michx., s m Rhynchospora caduca Elliott, MM 7127; LBSH 32150 Rhynchospora corniculata (Lam.) A. Gray, SEWP 231 +Rhynchospora debilis Gale, LBSH 32140; LBRH 32087 Rhynchospora elliottii A. Dietr., LBSH 32214 Rhynchospora fascicularis (Michx.) Vahl, LB 32479 Rhynchospora globularis (Chapm.) Small var. globularis LBSH 32132, 32220 Rhynchospora glomerata (L.) Vahl, SW 98; LB 32472, 33019 Rhynchospora inexpansa (Michx.) Vahl, MM 7118 Rhynchospora mixta Britton, MM 7074; LB 33038 Rhynchospora plumosa Elliott, LB 32022; LBRH 32078; MM ; SW 109 Rhynchospora pusilla Chapm. ex M.A. Curtis, LBSH 3214 Scirpus cyperinus L., LB 32164 Scleria oligantha Michx., LB 31995; MM 7 Scleria trigiomerata Michx., MM 7192; LB Ge LBJL 32593; SW 120 655 Cyrillaceae Cyrilla racemiflora L., SW 22; MM 7121 Dennstaedtiaceae Pteridium aquilinum (L) Kuhn, SW 96; MM 7133 Dioscoreaceae Dioscorea villosa L., GW 2288 Droseraceae Drosera brevifolia Pursh, SW 27; MM 7654 Dryopteridaceae Athyrium ee (L.) Roth var. asplenioides (Michx.) Farw., MM 7 ene sensi L, MM 7228; LBSH 321 d B 31937 Ebenaceae Diospyros virginiana L., LB 32482 Erica iu oe racemosa (L.) A. M ~ 359 Lyonia ligustrina (L) DC, MM 7 Lyonia mariana (L) D. Don, dn 416 Monotropa uniflora L., SW 89; MM 7117; LB 33035 Rhododendron canescens (Michx.) Sweet, WP 112 e viscosum (L) Torr, SW 18; MM 7122; LBSH DUE arboreum Marshall, PH 005 Vaccinium corymbosum L., SW 19, 130, 45; LB 31950 Euphorbiaceae Acalypha gracilens A. Gray, SW 99; GW 2255 Chamaesyce maculata (L.) Small, SEWP 247 Chamaesyce nutans (Lag.) Small, SEWP 251 Croton capitatus Michx., MM 7200 Croton glandulosus L. var. septentrionalis Muell.-Arg., SEWP 270 Croton monanthogynus Michx., SEWP 259 Croton willdenowii G.L. Webster, LB 32490 Euphorbia corollata L., LB 32807 Fuphorbia dentata Michx., SEWP 254; | B 32460 M d caroliniensis SSC LB 32475 *Phyllanthus urinaria L., M 1 Sebastiania fruticosa (W. Em Fernald, SW 14 *Triadica sebifera (L.) Small, SW 01 Fabac ias NUBE Durazz, MM 7 Centrosema virginianum (L.) rn GH 32604 Cercis canadensis L. var. canadensis, SEWP 271 Chamaecrista fasciculata (Michx.) Greene, SEWP 276; LBJL 32605 Chamaecrista nictitans (L) Moench, LB 32461, 32179, 32486, 32806 Crotalaria sagittalis L, SW 65; LB 32488 esmodium ciliare (Muhl. ex Willd.) DC., SEWPLB 497 Desmodium glabellum (Michx.) D 2337 yth Galactia volubilis (L) nom LB 32489 Glottidium vesicarium (lacq.) Harper, SEWP 43 *K a striata (TI b) Schlindler, LBSH 32128 656 *Lespedeza cuneata (Dum-Cours.) Harper, SEWP 348 Lespedeza virginica (L.) Britton, LB 32808 *Medicago lupulina L., LB 31941 Mimosa end (Small ex Britton & Rose) B.L. Turner, LBSH 3211 Rn latifolia Nutt., LBRH 3 Strophostyles umbellata (Muhl. ex uS Britton, SEWP 347; LBSH 32118; LB 32809 Stylosanthes biflora (L.) Britton, Sterns, € Poggenb., ~ 235 *Trifolium campestre Schreb., MM 7151; LBSH 3215 *Trifolium incarnatum L., SW 124 *Trifolium lappaceum L., LB 32172 Wisteria frutescens (L.) Poir., LB E D. *Wisteria sinensis (Sims) Sweet, (EN *Vicia sativa L. subsp. nigra (L.) len LB 32002; MM 7605 = > Fagaceae Castanea pumila (L.) Mill, SEWP 408; LB 33021 Fagus grandifolia Ehrh., SW 09; MM 7075 Quercus alba L., SW 06; MM 7077 Quercus falcata Michx, SW 52; MM 710 Quercus | Bartr. ex Willd., an 54, 55; LB 33028; MM Quercus laurifolia Michx., SEWP 344 Quercus margaretta Ashe ex Small, SEWP 424 Quercus michauxii Me ~ 7229; SW 44a Quercus nigra L, MM 7 Quercus phellos L., E. 484 Quercus velutina Lam., LBRH 32084 Gentianaceae +Bartonia texana Correll, SEWP 340; LB 33010 *Centaurium pulchellum (Sw.) Druce, SW 108 Sabatia campestris Nutt., LB 32169 Geraniaceae Geranium carolinianum L., WP 118 Grossular Itea a L SEWP 403a; LB 32024, 32/84; MM 7660 Haloragaceae Proserpinaca palustris L., SEWPLB 489 Hammamelidace Hamamelis Kee = MM 7054 Hamamelis virginiana L, MM 7217; GW 2289 Liquidambar styraciflua L, MM 7053 Hyperic Gd Ge (Grev. € Hook) Torr. & A. Gray, GW 2259; LBJL 32617 Hypericum galioides Lam., MM 7207; GW 2257 Hypericum gentianoides (L.) Britton, Sterns, € Poggenb., LB- SEWP 31896; SW 95 Hypericum hypericoides (L) Crantz, SW 73; MM 7220; LBRH 32063; LBJL 32599 Hypericum mutilum L., SW 66, 25 Triadenum walteri (J.G. Gmel.) Gleason, SW 35, 85 Iridaceae isyrinchium campestre E.P. Bicknell, SEWP 543 Sisyrinchium sagittiferum E.P. Bicknell, LB 31952 Sisyrinchium rosulatum E.P. Bicknell, LB 32011; MM 7684, /680; SW 114 ias aquatica (Michx. f.) Nutt., SEWPLB 495 Carya texana Buckley, SEWP 412 Carya tomentosa (Lam. ex Poir) Nutt., SW 28 Juncaceae Juncus acuminatus Michx., LB 32005; LBRH 32069; LBSH 32142a Juncus anthelatus Ge R.L. Brooks, LB 31997 Juncus coriaceus Mack, Juncus diffusissimus Get Si 32070, 32077; LBSH 32125; Juncus effusus L, LB 31938 Juncus marginatus Rostk, MM 7130; LBRH 32049 Juncus repens Michx., SEWP 355 Juncus scirpoides Lam., e 32799 Juncus tenuis Willd., LB 32020 Juncus validus Coville, MM 7167; LBRH 32088 Luzula bulbosa (AW. Wood) Smyth, LB 31935 Lamiaceae Hyptis alata (R , SEWP 404 Lycopus rubellus Moench, SEWP 398 Lycopus virginicus L, MM 7227; 80; LB 32797 Monarda citriodora Cerv. ex Lag., MM 7203 Monarda punctata L. var. punctata, LB 32492 *Perilla frutescens Se Britton, SEWP 267 Prunella vulgaris 0 Pycnanthemum e Torr. €: A. Gray, SEWPLB 499 Pycnanthemum tenuifolium Schrad., LB 32176 Salvia lyrata L., MM 7096 Scutellaria elliptica Muhl. ex Spreng., SEWP 239; LB 32013 Scutellaria integrifolia L, LBRH 32043 tachys crenata Raf., LB 32008 Teucrium cubense Jacq., SW 69; MM 7157 Trichostema dichotomum L., SW 77; LB 33020 feel DS Lauraceae Persea borbonia (L.) Spreng., SW 08, MM 7088 Persea palustris (Raf) Sarg., SEWPLB 480; LB 32785 Sassafras albidum (Nutt.) Nees, SEWP 241 iliaceae ae curtissii Rose, SW 39, 97, 82; LBJL 32594 Lilt ium EE Por, LBSH 32136 (L) Britton, MM 7216; LB 33022 = Linaceae C (Pla cl » Fer | ald, MM 7142 Linum striatum Walter, MM 7174; LBSH 32146 Loganiaceae Gelsemium sempervirens (L.) Aiton f, SW 07; MM 7067; LB 33008 Mitreola petiolata (J.F. Gmel) Torr. & A. Gray, SW 75; LB 32802 Lygodia "een p (Thunb. ex Murray) Sw., SW 87; MM Lythraceae *Cuphea carthagenensis (Jacq.) J.F. Macbr, SW 30; WRC 11400 Brown et al., Vascular fl it, Big Thicket *Cuphea glutinosa Cham. & Schlecht, LB 3246 Rotala ramosior (L.) Koehne, LB 32789 Magnoliac Magnolia ree L, MM = Magnolia virginiana L., SW 1 Malvaceae Sida rhombifolia L., MM 7171 Melastomataceae Rhexia mariana L., MM 7149 Meliac geg ee? rach L, SEWPLB 504 Moraceae Maclura pomifera (Raf) C.K. Schneid., GW 2296 Morus rubra L., SEWP 266; MM 7134 Myricaceae Morella caroliniensis (Mill) Small, SW 21; MM 7090; LBSH 3212] Morella cerifera (L.) Small, LB 32790 Chionanthus virginicus L., SEWP 409 Fraxinus americana L., MM 7213, 7063; GW 2252; SW 113 Fraxinus caroliniana Mill., SEWP 361; LB 32786 Fraxinus pennsylvanica Marsh., LB 32795 igustrum sinense Lour., MM 7146 Onagraceae Ludwigia alternifolia L, SEWP 261, 236; LB 32168 Ludwigia leptocarpa (Nutt) Hara, SEWP 358 Ludwigia pilosa Walter, SEWP 406; LB 32796, 33017 Ludwigia palustris (L.) Elliott, LB 31993 Oenothera biennis L., SEWP 392 Oenothera laciniata Torr. & A. Gray, LBRH 32052; SW 106 Oenothera linifolia Nutt., LBRH 32056 Ophioglossace Botrychium Se (Sav.) Underwood, SW 78 Orchidac an praecox (Walter) S. Watson., SW 41; LBRH 32066, 81 Spiranthes vernalis Engelm. €: A. Gray, SW Tipularia discolor (Pursh) Nutt., MM 7219, id LBJL 32598 Orobanchaceae Epifagus virginiana (L.) W. Bart, SW 90 Osmundaceae Osmunda cinnamonea L., SW 2 Osmunda regalis L. eut (Willd.) A. Gray, SW 13 Oxalidaceae Oxalis corniculata L.var. wrightii (A. Gray) Turner, MM 7095; LB 31951; SW 117 *Oxalis debilis Kunth var. corymbosa (DC.) Lourteig, MM 7664 Passifloraceae Passiflora lutea L., MM 7060 657 Phytolaccaceae Phytolacca americana L, MM 7110 Pinaceae Pinus echinata Mill., SEWP 437 Pinus taeda L., MM 7093 Plantaginaceae Plantago virginica L., MM 7094; LBRH 32079 Platan Sane MS L., SEWP 363 Poaceae Agrostis eee (Walter) Britton, Sterns & Poggenb., LBRH 32085; MM 7675 o 5 cayophyle L. var. capillaris | inde id 32054 rns, & Poggenb,, MM 71 96. Andropogon L, SW 101 ECH purpurascens Poir. var. purpurascens, MM 7191; LB 41 hundana gigantea (Walter) Mi Ad de 72 Idi) K 32076 Celarier & Harlan, LB 3301 8 *Briza minor L., SW 62; MM 7152 *Bromus catharticus Vahl, LBRH 32072; MM 7658 Chasmanthium laxum (L.) H.O. Yates, MM 7104 Chasmanthium sessiliflorum (Poir) H.O. Yates, SEWPLB 498 *Cynodon dactylon (L.) Pers., SEWP 246 Dichantheli iculare (D Poir) Gould & C.A. Clark var. (Fish. & Mey) aciculare, LB 32459, 33000 Dichanthelium acuminatum (Sw.) Gould & CA Clark var. acuminatum, LBRH 32089; LBSH 32142 Dichanthelium ~ (L) Gould & C.A. Clark, MM 7165; LB 31990, 3218 tatum LBSEWP 31891; LB 3194 ag Dichanthelium dichotomum (L) Gould subsp. dichotomum, SEW : MM 7137, 7071; LBSH 32134 Dichanthelium dichotomum (L) Gould subsp. microcarpon (Muhl. ex Elliott) Freckman & Lelong, LBRH 32083; LB 171 PE e ln L VI | commu- Dichanthelium dichotomum (L.) Gould ~ roanokense Ashe) Freckman & Lelong, LBSH 321 mm laxiflorum (Lam) Gould, ue 31890 ) Gould var. oligosanthes, LB 3201 8, 32471; LBRH 32064 It) Mohlenl M 7136 Dichanthelium scabriusculum (Elliott) Gould S C.A. Clark, Dichanthelium scoparium (Lam) Gould, MM 7218; LB 33005 Faral el Jr LIA DI | q, | DD) 32044; LBSH 32138, 32123a, 32121a Digitaria ciliaris (Retz.) Koel, GW 2330; LB 32476, 32458 *Digitaria ischaemum (Schreb.) Se MM 7232 *Eleusine indica (L.) eb mE Elymus virginicus L, LB 3 Eragrostis hirsuta und less LB 33040 Eragrostis refracta (Muhl.) Scribn., MM 7230 Eragrostis spectabilis (Pursh) Steud., LB 33025 — ~~ 658 Hordeum pusillum Nutt., LB 3201 Leersia virginica Willd., LBSEWP ep 892; MM 7195 *Lolium perenne L., LBRH 32073 Melica EES Walter, MM 7168 t L) P. Beauv. subsp setarius (l YM - Ekman, SEWPLB 483 Panicum anceps Michx., LB 32485 Panicum brachyanthum Steud., MM ee Panicum rigidulum Bosc ex Nees, SEWP 396 Panicum verrucosum Muhl., ipi E ~ ae 1402; LB 33006 *Paspalum dilatatum Poir., LB 3 Paspalum langei (E. Fourn.) er ae 494 *Paspalum notatum Fluegge, MM 7158 Paspalum plicatulum Michx., SW 64; MM 7190 Paspalum setaceum Michx., LB 32478 *Paspalum urvillei Steud., MM 7202; SW 70; GW 2328 Phalaris caroliniana Walter, LBSH 32144 Piptochaetium avenaceum (L.) Parodi, LB 31992 *Poa annua L., WP 117 BCE indica (L.) Chase, LBSEWP 31897; SW 04; WRC HC ml D q ft | Larra t, LB 32003 Schizachyrium scoparium (Michx) Nash, LB 32803 Sphenopholis intermedia (Rydb) i Rydb., LB a MM 7656 ibn., LBSH 32145 Sporobolus indicus (L.) R. Br, SW 102; GW e Tridens flavus (L.) Hitchc. var. flavus, SW 103 Vulpia octoflora (Walter) Rydb., LBRH 32057 Polygalac Ania mariana Mill., LBJL 32615 Polygala polygama L., LBRH 32045 Polygonacea Brunnichia ovata namen Shinners, SEWP 402a Polygonum punctatum Elliott., S 485 Rumex hastatulus Baldw., LBRH 32055 Polypodiaceae Pleopeltis polypodioides And & Wind! bsp. michauxi- ,GW 53; MM 7105; SEWP 402b; SW 81 Primulaceae *Anagallis arvensis L., MM 7657 Samolus valerandi |. subsp. , 31888 florus (Raf) Hulten, LBSEWP Rhamnaceae Berchemia scandens (Hill) K. Koch, MM 7057 Frangula caroliniana (Walter) A. Gray, MM 7082; LB 33026 Ros Crataegus iid Pope MM 7156 ex Hook, LBSH 32135 SC spathulata VEM MM 7156, 7064 *Duchesnea indica (Andr Focke, MM 721 5 Prunus caroliniana (Mill) Aiton, MM 7068 Prunus serotina Ehrh., SEWP 426 Rubus argutus Link, SEWPLB 482 Rubus trivialis Michx., MM 7144 eae Cephalanthus occidentalis e ~ 7154 Diodia teres aner, pu Diodia Galium pilosum d pie LB 492 Houstonia pusilla pus LB 31933 Mitchella repens L., MM 7066 Oldenlandia boscii Bo Chapm., SEWP 364; GW 2256; LB 3194 Misi soi L., SEWPLB 486; LB 32810, 32999 *Richardia s , SEWP m MM 7682, SW 107 Keen arvensis L., WP Stenaria nigricans (Lam.) ei SEWP 233 Saururaceae Saururus cernuus L, MM 7132 Scrophulariaceae Bacopa monnieri (L.) Pennell, SEWP 243; MM 7681; SW 11 Gratiola pilosa EE e e Gratiola nl crustacea e : Muel, D 33014 Lindernia dubia (L.) Penn., LB 3 iris acuminata ES m SW 76; MM 7223; PM LB 32788 ee umbrosum (J.G. Gmel.) Blake, SEWP 352; MM 7667 Mimulus alatus Aiton, LB 32787 m texana (Scheele) Sutton, LB 31944; | BRH TN Se L, MM 7221; e 79; GW 3360a, 2254 *Veronica arvensis L., LB 319 Smilaca Smilax man -nox L., SEWP 268 Smilax glauca Walter, MM 7145 Smilax laurifolia L., SEWP 417; SW 24 Smilax pumila Walter, MM 7115, 7070 Smilax rotundifolia L., SEWP 274 Smilax smallii Morong, MM 7086 Solanaceae Solanum carolinense L., MM 7173; LBRH 32068 Symplocaceae Symplocos tinctoria (L.) L. Her, MM 7116, 7113 Styraceae Halesia diptera Ellis, SEWPLB 505 Styrax americanus Lam., SW 04 Thelypteridac Thelypteris Ge Can CV. Morton, LB 32480 Ulmaceae Celtis laevigata Willd., GW 2335 Michx., SW 56; GW 2334 Ulmus americana L., SEWPLB 493 Urticaceae Boehmeria cylindrica (L.) Sw., MM 7208; SW 37; LB 3021 Valerianacea Valerianella ae (L) Dufr, WP 115 E al nm LF Brown et al., Vascular fl k Unit, Big Thicket - Valerianeila radiata (L.) Dufr. f. parviflora (Dyal) Egg. Ware 3205 naceae allicarpa americana L., MM 7084 *Glandularia pulchella (Sweet) Troncoso, SEWP 256 GA nodiflora (L ) Greene, SW 9 bena oras“ iensis Vell., SEWP an Verbena halei Small, MM 7143 *Verbena rigida Spreng., SEWP 257 Violaceae Viola lanceolata L., MM 7092; LB 33013 Viola palmata L, MM 7159; GW 2295; PM 05 659 Viola primulifolia L., SW 1 E Viola sagittata Aiton, GW 2 Viola sororia Willd., GW SE E 31986 Vitac Moss arborea (L.) Koehne, SW 2 Parthenocissus quinquefolia (L LE SCH 250 Vitis aestivalis Michx., MM Ge Vitis rotundifolia Michx., MM 705 Xyridac p fr cl var. difformis, LBSEWP 31893, 31898; 7204, a e Rich., SEWP 419, 354, 328; LB 32462 ÁDDENDUM J Isotria verticillata (whorled pogonia) and sugar maple Ge DER SE m or Ácer leucoderme) are listed for the unit on the Beechwoods Trail Guide, a e at the trail entrance, but ve failed to collect them during our two- year field survey, dud Seef not find herbarium material of same. Geraldine Watson (2006) also indicated that sugar maple (Acer barbatum) is making a remarkable recovery in this unit with the removal of cattle grazing. NOTES Carex section ovales. The specimen in this collection was too immature for specific identification. The only other Carex in section ovales was C. brevior. Perhaps this was an immature C. brevior. Bartonia texana. Currently Beech Creek is the only Big Thicket unit to have this rare Texas endemic. We made two collections, one in a sphagnum mat along the Beechwoods Trail, and one far from any trail, near the intersection of Beech Creek with Little Beech Creek. Bradburia pilosa. We are following Semple (2006). An equally correct name is Chrysopsis pilosa Nutt. Rhynchospora debilis. This is the third collection for Texas and the first for the Pineywoods. See Brown et al. (2007) for additional information. Xyris jupicai. There is no consensus as to the native or introduced status of this yellow-eyed-grass. DISCUSSION There are 103 families and 229 genera represented for the 470 taxa (418 native) on the Beech Creek Unit list. Eight families: Poaceae (59 taxa), Asteraceae (53 taxa), Cyperaceae (40 taxa), Fabaceae (26 taxa), Lamiaceae (15 taxa), Euphorbiaceae (13 taxa), Scrophulariaceae (11 taxa), and Juncaceae (11 taxa) account for 48.9% of the taxa collected. The largest genera are: Carex (15 taxa), Dichanthelium (13 taxa), Rhynchospora (11 taxa), Juncus (10 taxa), Quercus (9 taxa), and Cyperus (7 taxa). While it is the case that our list is i plete, as are all floras, and that a few taxa reported here may no longer exist on the unit, specimens of most taxa that have grown in the Beech Creek Unit during the past quarter century have been collected (probably 954 96). We estimate that the Beech Creek Unit has about 425 to 450 native species/taxa. — ACKNOWLEDGMENTS This study was supported in part by a National Park Service Cooperative Agreement No. CA 14001004 to Paul Harcombe and in part by an All Taxa Biological Inventory (ATBI) grant to the first author from the Big Thicket Association of Saratoga, Texas. The page charges were paid for from this grant. The first author appreciates Deanna Boensch and Lily Zhou, Big Thicket fire ecologists, for facilitating his passage across private land to the northeast corner of the unit and then walking with him along Little Beech Creek south 660 | t tanical Insti Texas 2( to near the confluence with Beech Creek. Eleven new additions to the species list were collected on this all-day hike. Thanks also to Ray C. Telfair Il and Eric Keith for many helpful suggestions derived from their careful reading of the paper. REFERENCES AJiLvsai, 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, W.W. Pruess, I.S. ELsix, 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. MacRogerts, M.H. MacRoserts, PA. Harcomse, W.W. Pruess, 1.5. Das, and S.D. Jones. 2006a. Annotated checklist of the vascular flora of the Big Sandy Unit of the Big Thicket National Preserve, Polk County, Texas. Sida 22:705-723. Brown, LE B.R. MacRoserTs, M.H. MacRoserts, PA. Harcomee, W.W. Pruess, I.S. Er sik, and S.D. Jones. 2006b. Annotated checklist of the vascular flora of the Lance Rosier Unit of the Big Thicket National Preserve, Hardin County, Texas. Sida 22:1175-1189. BROWN, LE E.L. Kern, DJ. Rosen, and J. Liccio. 2007. Notes on the flora of Texas with additions and other significant records. J. Bot. Res. Inst. Texas 1:1255-1264 DEsHoTELS, J.D. 1978. Soil survey for the Big Thicket National Preserve, Texas. USDA Soil Conservation Service, College Station, Texas. Dicas, G.M., B.L. Lipscome, M.D. Reep, and R.J. O'Kennon. 2006. Illustrated flora of east Texas. Sida, Bot. Misc. 26:1-1594. Harcomee, PA. and PL. Marks. 1979. Forest vegetation of the Big Thicket National Preserve. Unpublished report: U.S. Park Service, Santa Fe, New Mexico. Harcomee, PA., J.S. GLITZENSTEIN, R.G. Knox, S.L. OrzeLL, and E.L. Bripces. 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. Weer, and PM. Montcomery. 1997, Vascular plants EE Univ. Texas Press, Austin. KARTESZ, J.T. 1999, A synonymized checklist and atlas with biol for the vascular flora ofthe United States, Canada, and Greenland. 1st ed. In J.T. Kartesz and CA Meacham. Synthesis of North American flora. Version 1.0. North Carolina Botanical Garden. Chapel Hill. MacRoserTs, B.R., M.H. MacRoserts, 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, P.L. and PA. HarcomBe, 1981. Forest vegetation of the Big Thicket, southeast Texas. Ecol. Monogr. 51:287—305. Peacock, H.H. 1994, Nature lover's guide to the Big Thicket. Texas A&M Press, College Station. SempLe, J.C. 2006. Bradburia. In: The flora of North America. Vol. 20. Oxford University Press. New York. TURNER, B.L., H. NicHots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas, Vol. 1: Dicots, Vol. 2: Ferns, , monocots. Sida, Bot. Misc. 24. NM K EY, JE 2006. The pineywoods. In: G.M. Diggs, B.L. Lipscomb, M.D. Reed, and RJ. O'Kennon, Illustrated flora of east Texas. Sida, Bot. Misc. 26:1-1594. Pp. 76-106. Watson, G.E. 1979. Big Thicket plant ecology: an introduction. Big Thicket Mus. Publ. Ser. No. 5, Saratoga, Texas. Watson, G.E. 2006. Big Thicket plant ecology, an introduction, 3rd ed. University of North Texas Press. Denton, Texas. WATSON, G.E. 1982. Vegetational survey of Big Thicket National Preserve. Mi graphed unpublished report. Big Thicket National Preserve, Beaumont, Texas. EARLY HISTORICAL REFERENCES TO THE BIG THICKET, TEXAS 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 W, f 1860 ref to the Bi g Thicket All place the Big TI icket the San Jacinto River west or at the western edge ep where edern accounts place it. Two d , E d RESUMEN Citamos cuatro lc anteriores a 1860 del Big Thicket. Todas colocan el Big Thicket en o cerca de San Jacinto River al oeste o en los modernos. Dos lo describen como un matorral, y otros dos como bosques abiertos There is much disagreement on what should be called the Big Thicket (Cozine 2004; Diggs et al. 2006). The broadest conception has it encompassing about 3.5 million hectares in an area extending from the Brazos to the Sabine rivers and from Nacogdoches to Beaumont. The narrowest conception is the “Hunters Thicket” encompassing only 130,000 hectares mostly in Hardin County. Today, the most generally ac- cepted view corresponds more or less to a compromise between the Biological Survey definition of Parks and Cory (1938) and the Ecological Area definition of McLeod (1971). In this view it encompasses more or less Jasper, Newton, Tyler, Hardin, Polk, San Jacinto, Montgomery, and northern Liberty counties: about 1.8 million hectares (see Diggs et al. [2006] for a detailed discussion of literature and maps of the various locations of the Big Thicket). Del Weniger, the eminent ecological historian of Texas, argued that the phrase “Big Thicket” was not old but dated from only the mid-nineteenth century and that today's location of the Big Thicket is not where it originally was (Weniger 1984). The two earliest (pre-1860) accounts he was able to find which used that phrase were by Don E.E. Braman and J. De Cordoba, from 1857 and 1858, respectively. These accounts place the Big Thicket on the San Jacinto River or between the two forks of the San Jacinto River in what are now San os TOM MEN and pes Me counties, to Aa west or at dps western edge of where modern te it (county | ange bly since the 1850s) (Fig. 1). These two sources do not describe the Big Thicket as an SEM forest but as a highly desirable region of mostly large, dense timber with fertile and productive soil that was already extensively settled by the late 1850s. Diggs et al. (2006) recently described a third pre-1860s account of the Big Thicket. This is in A Stray Yankee in Texas, published in 1853 by Philip Paxton (Paxton 1853). As Diggs et al. (2006) only briefly men- tion this reference, additional comment seems warranted. Philip Paxton (a pseudonym for Samuel Adams Hammett, 1816-1865) was born in Connecticut and moved to Texas in 1835 where he remained until 1848 working in and around Montgomery and Houston. Upon his return east, he wrote serious and humorous magazine articles about Texas. In 1853 he published these as A Stray Yankee in Texas (Hoole 2007). Paxton's account of the Big Thicket is interesting for two main reasons. First, he places the Big Thicket on the San Jacinto River, which, as in the accounts of Braman and De Cordoba, is west of or on the western edge of where current descriptions locate it. Second, unlike Braman and De Cordoba, Paxton describes the Big Thicket as a thicket. According to him, the Big Thicket, while having "the finest timber in the world" is *a thicket so dense that even in that Eon. of uM forest, it is known—par excellence—as the big thicket...." He continues: “This spot had been aptly named the “big thicket.' Immense bamboo briers, like vegetable Pythons, | nie Das Inst. Texas 2(1): 661 — 663. 2008 fall Dos H Inm LI rr iT. a TE h A d e e e Newton 4 ^ Washington "€ s” ` 1 " H Fd KN Te m. S pm A IN Colorado 4 ,^ i E v SN q % A Ki a ^. T F e Vë Kc pti Ki ^^ "o Wharton ^ e | "d BR e " be * 4 j q Y * ^ Victoria |; i Matagorda . 1 `, Gulf of Mexico : ee on M ot | | j Calhoun à p^ à Ka P a4 Refugio "f Fiz 1 E L LT L a, "2/701 me E sl Big TI . a FL FEN | J 1 | a. dd | AN E al nº hg! | Im i | 1.1 J ) (Fig lified from Diggs et al. 2006, Figure 8) twined and intertwined, crossed and recrossed, from tree to tree, and shrub to shrub, forming a natural trellis-work for the thousand and one wild and beautiful vines that abound there.” Further, he writes: “The first part of our journey was effected on horseback; but after proceeding some half a mile into the ‘timber, this mode of progression was suddenly brought to a period by the dense undergrowth, and we were reduced to a very natural and primitive style of locomotion.” Finally, he adds: “We now had to contend for every step we gained; knife and hatchet were in constant requisition, and for one hour we pressed on in Indian file as fast as we could.” Paxton describes or refers to such plant habitats as “cypress-brakes,” “palmetto swamps,” “cane-brakes,” “sloughs,” and “prairies.” He mentions individual plants: “magnolia grandiflora,” “bamboo,” 3 tt ant i AAR. DL ts, Hist salen tec Big Thid di 663 » c E 231 LL “passion vine,” “cypress vine,” “morning-glory” “trumpet-creeper,” “cypress,” “hanging-moss” (Spanish moss), and “cane,” and animals: “log-cock” (Pileated woodpecker), “paroquet” (Carolina parakeet), “mocking-bird ” “moccasin snake,” “alligator,” “bear,” and “fish.” With the aid of Parker’s bibliography (1977), we located another early account. This occurs in an article “The Big Thicket Ruins” in The Texas Republic, 3 May 1856. The relevant passage reads: “A man by the name of Smith, reported to the Washington American about the First of April, that he had recently discovered in the Big Thicket on the San Jacinto some twenty-five miles from Montgomery, an ancient ruin....” (italics in original). The article goes on to describe these ruins as being in “a dense thicket almost impenetrable.” Thus, all four of the earliest known references to the Big Thicket place it on the San Jacinto River or between the forks of the San Jacinto River on the western edge of the generally accepted conception of the Big Thicket (Diggs et al. 2006), or as Weniger (1984) stated: “this places the Big Thicket in the area of present Montgomery and San Jacinto counties, roughly between Conroe, Cleveland, Coldspring, and Huntsville,” which is basically what is today the Sam Houston National Forest. To our knowledge, there are no early accounts that place the Big Thicket in Hardin, Tyler, Jasper, Newton, Liberty, or Polk counties or on the Sabine, EEN or Trinity rivers where PES scholars bis it and where the Big Thicket National 1. But, both Paxton’s and Smit] t t Weniger's finding that the Big Thicket was originally described as an open and hospitable land. Both accounts clearly describe the Big Thicket as virtually impassable. Why and when the phrase “Big Thicket” came to be applied to tl tern region ins unknown. If anyone knows of other pre-1860 references to the Big ‘Thicket or has information on the evolution of the concept of the “Big Thicket,” we would appreciate hearing from them. ACKNOWLEDGMENTS We appreciate the reviews by Guy Nesom and George Diggs. REFERENCES Cozine, J. 2004. Saving the Big Thicket: From exploration to preservation, 1685-2003. Univ. North Texas Press, Denton. Dices, G.M., B.L. Lipscoms, M.D. Reco, and R.J. O'Kennon. 2006. Illustrated flora of east Texas. Sida, Bot. Misc. 26:1—1594. Hoote, W.S. 2007. Hammett, Samuel Adams. The handbook of Texas online. www.tsha.utexas.edu/handbook/ online/articles/HH/tha42.htm| McLeon, CA 1971. The Big Thicket forest of East Texas. Texas J. Sci, 23:221-233. Parker, L.W. 1977. The Big Thicket of Texas: a comprehensive annotated bibliography. Sable Publication, Arling- ton, Texas. Parks, H.B. and V. Cory. 1936. Biological survey of the East Texas Big Thicket area. Texas Agric. Exp. Sta., College Station. Paxton, P. 1853. A stray Yankee in Texas. Redfield, New York. Wenicer, D. 1984. The explorers’ Texas: the lands and waters. Eakin Press, Austin. 664 [| [| fal Dos H In Li rr Er ms, 147 BOOK NOTICE WAYNE A. SINCLAIR AND HOWARD H. Lyon. 2005. Diseases of Tress and Shrubs, Second Edition. (ISBN 978- 0-8014-4371-8, hbk.). Cornell every Press, Sage House, 512 East State Street, Ithaca, New York 14850, U.S.A. (Orders: ww cornell.edu, 1-607-277-2211, 1-800-688-2877 fax). $89.95, 660 pp., color plates, companion CD- ROM, 9 1/4" x 12 3/8". Contents: About this book How to use this book Dedication and acknowledgments Diseases caused by fungi " : Et 1 e hliehe ] o 14] 1 Cr Ve: fan] J ASA. Diseases caused | A (Foli d q At haa] ; & , , Root diseases; Systematic es) Diseases caused S E (Foliar diseases; Rusts; Trunk and limi : Canker-rots; Root diseases) Diseases caused by Oomyco pu diseases Viral diseases Diseases caused by nematodes Plant- Ee algae a plants Declin Declines Damage by toxic agents Mineral deficiencies Damage by drought, heat, freezing, excess water Miscellaneous injuries and disorders Restoration of OAM and ini after i RA or infection 1 “11 d plants VV SELL VÀ ao References Index J. Bot. Res, Inst. Texas 2(1): 664. 2008 THE BIG THICKET OF TEXAS AS FLORISTICALLY UNIQUE HABITAT 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 The Big Thicket is floristically closely related to tl ] United Stat Using Species that are dues to the Big Thicket region A À zw = al 11 I in Texas, we find d e most are eH adapted The Big T y , geologically, phys- ; 1 J Tue luci AA + > 1 1 4 4 i lii iographica y, Lë . r5 Lal DÉI : ] à dies ERES H +h: limit in southeastern Ti RESUMEN El Gran Matorral está muy relacionado A. con a sureste de los o Unidos. Usando especies que son únicas en la región del Gran Matorral en Texas, | rí aptadas a la humedad. El Gran Matorral es el área más occidental de Texas y tiene continuidad edáfica, See, Falo grafica, y imd ia con el sureste de Los Estados Unidos. Muchas es- pecies orientales tienen su límite de distribución en el sureste de Texas debido a la alta precipitación y condiciones de humedad que caracterizan esta área. INTRODUCTION The Big Thicket has been described as an area where many floras converge (see MacRoberts & MacRoberts 2004, 2007; Diggs et al. 2006 for discussion and literature). This characterization has been questioned (MacRoberts & MacRoberts 2004a; MacRoberts et al. 2007). The affinity of the Big Thicket flora is almost entirely within the eastern and especially southeastern U.S.A. (Diggs 2002; Diggs et al. 2006; MacRoberts & MacRoberts 2003, 2004a, 2007; McLaughlin 2007) The present study originates from a statement in Diggs et al. (2006:171): “While clearly part of the southeastern U.S., the Big Thicket represents the southwestern extreme of the Southeastern Mixed Forest Province and the Outer Coastal Plain Mixed Forest Province.... Hundreds of species that occur in an arc from Virginia or North Carolina south to Florida and west to Texas reach the southern or southwestern limit of their ranges in the Big Thicket, or are confined or largely confined in Texas to the Big Thicket area. The reasons for their occurrence in the Big Thicket include the high rainfall and overall mesic conditions and the presence of specialized habitats. . As part of a continuing effort to describe and understand the flora of the Big Thicket region (Parks & Cory 1936; Watson 1975; Ajilvsgi 1979; Marks & Harcombe 1981; Bridges & Orzell 1989; Harcombe et al. 1993; MacRoberts & MacRoberts 2004a; MacRoberts et al. 2002, 2007; Brown et al. 2005, 2006a, 2006b, in press; Diggs et al. 2006), we concentrate in this study on what is unique floristically to the Big Thicket region in Texas and how this flora relates to abiotic conditions. THE BIG THICKET REGION Na 4 E 11 1 D Lt: 11 1 ] Southeastern Texas (in which the Big Thicket ) y, geologically, and phy y related to and continuous with the remainder of eastern Texas and western Louisiana (ennenen 1938: Godfrey et al. 1973; Groat 1984; Deshotels 1976, Aronow 1981; Bureau of Economic Geology 1992; Shelby et al. 1992; and Diggs et al 2006). What is unique to the Big Thicket region within Texas is its humid subtropical climate (Bomar Atl SEET is receives more rainfall than any other area of Texas, northwestern Louisiana, A g. 1) (Owen & Schmidly 1986; Bomar 1995; Goins & Caldwell 1995; Diggs et al. 2006). Rainfall ranges hom: 122 to 152 cm, making southeastern Texas the westernmost area J. Bot. Res. Inst. Texas 2(1): 665 — 671. 2008 666 Jo ti tanical Insti Texas 2(1) 97 Fic. 1. A of the eastern United States to receive such large amounts of precipitation (Diggs et al. 2006). Additionally, precipitation is well distributed through the year, the difference between driest and wettest month being only a few centimeters (Bomar 1995; Diggs et al. 2006). The only significant variation in elevation in the Big Thicket region is connected to the Willis and Fleming formations to the north where elevation rises to 110 meters and where water runoff is fairly rapid. To the south on the Lissie and Beaumont formations there is little or no relief and the area is poorly drained. Because of high precipitation and flat terrain, soils waterlogged, and water may stand for long periods of time in some areas, e.g., wetland pine savannas, bald- cypress sloughs, and palmetto-hardwood flats (Bridges & Orzell 1989; Marks & Harcombe 1981; Harcombe et al. 1993; Diggs et al. 2006). Additionally, because of high rainfall, the soils—Ultisols and Alfisols—are often deeply weathered. The relatively low latitude (approximately 30 degrees N) and proximity to the Gulf of Mexico result in uniformly high temperatures and a long humid growing season (Bomar 1995). South- MacRobert I MacRol ts, The Big Thicket, floristi ily ni habitat 667 eastern Texas has the highest net above-ground primary productivity in Texas (Owen & Schmidly 1986), a net primary productivity comparable to the southeastern United States. METHODS Using the plant distribution maps in Turner et al. (2003) and Diggs et al. (2006), we developed a list of 98 native species “unique” to the Big Thicket region in Texas. This list consists of all species that are either confined entirely to the Big Thicket in Texas (wherever else they occur in North America) (e.g., Spiranthes longilabris Lindl.) or that are concentrated in but not totally confined to the Big Thicket in Texas (wherever else they occur in North America) (e.g., Xyris scabrifolia Harper). Using sources available (e.g., Diggs et al. 2006; MacRoberts et al. 2002), we determined in which habitats (communities) these species occur. Some species occur in more than one community; consequently, we divided their representation between those communities. Using USDA (2007), we determined the wetland status of each species. For those with no wetland rank, we either assigned one based on our personal experience, or, if we did not know the species adequately, put them in the “unknown” category. Using Kartesz and Meacham (2005) and Flora of North America (1993-2006), we plotted the North American distribution of the species to determine their rela- tionship to other regions. In order to put the Big Thicket into a larger context, using Turner et al. (2003) and Diggs et al. (2006) we developed a random selected list of 115 native species confined to eastern Texas but that were evenly distributed over the area, not just the Big Thicket, irrespective of wherever else they occurred in North America. As with the Big Thicket list, we determined, where possible, in which habitat (communities) these species occurred and their wetland status. Using the same sources as for the Big Thicket list, we plotted their North American distribution. For the purposes of the present study, the Big Thicket is defined as Jasper, Newton, Hardin, Tyler, Polk, San Jacinto, and northern Liberty counties. This definition corresponds closely with the “biological” definition of the Big Thicket and is the same as that used by Diggs et al. (2006). Eastern Texas is defined as the area of Texas east of about 97 degrees longitude. RESULTS We found 98 species that are either unique to the Big Thicket or that are concentrated in it within Texas (irrespective of where they occur outside Texas). The community preferences, where they could be deter- mined, are given in Table 1: 65 percent of the species favor wetland communities. The wetland status of these species, where known, is given in Table 2: 67 percent are obligate or facultative wetland species. The North American distribution of these species as a percentage of the total is given in Figure 2: 96 percent occur in Louisiana, 94 percent in Mississippi and Alabama, 91 percent in Georgia and Florida, 85 percent in South Carolina, 81 percent in North Carolina, and 56 percent in Virginia, all of which are coastal states. Percentages drop off precipitously for all other states, regions, and provinces. The eastern Texas sample had 115 species: 19 percent favored wetland communities (Table 1) and 28 percent were obligate or facultative wetland species (Table 2) (the discrepancy between the two figures is probably due to the high number of unknowns and the high number of plants undesignated by USDA [2007]). The North American distribution of these species is given in Figure 3. The distribution is mainly in the South, but also the East in general. DISCUSSION In the past, the Big Thicket has been characterized as an area in which the floras of many regions converge. In a recent study (MacRoberts & MacRoberts 2007), we found that the Big Thicket is almost entirely south- eastern with a flora composed of species typical of Louisiana, Mississippi, Alabama, and the East in general, not of species typical of western or central states. The only exceptions to this are prairie species (Brown et al. 2002). However, the Big Thicket is not the only part of the West Gulf Coastal Plain that has prairies. 668 t tani Institute of Texas 2( TABLE 1. Habitat ( ity) preferences of Big Thicket and eastern Texas species. Big Thicket East Texas Community/habitat No. 96 No. 96 Bog/Wetland pine savanna 38.5 39 Ss 5 Upland longleaf pi Mixed woods 130 13 295 26 Pond/Marsh/Lake margin 13.0 13 90 8 Stream course 6.5 7 3.03 Baygall/Seepage slope 6.0 6 40 3 Beech/Wet woods 6.0 6 7.0 6 Mesic Hardwood-Pine forest 6.0 6 30.0 26 Xeric sandylands 1.0 1 9.0 8 Unknown 8.0 8 18.0 15 Total 98.0 100 115 100 En ) Ri aL, A - LXI AR ED''. Thiel 4 Prairi cattered theastern Oklahoma, southern Arkansas, western Louisiana, and eastern Texas (MacRoberts et al. 2003; MacRoberts & MacRoberts 2004b). In the present study, we have extended our earlier analysis of the Big Thicket flora by isolating species nd ining tl hysi hic, and climatic + edaphic. geologic L LEN CH Q T unique to the region in Texas a phy [am] AA naL A JAS B.L 4 , The Big Thicket, e a ye lly D [| L’ a 669 Taste 2. Wetland status of Big Thicket and eastern Texas species (see USDA [2007] for explanation of status designations). Big Thicket East Texas Status No. % No. OBL (obligate wetland) 30 EN 13 11 FACW (facultative wetland) 35 36 20 17 FAC (facultative) (intermediate) 14 14 37 32 FACU (facultative upland) (dry) 12 12 30 26 UPL (upland) ‘very dry) 5 5 13 11 own 2 2 2 2 Total 98 100 115 100 En 1 Al ah A H l 24s: TA) E A T H £ E Assel Ca? E J E 5 he uniqueness of the region. As might be expected, considering that southeastern Texas is humid subtropical and the wettest area in Texas, the plants are largely wetland species. Also not unexpected, considering that the southeastern United States has virtually the same climate as does southeastern Texas and that both areas are edaphically, geologically, and physiographically similar, plants that are unique to the Big Thicket in Texas are also found in Louisiana, Mississippi, Alabama, Florida, Georgia, Virginia, and the Carolinas, all of which are coastal states with high rainfall and thus deeply weathered soils. This conciusion is supported by our analysis of the eastern Texas plant group. While two-thirds of 670 J l [| Eal Dos H In LL) de e a f Texas 2( the species confined to the Big Thicket region are wetland species, only one-quarter of the eastern Texas group are. Additionally, the North American distribution of these species is significantly different from the Big Thicket group. Instead of hugging the southern coastal states, they are distributed across eastern North America. Further support for the thesis that the Big Thicket is characterized ee by wetland plants is e fered by Diggs et al. (2006:169), who found that the Big Thicket is relatively rich in monocots when compare to other areas of Texas. They speculate that this difference probably Ke to high percentage of mesic to wet habitats favored by many monocot species. Another indicator of the importance of precipitation in the Big Thicket is the presence of drought- intolerant American beech (Fagus grandifolia Ehrh.), which reaches its westernmost limit in the United States in Montgomery and San Jacinto counties (McLeod 1975; Diggs et al. 2000). In conclusion, southeastern Texas and thus the Big Thicket is the westward abiotic extension of the southeastern United States and thus also its westward phytogeographic extension. Many southeastern species reach their distributional limit in southeastern Texas because of its high precipitation and edaphic conditions resulting from that high precipitation. Thus, as Diggs et al. (2006) point out, the Big Thicket represents a biological boundary and is therefore important for a number of reasons. For example, populations at the margin of a species’ range are often | ble resource. Also, peripheral populations are often more unique genetically and represent an irreg sensitive to environmental change and can act as ecological indicators. More locally, the Big Thicket is important because it is a unique area within Texas, and the Big Thicket National Preserve is one of the few relatively large protected areas in the entire West Gulf Coastal Plain serving as a refuge not only for species and communities in Texas but the entire area as well. ACKNOWLEDGMENTS Thanks to George Diggs and Guy Nesom for their helpful comments. REFERENCES Awuer, G. 1979. Wild flowers of the Big Thicket, east Texas, and western Louisiana. Texas A €: M Press, College Station. Aronow, S. 1981. Notes on the geologic units: Big Thicket National Preserve. Unpublished Manuscript, Big Thicket National Preserve, Beaumont Bomar, GW. 1995. Texas weather. 2" ed. University of Texas Press, Austin. BRIDGES, E.L. and S.L. OrzeLL. 1989. Longleaf pine communities of the West Gulf Coastal Plain. Natural Areas J. Brown, LE K. HiLtHouse, B.R. MacRoserts, and M.H. MacRoseats. 2002. The floristics of Windham Prairie, Polk County, Texas. Texas J. Sci. 54:227-240. BROWN, LE B.R. MacRoserTs, M.H. MacRoserts, PA. HarcomBe, W.W. Pruess, LS ELsix, 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, I.S. ELsik, and S.D. Jones. 2006a. Annotated checklist of the vascular flora of the Big Sandy Creek Unit of the Big Thicket National Preserve, Polk County, Texas. Sida 22:705-723. BROWN, L.E., B.R. MacRoserTs, M.H. MacRoserts, PA. HARcoMBE, W.W. PRusss, l.S. ELsix, and S.D. Jones. 2006b. Annotated checklist of the vascular flora of the Lance Rosier Unit of the Big Thicket National Preserve, Hardin County, Texas. Sida 22:1175-1189. BROWN, L.E., B.R. MacRoserts, M.H. MacRoserts, PA. HARCOMBE, W.W. Pruess, l.S. Esik, and S.B. WALKER in prep. Annotated checklist of the vascular flora of the Beech Creek Unit of the Big Thicket National Preserve, Tyler County, Texas. AacRobert | MacRoberts, The Big Thicket, a floristically unique habitat 671 d d Dices, G.M. 2002. East Texas as a unique habitat. In: Native Plant Soc. of Texas. Special habitats. Pp. 53-66. 2002 Symposium Proc., Houston, lexas. Diccs, G.M., B.L. Lipscoms, M.D. Been and RJ. O'Kennon. 2006. Illustrated flora of east Texas. Sida, Bot. Misc. 26:1-1594, FENNEMAN, N.M. 1938. Physiography of eastern United States. McGraw-Hill, New York. FLORA OF Nort+ AMERICA EDITORIAL Committee (eds.). 1993-2006. Flora of North America North of Mexico. Vols. 1, 2, 3, 4, 5, 19, 20, 21, 22, 23, 25, 26. Oxford Univ. Press, New York and Oxford. Gobrrey, C.L., G.S. McKee, and H. Oakes. 1973. General soil map of Texas. Texas Agric. Exp. Sta., College Station. Goins, C.R. and J. M. CaLoweLL. 1995. Historical atlas of Louisiana. Univ. Oklahoma Press, Norman. Groat, G.G. 1984. Geologic map of Louisiana. Louisiana Geological Survey, Baton Rouge. Harcomee, PA, J.S. GurzeNsrEIN, R.G. Knox, S.L. OrzeLt, and E.L. Brioces. 1993. Vegetation of the longleaf pine region of the West Gulf Coastal Plain. Proc. Tall Timbers Fire Ecology Conference 18:83- 104. Kartesz, J.A. and CA MEACHAM. 2005. Synthesis of North American flora. Version 2.0. North Carolina Botanical Garden. Chapel Hill. MacRoserts, B.R, M.H. MacRober7s, and LE 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. MacRoserts, M.H. and BR MacRoserts. 2003. The east-west transition of flora in Texas: a biogeographical analysis. Sida 20:1693-1700. MacRoserTs, M.H., B.R. MacRosenrs, 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. Pp. MacRoserts, M.H. and B.R. MacRoserts. 2004a. The Big Thicket: typical or atypical. East Texas Hist. Assoc. 42:42—5]. MacRoserts, M.H. and B.R. MacRoserts. 2004b. West Gulf Coastal Plain prairies: A first approximation at a synthesis. In J. Randall and LC Burns, eds. Proceedings of the Third Eastern Native Grass Symposium. The North Carolina Botanical Garden, Chapel Hili. Omnipress, Madison, Wisconsin. Pp. 5-21. MacRoserts, M.H., B.R. MacRoserTs, and R.G. Kauwskv. 2007. Vascular plant species/area relationships (species rich- ness) in the West Gulf Coastal Plain: A first approximation. J. Bot. Res. Inst. Texas 1:577—583. MacRoserTs, M.H. and B.R. MacRoserts. 2007. Phytogeography of the Big Thicket, east Texas. J. Bot. Research Inst. Texas (in press). Marks, PL. and PA. Harcomee. 1981. Forest vegetation of the Big Thicket, southeast Texas. Ecol. Monogr. 51:287-305. McLAUGHLIN, S.P. 2007. Tundra to tropics: the floristic plant geography of North America. Sida, Bot. Misc. McLeop, CA 1975. Southwestern limit of Fagus grandiflora Ehrh. Texas J. Sci. 26:1/9-184. Owen, J.G. and DJ. ScamibiY. 1986. Environmental variables of biological importance in Texas. Texas J. Sci. 38:99-117. Parks, H.B. and V. Cory. 1936. Biological survey of the east Texas Big Thicket area. Texas Agric. Exp., Sta, College Station. SHELBY, C.A., M. K. McGowen, S. ARcNow, W.L. Fisher, L.F. Brown, J.H. McGowen, C.G. Groat, and V.E. Barnes. 1992. Geologic atlas of Texas; Beaumont Sheet. Map. Turner, B.L., H. Nichols, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Sida Bot, Misc. 24:1 -888. USDA NRCS. 2007. The PLANTS Database (http://plants.usda.gov). Nationa! Plant Data Center, Baton Rouge, LA 70874-4490. Watson, G. 2006. Big Thicket plant ecology: an introduction. University of North Texas Press, Denton. 672 J | of the Botanical R hl BOOK NOTICE RICHARD H. WARING AND STEVEN W. RUNNING. 2007. Forest Ecosystems: Analysis at Multiple Scales, Third Edition. (ISBN 978-0-12-370605-8, pbk.). Elsevier Academic Press, 30 Corporate Drive, Suite 400, Burlington, Massachusetts 01803, U.S.A. (Orders: www.books.elsevier.com, 1-800-545-2522, 1-800- 568-5136 fax, Elsevier, Order Fulfillment, 11830 Westline Industrial Drive, St. Louis, Missouri 63146, U.S.A.). $69.95, 420 pp., maps, figures, tables, some color plates, 7 1/2" x 9 1/4" Contents: Preface to the Third Edition Preface to the Second Edition Preface to the First Edition l. Forest Ecosystem Analysis at Multiple Time and Space Scales Section I. Introducti to Ánalvsis of S ] Corl Cat ` y Water, Carbon, and Minerals through Forest Stands 2. Water Cycles 3. Carbon Cycle 4. Mineral Cycles Section II. KE to poo Scans d Function 6. __ ibusceptibility Es Response of Forests to Disturbance Section TIT Tus a | Aë a Pal Ab all C Jz Tc a2 Pa y I Modeling d Regional Ecosystem Analysis 8. Regional and landscape Ecological Aca sis 9. The Role of Forests in Global Ecology 1c NE AA I J Ff. 10. Advance in Eddy-Flux Analyses, Remote Sensing, and Evid f Climate Change Epilogue Bibliography Index From the back cover.—" New to P» edition: New companion website includ leling soft re cp and video clips; revised section on import g l scale analyses; nue an DEER to globa l scal y g ised color tailed n of predict across 65 eco-regi led models, equations V I r Pi o H In H 1 , 1 1 3 1 te J S1 Í£..11 J J; F +l +l = | 1 1 graphs, ples ] e Lu 1 J WIT LESCALCIL J. Bot. Res. inst. Texas 2(1): 672. 2008 SEED GERMINATION RESPONSE OF ZIZANIA TEXANA (POACEAE: ORYZAE) TO SOIL INUNDATION M.L. Alexander E. 5. Rn ana HUNE service ana ! Technology Center 500 E. McCarty Lane San Marcos, Texas 78666, U.S.A. ABSTRACT Although the current Eon of Se texana ino wild rice) supports the theory that this speci flowing water for stable existenc I g | k within the San M Ri Hays County. TX, U.S.A This may mean that, at times, Z. texana ade itl flowing water or germinated in saturated, but not i ils. In this study, T determine if 7 Ff E ot fi] a E j +1 1+1 ] 3 1] E L o e PE Ei o Dd dees | SI mi i + £41 E: J "E í Jc “1 à Es des Los f oO DIN cl à eo o I 2 4 o "E S a = en 1 E 41 1 1 1 1 [. + ] Fiel: 1 ] those tested in only saturated soils. y may help y g I RESUMEN l5 acc A 14 ea S " f. DN 4 x "Tus J $ RE 14 1 “ A I [e T L D requiere agua corriente para su existencia GER eege planta creciendo de rib ib n el Río E zs 1 = ES 1. D d 1 y mA TX, US p jue, g Kë g o germinó ti i dad ER este dudo d EM d : dad 1 avi e , L F " Lee TL at J 1 Dh H = = : 14.4 | Haste J H i A ] “11 germinacion | E | B g g 1 J J F. 3 f. lea É f^ MIN 11 1 J J T os resultados r L © L X ] " 28:3 A J 1 r € 1 4 1 € INI RODUCTION Ecophysiology of seed germination is very important for the conservation of endangered species as it is a key factor of reproduction and will therefore strongly aid in restoration strategies. For aquatic and wetland plants, it is essential to know if germination rates are affected by soil water-levels (Sifton 1959; van der Valk 1981; Keddy & Ellis 1985; Kellogg et al. 2003). This information will help in making decisions regarding where in an aquatic system seeds should be planted. There are three species in the genus Zizania L. in North America: Z. aquatica L., Z. palustris L., and Z. texana Hitch (Terrell et al. 1997). Each of these species grow in aquatic environments, but Z. texana may require flowing water for stable existence (Power 1996; Poole & Bowles 1999), while the other species can be found in stagnant water on the edges of lakes or along muddy shores of slow-moving streams that are subject to constant changes in water levels (Archibold & Weichel 1986; Pip & Stepaniuk 1988). Zizania texana is a federally listed endangered species endemic to the San Marcos River, San Marcos, Hays County, Texas, U.S.A. The distribution of Z. texana is restricted to the upper 2.4 km of the river, within mean water depths ranging between 0.72 to 0.83 m, and mean velocities ranging between 0.38 to 0.93 m/s (Poole & Bowles 1999). Historical documents that portray Z. texana growth in the San Marcos River describe the plant as growing bank to bank, including irrigation waterways (Silveus 1933; Terrell et al. 1978). The San Marcos River is spring-fed and river flow relies on the spring outflow as well as runoff. When outflow and runoff are low, such as during times of drought, the area near the banks may not always be covered by water. This may mean that, at times, Z. texana grew without flowing water or germinated in non-inundated soils. The purpose of this study was to evaluate the response of seed germination of Z. texana to soil inundation. For this study, I test the hypothesis that Z. texana seeds need to be in inundated soil for successful germination. J. Bot. Res. Inst. Texas 2(1): 673 — 676. 2008 674 tani itute of Texas 2(1) MATERIALS AND METHODS Approximately 3,375 cm? of soil consisting of one third pea gravel to two thirds of an equal mixture of compost, top soil and sand was placed into each of M 45 x 30 x 13 cm clear plastic rectangular containers. Soil mixtures were saturated with water from the E d ifer (the same water source as the San Marcos River) until the point at which pooled water just began to become visible on top of the soil. Seeds used for the study were collected randomly from container grown plants growing in the San Marcos National Fish Hatchery and Technology Center that were initially collected from the San Marcos River following tech- niques recommended by Richards et al. (2007) for the best genetic representation of the wild population. We used a total of 50 seeds for this study. Because this endangered plant has such a small population size, there are only a small number of seeds available for scientific studies. The seeds were collected four months prior to the experiment, in September 2006, and preserved in a refrigerator at 4 °C in moist paper towels within sealed plastic bags, allowing for optimal germination conditions (Rose & Power 2001). Lemna and palea were kept intact as their presence does not change germination probability and as it also mimics the natural condition in this species (personal observation). Five seeds of approximately equal size were placed equidistant to each other (5 cm apart) in the containers of saturated soil at a 4 cm depth. All containers were placed into a growth chamber set to a 12-hour photo period at 20 — 23 ?C. Water was added slowly to the five containers receiving the inundated treatment until it covered the soil with 2.5 cm of water. Each inundated treatment was paired next to a non-inundated treatment within the growth chamber. Every 24 hours, any water loss within the 10 containers was compensated for, seeds were visually examined for signs of germination, and the total number of germinated seeds was recorded. Seeds were considered to have germinated when the radical had visibly emerged from the pericarp. Since Power and Fonteyn (1995) found that most seeds of Zizania texana germinate within a 10-day period, following a 4-week period of daily visual examinations the study was terminated. A paired t-test was run in order to determine if there is a significant difference between the germination success rates of the two treatments, non-inundated and inundated (analysis performed using SYSTAT v.10 statistical package). RESULTS Sixteen of the 25 total seeds in inundated soil and four of the 25 total seeds in non-inundated soil germinated within the 4-week period (Fig. 1). Germination success S ter in inundated soil than in non-inundated soil (paired t = 4.7, df = 8, p = 0.009; Mean difference = 2.4; : 95% C.I. = 0.98 to 3.82). DISCUSSION In this study, more Z. texana seeds germinated in inundated soil than in non-inundated soil. This may be because Z. texana seeds are recalcitrant and do not survive for long outside of water (Horne & Kahn 2000). The seeds may more readily germinate in inundated soil than in non-inundated soil because Z. texana are so sensitive to desiccation. Zizania texana is considered a perennial grass (Terrell 2007). In its current distribution within the San Marcos River, Z. texana plants are, for the most part, submerged year round (personal observation) and rarely flower (Emery 1966, 1977). There is a possibility that the species has an annual form as well in certain environments. This species is a CO, obligate (Power & Doyle 2004). It may remain in perennial form if growing in current velocities such that the boundary layer surrounding the leaves is small enough to take in CO, for photosynthesis. However, if in stagnant, or slow moving currents, an individual plant may become emergent in order to take in CO, from the air (where it is much more abundant), flower, seed, and then die (i.e., annual growth form). If this species were to grow in its historic range of bank to bank, it may be possible that some of the plants would become annuals and grow in areas of low flow and shallow water. Because the seeds do not germinate as well in non-inundated soil, the San Marcos spring-flow has to continue to be high enough (>100 cubic feet per second) that the soil at the banks' edge is consistently inundated. Currently, Z. texana is inhibited from growing along the shallow, low flow banks because Colo- Alexander, Seed germination in Zizania texana 675 5 q 4 Q Qu) V^ "d E B 3 = y é E 5 ? lle Cé 1 0 m Inundated Non-inundated Treatment Fic. 1 inundated soil), casia esculenta (wild taro) has invaded and dominated the edges of the San Marcos River (Akridge & Fonteyn 1981; Nelson & Getsinger 2000). The results of this study will aid in future restoration efforts for this endangered aquatic plant. With data that supports the planting of seeds in inundated soil, all areas chosen for planting should be covered in water during average spring flows. Restoration efforts of Z. texana should include planting in both low and high flow areas to support both perennial and annual forms of the plant. This would have to include the removal of C. esculenta in order to make the river-edge habitat available for the endangered Z. texana. ACKNOWLEDGMENTS I would like to thank P. Grant, T. Wycoff, and E. Pratt for their help with data collection. The views expressed in this paper are the authors and do not necessarily reflect the view of the U.S. Fish and Wildlife Service. REFERENCES Axrioce, R.E. and PJ. Fonteyn. 1981. Naturalization of Colocasia esculenta (Araceae) in the San Marcos River, Texas. SouthW. Naturalist 26:210-211. 676 ARCHIBOLD, O.W. and B.J. WEICHEL. 1986. Variation in wild rice (Zizania palustris) stand tl Saskatcl Canad. J. Bot. 64:1204—121 1. Emery, W.H.P. 1966. The decline and threatened extinction of Texas wild-rice (Zizania texana Hitch.). SouthW. Naturalist 22:393-394, Emery, W.H.P. 1977. Current status of Texas wild-rice. SouthW. Naturalist 22:393-394, Horne, FR. and A. Kan. 2000. Water loss and viability in Zizania (Poaceae) seeds during short-term desiccation. Amer. J. Bot. 87:1707-1711. Keooy, PA. and T.H. Eus. 1985. Seedling recruitment of 11 wetland plant species along a water level gradient: shared or distinct responses? Canad. J. Bot. 63:1876-1879, KeLLOGG, C.H., S.D. BRDGHAM, and S.A. LeicHr. 2003 Effects of water level, s! | fi On ination and gro th of freshwater marsh plants a long a simulated successional gradient. J. Ecol. 91:274— -282. Netson, L.S. and K.D. Gersincer 2000. Herbicide evaluation for control of wild taro. J. Aquatic Pl. Managem. 38:70-72. Pip, E. and J. STEPANIUK. 1988. The effect of flooding on wild rice, Zizania aquatica L. Aquatic Bot. 32:283-290. Poote, J.M. and D.E. Bowies. 1999, Habitat characterization of Texas wild-rice (Zizania texana Hitchcock), an endan- gered aquatic macrophyte from the San M bd River, TX, USA. pl Geen 9:291-302. Power, P. 1996. Effects of current velocit | composition on growth of Texas wild-rice (Zizania texana). Aquat. Bot. 55:199-204. Power, P. and R.D. Dove. 2004. Carbon use by the endangered Texas wild rice (Zizania texana, Poaceae). Sida 21:389-396, Power, P and PJ EE 1995. Effects EE ti | substrate on seed germination an cl seedlina growth of Texas wild! ice (Zi ) 1 SouthW Naturalist 40:1—4. RicHARDS, C.M., AR ANTOLIN, J. Poote, and C. WALTERS 2007. Capturing genetic diversity of wild population for ex-situ conservation: Texas wild rice (Zizania texana). Genet. Resources Crop. Evol. 54:837- 848, Rose, F. and P. Power 2001. Maintenance of conservation population of Texas wildrice (Zizania texana). U.S. Fish and Wildlife Service, San Marcos. SIFTON, J.B. 1959. The germination of light-sensitive seeds of Typha latifolia L. Canad. J. Bot. 37:719-739. Siveus, W.A. 1933. Texas grasses. The Clegg Co, San Antonio, TX. TeRRELL, EE 2007. Zizania L. In: Flora of North America Editorial Committee, eds. Flora of North America. Oxford University Press, New York. Pp. 47-51. TerReLL, EE. W.H.P. Emery, and H.E. Beaty (1978) Observations on Zizania texana (Texas wildrice), an endangered species. Bull. Torrey Bot. Club 105:50-57. TERRELL, EE. PM. Peterson, J.L. Reveal, and M.R. DuvaLL. 1997. Taxonomy of North American species of Zizania (Poaceae). Sida 17:533-549, VAN DER VALK, A.G. 1981. Succession in wetlands: A Gleasonian approach. Ecology 62:688-696. PREDICTING VIOLA GUADALUPENSIS (VIOLACEAE) HABITAT IN THE GUADALUPE MOUNTAINS USING GIS: EVIDENCE OF A NEW ISOLATED POPULATION Timothy C. Mullet Fred Armstrong! Department of Biology Resource Management Office Sul Ross State University Guadalupe Mountains and NP Alpine, lexas 79832, U.S.A Salt Flat, TX 79847, USA. Benjamin Zank Christopher M. Ritzi ie Ee and Science Department of Biology at Smoky Mountains NP Sul Ross State University in Ges 37 U.S.A. Alpine, Texas 79832, U.S.A. ABSTRACT Viola guadalupensis i demic of the Guadal M ins in West Texas. Isolated t li tone, rock face, this species was known only to exist at a si gle location along the East Rim of Frijole Ridge within Guadalupe } ins Nati ] Park (GMNP) Despite traditional y ingly ] t, researchers had ! ble to locat een Its small population size, genetic isolation, and lack of significant habitat inf this speci Inerable to exti Conse CARLA the Resource T fe Tee [ CCMAKTD 1 E 1 j E se + CT ] tal + 1 7 serve this species’ viability. Geographical Information Systems (GIS) have previously | ful licti itable habi f rare species; therefore, we developed a GIS-based habita model — generated from the elevation, aspect, and ] ific to the locality — that displayed a map predicting suitable V. guadalupensis habitat throughout the vail We ground- mitad nine of e 14 locations produced by the habitat model. One location accurately poa Gg type locality, while a second location revealed a new pee ion V. yaa ae within a steep, slot-canyon d g g Ridg Tue ndi were ee to E gece T D 1 TENES | T for Y [s] C o JL ZE ad discovered a new EG id M i n SUUM efforts, t ful f GIS-I l habit lels f ying pecies i reaffirmed, | g Key Wonps: Geographical Information System, GIS, Violaceae, Viola guadalupensis, Guadalupe Mountains, habitat model, validation RESUMEN Vin] 141 : 4 : Jan] = rn 14] D a Texas ome =] n p A A esta o E E r ? = £f 1 | 1: ] 1 ] PERI q m E Bn Vë e T The | 1 1.1 AT qaod Tai 1m 1. ZORANTT A L D E L M Si J=] ] 1 1 ] y E ] 1 ] Lal T 1 J 1 E 1 E I E e f € a + A LI E 1 Lm | - + Av la fale E WE GE y Le 1 Tulsa Ee 1 e I. E 1 5 14 E t d oc la TD AA SEE AMI RARMTD 1 : 1 ] i J ] J ? o E EE i o lak t 1 ] 7 Lidad A + T (cer qus To E fOTCNI e I L [e] " q É Lal ] 4 11 ] 11 1,1 1, Lal 1 ] MIC le F L r Dt eMe cA H E : m f des Leef 1923: 1 1 Lal ] J 17 partir de la altitud, aspecto, y p I I | pa que T Ve guadalupensis en todo el parque. Compro! bre elt le las 14 localizaci producidas por el modelo de hábitat. Una 1 1 I ] j I dad j- 4, po, a | g Ja] 1 1 I 1.] A guadalupensis cl A A Rh t A A : la Ele D cC ] 1 1: 1 o r J 4 o i CN d.d : : E WI : ] J r^ DI A : 141 1: osa ción de V. guadalupensis y | predij la localidad tipo y descubrió población, a p le | f tradicionales d imiento, se reafirma la utilidad de lo hal 1 ] STE ] £33.13 A : " A + J E E E I | "E (che I» e E Habitat models developed from Geographical Information Systems (GIS) are useful tools for predicting the distribution of rare species, decreasing the ineffectiveness of random surveys, providing potential sites for relocation, and aiding the overall development of conservation management plans (Vogiatzakis 2003; Imm et al. 2001; Wu & Smeins 2000). Van Manen et al. 2005) developed a predictive GIS-based habitat model "Correspondent: fred_armstrong@nps.gov 1 Rat Dec Inet Tavac 2(1): ATI — 684. 2008 I fal n.a H Inm LI "LET £T STE A! i^ 7 678 derived from the variables of known plant locations for a variety of species in Shenandoah National Park, VA. The efficacy of their predictive model at locating species of interest was 4.5 to 12.3 times more efficient than random surveys. Husveth (2003) used a baseline GIS model generated from observed plant locations, surficial geology, and major soil associations to predict likely sites for rare plant species in a 15,378 ha plot of Anoka County, MN. Field validation surveys (i.e. ground-truthing) of this model revealed additional rare plant populations as well as high to medium quality habitats for rare species. Although the predictive capabilities of GIS-based models are only as good as the variables used to generate them, they can provide useful insight into the dependency of rare species on habitat variables. The Guadalupe Mountains violet (Viola guadalupensis Powell & Wauer) is a rare and unique species of wildflower endemic to the highest elevations of the Guadalupe Mountain range in West Texas. The species was described in 1990 within Guadalupe Mountains National Park (GMNP; Fig D, Culberson County, TX, and since its discovery has only been recorded from a single location along the East Rim of Frijole Ridge (Fig 2; Powell & Wauer 1990). At its type locality, this perennial, yellow-flowered violet grows in mats of Bp to e individuals rooted in small openings of a northwest-facing, dolomitized, limestone rock face y 2,600 m above sea level (Powell & Wauer 1990). The plants are well shaded by two large Beet firs (Pseudotsuga menziesii in ) and pow in association with rock spi (Petrophytum caespitosum [Nutt.] Rydb.), Guadalupe valerian Steyernm.), small rocklettuce (Pinaropappus parvus Blake.), Guadalupe leastdaisy (Chaeto- pappa hersheyi Blake.), littleawn needlegrass (Achnatherum lobatum Swall.), and a species of sedge (Carex sp.; Powell & Wauer 1990) Little is known about this species’ natural history or the specific habitat characteristics needed to maintain its viability. Speculation has been made of its dependency on microclimatic conditions and substrate type; however, these data have only recently been recorded (F. Armstrong, pers comm.). Average temperature from January 2003 to January 2004, at the type locality, was approximately 12.78°C. Relative humidity during the same time interval ranged from <10 to 100% and daytime light intensity during the ENDE season E a July 2002) averaged 0.22 log lum/m* (F. Armstrong 2004, unpubl. report). counts, attempts to germinate see dlings in the laboratory discovered that chilling seeds and treating them with De MW: allowed for successful germination (n = 5; C. Blaxland, pers comm.). Seedlings in potting soil became chlorotic, but when lime was added they regained green coloration indicating an affinity for alkaline substrate e PLE pers comm.). Although baseline data have been collected, no intensive ive analysis or sig t research has been done at the risk I of damaging the population. Thorough investigations to locate additional populations in seemingly suitable habitat have been con- ducted along 8 km of the East Rim of Frijole Ridge from the top of Bear Canyon to Lamar Canyon (Powell & Wauer 1990). Despite these efforts, no other populations were found. Viola guadalupensis is currently categorized as a National Park Service Species of Concern, yet it appears to be in danger of extinction based on its small population size and lack of significant habitat information. Powell and Wauer (1990:1) had even commented that V. guadalupensis “is an immediate candidate for endangered status” for these very reasons. With this in mind, the Resource Management Office of GMNP has implemented a plan to collect seeds and germinate them in a laboratory setting with the intentions of introducing V. guadalupensis to other suitable locations within the park. The objective of this study was to locate suitable V. guadalupensis habitat sites within GMNP, using a GIS-based habitat model generated from variables unique to the type locality, and validate those locations with ground surveys. METHODS Study Area.—Our study was conducted within the 34,971 ha of GMNP, Culberson County, TX. The Gua- dalupe Mountain range is a unique biological sky island consisting of Chihuahuan Desert vegetation (Agave, Mullet et al., GIS-based distribution of Viola guadalupensis 679 Legend N [|| | Political boundaries rd c Texas aale Mountains NP [= ZS RE D 175 350 700 Kiometers Lë 2, Ec i aor — Los ee: ia Ed 4,008 3,000 Meters Culberson Co. { | Fic 1 CL | J FPE a E e 21.1 aa A a: klag’ Lm LI A yo FP I! MEI J ELS AR FATTI A SÉ Yucca, Larrea, Acacia, etc.) at lower elevations (< 2,000 m) and mixed-conifer woodlands of Pseudotsuga menziesii, southwestern white pine (Piris oA Engelm.), ponderosa pine (P. ponderosa Laws.), pinyon pine (P. edulis Engelm.), and 9, p.) at higher elevations (> 2,000 m). This unique mountain range of the Chihuahuan Desert is the past limestone uplift in the SECH United States, formed 250 million years ago by an ancient reef system (Murphy 1984). Mountain summits excee d heights of 2,600 m, shadowing deep canyon systems containing a variety of mountain springs. A 19-year average of temperature and precipitation collected at the 2,455-m elevation automated weather station documented average winter lows of -1.7?C, average summer highs of 23.9?C, and average annual precipitation of 45.0 cm (National Park Service 2005). Monitoring the type locality.—The condition of the type locality was monitored on 26 May 2006. We compared the counts of individual V. guadalupensis plants to a detailed line drawing of individual V. guadalupensis plant locations and numbers produced by the Guadalupe Mountains Resource Management Specialist during inventories conducted in 2001, 2002, and 2004. To avoid over estimating the population size, individual plants were conservatively tallied and then averaged between counts for consensus. GIS model development.—Based on the fact that V. guadalupensis had not been observed by park staff or researchers in any of the mesic regions surveyed within the park, we speculated that this species may have specific topographic habitat preferences that include 1) elevations with overall cooler, moister conditions, 2) aspects with minimal hours of direct sunlight, and 3) vertical rock versus horizontal soil substrate as indicated at the type locality. The National Park Service Intermountain Regional GIS Office, in 680 Journal of t tanical h Insti Texas 2( Guadalupe Mountains NP Legend Elevation (m) 2.662 1,559 1.250 | 2,500 5,000 Meters Ex 9 Natailed al es Unf fab. East DE | Frij | Ridg ( +1: lin hl k) f (. (al I M tal Nati IP k, Culberson Co., TX conjunction with GMNP, performed habitat modeling based on these variables to narrow the search areas for potential habitat and suitable locations for establishing additional colonies. GIS modeling was performed with ESRT's ArcMap Ver. 9.1 software (available from http://www.esri.com/ software/) incorporating the following parameters in order of priority: 1) elevation bands from 7,800 to 8,000 ft (feet were used for the ease of visually correlating potential sites to USGS topographic maps printed with 40-ft elevation contour intervals), 2) a northwest aspect range from 300 to 330 degrees, and 3) slopes >70 percent. Elevation, aspect, and slope were all derived from a 30-m digital elevation model (DEM) of GMNP obtained from the National Park Service, Natural Resource GIS Data Store. Using ESRI's ArcToolbox Reclassify tool, DEM cell values were used to separate elevation and aspect into three classes (Class 1 = elevations between 3,500 and 7,800 ft and aspects O to 300 degrees, Class 2 = elevations from 7,800 to 8,000 ft and aspects 300 to 330 degrees, and Class 3 = elevations > 8,000 ft and aspects 330 to 360 degrees). The Raster Surface Slope tool was used to divide slope values into two classes (Class 1 = slopes 0 to 7096 and Class 2 = slopes >70%). Each class of elevation, aspect, and slope was then assigned a model value of 3, 2, 1, or 0. A model value of three was assigned to the elevation parameters met by Class 3, while model values of 2 and 1 were assigned for the aspect and slope parameters in Class 2, respectively. An assignment of 0 represented all other non-target cell values for every class. All four model values were unioned together producing a model with seven categories from 0 to 6 dis- played as a digital predictive habitat map. Category O displayed all non-target parameters; categories 1 and 2 displayed only slope and aspect, respectively. Category 3 displayed criteria for elevation or a combination Mullet et al., GIS [ O ti DTN g m A T of slope and aspect. Category 4 displayed areas where a combination of elevation and slope criteria were met, while Category 5 req ted areas of both elevation and aspect. Finally, Category 6 displayed locations where all three parameters of elevation, aspect, and slope were met (Table 1). Model Validation.—Field validation surveys of predicted V. guadalupensis sites were conducted May through July 2006. We used Universal Transverse Mercator (UTM) coordinates digitized in ArcMap to find predicted sites where all three criteria were met (i.e. Gatto 6). A radius of 200 m was investigated around each UTM location to compensate for any inacc es of positioning due to tellite geometry. The specific characteristics used to determine suitable habitat of predicted eai in the field consisted of north to northwest-facing rock walls with crevices and vegetation similar to that of the type locality, a physically cooler ambient temperature, and shaded areas with sufficient canopy cover to protect the site from direct sunlight. Rock faces <3 x 3 m were ruled out as insufficient space for colonization due to the lack of potential dispersal of plants within the area. RESULTS Status of the type locality.—Compared to previous inventories, 20 new individual pans (n = 94) were recorded in 2006. Although no Ee were observed to be fl ing, several pl d No conspicuous diff in veget ition were observed when cornered to detailed ER draw- CICIICCS 111 VESELA Came: ings and photographs from previous inventories. Model validation -- The model displayed 14 locations meeting all three criteria in Category 6. We investi- gated a total of nine locations consisting of three sites near Guadalupe Peak (WGP, NGP1, and NGP2), two near Bush Mountain (BMD1 and BMD2), one north of the Bowl (NOB), one near South McKittrick Canyon (SMK), and two locations on Frijole Ridge. Five locations were omitted from investigation due to inacces- sibility and by request of the GMNP Resource Management Office, therefore, no site codes were assigned to these locations. Two of the nine investigated locations predicted by the model on Frijole Ridge revealed populations of V. guadalupensis. One of these locations was the type locality (designated VIGUL). The second location revealed a new, isolated population of V. guadalupensis in a steep, slot-canyon drainage (approximately 2.1 to 2.4 m wide), on a 35 degree, northeast-facing limestone wall. The site (designated VIGU2) is located off a plateau on the East Rim approximately 2 km northeast of the type locality. We observed 25 individual V. guadalupensis plants at VIGU2 on 26 May 2006, growing largely in a monoculture. Several V. guadalupensis plants were located approximately 1.5 m overhead, making an exact count difficult. Although no seed capsules were observed on non-flowering plants, a number of violets still possessed flowers. All violets had noticeably longer petioles and grew predominantly as individual plants rather than in the mat clusters characteristic of violets at the type locality (VIGU1). Associated plant species of VIGU2 included Valeriana texana, bedstraw (Galium sp.), and red alum- root (Heuchera rubescens L.). Violets were shaded by the steep, canyon walls and a canopy of western hop- hornbeam (Ostrya knowltonii Cov.) and P. strobiformis. Other surrounding vegetation consisted of mountain spirea (Holodiscus dumosus Nutt.), Pseudotsuga menziesii, and Petrophytum caespitosum. Three of the seven sites without violets (WGP, BMD2, and SMK) were considered to be suitable for introduction upon comparison to the type locality. The remaining four locations (NGP1, NGP2, NOB, and BMDI) were considered unsuitable, possessing sparse vegetation and fully exposed rock faces atypical of either violet location. Twenty-two percent of all survey sites predicted by the model had extant populations of V. guadalupensis. Three of the nine sites (3396) were considered to be suitable sites for introduction. Overall, this model had 5696 accuracy for predicting suitable or existing V. guadalupensis habitat. Potential locations for introduction.—The first site considered to be suitable was located around WGP and appeared to be very similar to that of VIGU1. This site (designated WGPI) included a steep rock g iesii and Ostrya knowltonii trees approximately 40 m down a p—> face, shaded by several small P 682 | t tanical i Texas 2(1) TABLE 1; e Ee cana non target lel | ters to theii iated category lto g te the Viola gua i ins N | Park. The desired lel ters of categories 1 through 6 included slopes » 7096, mes between 300 and 330°, and elevations >8,000 ft, respectively. Non-target parameters of Category D included all criteria outside the desired parameters. Category 6 was used to identify potential Y guadalupensis habitats throughout the park Category Model parameters 0 Non-target parameters 1 Slope 2 Aspect 3 Elevation or slope and aspect 4 Elevation and slope 5 Elevation and aspect 6 Elevation, siope, and aspect steep drainage. The ambient temperature was physically cooler in the early and late afternoon (1120 h and 1619 h). This rock wall faces north to northwest with a number of crevices and associated vegetation similar to VIGU1. The location is approximately 15 to 20 m long and 5 to 8 m tall with an elevation of ap- proximately 2,500 m. The second potential site (BMD2) possessed very tall and wide north- and west-facing cliff walls that appeared to be suitable habitat, however, as investigation of the area progressed into the afternoon (1300 h), the rock walls were observed to be more exposed to direct sunlight than earlier that morning (1100 h) as a result of sparse py cover. However, one rock wall, in particular, was considered suitable because it met all desirable criteria outlined by the model and remained shaded by Pseudotsuga menziesii and Quercus sp. at the time of investigation. Finally, the third potential sit South McKittrick Canyon (SMK) possessed a number of small rock walls (between 3 x 3 m and 5 x 5 m) with one single location (designated SMK2) significantly shaded by Ostrya knowltonii, Pinus ponderosa, Pinus strobiformis, and Pseudotsuga menziesii. Data loggers, programmed to record light intensity, relative humidity, and temperature, were placed at VIGU1, VIGU2, WGP1, BMD2, and SMK2 for future comparison of microclimate data. DISCUSSION ifi of GIS-1 ] predicti deling.—The results of this study indicate two essential aspects e understanding the sentando of this predictive model when compared to previous ground surveys and the habitat requirements of V. guadalupensis. First, the GIS-based habitat model caed iiis type locality and an additional population of V. guadalupensis along Frijole Ridge, as well as three potentially suitable habitats for introduction west of this region. Since its discovery in 1990, extensive ground surveys of seemingly suitable habitat have been unable to locate additional populations (Powell & Wauer 1990). Based on our results, this study supports the relative usefulness of GIS-based habitat models for predicting the distribu- tion of rare species consequently ee Se EE of random surveys. Second, V. g pla; y t lopes, northerly aspects, and high elevations on Frijole Ridge. Tes lts substantiate ou e that V. puadalupensls have specific topographic habitat preferences toward the variables - to develop our model. It is important to consider the fact that violets at VIGU2 were positioned on a northeast aspect rather than a northwestern aspect characteristic of VIGU1. This observation might indicate that aspects ranging from 300 to 45 degrees (northwest to north- east) may be a better representation of this species’ aspect preference than those strictly between 300 and 330 degrees. Model efficacy.— Considering two out of the nine predicted locations represented violet populations, while the remaining seven did not, suggests that 1) the variables used to develop the model were not spe- cific enough to locate habitats strictly containing V. guadalupensis or 2) V. guadalupensis is restricted only to Mullet et al., G1S-! | distributi fViol g lal i 683 these two locations on Frijole ridge and may have not had the opportunity to disperse to other areas. It is also important to take into account that three of the five locations not ground-truthed in this study were also located along Frijole Ridge. It is possible these locations may possess populations of V. guadalupensis. Further investigation into these areas is suggested in order to allow for a more complete model evaluation and to confirm the distribution of V. guadalupensis using this GIS-based habitat model. Because the efficacy of a predictive model is dependant on the variables used to generate it, accurate information of habitat requirements for a species is necessary for the devel t of an accurate predictive model. Cherrill et al. (1995) stated that the rarity of a species greatly us a model's efficacy to predict species-habitat distributions. Their model was generated from variables of known locations to predict rare and common species compared to traditional field surveys. Their results found a higher rate of accuracy for predicting common species than those of rare species. These results were dependent on the amount of information available for each species (Cherrill et à 1995). Additional EE -habitat information or V. guadalupensis (e.g., microhabitat conditions) will likely enabl g generate model for predicting V. guadalupensis distribution and abundance. Conservation and management.—Proper management of the two V. guadalupensis sites is vital for its continued existence within the Guadalupe Mountains. The distance between VIGUI and VIGU2 (2 km) suggests that gene flow between these populations is unlikely. Because the most promising mechanism of seed dispersal across such distances is wind (Davies et al. 2004), this mechanism is improbable due to the density of vegetation around VIGUI and the steep, narrow, rock walls surrounding VIGU2. These condi- tions likely protect these areas from wind rather than ae to the use of wind- PRSE eed ia sae ] Er: cana Consequently, tandi | d por for its conservation. mr we suggest that p— of violets into additional hould representatives from both locations to improve genetic diversity within new areas to prevent an additional bottleneck effect. Although the locations of this species are far from human impact, several environmental factors may threaten the persistence of these populations. The accumulation of ladder fuels within the park has cre- ated conditions suitable for catastrophic wildfires along the East Rim. This could result in the burning of V. d niin irn habitat or a reduction in the number of shade trees surrounding the area causing fatal heat exposure (F. Armstrong, unpubl. report). Predation of leafs and seedpods by small mam- nd. has been Na to occur at VIGUI (F. Armstrong 2002 and 2004, unpubl. field notes). Even though there has been no direct observation of this activity, seed predation will likely affect the reproductive E 1 A LICLUUE success of individual plants, reducing overall genetic diversity and dispersal. Continued monitoring of both V. guadalupensis sites and comparing the long-term microhabitat condi- tions of known violet sites to potential introduction sites will provide a better understanding of specific habitat requirements for introduction and habitat management. The two populations of violets, as well as the three uninvestigated sites on Frijole Ridge, should also be considered during fire management planning to prevent unfavorable conditions due to catastrophic wildfires along the East Rim. The habitat model developed by the National RUE ala to predict potential V. guadalupensis habitat has provided critical baseline data for additional i d research within GMNP. Due to the suc- cess of finding an additional population and other MN sites for introduction, resource managers can utilize this information for continuing plans to proliferate and conserve V. guadalupensis populations with potential success. ACKNOWLEDGMENTS We are grateful for the support of the Guadalupe Mountains National Park and the National Park Service, DOI who funded this project. Sincerest thanks to the significant contributions made by C. Blaxland for her work to successfully germinate and propagate Viola seeds. Special thanks to field inventory and survey members A Ames, C. Blaxland, D. Ferris, I. Khan, M. Lommler, W. Lucas, and A. Swanson. We express I Lal D H ID LI ror ET 684 } t lexas 2(1) additional appreciation to Joyce Maschinski and Justin Williams for their supportive comments and ben- eficial edits of this manuscript. REFERENCES CHERRILL, AJ., C. McCuean, P. Watson, K. Tucker, S.P. Rushton, and R. SANDERSON. 1995. Predicting the distributions of plant species at the regional scale: a hierarchical matrix model. Landscape Ecol. 10:197--207. Daves, S., A. Wurre, and A. Lowe, 2004. An investigation into the effect of long-dist | dispersal on organelle population genetic structure and colonization rate: a model analysis. Heredity 93:566--576. HusverH, J.J. 2003. Ecological surveys of rare plants and plant communities in eastern Anoka County, Minnesota. Conservation Biology Research Grants Program, Division of Ecological Services, E Department of Natural Resources. http//files.dnr.state.mn.us/ecological_services/nongame/projects/consgrant_re- ports/2003_Husveth_sign2 (Accessed October 2006). Imm, D.W, H.E. SHEALY JR, KW. Moon, and B. Couns. 2001. Rare plants of southwestern hardwood forests and the role of predictive modeling. Nat. Areas J. 221:39--49. NATIONAL Park Service, 2005. Fire Management Plan, Guadalupe Mountains NP, Texas. PoweLL, A.M. and B. Wauer. 1990. A new species of Viola (Violaceae) from the Guadalupe Mountains, Trans-Pecos Texas. Sida 14:1--6. VAN MANEN, FT, J.A. YOUNG, CA THATCHER, W.B. Cass, and C. ULrey. 2005. Habitat models to assist plant protection efforts in Shenandoah National Park, Virginia, USA. Nat. Areas J. 25:339--350, VoaiATzAKis, I.N. 2003. GIS-based modelling and ecology: a review of tools and methods. Geographical Paper no. 170. Department of Geography, The University of Reading, Whiteknights, U.K. http://www.geog.rag.ac.uk/ Research/Papers/GP170 (Accessed: October 2006). Wu, X.B. and FE. Smeins. 2000. Multiple-scale habitat modeling approach for rare plant conservation. Landscape Urban Planning 51:11--28. ANNOTATED VASCULAR FLORA OF THE DEAD HORSE MOUNTAINS, BIG BEND NATIONAL PARK, TEXAS, WITH NOTES ON LOCAL VEGETATION COMMUNITIES AND REGIONAL FLORISTIC RELATIONSHIPS Joselyn Fenstermacher A. Michael Powell Sul = State University Sul = State University -64 ox C-64. Alpine, m Era 0001, U.S.A. Alpine, e 79832-0001, U.S.A jfen1580sulross.edu Joe Sirotnak Martin Terry PO. Box 29 Sul Ross State University Box C-64 Box C-64 Big Bend National Park, Texas 79834-0029, U.S.A. Alpine, Texas 79832-0001, U.S.A. ABSTRACT mE ne Horse Mountains of Big n Nene nd are one of bu least UMEN areas in Texas. boi is the us HOME m of VIIICVLLIVSIL APL specimens. Field work was performed between August 2003 mi September 2006. The MS area covers 176, 800 acres (71, 548 hect- ares; 276 mi?, 715 En Wee a 4100 ft (1250 m) rang evation. Non-v eA perimien (bry ope tes, P a ae and d led in the final lysi eae d I herbari I 12212 previously-coll | fl ists of 662 speciesi udi Ee and o families. The best represented families are the Ast (80 ies) P (75 jes), Fal (57 ies), (34 species), Cactaceae Di EE Pterid (20 SPECIES) aid Boraginaceae (20 species) Seyn meria falcata v var. x falcata w Was Ge de E e time in the 1.21 : 21 to the United States. ebe taxa increased by 12% and 55 i ] lin tl ly for the first time. Twenty-four non-native species are known from the study area; three are state-listed noxious weeds, and six are considered invasive. The number e known Texas un in h^ Dead Horse Ma ari to 11; seventeen ^ dii are Ron rare. "d Dead Horse e differs from hw an inrreaced relative 1mmorrance of monocots. and the elevatio ominant kaf À L a ASALLA LALALA with the Asteraceae. These and. other fact t tions t 1 peripheral floristic areas, ranging f the Mogollon Rim and the Great Plains to South Texas and cambie Mexico. RESUMEN T c: 1.1 æl 11 AM í del Big Bend National P 1 1 1 po A Tejas. Est 1 I DH . ig 1 ÍT 1 + 1 E : "n J TaT existentes en herbarios regionales. La inv qd fue pena entre e Agosto 2005 y SC 2006. La zona de la investigación pus 176.800 acres n 548 ipe b: M mi?, 715] e E g I Tas Dien final Tinrante el ectiurli art al, 1 di especimenes ] 1 1 Vade d Jat 1 2 + 9915 Fd s: t = + T fT Uc pralitas F E T E DW E " ] f. las Ást (80 especies), Poaceae (75 especies), 662 especies de 344g géneros y 91 familias EE foe peu LM (34 especies), Cactaceae G4 ales Geesen e eae, y Bonaeiiate4e (20 DR U H I primera vez en nidos, yf € uds de Bowie lección adicional (Galacti ) pued peci pecie d nocida en los E U. El número da tasa st ] tad 12%, y 55 esp f ] tad l na del por primera vez Se reconocen 24 t la zona del estud i l hiert El número cies A de Pa qe se geg nidi en la Sierra e Muerto subió a 11; diecisiete especi id La flora Cu PER euer es T = 1 7 t T H 4 = ke r £ L L ae de las Monocotiledóneas, y por la el ión de las P hasta el punto de ser co-dominant las Asteraceae. Estos y otros A flori tér] tendiénd desde el Mosollon Rim en Arizona v los Sé o P Grandes Eines dt los E.U., hasta el sur de Tejas y México — | Rot Rac Inct "TU 2(1): 685 — 730. 2008 686 | | tanical h Instit Texas 2( INTRODUCTION The Dead Horse Mountains (DH) of Big Bend National Park (BBNP) create a remote and forbidding view- scape on the West Texas park's eastern skyline (Fig. 1). The rugged limestone escarpments offer no reliable surface water, nor infrastructure allowing quick access to the interior, leading to claims of it as being un- charted wilderness (Wood et al. 1999). These logistical challenges, in addition to rugged topography and extreme climate, have left the area one of the least botanically understood areas in Texas and the northern Chihuahuan Desert Region (CDR). Much collecting has taken place in the area over the past 100 years. The height of early scholarly investigations came in the 1950s-1960s, and included a SRSU master's thesis by Rogers dd a EEN E excellently cl terized field report by Wells (1965), and collections by numer- Sul Ross State University (SRSU) botanist Barton Warnock. The largest coherent body of collections was ade for a report to the Nature Conservancy (Amos and Giles 1992), which at the time owned a part of the study area that is now part of the Texas Parks and Wildlife's Black Gap Wildlife Management Área. The majority of vouchers made through these efforts came from more easily-accessible areas, near roads and trails peripheral to the end DE ER range, without widely sampling the range of niche habitats existing in the area. Despite a rat! torical interest in this area, no overall floristic characterization has been made for the Dead Horse. Several lists of local flora exist, including more than one in use for Big Bend National Park; however these are based on sightings, literature reports and, to an unknown degree, herbarium specimens (Mahler 1971; National Park Service 1996; Louie 1996; Clelland 2001; Worthington 2001; Alex et al. 2006). A floristic analysis of the Dead Horse will create an important context for Big Bend regional floristics. The DH are located in the Trans-Pecos Mountains and Basins vegetational area (Hatch et al. 1990) but phytogeographic connections have been noted with other Texas floristic areas including the High Plains, Rolling Plains, Edwards SECH and the South Texas Plains, also known as the Tamaulipan Thornscrub region (Powell 2000). C | to more northerly vegetational areas are evidenced by the populations of aspen (Populus tremuloides), Douglas-fir (Pseudotsuga menziesii), and other Rocky Mountain species growing well south of their normal distributions. Floristic links also reach south to the Sierra Madre Oriental and Occidental, expanding to the greater CDR of Mexico (Larke 1989). The graminoid flora even has an affinity to the Great Plains grasslands (Powell 2000; Christie 2006). Because there are so many regions intergrading in the local area, there is high potential in the study area to discover species out of their expected ranges. Furthermore, the DH is topographically complex and the resulting microclimates may harbor species new to science. Already the area shelters several little-known species: Andrachne arida is known from only a few localities in the CDR and has not been seen in the DH since the 1960s, while the purported Dead Horse endemic Hedyotis pooleana has not been observed since the original 1985 collection. Other species are as- sumed to occur in the study area based on their known habitat, but are not well understood, being known only from a few collections in a restricted area. One such example is Bouteloua kayi; previous to this study it was not known to occur in the study area but was discovered during the current work. s environmental complexity, in Boti topography and regional climate regimes, no doubt has helped f the CDR, but instead of being widespread, endemics are mostly ely distributed (Brown 1994). indeed: in Trans-Pecos Texas, Brewster County tallies the highest numbers of endemic (TAM-BWG 2007) and rare plant species (Alex et al. 2006; Poole et al. 2007). Often, however, species of concern are little known beyond data taken at their original collection site. Thus, targeted field studies in Chihuahuan Desert areas like the Dead Horse have a high potential for new floristic discoveries. The main objective of this project was to create a collective body of work describing the floral composi- tion of the DH. Goals to accomplish this objective included: (1) developing an annotated flora for the study area through documenting existing voucher specimens in regional herbaria, and through new collections from a wide variety of underrepresented areas and habitat niches in the range; (2) gaining more knowledge about little-known species that occur in the range, includi ies known regionally but not yet recorded over 687 T ri 47 ER ERE DA Fic 1 Ld | Fa Tu AA FI Li F1 ly LAF A | I rf A I I ye E al ge, LS e m | sil EL E PN ANS. Bs A n 2 P = s E. CAM and RY Een A | L EA Stack Gap Wadia | | re EA i Management Area ; e e Bu Headquarters a H " que! ns OL. NATIONALPARK Š IETARI 3 CLOCK MESA AS Ho POETA ata ers Pree poe im d D en ae = / ROSILLOS uum D s. BOhTH ETT RANCH OH ata ro] x ——— "'CHISOS | MOUBTAINS ` BIG BEND NATIONAL PARK Ep zent. "eh? € A ZER > “th une d a MU, = 1 "m IT Col , Pini: road e e Picnic wa station dea bigh- EJ a z OLPC. vihiichós ordy} Leg end ipod SA -guidog wail LI (Deo 5 == en ower Fund owner's SANTA ELENA ZAWYI "d t " 4 Y E + Pi i "Y P DH m AIS AN In L FARINE MAMA, “ak aL POE | Sind TL nm nu LA Eu LL IP HE: Lif He: ) y " and “Sierra del Caballo Muerto." 688 within the study area; and (3) documenting the occurrence e E Se for the study area, whether rare, t Subsequent disjunct, or invasive, and providing new insight into their abitat requi analysis was used to interpret the Dead Horse flora in a larger regional context within the Trans-Pecos and also the larger southwestern United States. The study was conducted as the first author's MS thesis research (Fenstermacher 2007). The full thesis manuscript, the entire voucher collection database, and many related photos can be found on the Cactus Conservation Institute, Inc. website: http://www.cactusconservation.org/ CCLI/botany/fdhm.html. Site Description The DH are located in the Big Bend region of western Texas, in Me ROSE County about 80 mi south of the town of Alpine (Figs. 1, 2). The study area boundary starts in t tern part of the national park at Dog Canyon, continues south along the western foothills of the DH, and extends east into a portion of Texas Parks and Wildlife's Black Gap Wildlife Management Area (BGWMA), including a few parcels of private ranchland (Fig. 3). The Rio Grande marks the southern limit. The DH are a smaller, northern extension of the impressive Sierra del Carmen of northern Mexico, exhil imilar physical characteristics: northwest- trending ridges alternating with alluvial basins, and generally — arid, rocky terrain. Extending 31.5 mi (51 km) N/S and an average of 8.5 mi (14 km) E/W, the study area covers approximately 176,800 acres (71,548 ha; 276 mi?, 715 km?) and ranges in elevation from 1800 ft (549 m) at the Rio Grande to 5840 ft (1780 m) at Sue Peaks in the heart of the range. The only major physical feature that trends east-west in the study area besides Boquillas Canyon along the Rio Grande is the 14-mile-iong Ge 5 km) EE SE that bisects the DH. There are l major drainages in the study area, but tl Any PISCIS Ms occurs usually flows overland into the dry arroyos, Sie powerful but temporary floods d Rio Grande. Away from the river, surface water is found only unreliably in tinajas, i.e., stone GE eroded usually out of arroyo or canyon beds with bedrock outcrops. The study area experiences hot summers and cool-to-cold winters. The average annual high is 85?F while the low averages 53?F with an average annual maximum of 100? F/38? C and minimum of 33? F/1? C. Large diurnal temperature ranges are common, with a mean of about 32? F/0? C and a maximum of about 50? F/10? C. EEN peaks between May and October, the 10 in/25 cm yearly average falling usually as late afternoon t that range in intensity and often create hail (temperature data from National Park Service, Big Bend National Park, unpublished data). The DH are composed primarily of lower Cretaceous limestone deposited in a deep sea environment ca. 180 mya. As the area was close to the shoreline of the great interior sea, and as sea levels fluctuated, some shales and clays were deposited, reflecting the changing environment. During the Cenozoic (ca. 30 mya), massive blocks of Del Carmen and Santa Elena Limestone were uplifted and tilted along fault lines, resulting in the basin and range morphology seen today: gradual western slopes and steeper escarpments to the east, ge with alluvial basins in between (Maxwell 1979). DH soils are mostly shallow with a high pene f rock fragments (mostly calcium carbonate). Boc nen are frequent and most slopes are steep. There are small, localized areas on alluvial upland ] ghtly deeper, while deep, loamy soils can be found in some basins and near the Rio Gránde. The highest elev STE have soil darker in color because of its high organic content and may be moist more often than dry (Cochran & Rives 1985). Human History and Disturbance Regimes People were living in the Big Bend area as early as the late Paleo-Indian period (12,000—6500 BC) and there is abundant evidence of habitation in the Dead Horse dating from the Archaic period (6500 BC-ca. 100 AD; Alex 1990). The periphery of the DH was most probably used as a travel corridor throughout history until more permanent settlement occurred in the Historic era (1535 AD— Pesos after the threat from hostile Indians was removed, panne and EE activities increased subst ly (Tyler 1975). Human-induced g t have affected DH to a ue degree nice BIRDS Pans resource | f Euphorbia antisvohiliti Boutel lto [m] L wi L cinnabar ore mining, fire, and recreational p it has hiking, — and river "V However, the Fenstermacher et al., Vascular flora ofthe Dead Horse Mountains Fic. 3. Dead Horse Mountains study area locations: (1) Dog Canyon, (2) Dagger Mountain, (3) Stuarts Peak, (4) Old Ore Road, (5) Brushy Canyon, (6) Sue Peaks, (7) Telephone Canyon, (8) Ernst Tinaja, (9) RGV, (10) Boquillas Canyon. [| | fal n. a In LI rr ET [a * F 690 A Ds E BT a Gg d m - E - " J " n Fic. 4a—b. E les of different tat | (psc), lower Telephone Canyon Lu - trail area. B. Sotol-Y (sya), up] i y Fenstermacher et al., Vascular flora ofthe Dead Horse Mountains 691 Vy ni | "NT, | ub: ge H 13 fe, Ne ^ e Mid 5 de, REX oa - e. Gr Jed SÉ Se E di || E FILE A End! tals al [a1 IU HA Aun A J d C Chaparral (cup), | Brushv C y D. Desert IG. xamples o g Canyon (oca), east of Dagger Mountain. Journal of the Botanical Research Institute of Texas 2(1) Das Po” e q a LE m Ge ZAS Tu bis AH e eo” ; We ` d ta GC Ta ch A d LN: d » Ir T hs N i» d ^ - = " wt d vea =. Phe $ A cis ddp: Een EL eg dia ee ah e Dei q "duds e vw " d e a diee m. T: VUN I" "cra DW. | nm. We ql DON E CN ee E, a T 5. UE Y Wie a. CN ^ xe KI E dy, R LES d y hy E Mie A A A é Ce : AU. A ENS. AN S e M: “> y. Fic. 4e—f. Examples of different vegetation zones within the Dead Horse Mountains study area. E. Rio Grande Riparian Corridor (rio), Boquillas Canyon. F. Mixed Desert Scrub and Sotol-Yucca Grassland, Stuarts Peak. Fenstermacher et al., Vascular flora of the Dead Horse Mountains 693 H Fic. 4g—h. E les of different tati ithin the Dead Horse Mountains study G. Sotol-Yucca Grassland, Sue Peaks. H. Chaparral, Sue 694 | | tani ti f Texas 2( most damaging and far-reaching would likely be the i d distribution and dominance of invasive plant species. Tamarix spp. and Arundo donax have expanded their range along the Rio Grande, creating miles and miles of dense thickets lining the riverbanks. Pennisetum ciliare is well established in the lower elevations of BBNP, including the extreme southern end of the study area. However, it is not as dense as it has become in other low-elevation areas, perhaps being held in check by other non-native species, burros and horses. The livestock enter the park from Mexico, trampling vegetation and breaking up biological crusts, increasing the erosion potential in many places along the river corridor. Fire may be a significant natural factor driving the study area ecology, creating the characteristic patchiness of higher elevation vegetation. Human-caused fires are limited to the developed areas but the interior puns are mainly lightning strikes, burning an individual yucen in more DATED rocky areas, but nages or at higher elevations wl denser. Henrickson and Johnston (198 6) suggest that the unique Yucca Woodland community at higher CDR elevations may be maintained by fire, favoring the semi-succulent monocots over more woody species. There has been little suppression effort in the DH, so current states should be considered normal. General Seg Aon PAGNO Many | ion classificatio | ] t to the region. For the CDR asa whole, there exist excellent overviews p basic and detailed — structure (Morafka 1977; Henrickson & Johnston 1986, 2004; Brown 1994; Powell & Hilsenbeck 1995). Descriptions in several local floras (Rogers 1964; Butterwick & Strong 1976; Amos & Giles 1992; Hardy 1997) demonstrate the degree of local diver- sity possible within hierarchies created by the above publications. Beyond a large-scale tendency towards homogeneity (Brown 1994), the DH and larger CDR have basic plant-community types that intergrade and change in abundance and dominance in response to the complex physiography. Perhaps Whittaker, as quoted in Takhtajan (1986), has offered the best explanation for general Chihuahuan Desert vegetation patterns: in warm, semi-arid deserts there is no clear convergence of dominant forms as can be found in cool semi-deserts. Warm deserts may be too dry to support arborescent life-forms or consistent canopy cover. However a diversity of non-arborescent growth is supported in pockets of higher moisture availability, maintained by the right combination of aspect, slope, elevation, and substrate. Larger plants, such as shrubs and Yucca spp., can also create special microclimates. Many species take advantage of the shade and wind protection that these larger nurse plants provide. This could be considered the canopy of the CDR—low and discontinuous, but eeng Two vegetation maps, prod ling, generally include the study area (Plumb 1991, 1992; Wood ¢ et al. 1999) bur are limited by ae E GE and ground- truthing. However they are useful to form an overall impression of the relative importance of cover types. Based on a consolidated version of Plumb's (1991, ee vegetation maps, Lechuguilla i covers e 4% of the study area. This vegetation type includes Agave lechu guilla, Opuntia spp., Eu phorb 1, and many species of grasses. AOS ees important in the study area is the High Desert Grascland vegetation at 45.1% coverage, wl pecies like Nolina erumpens, Dasylirion leiophyllum and Yucca spp. are codominant with graminoids. METHODS The herbaria of institutions that sponsored botanists known to SEN in SE dii: aiu in AA to those Angelo State geographically close to the study area, were d and when p University (SAT), the Botanical Research Institute of Texas (BRID, the New York nica] Garden (NYBO), Sam Houston State University (SHST), Sul Ross State University (SRSC), the Texas A&M University Biology Department herbarium (TAMU), Texas A&M's Tracy herbarium (TAES), Texas State University (SW T), Uni- versity of Kansas (KANU), University of Texas at Austin (TEX-LL), the University of Texas at El Paso (UTEP), and Big Bend National Park (BIBE). Sheet-by-sheet searches were conducted at SRSC and BGWMA. Fenstermacher et al., Vascular flora of the Dead Horse Mountains 695 All species found in flower were collected during every trip. Collection of rare species or plants of uncertain identity and/or distribution was guided by BBNP protocol: no more than 10% of the individuals of any viable reproductive population of a rare plant were harvested. Most collecting was done within the National Park, due to easier and quicker access to more areas. Access to the study area was mainly on foot using cross-country navigation but two rough dirt roads and two maintained trails did provide effective access to more interior areas of the tange: Two river trips allowed limited Md access. A few extended, overnight trips were enabled by water je ly transported into tl try by NPS trail crew personnel and packstock. Sampling areas were chosen based on intricacy of topography such as deep and/or sinuous canyons, unvisited but potentially unique areas such as the higher peaks, and otherwise intriguing spots identified on USGS 1:24,000 topographic maps. Search methodology was a directed meander (Goff et al. 1982) at each locality visited, focusing on promising microhabitats and moving from patch to patch as the observ- able species diversity was documented. Field notes and resulting label data included collection location, habitat, habit, description of notable plant features, relative abundance, and associated species. UTM coor- dinates were taken on site or approximated post-field using digital elevation model data through ArcView 3.2. Specimens were identified using multiple sources, including Correll and Johnston (1970), Warnock (1970, 1974, 1977), Flora of North America Poaceae and Asteraceae volumes (FNA 2003b, 2006, 2007), Powell (1998, 2000), Yarborough and Powell (2002), Turner et al. (2003), and Powell and Weedin (2004). Specimens were determined by B.L. Turner of Pu LL vo A. ee Powell of SRSC. The atlas of Turner et al. (2003) was the pr imary source for an ti l data, and two online sources (ITIS 2007, MBG 2007) were followed for author ER b vee of the Vascular Plants of Texas (Correll & Johnston 1970) provided the fund tal taxon g for this work. However, numerous other sources were utilized for the most up-to-date t ic i tations, most notably Powell (in prep.) and Powell et al. Gn ms.) for non-woody vascular species and — the Flora of North America for Poaceae and Asteraceae treatments (FNA 2003b, 2006, 2007), and Powell and Weedin (2004) for the Cactaceae. Voucher specimens made during this project deposited at SRSC; selected duplicates were sent to BIBE, TEX-LL, and BRIT. RESULTS AND DISCUSSION Between August 2003 and See SE a p of 49 days and approximately 400 hours were spent in the field. The 1549 voucher those three years, plus four observations without vouch- ers, represent 84 families, 283 genera, and 490 p dee totaling 493 taxa at or below the species level. The herbaria search revealed an additional seven families, 58 genera, and 173 species (178 taxa) that were not encountered during the current study. Similarly, the current study documented a itiona families, 45 genera, and 120 species (121 taxa) beyond the historical baseline found in the herbaria search. Fifty-five of those additional 120 species were new records for the DH, representing a 1096 increase in known taxa. Another 65 species (67 taxa), though not previously represented by vouchers, were common and suspected to occur and thus not considered novel records. In sum, the total known flora of the DH to date consists of 91 families, 344 genera, and 662 species (671 total taxa). The Asteraceae is the most prominent family in the DH, tallying 80 species. Other important families include the Poaceae (75 species), Fabaceae (52 species), Euphorbiaceae (34 species), and Cactaceae (31 spe- cies). The eight most species-rich families comprise 5096 of the flora while 24 families are represented by a single taxon. The most abundant genera represented in the DH are Chama d Dalea at 13 species each A taxonomic breakdown of species found in the DH shows that sadios make up the largest propor- tion of the flora at 78.9% (Table 1). Gymnosperms comprise the smallest portion of the flora after monocots and the ferns and fern allies. Perennial herbs were the most abundant life form encountered, followed by annual herbs, shrubs, subshrubs, succulents, vines, and trees (Table 2). The annotated list of species found in the DH provides the following information for each taxon: native status, new records, rare species status [| [| ful n.a DH In LI PR LE Tarima TEAM) 696 TABLE 1. Taxonomic composition of the flora of the Dead Horse Mountains. Taxonomic group Families Genera Species Total Species Native Non-native Ferns and fern allies 3 7 23 0 23 Gymnosperms 3 3 7 0 7 Monocotyledons 10 57 100 7 107 Eudicotyledons 75 277 508 17 525 Totals 91 345 638 24 662 TABLE 2. Summary of the Dead H M tain flora by life form (USDA 2007). The semi-succulent members ofthe Agavaceae, Liliaceae, and li included in the shrub categ viai taxa can be either trees or shrubs depending on the habitat. These divid een the tree and sl shrub categories Twenty-eight taxa were listed by USDA as both annual and perenridi herbs. TE were result evenly — the two herb categories. Graminoid species were included under herbs. Life Form Number of Percent Species of Flora Trees 27 4.1 Shrubs and semi-succulents 97 14.7 Sub-shrubs 55 8.3 Vines 29 44 nual herbaceous 1 0.2 Gen herbaceous 24 3.6 Wood 4 0.6 erbs 420 63.4 Perennial 277 41.8 Annual 143 21.6 Cacti and succulents 34 5.1 and ranking, common name, selected synonyms, relative abundance in the study area, life form, vegetation community, elevation, and all known vouchers. Herbarium search It is certain that more DH collections exist in regional herbaria. The collections of one botanist who made significant collections in the BGWMA reside at BRIT, but are not electronically databased. Unfortunately, time and funding constraints precluded a sheet-by-sheet search of that institution for this project. Digital records are a boon to this type of research; however, when in existence they should not be assumed to be complete or entirely accurate. For example, in researching the holdi t SAT, an onli h of the Floral Consortium of Texas website revealed 64 vouchers, but an onsite secs of SAT’s internal digital database resulted in 245 collections, with only three overlapping with the consortium’s results. Added to that dis- crepancy is the fact that not all of the holdings at TEX-LL are included in their online searchable database. Given that that is the main repository in the state, there is a high probability that many DH vouchers were not accounted for through the herbaria search. Compras f collection efforts in the study area, including those during the past 15 years, there was a significant rise in peda found, without any evident leveling of return for effort. This is to be expected, with much new territory having been covered in more recent years by the Amos survey of Brushy Canyon in addition to the current project. However, the return of new species per year of effort during the course of this study began to taper off and fewer new species were found than remained unseen from the historical Fenstermacher et al., Vascular flora of the Dead Horse Mountains 607 baseline. S historicall llected were not revisited, such as tl inland riparian habitats of the 1 Rio Grande Village area, he species may still be extant or perhaps extirpated due to the ging nature of the river corridor. More localized examination of return for effort tells a mixed story: after three visits to the same area of mixed, mid-elevation habitat, there was no reduction of return for effort. However, after two SC to the high-elevation Sue Peaks environment, the return of new species decreased by about 50%. he differences may lie in the types of habitat visited, but one must also keep in mind the vagaries of desert flora—life strategies built around responses to ephemeral rain, general seasonal/temperature restrictions, ane even bi Enn cycles, all serve to limit the flora prem at any given time. Certainly more species await in the study area; collectio hout most of the Dead Horse range, but some interior locations such as Margaret Basin, the upper Strawhouse trail, the eastern end of Telephone Canyon, and areas interior to Boquillas Canyon remain under collected. Despite this, however, it seems clear that the majority of the flora has been documented. Vegetation Patterns Henrickson and Johnston (1986) comment on the high degree of intergradation between vegetation types in the CDR and the difficulty of placing many areas into an exact classification. This can result in many Sech ee pee cen, as in EE and Plumb's pius) work. In the DH, six vegetation types are clearly 1 levati ilability. These types are similar to ones identi- fied by edison and EECH Ge CR CDR Other CDR ane may have only marginal representation in the DH and are discussed as unique habitat areas. The three dominan ttypes follow elevational zones, with Mixed Desert Scrub (psc) at lower to mid elevations (Figs. 4a, f), Sotol-Yucca Grassland (svo) at mid-to-high elevations (Figs. 4b, f, g), and Chaparral (CHP) EE in i pathos at the highest EE within the sy (Figs. 4c, h). The remaining three vegetation ty] E related areas, i , canyons, and the Rio Grande corridor. The Sandy Desert Arroyo (DAR) and Desert Canyon (ca) Ee el with and intersect mainly the lower elevations of psc but can allow intermixing of higher- and lower-elevation species in steep, protected drainages on mountain slopes (Fig. 4d). The Rio Grande Riparian Corridor (rio) is the only vegetation type that occurs in association with permanently flowing water and mainly occurs within Boquillas Canyon: (Fig. te), ELA the slightly inland, developed areas around RGV have significant riparian habitats that rsity ofthe st study area. Although few plants are seen throughout the range at all elevations, Ger ECH and Aristida spp. are possibly the most ubiquitous. Mixed Desert Scrub.—The psc vegetation type extends from the lowest elevations to about 4000 ft (1220 m), nearing the higher plateaus of the Dead Horse. The lowest elevati ] flattest terrain of the park are dominate d by Larrea tridentata, which is seen most often on low, foothill approaches and interior basins of the DH. Intermingling at lower elevations are Opuntia aggeria and O. grahamii, Parthenium incanum, Nama hispidum, and Acacia neovernicosa. Flourensia cernua is often cited as being important, even co-dominant, in b ecological zone ES the o but it was any seen. DE posue various shrubs, succulents, and I I d l include: Tiquilia greggii, Prosopis STEEN Condalia sp., Zizipl btusifolia, Buddlei iifolia, I hyll 1L. candidum, Vigui- era stenoloba, and Chrysactinia mexicana. Iutergrading with jower-elevarian species is Dasylirion leiophyllum, which begins to occur as low as oo ft (1066 m), but extends to the oe elevations of the range. Dense stands of the rhizomatous Ag illa and Hectia texensis occur sporad dically and d usually include more obvious occurrences of the a common species: Selaginella lepidophylla, Selaginella wrightii, Astrolepis spp., Argyrochosma microphylla, Calliandra iselyi, and Opuntia rufida. Biological soil crusts of varying degrees of development occur in most inter-rock spaces, harboring various species of cyanobacteria, lichens, and in the right environment even mosses. Desert Arroyo.—Many of the general psc species overlap into the dry watercourses intersecting the region; the two xeric-riparian vegetation types are generally distinguishable according to the amount of moisture available. par, or Sandy Arroyo Scrub as named by Henricl and Johnston (1986), occurs in more open, low-elevation drainages incised into the desert flats. TI gi ich with perennial plants, which 698 t tani titute of Texas 2( often grow into impenetrable thorny thickets across the wash. Fallugia paradoxa and Brickellia laciniata are commonly present in the wash cobbles of larger drainages, while Prosopis glandulosa, Guaiacum angus tifolium, Forestiera angustifolia, Celtis pallida, Lycium berlandieri, and Porophyllum scoparium are common throughout. Desert Canyon.—In rockier habitats, larger drainages, and within canyons that provide partial shade, the pca vegetation type includes additional species such as Berberis trifoliolata, Fraxinus greggii, Diospyros texana, Eysenhardtia texana, vids inii deg SPP. Acacia roemeriana, Schaefferia cuneifolia, Rhus virens, Tecoma stans, Clematis drummon , and Maurandya antirrhiniflora. Common wash- associate grasses are Bothriochloa laguroides, Heteropogon contortus, Setaria ina and Aristida spp. More water-dependent species such as Ung E 1 and Chilopsis linearis occur infrequently, restricted to the largest drainages. Juglans mi pa is not a typical DH species within BBNP; it was seen only once in the National Park at the north end of the range, but as reported by Amos and Giles (1992), it was observed during the cur- rent study to be much more dominant in BGWMAS Brushy Canyon. In the same Amos and Giles (1992) report, Cercis canadensis var. mexicana was listed as a dominant in Brushy Canyon, it was not observed in the National Park portion of the DH. Canyon walls throughout the range provide a relatively consistent but limited habitat for cliff-dwelling species including Cirsium turneri, Perityle aglossa, and Perityle bisetosa var. scalaris. Rio Grande Riparian Corridor.—A more traditional riparian woodland M ed SE exists nd along the Rio Grande, e the consistent water table can support native speci hasP Salix spp., Bacc licifolia, Pl ites australis, and tl t d and invasive Arundo dona "€ Dude spp. Á a solitary individus Ba more drenan genera (e.g., Salix, Baccharis, and Tamarix) occur in at least one other place in the DH interior: above Ernst Tinaja, where their water-hungry roots may have found a perched water table. Also important along the Rio Grande are Acacia farnesiana, Nicotiana glauca, Prosopis glandulosa, and Chilopsis linearis. The major, non-native species have altered and will continue to alter the riparian landscape. Thickets of cane with abundantly dense rhizomes have solidified the river banks, channelizing the watercourse. This creates, in many cases, an irreversible cycle: the channel concentrates the river flow, which increases its velocity and thus erosive power, which, in turn, creates a deeper channel that does not allow flooding over its banks as easily as before. With over-bank flooding decreased due to channelization and a generally reduced flow/flood cycle due to upstream dams, dense canopies of Arundo donax and Tamarix spp. are al- lowed to establish, precluding the formation of native Salix spp. and Populus spp. bosques (Howe & Knopf 1991, Bell 1997). Sotol-Yucca Grassland.—Found on the higher elevation slopes and plateaus to the highest peaks, sY is the dominant vegetation above psc. The vegetational boundary seems to occur , atleast on the western side of the range, at around 5000 ft (1524 m), where the steep western slopes top out to form a main central plateau within the DH range. Stuarts Peak, though slightly lower at 4800 ft (1463 m), should also be included, due to its many unique Gen characteristics, o a eios lichen flora and, in the DH, the only known populations of two tl I ted vegetation— Philadelphus microphyllus and gedeien curviflium. 1: 1 1 A P AS IN E «I I ¿hh and srace_dnminated CALLA e CAU MALA ELI LALA AA " AL LR dl + isually VLLOLILIX.L AL understory. Also known as Toda, Sotolal, Ze Rosetófilo, this vegetation type is considered by some to be shrub-dominant (e.g., Henrickson & Johnston 1986), and so the term “grassland” is omitted in their official terminology. The DH do seem to have a Been: shrub ne BEES Dy Fi dps leiophyllum, Nolina erumpens, and Yucca spp., as the g However, the two life-forms seem to be codominant and the term me is used here to convey the importance of bunch grasses in the inter-shrub spaces. On the main DH plateau, Nolina erumpens becomes a significant, visual member of the svc creating, along with the Dasylirion leiophyllum, large masses of thick vegetation, standing in some places almost head high. Fenstermacher et al., Vascular flora of the Dead Horse Mountains 699 Common plants associated with the syc of the DH include Yucca torreyi, Yucca thompsoniana, Nolina erumpens, Fouquieria splendens, Parthenium argentatum, Bouchea spathulata, Leucophyllum spp., Bernardia obovata, Vauquelinia angustifolia, Mortonia scabrella, Zinnia acerosa, Coryphantha tuberculosa var. tuberculosa, Coryphantha echinus, Neolloydia conoidea, Epithelantha macromeris, Bouteloua curtipendula, Aristida spp., En- neapogon. desvauxii, Leptochloa dubia, and Muhlenbergia tenuifolia. True grasslands, though included in reviews of CDR vegetation types, are not functionally a part of the DH t. Areas that are clearly dominated by graminoids without a significant shrub tare more commonly associated with the higher plains around the towns of Marathon, Alpine, and Marfa i in the northern Big Bend. In the DH, grasses are important components of the shrubby slopes and higher plateaus, and at higher elevations, a oe dime m are important components of the vast grama grasslands to the north do occur. However, small areas dominate d by grasses occur only on the scale of remnant patches rather than functional ecosystems; they are not dense or contiguous enough, nor dominated enough by the typical grassland species Bouteloua eriopoda and Bouteloua gracilis, to be considered a true grassland. This seems to follow a general trend of grama grasslands being established on igneous soils, whereas limestone substrates in similar conditions will support only desert-scrub assemblages (Henrickson & Johnston 1986). Chaparral.—Small pockets of shrub thickets, or cup, occur within the syc on plateaus and eastern slopes associated with the high EE M the Sue Peaks area. Small and sometimes dense clumps of Quercus pungens, some Q ind Quercus grisea, Cercocarpus glaberoides, Berberis trifoliolata and Rhus virens associate with large Dasylirion eia anid Nolina erumpens, all overtopped by the occasional but conspicuous Yucca faxoniana. Expected but less frequently seen species include Fraxinus greggii, Garrya ovata, Ptelea trifoliolata, and Muhlenbergia parviglumis. Oaks and other more mesic species occur also at the highest summits and in more protected parts of drainages, even in steep drainages and lower elevations of the western slopes. These micro-niche species include Malacomeles denticulata, Quercus pungens, Fendlera falcata, Fraxinus greggii, Prunus havardii, and Ina ida spp. Junipers, including Juni huil hotii, occur sporadically on the high plateaus amongst the chaparral clumps and, very panes i in ieee, -bwendevatin canyons. Pinyon pines are rare, documented only at a few eastern-slope locations. These species are probably relicts from the previously extensive juniper-pinyon-oak woodland covering the region between 22,000 and 11,000 years ago. The main pine of the DH is Pinus remota, a limestone specialist (Powell 1998). Once more widespread, Pinus remota in the Trans-Pecos is limited to isolated pockets, being more abundant on the Edwards Plateau and in Mexico. Records of Pinus cembroides in the DH are unusual; it is a higher-elevation taxon, although the two species are known to occur to BERE Remum in the Big Bend region (Powell 1998). Unique Habitats.—Several encountered during the course of the study; only the sand dunes at the mouth of Boule Canyon have been explored rather thoroughly. Though not extensive enough to support a full representation of Henrickson and Johnston's (1986) Sand Dune Scrub vegetation type, they have been well botanized and do provide a unique habitat for a few species including the rare Chamaesyce golondrina and the sand-loving species Heliotropium convolvulaceum, Corispermum americanum, and Dalea terminalis. Several other unusual vegetative would no doubt profit from further examination. An isolated bottomland of Pleuraphis mutica covers the western edge of Ernst Basin, in stark contrast to the sparsely vegetated, more elevated flats over the rest of the basin. This tobosa grassland area may rep- resent a small inclusion of Tornillo soils, which was previously unknown in the DH and is interesting not only because Pleuraphis mutica occurs in distinctly limited areas of BBNP, but because several other species were collected only there: Malvella leprosa, Leptochloa panicea, Chamaesyce serpens, and the Texas endemic Lycium puberulum var. berberioides. The possibility of similar habitats, i.e., the existence of Tornillo soils, in other large basins in the DH should be explored, in particular at the northern end of the Ore Terminal trail, in Margaret Basin, and along the Strawhouse trail. The vegetation in a high saddle (5300 ft, 1615 m) just north and west of Sue Peaks was completely different from the immediate, Rosetófilo-type surrounding community. The soil was finer, deeper, and less 700 t tani i Texas 2(1) rocky than the surroundings; old wire, pieces of wood, and prehistoric lithic scatter including point bases and partial tools was found on the ground surface. Many plant species found there are reminiscent of the tobosa grassland area north of the bid Mountains in the Harte Ranch area of BBNP, Pleuraphis mutica, Panicum obtusum, and Hoff in particular. Other species diia ER ES inique com- L C heh "1 ptandrus, A adiado E Aida PEN E YE ; and munity included Sporobo l [S dit video S The other habitats deserving of more attention are at the highest elevations, especially Sue Peaks and Stuarts Peak. Each has different characteristics making them interesting. Stuarts Peak is a rather narrow ridge, with not much area formi ng the actual summit However, many plants li | there were surprising to find in such a seemingly arid location, including Achnatherum curvifolium and Philadelphus microphyllus. The lichen and moss flora on the summit ridge was impressive in its diversity, which was not observed to the same extent anywhere else in the study area. Also notable was that the summit ridge was one of the few places encountered where the cactus Neolloydia conoidea was growing densely enough as to preclude non- damaging travel. The Sue Peaks area exhibits a well developed cup community with shrub oaks being very common in EIA mane DH Gegen area. The peaks themselves are rather bare of large shrubs, those being relegated to d ledges. Several species, ome only in this area, were observed and collected on hatami6um a trip following significant rains in the area: revicaulis, Ipomoea costellata, N i Dichromanthus cinnabarinus, and Mirabilis linearis. These species, in addition to others known only from the peaks or the surrounding area including Asplenium resiliens, Asclepias sperryi, Quercus mohriana, Fraxinus cuspidata, Quercus laceyi, and Pinus remota, indicate that conditions on the whole are relatively mesic and that under favorable conditions, additional unique species may be in evidence. Non-natives, Range Extensions, Rare Taxa, Notable Collections Non-native species constitute a small part of the DH flora. The USDA considers 24 species collected in the DH as exotic (USDA 2007). Six of those species are considered invasive, and three are state-listed noxious weeds: Arundo donax, Tamarix aphylla, and Tamarix chinensis (PTI 2006). Significantly, Pennisetum ciliare, an introduced grass from Africa, is not among the Texas-listed invasives; however, it is a demonstrated threat to native ecosystems throughout the southwest (e.g., TNC 2005) and is listed by the state of Arizona as a noxious weed (AZDA 2005). The fact that it is widely planted in Texas and northern Mexico as a range grass perhaps makes its listing unlikely. Eragrostis lehmanniana is a well-known invasive in the desert southwest, but its presence in the study area appeared to be minimal outside of the river corridor. It seems to require more moisture than is reliably present in the majority of the range, with isolated plants being found near an old header dam and in shady or sandy areas of large drainages. Overall, non-native species constitute 3.6% of the DH flora. Eleven species found in the DH are endemic to Texas, according to the Texas A&M Bioinformatics Working Group and updated taxonomic and distribution information (TAM-BWG 2006). The occurrence of Matelea sagittifolia in Brewster County is not currently reflected on the referenced website, but it was col- lected during the present study and also during a recent survey of the Lone Mountain area of BBNP (Fen- stermacher et al. 2006). Lycium puberulum var. berberioides is also not listed by the Bioinformatics Working Group as occurring in the county but should be; it has been a known dd a the NE flora for some time (Powell 1998). Hedyotis pooleana, though included in this DH-endemi ; i t listed by the Bioinformatics Working Group, perhaps due t nsensus its — (Turner 1995, 1997; Terrell 1996, 2001). Endemic species discount for 1.796 of the DH flora. Many noteworthy collections were made during the study period. The Wildlife Diversity Program of TPWD, in conjunction with the Texas Conservation Data Center of the Nature EECHER of Texas, tracks seventeen rare spectes d were collected in the DH (TPWD 2006). Several of tl ollecte new locations. Bout ly known only from one location east of the DH but it was found to be one of the dominant grasses at eben elevations near Sue Peaks. In the mid 1980s, Hedyotis pooleana was first collected at one locality in the DH and named as a new species. Subsequently it was not seen until the Fenstermacher et al., Vascular flora of the Dead Horse Mountains 701 present study during which it was found throughout the majority of the range in higher-elevation bedrock exposures. Andrachne arida was last collected in the DH in the 1960s, and during the study period it was relocated in the general vicinity of the historical collections. One small population of the diminutive legume Senna ripleyana was discovered, clarifying its status as having remained extant north of the international border since its last documented collection in northern Brewster County in the 1940s. Another historically col- lected taxon Sed ifolium was encountered in shallow soil pockets on the DH plateau. Regionally it was previously known Bg one collection in the limestone mountains of northern Brewster County (Warnock 1977) and has been recently observed on hilltops of the Cox and Guadalupe ranches, about 15 mi (24 km) west of Sanderson (M. Terry pers. observ.). One taxon, a member of the genus Galactia, as yet is not identified to the specific level. It does not fit descriptions of known Texan taxa (B. Turner pers. comm.) and awaits review when appropriate herbarium specimens become available. Populations have been found in several southern DH locations and the plant was encountered more commonly near the Rio Grande within side drainages of Boquillas Canyon. It is a sprawling, procumbent perennial from a central taproot, often growing vine-like within larger, surrounding vegetation. It has dense, silvery pubescence on all vegetative parts but is more pronounced on the leaves, those having prominent under-surface veins. The flowers are pink, the banner exhibits a green-yellow base, and the petals turn a deep blue-purple upon dessication. New to the United States, the Mexican taxon Seymeria falcata var. falcata was collected north of the border originally in 1994 (MT 137 SRSC), but it was not recognized as such mu Ene Ve the current study spurred a review of previous collections of its more widespread 2006). Eight species were newly recorded for Brewster County, according to Turner et al. (2003) and known herbarium specimens: Achnatherum curvifolium, Atriplex rosea, Argemone mexicana, Hedeoma serpyllifolia, Matelea sagittifolia, Maurandya wislizeni, Nama dichotomum, and Rumex maritimus. Three of these species, Atriplex rosea, Arg mexicana, and Maurandya wislizeni, were collected by NPS staff in conjunction with the restoration site at the mouth of Boquillas Canyon. Many of the newly collected species for the DH are taxa that normally occur in much less arid condi- tions at the highest elevations in the Trans-Pecos. Achnatherum curvifolium is a C3 grass previously known from a restricted range in the Guadalupe Mountains of Texas and New Mexico (Powell 2000), and the nearby Delaware Mountains (Worthington 24663 SRSC). de EH is deeg elevation-restricted species, previously known from the Hos pa 1G I ibution ranges from Colorado down through the Mexican highl into t te South America (Correll and Johnston 1970; Turner et al. 2003). In addition to the sparsely occurring Ee taxa of Juniperus and Pinus, several other species were found whose presence was not expected in such relatively arid conditions: Philadelphus microphyllus, and several other C3 grasses Achnatherum eminens, Achnatherum lobatum, Hesperostipa neomexicana, and Melica montezumae. The DH are virtually the only area in the park where the giant dagger, Yucca faxoniana, occurs. Interesting observations were made of dog cholla (Opuntia spp.) populations from the southern end of the study area. Dog cholla species of the southern Big Bend, most as part of the Opuntia schottii complex, have been of taxonomic interest for some time (Anthony 1956). The most recent attention in the literature documented that a long-believed hybrid, purportedly a cross between the tetraploids Opuntia grahamii and O. schottii, was finally determined to be a distinct species, Opuntia aggeria, based on morphology and its diploid nature (Ralston 1987, Ralston and Hilsenbeck 1989). During the current study, however, several observations and collections provide impetus for a fresh look a the taxonomy of southern Big Bend dog chol- las: (1) a triploid hybrid was documented in el Ernst eis area aa 635 SRSU; parents undetermined); (2) Diploid individuals with widely varying her 640, 1574 SRSC) may indicate that Opuntia aggeria has an even broader species content awe prea described by Ralston and Hilsenbeck (1989); (3) a tetraploid cholla was documented at Ernst Tinaja (Fenstermacher 691 BIBE) with morphology 702 ) t tanical Institute of Texas 2(1) dissimilar to both Opuntia schottii and O. grahamii, possibly suggesting a larger distribution of Opuntia densispina in the Big Bend than previously surmised (Ralston and Hilsenbeck 1992, Powell and Weedin 2004); (4) all collections and individuals examined of Opuntia grahamii were sterile, effectively producing no pollen (J. Fenstermacher unpub. data). These findings suggest that the taxonomy of the Opuntia schottii complex is not completely understood; more data from Mexican populations may allow further resolution of the problem. Floristic Analysis and Phytogeographic Connections As a portion of Big Bend National Park, the flora of the DH is 5496 of the park's listed total (NPS 1996) yet is found only on 2496 of the land area. Clearly, this eastern side of the national park with its riverine and high- elevation habitats is an important source of area id RA Manes with regional floras are hampered by their mostly out-of-date checklists, but exploring hips to peripheral floristi d the larger southwest region is possible. One of clearest results of analyzing the DH flora is that while the Asteraceae is the most dominant family E the Southwestern flora (McLaughlin 1986), the Poaceae is co-dominant with Asteraceae in the DH and ot thern Big Bend fl (Fenstermacher 2007). This suggests the connection to the Great Plains grasslands, moving into Texas through the High and Rolling Plains vegetation zones of the Panhandle region. The connection has also been quantitatively demonstrated by Christie (2006). The Fabaceae is also important in the Southwest, but more important locally in the DH. This family comprises a larger percentage of the species of the SW flora than in the DH, yet the DH have more fabaceous species (52) than the average for SW floras (28). In fact, only five DH genera make the Southwestern flora's top-twenty list, and they do not include Dalea, one of the two dominant genera of the DH. The local im- Poen of the Fabaceae, in addition to the many other resident thornscrub species like Aloysia gratissima, ophyllum spp., may indicate the importance of the Tamaulipan Thornscrub flora (WWF 2001). Indeed, for the few Trans-Pecos individuals of the more Tamaulipan species, Acacia berland- ieri, Croton torreyi, and Karwinskia humboltiana, the Rio Grande corridor has most likely provided a conduit of limited habitat that enabled these species to extend their distributions upriver into generally more arid surroundings. These connections to peripheral vegetational areas suggest that the CDR, and by extension the DH, are not simple extensions of the Sonoran region, as Cronquist suggests (1982). Rzedowski (1973) notes that both desert areas have several endemic genera (21 Sonoran, 16 CDR), which could suggest a long period of isolation and autochthonous development. In the same paper, comparisons of similarity index analyses show that the CDR flora has more affinity to the southern end of the Sierra Madre Oriental than it does to the Baja/Sonoran flora. Perhaps because there do not seem to be distinctive topographic limits to northern vegetative expansion, the limits of the CDR may be more climatically controlled (McLaughlin 1989): CDR flora shows more ai to coastal Peru than to the Great Basin (Rzedowski 1973). Climate as a limiting factor becomes iff t when realizing that the Great Basin, Sonoran, and Chihuahuan deserts are linked topographically and geologically as members of the Basin ena Range province. Individual species’ tolerances are probably what d d l changes in response to the changing climate. However, as McLaughlin (1986) points out, dies is no reason why, on a larger scale, there cannot be found coherent assemblages of species occurring in non-random patterns. Many floristic regions, delineated as places where the climate acts upon the vegetation (as opposed to more local vegetational areas, using delimiters like soils or geography), have been applied to the Southwest (Dice 1943; Cronquist 1982; Takhtajan 1986; McLaughlin 1986; Bailey 1998). The boundaries of these large-scale, ecological regions can be drawn strictly using climate and gross vegetational features (Bailey 1998), by using centers of endemism to create distinct groupings (Cronquist 1982; Takhtajan 1986), which could be called a “scarceness” model, or the focus could be on the degree of overlap between individual floras throughout a region (McLaughlin 1986, 1989). This latter model does not focus on the dominant or rare taxa; rather, it takes into account all extant species and is a method based on “commonness.” To s end, EE (1986, 1989) work has provided floristically-based evidence torical distinctions, intuitive or otherwise. Interestingly, the area of soutien er and es New Mexico, where Henrickson and John- Fenstermacher et al., Vascular flora of the Dead Horse Mountains 703 ston (2004) draw an arbitrary border of the CDR because of its perceived transitional nature, is shown by McLaughlin (1989) to be part of a distinct floristic area originally intuitively recognized by Dice (1943). McLaughlin's (1989) similarity indices identified an outlying connection between his so-called “Apachian” flora and the Davis Mountains region of the Big Bend, which Turner (1959) actually suggested in his floris- tic interpretation of the area. Also in support of this connection, the Davis Mountains, based on a subset of its flora (Larke 1989), has a family profile more similar to that of the Southwest (McLaughlin 1989), and is located more closely to the core of the Apachian element than to floras in the southern Big Bend. The Apachian floristic element is thought to be autochthonous but allied to the Sierra Madre Occidental (McLaughlin 1989). Perhaps during the expansions and contractions of vegetation communities in response to Pleistocene climate change, this Apachian center extended its influence into the Big Bend. As many as 25 glacial-interglacial periods have affected the southwest (Henrickson & Johnston 2004), allowing more temperate plant species to expand into the Big Bend region from both the north and the south. In the Chisos Mountains of BBNP, cuan menziesii and EE Ee occur with the Mexican montane species Cupressus arizonica and what is believed to be Pi (Powell 1998). In the DH, cooler-climate species that may be remnants of glacial expansion include Achnatherum curvifolium from the north and the Mexican pinyons from the south. la warmer times, the desert scrub and more tropical species expanded from the south. These climate fluctuations could have allowed for speciation through isolation, but there would have been more ities for diversification in the southern CDR, away from more extreme climate changes coming Gas, the continental interior (Van Devender 1986). Of course, any discussion of speciation must acknowledge that the process is not simply a function of time or climate. Life forms, breeding systems, and genetics are also critical in producing differential speciation abilities (Grant 1958). However, perhaps more influential than genetic isolation, extinction may have been a more effective mechanism than speciation in affecting today's floristic diversity in the Big Bend. More floristic study is warranted for the region. Beyond simply improving baseline knowledge of the Trans-Pecos flora, previous floristic work (McLaughlin 1986, 1989) should be revisited and improved by updating the original data, since the size of several Trans-Pecos floras have increased by 40-50% since the 1980s (e.g., Warnock 1967 in McLaughlin 1986 and Worthington 2001). Regionally, similarity index studies similar to those by McLaughlin (1986, 1989, 2007) and especially Christie (2006) could be used to elucidate in more detail the floristic influences on the Big Bend region and might help to identify more concretely the distincti g CDR provinces. In this context, the completion of the present Dead Horse Mountains flora constitutes a cca step towards a more complete understanding of West Texas and regional floristic relationships. ANNOTATED CHECKLIST OF VASCULAR PLANT TAXA Species are arranged by divisions following Raven et al. (2003): Lycophyta, Pteridophyta, Coniferophyta, Gnetophyta, and Anthophyta. The flowering plants are subdivided into the classes Monocotyledones and Eudicotyledones. Within these hierarchical groupings, taxa are listed alphabetically by family, genus, spe- cies, and infraspecific rank as appropriate. Beyond a basic reliance on the Manual of Vascular Plants of Texas (Correll & Johnston 1970), taxonomy and nomenclature were taken preferentially from several updated sources, including Powell (in prep.) and Powell et al. (in ms.) for non-woody vascular species and genera, the Flora of North America volumes for the Poaceae and Asteraceae (FNA 2003b, 2006, 2007), and Powell and Weedin (2004) for the Cactaceae. Online databases were consulted Ke currency, EE and Se EIS 2007; MBG 2007; USDA 2007). Turner et al. (2003) thef tal an tributi | data. Specimens found through the herbaria search were assumed to be correctly identified except those that gege to Turner et al. (2003) would have been far outside of their normal range. These were requested on loan, examined, and included or excluded from the final flora based on the resulting determinations (Fenster- macher2007). Native status, com (USDA 2007). Locally used common names from regional literature were included if not utilized by the USDA. 704 LI RT adi f Texas 2(1) li a +l TABLE 3. Index to | ¡0 | de -l Cua y aired. ACK AC. Koelling DM D. Miller JM J. Masters RE R. Eckhardt AD A. Denyas DS D. Seigler JMM JM. Milter RH R. Halfmann AG . Grimes [252 D. Smith JMP J.M. Poole RLH R.L. Hartman AL A. Freeman/A. Leavitt | DSC D.S. Correll JS J. Sirotnak RLU R.L. Ufkes AL2 A Lewis ECM . E.Castro-Mendoza |JT J. Todd RN R. Nichols AMP AM. Powell EGM D. Marsh KJC KJ. Castro RRI R. Rose-Innes AT A Traverse EJL E.J. Lott LCH L.C. Hinckley RS R. Studhalter B Blevins EW Eula Whitehouse LEB L.E. Brown SJ S. Jones BA B. Alex GJG G.J. Goodman LF . Fowle SMU ` SMU-DMNH BA2 . B. Amos GJ G. Jones LJT LJ. Toolin SAP S.A. Powell BAR BA. Ralston GLB G.L. Bradley LS L. Slauson 55 S. Sikes BCT BC Tharp GLW ` GL Webster MA M. Anthony TA T. Anderson BD B. Dodson HBC H.B. Correll MC M. Cole TE T. Emerson BDGL BDG. Leopold HBP H.B. Parks MO | MC. Johnston TW TJ.Watson BGH BG Hughes HC H. Croad MD M. Darrach TK Tony Klein BHW BH Warnock HCC | HC Cutler MLP ML. Powell TM T. Mollhagen BLT B.L. Turner HCH | HC Hanson MPG M.P Griffith TR T. Rogers BM B. Moon HKB H.K. Beuchner MSY | MS. Young TRV T.R. Van Devender BPM Billy Pat McKinney HL Hillary Loring MT M. Talbot TXDOT Tx Highway Dept. BR B. Rector HTF H.T. Fletcher MT2 M.Turner UTW UT. Waterfall BRM BR McKinney IMJ I.M. Johnston MWB | M.W.Bierner VB V. Bailey CA C. Adkins JAM JA. Moore MY M. Yuhas VLC L Cory CAB C.A. Bane JAS J.A. Steyermark OCW | OC Wallmo WBM WB. McDougall CMR CM. Rowell JE J, Clark OES O.E. Sperry WEH WE. Hall CN C. Newman JDA J.D. Allen PAS PA. Smith WFB WF. Barr DAL DA. Louie JD8 J.D. Bacon PC P. Cruze WFM WF. Mahler DAZ DA. Zimmerman JEA J.E. Averett PDW PD.Whitson WGD WG. Degenhardt DB D. Baker JF J. Fenstermacher PM P Manning WH W. Hodgson DB2 D. Benham JFW Weedin PM2 P McNeal WOD ` Justice WO. Douglas DF . Flyr JH J. Hard PVW . DV Wells WRC R. Carr DG D. Giles JJ J. Jernigan RAH | RA Hilsenbeck DHR D.H. Riskind JLB LL. Blassingame RB R. Baker = non-native; ! = invasive non-native; !! = noxious Various symbols precede the species entries: non-native (PTI 2006); A = observed but not collected; 8 = previously collected, but not during the current study period; + = first voucher of a common or expected taxon; # = newly documented taxon for the DH; © = species is shown to occur in the study area in Turner et al. (2003) but no voucher was found during the herbarium search; e = species is not shown to occur in study area in Turner et al. (2003) but previ- ous voucher was discovered during herbarium search; = new county record; species name = new park record; bold type = rare (TPWD 2007). An explanation of rankings for rare species can be found in Poole et al. (2007). Abundance categories were based on Palmer et al. (1995) and ranked as follows: abundant = dominant or co-dominant in one or more common habitats; common = easily seen or found in one or more common habitats but not dominant in any common habitat; occasional = widely scattered but not difficult to find: uncommon = difficult to find with few individuals or colonies but found in several locations; rare = very difficult to find and limited to one or very few locations or uncommon habitats. Any historical abun- dance observations taken from label data are placed within quotations. The terms used historically were not changed (e.g., from “infrequent” to “uncommon”), because the previous collector’s frame of reference or personal distinctions of scale categories is unknown. Many species may have a “rare” abundance within the study area but are not tracked by TPWD or TNC as being imperiled state- or worldwide and, thus, are not listed in bold and do not have a G or S ranking. Vegetation communities: psc = Desert Scrub; par = Desert Arroyo; DCA = Desert Canyon; RIO = Rio Grande Riparian Corridor; sy = Sotol-Yucca Grassland; cup = Chaparral. Two vegetation communities listed Fenstermacher et al., Vascular flora of the Dead Horse Mountains 705 together (e.g., srG/cHP) indicate that the species was found in an intermediate matrix of vegetation types. More detailed habitat/substrate information, specific collection localities, and other notes are included in the MS thesis manuscript (Fenstermacher 2007). All collections made during the current study are housed at SRSC. Selected duplicates were sent to BIBE, TEX-LL, and BRIT. All SOC glia in the annotated list are grouped by the responsible herbarium. One hundred and sixteen collect contributed to the flora of the DH; initials were used for the names taken from the databases and are listed alphabetically by the first letter (Table 3). LYCOPODIOPHYTA Selaginellaceae rion GE ips & d Spring, pisa nial, bsc, DCA, sva, 2200-4200 ft; BA? Se nes MC 158 ae) pe 18407, 10760, Er 15988, JF 253, MT 99, 420 (SRSC); BHW 10760, BHW & RRI 562, DSC & BHW 14980, RRI & BM 1316a (TEX-LL 8 Selaginella pilifera A. Braun, Resurrection plant perennial; A o wrightii Hieron., Wright spikemoss, occasional to mon perennial, psc, DCA, sys, 3500-4000 ft; WFM 43 Ger JF 458, 912, BHW 10827 & 18409, BHW & MCJ 15985 (SRSC); BHW 10827, DSC & BHW 14981 (TEX-LL). PTERIDOPHYTA Aspleniaceae +0 lla resiliens Kunze, Blackstem spleenwort, rare perennial, sv, cHP, 5680-5840 ft; JF 1279, 1993 (SRSC). Pteridaceae Argyrochosma limitanea (Maxon) Windham ssp. mexicana FRA ViAr II C | " E] | pu iunmneconm mon perennial herb, DCA, sva, CHP, 2120-5840 ft; JF 270, 432, 529, 675, 1504, 2023, 2198; AMP & SAP 6142 (SRSC). ociosa microphylla (Mett. ex ini Windham, Small leaf false cloakfern, occasional! nial, DSC, sva, CHP, 2780-5000 ft; JLB 2957, BA2 5392 SEN JF 183, 484, 527, 768, 1175, 1198, BHW 10822 (SRSC), BHW 10822, BHW € LCH BG-143 (TEX-LL). Astrolepis SE (Goodding) Benham E Windham, Co- chise scal DSC, DCA, SYG, CHP, 1800-5500 ft; WFM 58, BHW & Hen (BGWMA); PVW s.n. (KANUJ; JF 305, 953, 1232, 1257, 1546, 2098a, JF & MD 462, AMP & SAP 61434, 6160, MT 54, BHW 10829, BHW & LCH BG- 131, PENEN aso lee BHW e LL). Astrolepis integerrima (Hook. & Windham, Hybrid cloakfern, common geet DSC, SYG, 1900-4320 ft; BA? 5393, 5578, 5579, JLB 938, 2599, HC 1, 32, 34 (SAT); JF 206a, 433, 487, 680, 952, 1054, 1138, 1489, 1496, 1547, 2043, AMP & SAP 6143b, 6161, TR 88, 464, MT 55, BHW 8: MCJ 16068; (SRSC); DSC & BHW 14982a (TEX-LL) Astrolepis sinuata (Lag. ex Sw.) BERI & Windham, Wavy ca "ld , DAR, DCA, 3050-3560 ft; WFM s.n. (BGWMA); JLB 939, 5/6510 (SAT); JF 535,1125, 1156 RO. DSC & BHW 14982 (TEX-LL). + See windhamii Benham, Windham cloakfern, un- mon perennial, psc, svc, 2200-4700 ft; JF 1697, 1705, e 2202 (SRSC) Cheilanthes alabamensis (Buckl.) Kunze, Alabama lipfern, rare ebbe. DCA, SYG, 4650 ft; JF 2199, MT 229 (SRSC); MT 229 (TEX-LL). 8 Cheilanthes bonariensis (Willd.) Proctor, Golden lipfern, perennis aa ft e Tu GE Cl n lipfer | DSC, SYG, 3/00-4960 E y 767, 1201, 1501, MT 116, BHW 20629 (SRSC). Cheilanthes feei 1. Moore, Slender lipfern, occasional perennial, 50-4900 ft; JF 533, 758, 1127, 1200, 1500, 1708, 19, BHW 10765, 20605 (SRSC); BHW 10765 (TEX-LL). Cheilanthes horridula Maxon, Rough lipfern, occasional pe- rennial, DAR, DCA, RIO, 1850-3050 ft; CMR 2602 (BGWMA); BA2 5498, 5580, HC 36, 40 (SAT); JF 670, 1059, 1141, 1611, 2046, TR 465, BHW 18412, 20607 (SRSC); DSC & BHW 14977 (TEX-LL). Nee villosa Davenp. ex Maxon, Villous lipfern, uncom- n perennial, pca, 3560-3800 ft; JLB 2596, 3801 (SAT); JF n. BHW 10757 (SRSC); BHW 10757, DSC & BHW 14983 E LL). i E Lë L c | j I LE , fare perennial; JLB 2597, 2956 (SAT). S Notholaena greggii (Mett. ex Kuhn) Maxon, Gregg cloak on perennial, pca, 1800-4300 ft; CMR 2600 (BGWMA); JLB 2600 (SAT); BHW 10841, BHW 20897, BHW & LCH 16061, BHW & MCJ 16859 (SRSC); DSC & BHW 14978, DS 1096 (TEX-LL). Notholaena nealleyi Seaton ex 3.M. Coult., Nealley cloak fern, rare perennial, psc, pca, 3650 ft; JLB 3806 (SAT); JF 479 (SRSC). IL zl J l 4 A AA Ll, E re perennia al, DCA, 1850— 3560 EJE 603, 1088 (SRSC); DSC 14984, DSC 8 BHW 14979 (TEX-LL). Notholaena dei Maxon, Star cloak fern, occasional peren- nial, DSC, DAR, DCA, SYG, 2880-4360 ft; JLB 2595, KIC 176, HC 35, ECM 251 (SAT); JF 604, 867, 1126, 1007a, BHW 9157, BHW & LCH 15987 (SRSC); DSC & BHW 14985, BHW 9157 (TEX-LL). Pellaea atropurpurea (L.) Link, Purple clifforake, rare perennial, SYG, CHP, 3500-5680 ft; JF 1285, 1299a, 2200, BHW & LCH BG-144 (SRSC) Gi D 'H RAAT hn lar | AAA LI: EE H ex Kuhn, , fare perennial, sra, Cup, 2560-4900 ft; JF 1199, 2201, MT 119 (SRSC). BHW 559 (TEX-LL). CONIFEROPHYTA Cupressaceae Juniperus coahuilensis (Martínez) Gaussen [Juniperus eryth- rocarpa Cory], Rose-fruited juniper, rare shrub-tree, che, 4600-5470 ft; JF 1240 (BIBE); JF 1037, 1219, 1240, MT 237 (SRSC); BA2 5370 (SAT). 706 Juniperus pinchotii Sudw., Red-berry Juniper, uncommon shrub-tree, DCA, svc, CHP; 3050-5680 ft; PVW s.n. (KANU); JF 499, 1048, 1136, 1298, 1913, AMP & SAP 5135, BHW & LCH BG-157 (SRSC) Pinaceae 8 Pinus c Zucc., Mexican pinyon, rare tree, svc, ca. 4200-5300 ft; WFM 54 (BGWMA); BA2 5229 (SAT); BHW Pinus remota (Little) D.K. Bailey & Hawksw., Papershell pinyon, rare tree, sya, 5200 ft; JF 1249, 1310, MT 88, 258, AMP 5492 (SRSC) GNETOPHYTA Ephedrac 8 jul anita Berland. ex C.A. Mey., Clapweed, ent" shrub, psc, 3520 ft; CMR 17227 [BIB iiid SRSC: o a FF ] J O | ex $ Mo. LD Benson Rough jointfir n shr Ub, DSC, DAR, DCA, SYG, CHP, 1900-5250 ft; OES Ges ege BA2 5710, ECM 72, CMR 17227 (SAT); JF 349, 455, 474, 563, 578, 723, 745a, 806, 1003, 1237, 1438, 1485, 1665, 2052, JF & MD 583, OES 1365, 1743, BHW 1742 (SRSC). Ephedra trifurca Torr. ex S. Watson, Longleaf jointfir, common DSC, DCA, 1800-3500 ft; EGM 143, ACK 830 (BIBE); HCC 1856 (NYBG); CMR 14224, 17226 (SAT); JF 1388, BHW 47059 (SRSC); BHW & RRI 557a, 57b, 47059 (TEX-LL). ANTHOPHYTA, Monocotyledones Agavaceae HA Agave havardiana Trel., Havard century plant, rare shrub/ mi-succulent, sra, 5840 ft; South end of southern Sue Peak. Agave lechuguilla Torr, Lechuguilla, abundant shrub/semi- culent, psc, DCA, svc, 1800-5840 ft; BA2 5264, DB s.n. (SAT); JF 257, TR 78 (SRSC); MSY s.n. (TEX-LL). rr Amaryllidaceae Cooperia drummondii Herbert, Evening rainlily, uncommo perennial herb, psc, svc, cHP 3150-5840 ft; BA2 5686 Ge? JF 1102, 1984 (SRSC ^ nial herb, psc, svo, 3500-4840 ft JF 584, BHW 1080- 178 (SRSC). Bromeliaceae Hechtia texensis S. Watson [Hechtia scaricsa L.B. Sm], Texas false agave, occasional shrub/semi-succulent, rock, psc, 1800-4300 ft; DG 219 (BIBE); BA2 5253, 5273, DG 219, MC s.n. (SAT); JF 328, 1002, 1004, OES 1323, BHW 13036, 14047, 20625, 21206 (SRSC); AT 2242 (TEX-LL). Commelinaceae a erecta S e dayflower, rare perennial herb, DAR, Dsc, 2300-2400 ft; BRM 3032 (BGWMA); JF 1445, CW 507 (SRS S Tradescantia wrightii Rose & Bush var. glandulopubescens B.L. Turner, Wright spiderwort, rare perennial herb, 3800 ft; BHW 10775 (SRSC & TEX-LL). | | | E al m... a im LI Sl f Texas 2(1) Cyperaceae C iculata FJ. Herm. [C K | Schiede sedge, uncommon perennial, sve, 3000-5250 ft; BHW & LCH BG-136 (BGWMA); JF 2193 (BIBE); BA2 5219, 5305 (SAT); JF 772, 1193, 1236, 2193, MT 69, BHW 10821, 10847, BHW & LCH BG-136 (SRSC); JF 1236, BHW 10821 (TEX-LL). § Gusta mariscus (L.) Pohl ssp. jamaicense (Cra ic Kü See ft; BHW 13068 (SRSC). +0 Cyperus GE L, Royal flatsedge, rare perennial, DAR, 2400 ft; JF 1449 (SRSC). 8 Cyperus E Smooth flatsedge, “rare” perennial, DCA, 1800 ft; BHW 772 (SRSC). § Cyperus odoratus L., Fragrant flatsedge, “infrequent” annual or perennial, rio, 1800 ft; CAB 31 (BIBE); JS s.n., EGM 162m, BHW 13046, 21441 (SRSC); BHW 327 (TEX-LL). § Cyperus ide L., Bearded flatsedge, “rare to infrequent an 800 ft; BHW 774 (SRSC €: TEX-LL). S Bo. idis lata (L) Roemer & J.A. Schultes [Eleocharis caribaea (Rottb.) S.F. Blake], Canada spikesedge, "infre- quent” annual, rio, 1800 ft; BHW 13047 (BIBE €: SRSC); BHW E iod MESA: ES bad n bes "| | $ af , frequent” 3 AL P ual, RIO, , 1800 ft; HTF 288 (RSC). § Ee palustris (L) Roemer & J.A. Schultes [E cotar macrostachya Britton], Common spikerush, "freq perennial, ca. 3000 ft; BHW & MCJ 16673 (SRSC). § Fuirena simplex Vahl var. aristulata (Torr) Kral, Western umbreila-sedge, uncommon perennial, no, 1800 ft; BA2 4605 (SAT); BHW 21444 (SRSC & TEX-LL S Schoenoplectus pungens (Vahl) Palla [Scirpus americanus . var. longispicatus Britton], Common threesquare, perennial, OES 1350 (SRSC). Juncaceae 8 Juncus acuminatus Michx., Tapertip rush, perennial, 1800 ft; CMR 17236 (SAT). S Juncus bufonius L., Toad rush, rare annual, rio, 1800 ft; BHW C298 (SRSC & TEX-LL Liliaceae $ Allium kunthii G. Don, Kunth onion, rare perennial herb, sva, SCH 5500 ft, i i Eu Apud Trel., Sotol, semi- succulent DSC, DAR, , CHP, Ca. 3000-5840 ft WFM s.n, TR s.n. CUN. po d (BIBE); JF 860, OES Lf .R. (SRSC). Nolina erumpens (Torr) S. Watson, Foothill beargrass, shrub/ semi-succulent, sra, CHP, 4320 ft; BA2 5704 (SAT); JF 128, 821, AMP & SAP 5127, TR 130, MT 188, 190 (SRSC); BHW & LCH BG-167 (SRSC & TEX-LL). S Nolina texana S. Watson, Texas sacahuista, rare shrub/semi- succulent; 3500 ft; BHW & LCH s.n. (BGWMA). Nothoscordum bivalve (L.) Britt., Crowpoison, rare perennial herb; svc; BHW 10773 (SRSC). Collected in 1952 at the head E FO Camela S Y shrub/semi-succulent, DSC, DCA, 3520 ft; BA2& DG 4861, BHW 877 (BIBE); BA2 4861 (SAT); BHW 877 (SRSC); BHW 877, s.n, MSY s.n. (TEX-LL). Fenstermacher et al Vascular flora of the Dead Horse Mountains § Yucca faxoniana Sarg. Giant dagger yucca, occasional shrub/ mi-succulent, psc, sv, CHP, 3500-5840 ft OES 1322 (BIBE); BRM s.n. (BGWMA); SMU 162 (SAT); WH, TA, LS, & JP 9024, OWS 1322, MT 124 (SRSC); RN E-4-6-92, 5-8-92, BHW Lot 45 (SRSC & TEX-LL). Yucca rostrata Engelm. ex Trel., Beaked yucca, occasional ub/semi-succulent, osc, 2150-3400 ft; PVW s.n. (KANU); WH, TA, LS & JP 9025, JF 849, TR 129, MT 50, BHW & LCH BG-62 (SRSO); RN E-5-08-92, 4-6-1992 (TEX-LL). § Yucca thompsoniana Trel., Thompson yucca, occasional ois Se MSY s.n. TEX- LD. , Torrey yl ICCa, culent, DSC, DAR, DCA, SYG, 2100-3500 ft; SMU 164 n WH TA, LS & JP 9026, JF 683, 1723 (SRSC); BHW 9940 (TEX-LL). Le Orchidaceae # Dichromanthus cinnabarinus (Llave & Lex) Garay, Scarlet ladies' tresses, rare perennial herb, svc, 5500 ft; JF 1958 (SRSC). Poaceae 4% Achnatherum curvifolium (Swallen) Barkworth [Stipa curvifolia Swallen], SE ricegrass, rare perennial erb, sva, 5000 ft; JF 735a (SRSC). Achnatherum eminens Ge Barkworth [Stipa eminens Cav.], Southwestern needlegrass, rare perennial herb, psc, zu 3400—5300 ft; JF 366, 1990; BHW 1 10768 (SRSC); BHW 10768 (TEX-LL). II AR i D LC: A) m C i len], Littleawn needlegrass, uncommon n perennial herb, DSC, SYG, 0-5 ft; JF 760, MT 146, BHW 10771 (SRSC); 8 Andropogon glomeratus (Walt.) B.S.P, Bushy bluestem, perennial herb, rio, 1800 ft; CAB 30 (BIBE). +0 Aristida adscensionis L., Six-weeks threeawn, rare annual herb, svc, 5300-5840 ft; JF 1304, oe 1987 (SRSC). Aristida Ge Nutt. var. purpurea [Aristida roemeriana cheele], Purple threeawn, occasional perennial herb, DSC, DAR, DCA, 1800-3400 ft; JF 173, 325, 344, 1327, 1382, 2079; MT 71 (SRSC). ip^ purpurea Nutt. var. nealleyi (Vasey) Allred [Aristida uca (Nees) Walp.], Blue threeawn, common peren- Ge herb, Osc, DAR, DCA, Su, 2250-4800 ft; JF 486, 811, 935, 1178, 1192, 1536, 1537, CMR 5069, MT 70, 160, BHW 10758, 10811, 10816, 10835, BHW & LCH 8G-159, 191, 193 (SRSC); GJG & UTW 4628, BHW 10758, 10811, 10816, 10835, 13028 (TEX-LL). § es purpurea Nutt. var. wrightii (Nash) Allred, Wright eeawn, perennial herb; 3500 ft; BHW & LCH BG-159. oris ternipes Cav., Spidergrass, "frequent" perennial herb; 800 ft; BHW 20944, 20946; BHW & LCH BG-186 (SRSC). !+ Arundo donax L., Giant reed, abundant perennial herb, rio, DCA, 1800 ft; JF 2039 (SRSC). +0 Blepharidachne bigelovii (S. Wats.) Hack. Bigelow desert- grass, rare perennial: Jen DSC, eech JF 355 (SRSC). | bluestem, uncom- mon perennial herb, DAR, SYG, RIO, ers 4400 ft; BHW 779 (BIBE); JF 1534 (SRSC) Bothriochloa laguroides (DC.) Herter ssp. torreyana (Steud) 707 Allred & Gould, Silver Beardgrass, occasional perennial herb, DAR, pca, 2350-4360 ft; CMR 5096 (BGWMA); JF 358, 531,937, 1113, 1371, 1377, 1531, MT 198, 224, BHW s.n., BH & BLT 8320 (SRSC); BHW 21514, BHW & BLT 8320 (TEX-LL). 8 Bouteloua aristidoides (Kunth) Griseb., Needle grama, "fre- quent"annual herb, psc, uo, 1800 ft; BA2, DG, TE, & PC 45974, BHW 13034 (BIBE); BHW 12894, 13034 (SRSC). Bouteloua barbata Lag., Sixweeks grama, rare annual herb, RIO, DCA, 1800 ft; BA2, DG, TE, & PC 4597b (BIBE); JF 2096 SRSC). Bouteloua EE (Michx.) Torr. var. edet Gould BE Kapadia DCA, SYG, CHP, 1800- -5250 ft; JF 254, 1107, 1176, 1238, 1393, 1512, 1524, 2139, 2166, MT 38, 162, BHW 10780, BHW & LCH -190, BHW & BLT 8329 (SRSC); BHW 10780 (TE + Bouteloua potes e Torr, Black grama, rare perennial herb, sra, 4400 ft; JF 1533 (SRSC). talla trail are + SEH Ge Lag, Hairy grama, occasional "— nial , P, 3300-5500 ft; JF 1221 EE J 1027, 1032, 1221, es 1319, 1523, MT 46, 80, 131, 175 (SRSC). Bouteloua kayi Warnock, Kay grama, uncommon (6151) perennial herb, psc, sya, cup, 3500-5800 ft; JF 2186 (BIBE, SRSC, & TEX-LL); JF 296, 1169, MT 134, 182, 191, OCW 318, BHW 20621 (SRSC). Bouteloua ramosa Scribn. ex Vasey, Chino grama, abundant perennial herb, psc, DAR, DCA, SYG, CHP, 1800-4400 ft; CMR 5075 CUM JF 191, 338, 443, 930, 1160a, 1331, 1452, 1544, 1545, 2009, 2152, BHW 10853 (SRSC) Bouteloua trifida Thurb, Red grama, occasional to “frequent” ennial herb, psc, DAR, DCA, 1800-3360 ft; JF 341, 1398, 1406, 1474, 1476, 1543, 2144, BHW 12341, 21190, BHW& LCH BG-183 (SRSC); TRV, GLB & WEH 85-38 (TEX-LL). Cathestecum erectum Vasey & Es False grama, occasional perennial herb, psc, 1800-2450 ft; JF 1340, MT 121, 141, BHW 10813, 12900, BHW & e 16842 (SRSC); BHW 110813, Bi ERHI av. [Cenci M.A. Curtis], Coastal piss locally frequent annual or perennial herb, rio, pca, 1800-2400 ft; BA2, DG, TE, & PC 4601 (BIBE); AMP & SAP 5862, BHW 23037, BHW & MCJ 16844, BHW & BLT 8322 (SRSC); BHW 16844 (TEX-LL). Chloris virgata Sw, Feather fingergrass, rare annual herb, psc, 1800-3440 ft; JF 364 (SRSC); EGM 51-1129 (TEX-LL). ! Cynodon dactylon (L.) Pers, Bermudagrass, locally abundant perennial herb, pca, rio, 1800-2350 ft; BA2, DG, TE, & PC 4610 (BIBE); JF 1336 (SRSC); EGM 141, 51-1115 (TEX-LL). Dasyochloa ER ER KA EN Rydb. EEN pur L i fluffgrass, occasional perennial herb, DSC, DCA, 1 900-2760 ft; BA2, DG, TE, & PC 4595 (BIBE); JF 194, 678, 1338, 2025, 2049, did LCH BG- ! 98 (SRSC). Di A ttontop, uncom- mon perennial herb, DSC, DAR, DCA, SYG, CHP, 3000- 4000 ft; CMR 5095 (BGWMA); JF 335, 839, BHW 10801 (SRSC); BHW Sé Ge CES E her b, 1800 ft; BHW 775 (BIBE, SRSC, TEX-LL). +0 Digitaria pubiflora (Vasey) J. Wipff [Digitaria cognata , fare annual 708 J.A. Schultes) Pilger var. pubiflora Vasey ex L.H. Dewey], Carolina crabgrass, rare perennial herb, pca, 1800 ft; JF 212 T (RC). * Echinochloa colona (L) Link, Junglerice, uncommon an- nual herb, DAR, uo, 1800-3440 ft; DG & TE 4611 (BIBE); JF 1330, EGM 139, BHW 793 (SRSC); EGM 51-1113, BHW 795 (TEX-LL). § Echinochloa muricata (Beauv.) Fern, Rough barnyardgrass, "rare" annual herb, rio, 1800 ft; BHW s.n. (SRSC), BHW C299 a Lam ray Baan occasional perennial herb, DAR, DCA, SYG, 1800- 5000 0 ft: JF 193, 231a, 347, 1510, 1551, 2097, BHW 10770, 12897, BHW & MCI 16865 (SRSC); BHW 10770 (TEX-LL). Eragrostis intermedia A.S. Hitchc, Plains lovegrass, rare pe- rennial herb, psc, pca, 2500-3800 ft; JF 1894, BHW & MCJ 15984 (SRSC). | REG lehmanniana Nees, Lehmann lovegrass, uncom- on perennial herb, par, 2900-3440 ft; CMR 16950 (BG- wun Jr Ar 313 Á 06, MT 202, BLT 22- is (SRSC). Steud, Tufted loveg- rass, "Wiener ang herb, RIO, 1800 ft; BHW 785 (BIBE); BHW C595 (TEX-LL). Erioneuron avenaceum (Kunth) Tateoka, Shortleaf woolygrass, ncommon perennial herb, psc, sra, 3560-4400 ft; JF 515, 1532 (SRSC); BHW 10781 (TEX-LL). Erioneuron pilosum (Buckl.) Nash, Hairy woolygrass, common nial herb, psc, DAR, DCA, SYG, CHP, 2150-5800 ft; CMR 5089 (BGWMA); JF 756, 850, 1031, 1057, 1179, 1225, 1251, 1271, 1992, 2160, JF & HL 1905, MT 44, 73, 174, 197, BHW 10781, BHW & LCH BG-75 (SRSC); BHW 10789 (TEX-LL). # Hesperostipa neomexicana (Thurb. ex Coult) Barkworth [Stipa neomexicana (Thurb. ex Coult.) Scribn], New mexico c s rare perennial herb, svo, 5000 ft; JF 735 ORS n contortus (L.) Beauv. ex Roemer & J.A. Schultes, anglehead, occasional perennial herb, DAR, DCA, DSC, 1800- pes JF 345, 574, 1396, 1473, 1596, 2138, MT 176, d 13074, d a BG-189 (SRSC). i N letop, uncom- mon perennial herb, DAR, SYG, 3750- 5800 ft; “OCW 319 (BGWMA); JF 233, 299, 1046, 1055, 1268, 1516, 1528, MT 231, OCW 319 (SRSC) 8 Leptochloa fusca (L) Kunth ssp. uninervia (J. Presl) N. Snow [Leptochloa uninervia (J. Presl) AS Hitchc. & Chase], Mexican sprangletop, “infrequent to frequent" annual or perennial herb; rio, 1800 ft; HTF 289, BHW 13041 (SRSC); EGM 140, 51-1114 (TEX-LL). # Leptochloa panicea (Retz) Ohwi ssp. mucronata (Micha. C tochloa mucronata (Michx.) Kunth], Mu- cronate sprangletop, rare annual or perennial herb, psc, 2400 d JF e (SRSC). 2c d ET ae YET)? + Melica montezumae Piper, Montezuma melicgrass, rare perennial herb, osc/oca, 4040 ft; JF 721 (SRSC) # Muhlenbergia arenicola Buckl, Sand muhly, rare perennial herb, sva, 5300 ft; JF 1253 (BIBE); JF 1253, 1991 (SRSC). $ Muhlenbergia dubia Fourn. ex Hemsl, Pine muhly, occasional D | herb, par, sra, 3500-5500 ft; JF 734, 762, 1217, 1261, 2180 (SRSC). Possibly the dominant grass above 5000 ft. SES HI IB L Li perennial herb, psc, Dar, 2900 ft; CMR 16956 (BGWMA); JF 207 (SRSC). 4- Muhlenbergia rigens (Benth.) A.S. Hitchc, Deergrass, rare P erennial mu DAR IG: ich ft JF 1527 (SRSC). DCA; OCW 316 (BGWMA & SRSC e # Muhlenbergia spiciformis Trin. [Muhlenbergia parviglumis Vasey], Longawn muhly, occasional perennial herb, sy, cHP, 4280-5840 ft; JF 1053, 1216, 1243, 1262, 1303, 1513, 1525 (SRSC). Muhlenbergia tenuifolia (Ku nth) Trin., [Muhlenbergia MONGA Buckl, S hly, | herb, DCA, SYG, CHP, 5000 ft; OCW315b (BGWMA & SRSC); JR 845, 1191, aaa 1515, 1526, 1930, PEN TOZ IO RAC) C, DAR, DCA, SYG, CHP, 2400- 5150 ft; CMR 5091 (BGWMA); Bem 12893 (BIBE); JF 195, 575, 1221a, 1229, 1395, 1456, 1519, 1921, MT 81, 211, BHW & LCH BG-192 (SRSC). § Panicum hirticaule J. Presl, Mexican panicgrass, uncom erb, psc, 1800-3800 ft; BHW 12896 (BIBEJ; B 10795 (SRSC & TEX-LL). Panicum obtusum Kunth, Vine mesquite, rare perennial herb, sv, 5300 ft; JF 1997, TR s.n. (SR Pappophorum bicolor Fourn, Pink Gees rare to'infre- quent" perennial herb, DAR pca, 3050-3700 ft; JF 1105 (BIBE & SRSC); BHW 20629, BHW & LCH BG-166 (SRSC). ere vaginatum Buckl. Whiplash pappusgrass, uncommon perennial herb, psc, DAR, 2350-3480 ft; JF SE (BIBE & SRSC); JF 1095, 1368, 1412, BHW & LCH BG- 131 (SRSC): * Pennisetum ciliare (L) Link, h Se es Buffelgrass, overall rare, but locally al erb, DSC, DAR, DCA, Rio, 1850—2500 ft; BA2 4323 (BIB; p 1055, 2222; AMP 5861 (SRSC § Phragmites australis (Cav.) Trin. ex Steud, Common reed, locally abundant perennial herb, rio, 1800 ft; WFM s.n. (BIBE). LI oe pee a LI Hei = kl) Benth.], Tobosa | herb, DSc, DAR, SYG, 2350-5300 i um TE 4614 (BIBE); JF 1255, 1399 1435, 1478 (SRSC). Poa bigelovii Vasey & Scribn, Bigelow bluegrass, uncommon annual herb, psc, ocd, 3560 ft; JF 516, 537, BHW 21134, 47056, BHW & MCJ ee Gen (SRSC); BHW 21 134 (TEX- LL). tn, Wolfs svG, cHP, 3800- 5000 ft; JF SCH MT 39, BHW 10769 (SRSC): BHW 10769 (TEX-LL). É Rabbitfoot grass, locally occasional annual herb, RIO, 1809 ft HTF 290 BO. ichx) Nash, Little bluestem, rare +0 GE (Nutt) C.G. Reeder, Bristly il, rare pe- nial herb, sya, cur, 5150 ft; JF 1230 (SRSC). DH plateau. perennial herb, DCA, 3500- 4650 ft; JF 1950 (BIBE); JF 1939, 1950, OCW 317 (SRSC) rare annual her bs psc, 2960 ft; JE 217 (SRSO). Setaria leucopila (Scribn. € Merr.) K. Schum, Streambed ristlegrass, occasional perennial herb, DSC, DAR, DCA, SYG, 2400-5300 ft; JF 356, 1134, 1416, 1988, MT 102 (SRSC); EGM 164 (TEX-LL). S Setaria parviflora (Poir) Kerguélen [Setaria geniculata P. e 'arsh bristlegrass, locally common perennial herb, Rio, 1800 ft; CAB 32, BHW 13053 (BIBE); BHW 13053 (SRSC); EGM 51-1112, BHW 13053 (TEX-LL S Setaria reverchonii (Vasey) Pilger, [Setaria ramiseta (Scribn.) Pilg.], A EIN ee een perennial herb; 1800 ft; BHW & MCJ 16856 (SRSC) + Ge Gees (L.) Pers, Johnsongrass, rare perennial AR, 3440 ft; JF 362 (SRSC). Sorbo airoides (Torr) Torr, Alkali sacaton, uncommon nial herb, psc, sra, 1800-5300 ft; JF 1254, 1542, BGH SS Se See cyan (Torr) Gray, Sand dropseed, uncom- nial herb, psc, DAR, DCA, SYG, 1800-5300 ft; JF 352, 1999 2050, 2146, d b en sey) R ydb, NA A 709 perennial herb, psc, DAR, DCA, 2150-4360 ft; BRM 3138, CMR 5093 (BGWMA); BA2 5198, 5350, JF 1517, 1915, MT 65, BLT 22-145, BLT & BD 23-177, BHW & LCH 160 (SRSC); BLT 22-145, Bala LCH 8G- 87, BG-1 d ii E uncom- mon on perennial herb, DSC, DAR, DCA, , 2100- 3050 ft JF. MISS, 1, TRV s.n, BHW 21195 (SRSC nm warnockii B.L. Turner, Warnock water-willow, SC sional subshrub, psc, DAR, DCA, 1800-3720 ft; BHW BG-137(BGWMA); AL 161, 162, CMR 17231, a (BIBE); JF 229, 687, 1667, 2053, 2087, MT 152, 181, 227, BLT 22-149, BLT & BD 23-178, BHW 10786, 13063, 20410, BHW & LCH BG-137, BHW & MCJ 16827 (SRSC); BLT 22-149, BLT& BD 23-178, BHW 10786, BHW & LCH s.n. Ruellia parryi A. Gray, Parry wild petunia, common subshrub, DSC, DAR, DCA, 1800-4600 ft; SMU 48 (BGWMA); JF 202, 434, 520,829, 945, 2042, 2113, AMP & SAP 5136, MT 27, BLT 22-144 (SRSC); BLT 22-144 (TEX-LL). Amaranthaceae *+ Amaranthus blitoides S. Wats, Mat amaranth, rare annual HIE DAR 3000 TE JF 218 (SRSC). "frequent", perennial herb, rio, 1800 ft: BHW 8 MCI 16845, 16853 (SRSC). § Sporobolus pyramidatus (Lam. A.S. Hitchc, Madagascar dropseed,'infrequent" annual or perennial herb, rio, 1800 E iid 786 (SRSC). ibn, Big sacaton, perennial herb; EGM 123 (SRSC); EGM 123, 5 2 1110 (TEX-LL). 8 Tridens O Hees & bo. Nash, Lovegrass tri ens, peren | ] ~t HN ZA A Audi Adi ^ KI | Slim tridens, com herb, DSC, DAR, DCA, SYG, , 1800- 5500 ft; JF 327 (BIBE); JE ee 208, 212, 327, 762a, 1017, 1039, 1174, 1177, 1260, 1369, 14 pe 1937, 2017, 2078, TR 25, MT 138, 161, BHW E 8, BHW & LCH BG-65, BG-179 (SRSC). ocho arizonica (Scribn. € Merr.) o dep & F. Zuloaga Me ommon annual herb, rio, DCA, 1800 ft; JF 2105, BHW Dio 20945, O 16855 (SR 50) CALA DCA; RRI 21160 (TEX-LL). ,annual herb, Typnaceae: D trail marannial herb, Die “1800 it FGM 155 (SRSC); BHW 835 (BIBE & TEX-LL). ANTHOPHYTA, Eudicotyledones oo j ightia, uncom- mon subshrub, geet crevices, rocky soil, DSC, DAR, DCA, 1800-2350 ft; BHW 12898 (BIBE); JF 648, 1342, 2061, BHW (SRSC). E 12898, 18408 (SRS + Carlowrightia serpyllifolia A. Gray, Trans-Pecos wrightwort, occasional subshrub, psc, oca, 3680 ft; JF 447, 607, 667, 972, 1013, 1343, 2016, 2080 (SRSC). e Dyschoriste linearis var. cinerascens (Henrickson & Hilsenb.) B.L. Turner, Polkadots, uncommon and "locally frequent" S. Wats, Carelessweed, annual herb, nio, . 1800 ft; EGM 125, 26, js eee rare annual herb, DCA, 1800 ft; JF 2107 (SRSC). S est torreyi (Gray) Benth. ex S. Wats, Torrey ama- Eu o annual herb, psc, 3800 ft; BHW 10796 (SRSC X-LL). e e E ex Standl, Arizona snakecotton, perennial herb, DSC, DAR, oca; BHW & BLT 8318, BHW & MCJ 16847, GLW 4477 (SRSC); DSC & HBC 30635, BLT & MT2 97-333 (TEX-LL). Tidestromia lanuginosa (Nutt.) Standl, Wooly tidestromia, uncommon annual herb, psc, pca, 2850 ft; JF 942, BHW & MCI 16852 (SRSC). Tidestromia suffruticosa (Torr) Standl, Shrubby honeysweet, occasional subshrub, psc, oca, 1800-2760 ft; JF 149, 197, BHW 694 (SRSC); BHW 694 (TEX-LL). Anacardiac Rhus GE Engelm. ex A. Gray, Littleleaf sumac, rare shrub,bar, 2100-2400 ft; CMR 5076 (BGWMA); JF 1444, BHW & LCH BG-89 (SRSC). it Rhus trilobata var. pilosissima Engl, Skunkbush sumac, rare shrub, cup, 5680 ft; JF 1292 (SRSC). i: virens e ex A. Gray var. virens, Evergreen sumac, n to occasional shrub, pca, cue, 3050-5680 ha SCH pun (BGWMA); a 767a, 1040, 1135, 1182, 1295, 1508 (SRSC). Apocynaceae Lum ksii (L.D. Benson) L.D. B Mai cimicidum A. DC. var. crooksii L. Ben rare perennial herb, oca, 1850 ft; a o Telosiphonia macrosiphon (Torr.) Henrickson, Oe macrosiphon (Torr) A. Heller], Rocktrumpet, unco subshrub,psc, DAR, 2700-3350 ft; CMR 11625 CCS JE omg 933, 1481 (SRSC). 710 Aristolochiaceae E coryi I.M. Johnst, Dutchman pipe, occasional nnial herb, psc, DAR, DCA, RIO, za, 1800-4280 ft; BA2, Dc ph & B 4933 (BIBE); BA2 5716 (SAT); ins ps. 149], 2093, 2119, BLT 8 MT2 97-338, BHW T&MT2 97-338 (TEX-LL) Asclepiadaceae + Asclepias asperula (Dcne) Woods, Spider milkweed, un- common perennial herb, osc, 2560-3800 ft; JF 368, 1405, 1911 (SRSC). E Cham. & Schlecht, Zizot ilk | rare perennial herb, osc, DCA, 2300-2750 ft; JF 914 (SRSC); BLT & MT2 97-341 (TELL). # € sperryi Woods, Sperry milkweed, rare subshrub, SR See ft JF 1286, da FBO). , Bearded swal- LEE perennial haroaceous vine; JLB 870 (SAT). Cynanchum pringlei (Gray) Henrickson, [Cynanchum barbig- var. breviflorum Shinners], Pringle swallow-wort, uncommon to occasional perennial herbaceous vine, DSC, DAR, DCA, 3000-4240 ft; BA2 5315 (SAT); JF 256, 301, 1164, 1488, 2203, m BLT & BD 23-174, BHW 10785, BHW & LCH BG-133 a BLT & BD 23-173, BHW 10785, BHW & LCH BG-133 (TE Cynanchum racemosum itla cq.) Jacq. var. unifarium (Scheele) E. Sundell, [Cynanchum unifarium (Scheele) Wee Talayote, uncommon to locally common perennial her baceous vine, DAR, DCA, 2780—4040 ft; JF 1928, AL SOEN E 5317 (SAT); i 169 Ke ee Eno. rennial Escola vine, pla 1900 ft; JF 2057 REC § Matelea producta (Torr) Woods, Texas milkvine, perennial erbaceous vine, RIO, eons Sc E (BIBE). ommon perennial herbaceous vine, DAR, DCA, 3160- pe ft; BRM 3225, BHW & LCH BG-170 (BGWMAJ; JF 1938 (BIBE); BA2 5339 (SAT); JF 221, 1938, JF & MY 1317, AMP & APSA BHW& SCH BG-170 So, perennial herbaceous vi vine, DSC, 3350 ft; JF 1484 er 8 Sarcostemma crispum Benth, Wavyleaf twinvine, perennial NL ele vine, SC, de eg oe BA2 5365 (SAT). e adas GE e. Fri twinevine, "frequent on shrubs”, perennial herbaceous vine, RIO, osc, 1825 ft; BA2 & DG 5158 (BIBE); BHW 13050 (TEX-LL). Sarcostemma cynanchoides Dcne. var. hartwegii (Vail) Shin- ners, Hartweg twinevine, rare perennial herbaceous vine, psc, DCA, 1800-2350 ft: JF 1347 (SRSC); BHW 12904 (BIBE, SRSC, TEX-LL Sarcostemma torreyi (Gray) Woods, Soft twinevine, uncom- on perennial herbaceous vine, DAR, DCA, RIO, 1800-4650 ft; BA2 BER 5280, 5375 (SAT); JF 1956, 2000, TR 147, BHW 13050, BHW & LCH BG-165 (SRSC). Asteraceae f ti inata (Lag exD Don) B.L. Turner, Featherleaf de- , DSC, DCA, 1800-3700 + s | | | [11 17 ft; BRM 3147 (BGWMA); BA2 5217, 5440 (SAT); JF 147, 302, 459, 1339, 1353, 2115, OCW 342, BHW 10815, 20610, BH & LCH BG-181, BHW 8: MCJ 15899 (SRSC); BLT & MT 97-339, BHW 10815 (TEX-LL). Ageratina o E Gray) R. M. NDS & H. Rob, Ed wrightii A.G DCA, em, 4040-4900 ft: OCW s.n. (BGWMA); JF 722, En 210% ood e gë 2 Lud (TEX-LL). | perennial herb; BA2 5360 (SAT). $ Aphanostephus ramosissimus DC. var. humilis (Benth.) B.L. Turner & Birdsong, Plains dozedaisy, annual herb, DCA, RIO, Ea 2560 fu == VLC 18730 ee ee ludoviciqn | SYG, CHP, 305 0-5680 ft; BA2 5491 (SAT); JF 2220, 1152 (SRSC); BA2, DG & TE 5491 (TEX-LL). (ber DSC, DAR, DCA, # oe havardii Gray, Havard false willow, rare subshrub, A, 4050-4350 ft; JF 1933, 194] (SRSO. m SSC (Ruiz & Pavón) Pers., oe glutinosa Pers], Seepwillow, jara, locally common shrub, DAR, DCA, RO, 1800-4280 ft; EGM 12 (BIBE); Se E (SAD); JF 1345 (SRSO); BA2, DG & ET 5511 (TEX-LL). Se Baccharis salicina Torr. & Gray, Great plains false willow, RSC subshrub, BGH 643 Bahia absinthifolia Benth. var. dealbata (Gray) Gray, Dealbata ahi mon perennial herb, psc, DAR, 1800— ft; BHW 12888 (BIBE); JF 347a, 543, 810, 2109, MT 120, 235, 253, BHW 12888, GLW 4486 (SRSC); BA2 & TE 5301, BHW 12888 (TEX-LL). Bahia pedata Gray, Bluntscale bahia, uncommon annual herb, psc, DCA, 3050-3150 ft; JF 1102a, 1104 (SRSC); WFB 7 (TEX-LL). Baileya multiradiata Harvey & Gray ex Gray, Desert mari- gold, common to occasional annual herb, osc, DAR, DCA, 2400-3050 ft; BA2, DG, TE, & PC 4598, CN 1194; (BIBE); JF 32, 1455, 1466, MT 16 (SRSC). Brickellia coulteri Gray, Coulter brickellia, occasional subshrub, wash gravel, crevices in bedrock, rocky soil, osc, DAR DCA, Rio, 1800—3800 ft; JF 686, 1061, 1593, 1669, 2064; BHW 10834 (SRSC); DSC & HBC 30626, BHW 10834 (TEX-LL). iod eupatorioides (L.) Shinners var. chlorolepis (Woot. & andl) B.L. Turner, False boneset, rare subshrub, psc, DAR O ft; BA2 5510 (SAT & TEX-LL); JF 1041 (SRSC). Brickellia laciniata Gray, Splitleaf brickellia, occasional shrub, DAR, DCA, 1850—3050 ft; BA2 5483 (SAT & TEX-LL); JF 1084, 1146 (SRSC) e Brickellia aA var. conduplicata (B.L. Ges B.L. Turner, rickellbush, uncommon perennial her 4650 ft; ES p" DG 328, 329 (SAT); JF on 1949 E BA2 5455, BA2 & TE 5441 (TEX-LL) + Chaetopappa bellioides (Gray) Shinners, Manyflower least- daisy, rare perennial herb, psc, 2680 ft; JF 190 (SRSC). + Chaetopappa ericoides (Torr) Nesomn [Leucelene ericoides (Torr) Greene], Rose heath, leucelene, rare annual or pe- rennial EN SYG, 3000- 2 mar ie 29 ERC: Chl " +} VOZILI - je ilweed EE For 4 cool] TARA : - dr Fenstermacher et al., Vascular flora of the Dead Horse Mountains ll herb, rio, 1800 ft; BA2, DG, TE & B 4922 (BIBE); JF died ul SC). 1 | | | ay, /, Damianita, at higher BE DCA, SYG, CHP, 4080-5200 ft; SMU 52 (BGWMA); PVW s.n. (KANU); BA2 5297 (SAT); JF 572, MT 223 (SRSC); BA2 & TE 5297 (TEX-LL) Cirsium turneri iplb Cliff thistle, uncommon perennial herb, pca, 2920-4200 ft; AL 163 (BIBE); JF 187, 1121, 1704, AMP & SAP 5 126, MT 215 (SRSC); JF 1121 (TEX-LL 8 Conyza canadensis (L.) Crong, Canadian horseweed, annual herb; EGM 142 (BIBE). IA FTE DEE Ie: IP FL DC], Manyst | herb, psc, DAR, 2400-2520 ft; JF 1429 (SRSC); RRI 555 (TEX-LL). Dyssodia acerosa DC., [Thymophylla acerosa (DC.) Strother], Prickleleaf dogweed, common perennial herb, psc, DAR, DCA, 2150-4400 ft; BHW & LCH np (BGWMA); JF 512, 1588, MT 72, BHW & LCH BG-72, GLW 4447 (SRSC); BA2, DG & IE 5505 (TEX-LL). Dyssodia micropoides (DC) Loes., [Thymophylla micropoides (DC) Strother], Wooly dogweed, uncommon perennial herb, psc, 2350-2560 ft; Fa 317 (BIBE); JF 1401, 1684 (SRSC); BA2, DG & TE 5461 (TEX-LL). Dyssodia E (DC) B.L. Robins., [Thymophylia pen- tachaeta (DC) Small var. pentachaeta], Prickly dogweed, occasional perennial herb, psc, DAR DCA, Svc, 1800-5150 ft; CMR 5056 (BGWMA); BA2, DG, TE, & PC 4599 (BIBE); PVW s.n. (KANU); JF 192, 320, 460, 540c, 542, 576, 1033, 1085, 1226, 1228, 1927, 2108, HTF 1230, MT 62, 242, BHW 10817 (SRSC); BA2 & TE 5295, 5296 (TEX-LL). 8 Eclipta prostrata (L.) L., [Eclipta alba (L.) Hassk.], False daisy, “infrequent to frequent" annual or perennial herb, Rio, 1800-1825 ft; BHW 13039 BIBE SRSC, TEX-LL); BHW 13035 (TEX-LL). Erigeron flageilaris A. Gray, Trailing fleabane, uncommon biennial E 3300-4600 ft; JF 754, 799, AMP & SAP 5129, TASAS Erigeron Reeg Gray, Plains fleabane, uncommon perennial AR, 3460 ft; VLC 43989; JF 264, 373, MT 249 (SRSC); BA2 E TE BA2, TE & DG 5494 (TEX-LL). Erigeron tracyi Greene, [Erigeron colomexicanus A. Nels.], Running fleabane, occasional annual herb, psc, DAR, DCA, sva, 2880-5150 ft; JF 7025 (BIBE); JF 763, 1008, 1025, 1153, 1231, 1272, BLT & BD 23-170, BHW 10833 (SRSC); BLT & BD 23-170 (TEX-LL). 8 eis Sieg (Spreng.) C. Mohr, Running yellowtops, “fre- nnual herb, rio, 1800 ft; EGM 132, BHW 750, 13042 mum ES (SRSC); EGM 132, BHW 750 (TEX-LL). 8 Flourensia cernua DC, Tarbush, rare shrub; BA2 5520 (TEX- LL). § fs parryi (Gray) King & H.E. Robins, Chisos mountains ellbush, "infrequent" perennial herb; BHW 18403 Kä qs "ENS SYG, SC f JF ids 3 (SRSC) ¿annual or peren- | annual or perennial herb, par; BA2 & TE 5355 (TEX-LL). Gutierrezia microcephala (DC.) Gray , Threadleaf snakeweed, 711 subshrub DCA, DSC, SYG, 2500- 5840 ft JE 1324a, MT 45, 219, 252 (SRSC): BA2, DG, & b de TR ca MIES LL). + 2. Di ISby, B | J common subshrub, DSG, DAR, DCA, RIO, sYG, 1800-5100 ft; BA2, DG, B& TE 4921, JF 1411 (BIBE); GE (KANU); JF 318, 1411, 1417, TR 251, BLT & BD 23-179 (SRS Gutierrezia Vira d nid onde snakeweed, un- com shrub, D , SYG, 2400-4280 ft; JF 14110, OES 763 (SRSO) BLT A a a 179 SC LL). Gymnosperma glutinosum (Spreng.) Less., [Xanthocephalum glutinosum (Spreng.) Shinners], Tatalencho, gumhead, common subshrub, psc, DAR, DCA, svG, 2350-5300 ft; OCW 339 (BGWMA); JF 238, 321, 498a, 579, 712, 752, 1181, 1376, 1989, OCW 339 (SRSC); BA2 € TE 5323, BA2, DG & TE 5515 (TEX-LL) E Lisl H j A some Lv thal Ü Já (Gray) B , Pretty sneezeweed, annual herb, rio, 1800 ft; MWB & JEA 232a, 232c, 232d (TEX-LL). § Helenium microcephalum var. microcephalum, Small- head sneezeweed, "frequent" annual herb, no, 1800 ft; AL 322, Piera rra BHW SE SE OES & CN 504, BHW 13031 S Helenium e DC. var. ooclinium (Gray) Bierner, [Helenium ooclinium A. Gray], annual herb, rio, 1800 ft; MWB & JEA 232b, 232e (TEX-LL). Helianthus annuus L, Common sunflower, "frequent" annual herb, rio, 1800 ft; EGM 129, BHW 13045 (BIBE); JF 2263, BHW 13045 (SRSC). 8 Helianthus ciliaris DC, Texas blueweed, Ed frequent" erennial herb, rio, 1800 ft; BHW 13051 (BIBE & SRSC). it Heliopsis parvifolia Gray, Mountain oxeye, rare perennial herb, svc/cHp, 5680 ft; JF 1276 (BIBE & SRSC). S Isocoma pluriflora (Torr. € Gray) Greene, [Haplopappus heterophyllus (A. Gray) S.F. Blake; Haplopappus nios (Torr. € A. Gray) H.M. Hall], Jimmyweed, subshrub, p BHW 1076 (TEX-LL). en (Gray) y, Rag | , tagged marsh- elder, rare annual herb, psc, par, 3640 ft; JF 289 (SRSC); BA2, DG, & TE 5492 (TEX-LL). Jefea brevifolia (A. Gray) Strother, [Zexmenia brevifolia A. Gray], Shorthorn zexmenia, occasional shrub, psc, DCA, sva, 1800-4700 ft; PVW s.n. (KANU); JF 472, 708, 2132, CMR 5068, MT 106, BHW 10736 (SRSC); BA2, DG, & TE 5503, BA2 & TE 5298, DF 223, BHW 10763 (TEX-LL). + Laennecia coulteri (Gray) Nesom, [Conyza coulteri A. Gray], Coulter horseweed, annual horsetail, rare annual herb, DCA, ipia rt d iiid RSEN o (Kunth) Nees, Tahoka daisy, rare annual herb; Dar, 3480 ft; JF 544 (SRSO. Melampodium leucanthum Torr. & A. Gray, Blackfoot daisy, occasional subshrub, DSC, DAR, DCA, RIO, sYc, 1800-4450 ft; JF 228, 269, 490, 967, 2006, 2123, MT 4, 47, 58, 74, 238, BLT & BD 23-176, BHW & BLT 8328, GLW 4483 (SRSC); BLT & BD 23-176 (TEX-LL). + Packera millelobata (Rydb.) W.A. Weber & A. Lóve, Uinta rare perennial herb, svc/cHP, 5680-5840 ft; JF 1287, 1971 (SRSC). + 712 Partheni gentatum A. Gray, Guayule, locally uncommon to occasional, psc, svc, 2920-4880 ft; BHW & LCH BG-177 (BGWMA); JF 1024, BDGL s.n. (BIBE); PVW s.n. (KANU); JF 744, 1024,1139,1155,1183, 1901, JMM 888, 900, 903, AMP & SAP 4297, MT 49, BHW 10753 (SRSC); BHW 10753 (TEX-LL). Parthenium confertum A. Gray, Lyreleaf parthenium, uncom- mon to occasional annual or biennial herb, osc, DAR, DCA, 1850-5840 ft; JF 206, 359a, 524,852, 1068, 1083, 1433, 1907, DM & JC 380, 900, 904, MT 41, 254, BHW 20608, BHW & MCJ 10656 (S RSC); gt a 1E e (TEX-LL) Aariola DCA, SYG, 2300-5000 ft; pus 1023, OCW 511 e) t» E TE 5303 CEN LL). z E Te angustifolia, Limoncillo,* frequent” annual herb; BHW & BLT 8323 (SRSC); VLC 30209, DF 224 (TEX-LL). 8 Pectis papposa Harvey & Gray var. grandis Keil, Manybristle cinchweed, "frequent in sandy soil" annual herb, 1800 ft; BHW 286 (SRSC); BHW & BLT 8323 (TEX-LL). Perityle aglossa A. Gray, Bluff rockdaisy, occasional subshrub, cracks, psc, DCA, 1850-3560 ft; AL 141 (BIBE); JF 184, 530, 650, 685, 1005, 1007, 1073, 1350, 2014, SS 38, 931, BHW 20409 (SRSC); SS 931 (TEX-LL). Perityle bisetosa (Torr. ex A. Gray) Shinners var. scalaris A.M. Powell, Stairstep rockdaisy, uncommon to occasional (G2T1S1) subshrub, pca, sva, cHP, 3050-5680 ft; AL 146 (BIBE); JF 475, 608a, 1123, 1289, 1917, BHW 20604 (SRSC); (TEX-LL). Perityle vaseyi J.M. Coult, Vasey rockdaisy, rare subshrub, rocky soil; psc, DAR, 2080-2350 ft; JF 1367, SS 59, TRV & LIT 1816 _ RSC); > = (TEX-LL). (Nim Y Do cana Nutt American ke ene or r star-thistle, rare ual herb, psc, par, 3550-4420 ft; JF 507a, 843 (SRSC). Porophyllum scoparium A. Gray, Shrubby poreleaf, occasional UD, DSC, DAR, DCA, RIO, 1800-4000 ft; BRM 3132, 3227, CMR 11592, SMU 38 (BGWMA); JEA € TJW 170, JF 332, 47 1, 645, 1379, 2081, BGH 633, OES 1351, MT 184, BHW 18396, 20618, 21191 (SRSC); BA2 & TE 5336, BA2 & DG 5207, DSC 8 HBC 30633, DF 51 (TEX-LL). Psilostrophe gnaphalioides DC, Cudweed paperflower, rare o "infrequent" perennial herb, psc, 1900 ft; JF 2044, BHW 20627 (SRSC). id ipe tagetina (Nutt.) Greene var. cerifera (A. Nels.) B.L. Turner, Wooly paperflower, uncommon perennial herb, psc, DAR, 2300-3560 ft; JF 523, 605, 855, MT 17, 18, BHW 10788, 18390, GLW 4470 (SRSC); BA2 & TE 5362, TRV 85-37 (TEX-LL). + Senecio flaccidus Less. var. flaccidus, [Senecio douglasii DC], Threadleaf groundsel, rare perennial herb, psc, DAR, 2400-3560 ft; JF 513, 1431 (SRSC). *§ Sonchus asper (L.) Hill, Prickly sow thistle, annual herb, rio, 1800 ft; BA2 4320 (BIBE). *8 Sonchus oleraceus L, Sow-thistle, "frequent" annual herb, RIO; BHW 733 (BIBE & TEX-LL); GIE 209 (SR + Stephanomeria pauciflora (Torr) A. Nelson, Brownplume wirelettuce, rare perennial herb, pca, 2350 ft; JF 1358 (SRSC) 8 Symphyotrichum divaricatum (Nutt) G.L. Nesom, [Symphy- trict bulat ligulat (Shi ) S.D. Sundb, Southern saltmarsh aster, annual herb; EGM 134, BHW 30 (BIBE) Symphyotrichum EEN (Poepp. ex x Spreng.) hesem, Sundbi Sout! t It herb, rio, 1800 ft; JF 2037, BHW 13044 (SRSO). Tetraneuris scaposa (DC) Greene var. equ. o four e daisy, occasional perenn 2150-5840 ft; CMR 5088 ENA BHW um IRE: " 258, 524a, 614, 739, 755, 1052, 1234, 1247, 1374, 1403, 1970, MT 79, 233, BHW 10850, 12895, BHW & LCH BG-67 (SRSC); BA2 & TE 5300, 5434, BHW 12895 (TEX-LL). +1 annual herb, 3520 ft; ACK 826 (BIBE). Thelesperma longipes A. Gray, d npe. OC- casional perennial herb, psc, DAR, DCA, 1800-5150 ft; BRM 3140; CMR 5083, a co BHW ps (BIBE, SRSC, & TEX-LL); JF 761, 1233, 1394, 2125, BHW & LCH BG-73 (SRSC); BA2 & TE 5325, OES 734, BHW & LCH BG-73 Thelesperma megapotamicum (Spreng.) Kuntze var. mega- potamicum, Rayless greenthread, uncommon perennial herb, DSC, DAR, DCA, RIO, SYG, 1800-4800 ft; CMR 5084 (BG- WMA); JF 164, 262, 263, 1389, 2167 (SRSC); DSC & HBC 30634, JAM & JAS 3429, AMP & SAP 3610 (TEX-LL). § Thelesperma qe preco DS var. am- herb, DAR, DCA, RIO, 1800 le AMP & SAP 3610, BHW 21 189, GLW 4476 (SRSC). e Thelesperma simplicifolium Gray, Slender greenthread, uncommon perennial herb, pca, 1900-4180 ft; CMR 324, OCW 3243, BHW 8: LCH BG-68 (BGWMA); JF 865, MT 63, 151, 163, AMP & SAP 5135, OCW 324, BHW 21185, BHW & Trixis californica Kellogg var. californica, American trixis, occa- sional shrub, psc, DAR, DCA, 1800-4160 ft; JF 473, 649, 1062, 1079, 1314, 2003, 2114, BHW 10751, GLW 4455 (SRSC); BA2, Dh &TE adi MÉS LL). Cav.) Benth. & Hook. f. ex A. Gray, Golden crownbear | her! o, 1000-2560 M 138, pe 746 (BIBE); JF 1066, 2036 SEH Vguierad dentata (Cav) Spreng, Sunflower goldeneye, uncom- "perennial herb, DAR, SYG, CHP, SC 5150ft BHW& LCH BG- 15 (BGWMA); JF 291, 759, 1028, 1224, MT 234, BHW & LCH BG-155 (SRSC). Viguiera stenoloba S.F. Blake, Skeletonleaf goldeneye, com- on shrub, DSC, DAR, DCA, SYG, CHP, 1800-5000 ft; TR 56, CMR 16939, BHW & LCH BG-86 (BGWMA); JF 260, 1029, 1370, 2140, TR 56, MT 145, 173, BHW & LCH BG-86 (SRSC); BA2 & TE 5302 (TER LL). 8 Xanthisma spinulosum (Pursh) D.R. Morgan €: R.L. Hartman var. spinulosum, [Haplopappus spinulosus (Pursh) DC; Machaeranthera pinnatifida (Hook.) Shinners], Lacy tan- syaster, cutleaf goldenweed, "fairly common" perennial herb, psc, pan; GLW 4469 (SRSC); BA2 & TE 5389, BA2 & DG 5272 (TEX-LL). Xanthisma spinulosum (Pursh) D.R. Morgan & R.L. Hartman var. chihuahuanum (B.L. Turner & A L. Hariman) D.R. R.L. Hartman, [M val chihuahuana B.L. Turner 8: Hartman], Lacy tansyaster, oc- casional perennial herb, psc, DCA, Rio, 2300-3050 ft; JF 800, 958, 1130, 1373, MT 207, 210, BHW 18391 (SRSC); RLH € JDB 3504b, 3503 (TEX-LL). # Xylothamia triantha (Blake) Nesom, [Ericameria triantha (S.F. Blake) Shinners], Trans-Pecos desert goldenrod, rare subshrub, psc, DAR, 2400 ft; JF 1450 (SRSC). diis acerosa (DC) A. Gray, Spinyleaf zinnia, common pe- nial herb, psc, sva, 2760-4400 ft; BRM 3228, BHW & LCH SE (BGWMA); JF 153, 851, 1094, 1180, MT 5, 59, BHW 10837, BHW & LCH BG-142 (SRSC); BH2 & TE 5294, BHW 10837 (TEX-LL) erberidac + inis goe Meric, Agarita, algerita, uncommon to occasional shrub, psc, DAR, DCA, sva, 2880-4600 ft; JF 508, 528, 1001, 1036 (SRSC). re is (Cav) Sweet, D twill rss OCA, 1800-4280 ft; TR 140 (BGWMA); BA2 5234 (SAT); JF 1355, 1356, 2137, TR 140, MT 189 (SRSC). Tecoma stans (L.) Juss. ex Kunth, Yellow bells, occasional shrub, psc, DAR, DCA, 1800-3050 ft; TR 141 (BGWMA); BA2 5331 (SAT); JF 179, 1010, TR 141, MT 11 193, BHW & LCH BG-85, BHW & MCI 16866 (SRSC); BLT & MT2 97-347, BHW & LCH BG-85 (TEX-LL). Boraginaceae e Antiphytum heliotropioides A. DC, Mexican saucerflower, rare to"locally common’ shrub, DAR, pca, 4200-4350 ft; BA2 5220, 5313, 5453, 5465, 5600 (SAT); JF 1936 (SRSC). +o Cryptantha albida (Kunth) I.M. Johnst, New Mexico cryptantha, rare annual herb, oca, 1920-3000 ft; JF 646, 05 (SRSC). Cryptantha angustifolia (Torr) Greene, Panamint cryptantha, rare annual herb, psc, 2560-3560 ft; JF 522 (SRSC); RRI 552 (TEX-LL). a coryi LM. Johnst. [Cryptantha palmeri (A. Gray) Payson] Cory cryptantha, rare ver i psc, 3560 ft; JF 280 (SRSC); BLT & BD 23-164 (TEX-LL). Ao Cryptantha crassisepala (Torr. E A. e Greene, Thickse- GE annual herb, 1800 ft at the Boquillas Canyon restoration site. 8 ia mexicana (Brandegee) LM. Johnst, Mexican cryptantha, "infrequent" annual herb, 2300 ft; OCW 506 RSC). Heliotropium confertifolium (Torr.) Torr. ex A. Gray, Leafy neliotrope, occasional subshrub, psc, 1800-3800 ft; EGM 147, BHW 907, 940 (BIBE); JF 155, 367, BHW s.n., 940, 10848, BHW & si BG-76 (SRSC); BHW s.n., 10848; BHW & LCH BG-76 (TEX-LL). Heliotropium n (Nutt) A. Gray, Phlox heliotrope, locally common annual herb, mo, psc, 1800 ft; BA2, DG, TE, & PC 4608, BHW 13029 (BIBE); WRC & PM212200, JF 2232, AMP & SAP 5860, BHW 686, 13029, 16851, GLW 4474 (SRSC); DSC & HBC 30631, BHW 686 (TEX-LL). 713 var. curassavicum | Salt heliotrope, locally common subshrub, rio, 1800 ft; EGM 124, 943, CN 495 (BIBE); JF 2256, BCT 8802 (SRSC); BCT 8802, BHW 943 Es " I f L HA Heliotropium glabriusculum (Torr.) A. Gray, Greeneye heliotrope, perennial herb, 1800 ft, head of Boquillas anyon. 8 gal greggii Torr, Fragrant heliotrope, perennial herb, 18 § no precum bens Mill, Foursplks heliotrope, sub- shrub; BHW 1089 (TEX-LL). Lappula redowskii (Hornem.) Greene, IL. occidentalis (S. Wats.) Green] Flatspine stickseed, occasional annual herb, pan, pca, 1800-3560 ft; DB 22, TK 10, BRM 3124 (BGWMA); JF 427, 540a, 797, 2101, BHW 47057, 47059 (SRSC). +e Lithospermum incisum Lehm, Fringed puccoon, nar- rowleaf stoneseed, rare perennial herb, sra, 4360-4800 ft; BA2 5539, 5597 (SAT); JF 554, 1948 (SRSC). S Omphalodes aliena A. Gray ex Hemsl, Mexican navelwort, "frequent" annual herb, psc, DCA, RIO, 1800-2560 ft; BRM 3131 (BGWMA); JLB 1578, ECM 49 (SAT); AMP 5367, BHW 15991, d 8395 (SRSC); DSC: A el 30715 d Bs crinklemat, eeh subshrub, psc, 2200- 2400 ft; BA2 5340 (SAT); TR 4 (SRSC); TR 4, BHW 1230 (TEX-LL). ipe: gossypina (Wooton & Standl.) A.T. Richardson, Texas inklemat, occasional subshrub, psc, pca, 1800-3440 ft; JF a 2059, BHW 12899 (SRSC); BHW 909 (TEX-LL). Tiquilia T (Torr.) AT. Richardson, Plume tiquilia, occasional shru c; CMR 11634 (BGWMA); BA2 5270 (SAT); JF 359, 369; es 13, BHW 10800 (SRSC). 8 Tiquilia Seite (Torr) AT. Richardson, Rough coldenia, subshrub, psc, 1850-3520 ft; BA2, DG, TE, & B 4923, pues BHW 908, 909 (BIBE); BLT & BD 23-162, BHW id) 21692 rd ] ) A.T. Richardson, M kl mat, occasional subshrub, 1800-3400 ft; JF 152a,812, 1443, BHW 13072, 23038, GLW 4487 (SRSC). Brassicaceae Descurainia pinnata (Walter) Britton, Tansymustard, uncom- mon annual herb, psc, par, DCA, 3440-3500 ft; BA2 5587 (SAT); p 422, 617 (SRSC). 8 Dimorphocarpa wislizeni (Engelm.) Rollins, Spectacle pod, touristplant, locally common annual herb, pca, 3500 ft; BHW 47060 (SRSC & TEX-LL); VLC 43958, RRI 558 (TEX-LL). mE cuneifolia Nutt. ex Torr. & A. Gray, Wedgeleaf draba, asional annual herb, psc, DAR, DCA, sYG, 1800-5080 ft; E 429, 536, 566, 738, 802, BHW 18392, BHW E MU 15990 (SRSC); BHW & MU 15990 (TEX-LL). 5 Lepidium nire A.Gray var. angustifolium (C.L. Hitchc.) Rollins, Mesa pepperwort, perennial herb; HBP 1923 Lepidium lasiocarpum Nutt. ex Torr. & A. Gray var. wrightii (A. Gray) Thell, Wright pepperweed, uncommon annual herb, DAR, DCA, 3440-3560 ft; JF 619 (BIBE); BA2 734, 5540, 5586; RE 734 (SAT); JF 421, 565, 619 (SRSC). Mi iS camporum (A. Gray) Greene, Bicolored mustard, mmon perennial herb, psc, pca, svo, 2120-4280 ft; CMR 714 16047 (BGWMA); JF 492 (BIBE); JF 372, 492, 615, 681, MT 05BHW 208, 47054, BHW & HKB 47063 (SRSC); BHW 1208, Se (TEX-LL). Physaria fendleri (A. Gray) O'Kane €: Al-Shehbaz, Fendler bladderpod, occasional perennial herb, DSC, DAR, DCA, SYG 2200-5000 ft; JF 557 (BIBE); TR 105 (BGWMA); JF 200, 497a, 555, 557, 736, 1189, TR 139, MT 75, 104, BHW 47062 (SRSC). + Physaria gordonii (A. Gray) O'Kane & Al-Shehbaz, Gordon bladderpod, rare annual herb, sra, 4360 ft; JF 570 (SRSC). Physaria vaso ana ROI. Or Kane & i a 120-5840 ft; JF 497, 714, 1535, 1964, AMP & SAP 5133 a 8 Physaria TO (Gray) O'Kane & Ar SEH [Lesquerella purpurea (A ray) S. Watson], R j |, frequent” perennial he e BHW 8 MCI 15995 (SRSC). ey linearifolia (A. Gray) Rollins, Slimleaf plain- smustard, rare perennial herb, svc, 3500-5680 ft; BRM 3110 (BGWMA); JF 734, 1291, 2185, MT 42, 43, 246, BHW & LCH BG-154 (SRSC). 8 Selenia dissecta Torr. & A. Gray, Texas selenia, "frequent" an- pcA, 3500 ft; CMR 16410 (BGWMA); BHW 47056, grão MO aie hee) A A LL). nual herb, pca, 2560 E RRI 551 (TEX-LD. Streptanthus cutleri Cory, Cutler twistflower, uncommon (G2S2) biennial herb, psc, par, pca, 1800-3500 ft; JF 420, AMP 3570, 5366, AMP & SAP 4299, 5387, BHW 18399, BHW _& MCI 15992 (SRSC). (Cory) Rollins, T herb; OES 1288 (SRSC). § Thelypodium wrightii A. Sieg Wright thelypody, perennial erb, oca; BRM 3106 (BGWMA). al a actaceae Ariocarpus (sini (Engelm.) K. Schum, Living rock cactus, culent, psc, svo, 3350-ca. 4500 ft; BA2 5278, SC Se SAN; 2 274, 1472 e ie TERG) tt, Desert pincush- ion cactus, succulent, 4125 f MTSO me antha var. duncanii ae 7 E Pagan Duncan cory E rare (6311725152) succulent, psc, ca. 2000-4146 ftJ | JF & MD 612, AMP 5373, DS2 2228 (SRSC). contanto echinus (Engelm.) Britton & Rose var. echinus, [Mammillaria scolymoides Scheidw.], Sea-urchin cactus, a succulent, rocky soils, open desert and slopes, psc, svo, 3500-4360 ft; JF 593, 868, 1493, BHW 11061 (SRSC) # Coryphantha echinus (Engelm.) Britton & Rose var. robusta A.M. Powell, Multi-stemmed sea-urchin cactus, rare suc- gulen psc, 2400-3650 ft; JF 1440, 1494 (SRSC). Coryphantha sneedii (Britton & Rose) A. Berger var. albi- columnaria (Hester) A.D Zimmerman, Silverlace cactus, [Coryphantha albicolumnaria (Hester) Zimmerman; Corphantha sneedii var. albicolumnaria (Hester) A.D. Zimmerman (omnaoming, cited in nomen and Weedin (as ae C OC E: SM TAS succulent, psc, sva, 1800-4840 ft; DAZ 1541 (BIBE): JF 556, 587, 588 (SRSC). n.a H In LI rr nd ET BILE 127 compraria tuberculosa engem. A. SE var. tuberculosa, ctl 15, common succulent DSC, DCA, SYG, 1900- E oa (BGWMA); JF 273, 386, 550, 625, 781, 1300, MT 89 (SRSC). § Coryphantha tuberculosa (Engelm.) A. Berger var. varicolor (Tiegel) A.D. Zimmerman, [Coryphantha dasyacantha var. varicolor (Tiegel) L.D. Benson], Varicolor cob cactus, MEER BA2 d ek law cactus, occa- sional succulent, DSC, DCA, svG, 4360 ft CMR 5173 Deu. DB 470 (SAT); JF 591 (SRSC). Echinocereus dasyacanthus Engelm., Rainbow cactus, com- cculent, psc, svo, 1900-5700 ft; DB 469 (SAT); JF 556, 623 (SRS + Echinocereus enneacanthus Engelm. var. enneacanthus, Strawberry cactus, occasional succulent, psc, 1800 ft; JF 2148 (SRSC) Strawberry pitaya, common succulent, DSC, DCA, SYG, . 1800- 440 ft; JF 780, 2149 (SRSC). KN? (Hester) L.D. been Sclerocactus mariposensis (Hester) N.P. Taylor], Miss cactus, occasional ( ; ) suc- culent, psc, pca, 3300-3520 ft; CMR 5172 (BGWMA); JF 391, 624, AM o DS2 2221 (SRSC). OO MIES warnockii Be D Benson) eS é R Foster, [Sclero- ae A | occasional des DSC, DCA, 1 900-2560 ft; JF 2150, BHW 47459, BHW & HKB 47012 (SRSC). + GE antha micromeris a FA.C. Minas ex en FE £78 m0 rmn» 0m trn Cactus, DES un TD , Common b psc, sYG, 3560-4140 ft; JF 268a, 271, 586, 613, 6130, 1486 SRSC Ferocactus hamatacanthus (Muehlenpf) Britton € Rose var. matacanthus, Giant fishhook cactus, occasional suc- culent, DCA, svc, 4300-4900 ft; CMR 5174 (BGWMA); JF 590, 1507, MT 96 (SRSC) Glandulicactus uncinatus var. wrightii (Engelm.) Backeb., [An- cistrocactus uncinatus var. wrightii (Engelm.) L.D. Benson; Echinocactus uncinatus Galeotti ex Pfeiff.; Ferocactus uncinatus (Galeotti ex Pfeiff) Britton & Rose], Eagle claw cactus, uncommon succulent, psc, su, 3520-4040 ft; DS2 03b, JF 548 (SRSC) 8 Mammillaria heyderi Muehlenpf. var. heyderi, Heyder pin- cushion cactus, "rare" succulent, psc, 2500 ft; AMP SAP. Mis on 6 103 Cine olf ball cactus, succulent, DSC, Gs 3520 ft; d 180, 2222 Pin (SRSC). pple cactus, succulent; BHW 11065 (SRSC). Neoiloydia ENEE (DC) SEH Š BE a cone cactus, G, 3080-5200 ft JF 380, 782, DS2 2220 MT 52, 53, 122, B 1055, 11058 (SRSC). RE RE Ralston & misang; SC dog cholla, vations, psc, 1850-300 ft; pm p 691, 704, 1562, 1563, 1574, Ron 1576, 1620, BAR & RAH 136 (SRSC). aureispina (S. Brack &K.D. Heil) AM Powell 8: J.F. Weedin, [Opuntia aureispina (S. Brack & K.D. Heil) Pinkava €: B.D. Parfitt], Golden-spined prickly pear, GIS) succulent, rio, 1800 ft; JF 384, BGH 802 (SRSC). + Opuntia camanchica Engelm. & J.M. Bigelow, Comanche prickly pear, succulent, psc, car, 3560 ft; MA 848, JF 626 (SRSC). S Opuntia densispina, Densely-spined dog cholla, succulent; OK & JFW 53, AMP & SAP 6275 (SRSC). Opuntia dulcis Engelm, Sweet prickly pear, succulent, psc, 00—3560 ft; JF 628; AMP & SAP 6278; MPG 87, 88, 89 (SRSC). ame | ry | E I var engelman- hii, i Engelmann prickly pear, uncommon succulent, SYG, 4440 ft; JF 589 (SRSC). § Opuntia E var. lindheimeri (Engelm.) B.D. Parfitt & Pinkava, Texas prickly pear, succulent, osc, 1800 ft; AMP & SAP 6277, JFW 1171 (SRSC *8 Opuntia ficus-indica (L.) Mill, Indian fig, rare succulent; BGH 646 (SRSC). e grahamii Engelm, Graham dog cholla, occasional cculent, osc, 3000-3520 ft; CMR 5178 (BGWMA); MA 892 T JF. 335. 37. e 389, 783, 892, 1142a (SRSC). + , RIO, 1800 ft; BGH 841, JE 2040 (SRSQ). RE DC, Tasajillo, christmas cactus, occasional culent, osc, DAR, 3360-4120 ft; JF 388, 448, 816, WH, TA, E — ee Se: DSC, DAR, DCA, 1800- 3560 ft JF 67, BGH 803, WH, TA, LS & Me 8985 (5 RSEN succulent, psc, 1800-4600 ft; MAA- 3 A- -7, JF 385, BGH 801, 842, AMP & SAP 5824, 6089 (SRSC ). Opuntia sp., Prickly pear; BGH 800, 840; AMP & SAP 6276 (SRSC) Capparidaceae Zucc, Alltho E DCA, 2500- 3800 ft; WEM s.n, EE (BGWMA); JF SEH j & HL 1891, MT 8 (SRSC). Caprifoliaceae Lonicera albiflora Torr. & A. Gray, Western white honeysuckle, rare perennial woody vine, syG/cHP, DCA, 4280-5680 ft; BRM (BGWMA); PVW s.n. (KANU); BA2 5436 (SAT); JF 1296, AMP & SAP 5125 (SRSC). Caryophyllaceae § Arenaria benthamii Fenzl ex Torr. & A. Gray, Hilly sandwort, annual herb; BRM s.n. (BGWMA). Paronychia jamesii Torr. & A. Gray, James nailwort, uncommon perennial herb, psc, sya, 3800-5800 ft; BA2 5475 (SAT); JF 235, 765, 1269a, MT 33, 248 (SRSC). Celastraceae MORTON scabrela A. Gray, EEN Bis Grande tr) PWWsn (KANU); ra 1167, 2190, MT? (RSC). rar y, Desert yal IPON, 715 DSC, DAR, DCA, 1900-3300 ft; TR 126 (BGWMA); JF 676, 973, 1366, 2041, AMP & SAP 4302, BHW 15912 (SRSC); AMP & SAP 4302 (TEX-LL) Chenopodiaceae Atriplex canescens (Pursh) Nutt. var. canescens, Fourwing salt- ; ' mon shrub, DSC, DAR, DCA, SYG, 1800-4160 ft; BA2 5262 (SAT); JF 336, 861, 1091, 1410, 1895, 2143, MT 203, BHW & LCH BG-132 (SRSC); BHW & LCH BG-132 (TEX-LL). *e Atriplex rosea L, Tumbling saltweed, red orache, rare annual herb, rio, 1800 ft; AL 306 (BIBE); AL 307, JF 2236 (SRSC). # a berlandieri Moq, Pitseed goosefoot, rare ual herb, pan nio, 1800-3440 ft; JF 361, 616, 2122, 2238 dues Corispermum americanum (Nutt.) Nutt, American bugseed, locally common annua? herb, psc, uio, 1800-2100 ft; BHW 8330, JF 2221, 2224 (SRSC); DSC & HBC 30630, BHW 8330 TEX-L l# Salsola tragus L, Tumbleweed, locally common annual herb, RIO, 1800 ft; JF 2252 (SRSC) # Suaeda suffrutescens S. Watson var. suffrutescens, [Suaeda nigra (Raf) J.F. Macbr.], Desert seepweed, rare perennial subshrub, rio, 1800 ft; JF 2234 (SRSC). Convolvulac S Bonamia ovaifolla (Torr) Hallier f, Bigood lady's nightcap, "locally common" subshrub, psc, rro, 1800 ft; BA2, DG, TE, & PC 4551; JS 303 (BIBE); HBC 30628 (NYBG); BA2 5502 (SAT); BGH 622, DAL 40, AMP 3347, AMP & JH 5696, AMP & SAP 3609, 5859, s vr DSC & HBC 30628, AMP 3347, AMP & SAP 3609 (TEX-LL). conan oe (I.M. ll e F. o, & s ai. R 18 80 ft; CA & BA? 69, BA? 6191, SMU 41 a Gen (SAT); JF 154, 307, 435, 1075, 2062, 2131, MT 167, BHW & MCJ 16834 (SRSC). Convolvulus equitans Benth, Texas bindweed, uncommon perennial herbaceous vine, DAR, DCA, RIO, 1800-4350 ft; BRM 3149 (BGWMA); JF 1945, EGM 150 (BIBE); BA2 5309, 5470, 5720 (SAT); JF 181, 853, 1349, 1432, 1945, 2077, 2116 j £ L IAAF o CA Jj foot, rare perennial ps DAR, DCA, 3056-4600 ft; JF 859, 1122, AMP & SAP 5134 (SRSC). Evolvulus alsinoides (L.) L. var. dieta ale Slender dwarf morning-glory, occasional peren C, DAR, DCA, poe 3880 ft; TR 249 (BGWMA); Ge 5312, 5464 (SAD); JF Eod: 309, e pa Pes TR a ae Zeg (SRSC). nnual vine; BHW 10799 (SR fe ie 1994 (BIBE); JF 1969, 1994, MT 241 (SRSO). § Ipomoea lindheimeri A. Gray, Lindheimer morning-glory, "infrequent" perennial herbaceous vine, 3520 ft; MA & AD 874 (SRSC). Ipomoea rupicola House, Cliff morning-glory, occasional to ncommon perennial herbaceous vine, DSC, DAR, DCA, SYG, 716 3050-4800 ft; JF 248 (BIBE); JF 248, 308, 1103, 1186, 1196, MT 31, 144, 172, BHW 962, BLT & BD 23-167, OCW 352, BHW & LCH BG-148, BHW E MCJ 16828 (SRSC); BLT & BD 23-167 (TEX-LL). § Ipomoea tenuiloba Torr, Spiderleaf, rare perennial herba- ceous vine; MT 236 (SRSC). Crassulaceae Echeveria o A. Gray, Siempreviva, desert savior, rare syG/cHP, 5700-5840 ft; BRM 3224 (BGWMA); JF n pd BHW 10774 (SRSC # Sedum nanifolium Fród., [sedium robertsianum Alexander; 5 arvum Hemsl. subsp. e (Alexander R.T. Clausen], D succulent, sva, 5150 ft; JF 1215 (BIBE & SRSC). # Sedum wrightii A. Gray, Wright stonecrop, rare succulent, sva, 4200-5000 ft; JF 581, 1163, 1506 (SRSC). Mar” casional Crossosomataceae Glossopetalon spinescens A. Gray var. m [Forsellesia spinescens (A. Gray) Greene], Spiny grea mon shrub, sya, 4200-5000 ft; PVW s.n. O JF 1205, 1505, BHW 10843 (SRSC). Cucurbitaceae # Ibervillea lindheimeri (A. Gray) Greene, Lindheimer Pa berry, rare perennial herbaceous vine, psc, 3080 ft; J 1143 (SRSC). S Ibervillea tenuisecta (^. ey) Small, Slimiobe globeberry, psc, 3500-3800 ft; BA2 5368 (SAT); MA & AD 3500, BHW 10802, MT 206 (SRSC); BHW 10802 (TEX-LL). Ebenaceae ipe texana Scheele, Texas persimmon, common to onal shrub, pan, DCA, 1800-3840 ft; BRM 3146, CMR , 2088, WH, TA, Ls & JMP 8986 (SRSC). Euphorbiace EES monostachya cau Ara pa dd Torr.], on perennia erb, DSC, DAR, DCA, 1850- 3050 lis BA2 5237, 5334 (SAT); JF 1069, 1118, 1680, BHW 646, 21200 (SRSC); BHW 646 (TEX-LL). Acalypha phleoides Cav, [Acalypha lindheimeri Müll. Arg.], Shrubby copperleaf, occasional perennial herb, DAR, DCA, 2200-4800 ft; JF 1946 (BIBE); TR 68, CMR 11583 (BGWMA); BA2 5240, 5265 (SAT); JF 1108, 1128, 1149, 1946, JF & MY 1315, TR 68, MT 187 (SRSC); OCW s.n. (TEX-LL). Andrachne arida (Warnock € M.C. Johnst.) G.L. Webster, ns-Pecos maidenbush, rare but locally common (G251) shrub, se, 4200-4900 ft; JF 1170, 1194, 1207, 1499, 2164, 2177, BHW 10767, 1077; BHW & MCJ 16681, 16840, 16841 (SRSC). Bernardia obovata IM. Johnst, Desert myrtlecroton, occa- sional shrub, psc, DAR, DCA, 1920-4900 ft; CMR 5134, BHW & LCH s.n. (BGWMAJ; BA2, DG, TE, & B 4925 (BIBE); BA2 5250, BR 293 (SAT); JF 203, 496, 642, 715, 1093, 1212, MT 86, BHW 10832, BHW & LCH BG-178, BHW & MU 16833 (SRSC); BHW 10832, BHW & LCH BG-178 (TEX-LL). it Chamaesyce acuta (Engelm.) Millsp, Pointed sandmat, rare perennial herb, SYG, EE ub 1264 (SRSC). CH A. Gray) Small, Whitemar- gin sandmat, unc common RE herb, DSC, DAR, 1800-2560 ft; JF 1447 (SRSC); HBP & VLC 30177, OES 1540 (TAES). CI | (B y ton &Standl, Bristlecup sandmat, locally common subshrub, DSC, DCA, 1960-4600 ft; BHW & LCH s.n. (BGWMAJ; JF 261, 652, 943, AMR & SAP 5142, MT 32, 68, BHW & LCH BG-176 (SRSC). Chamaesyce ee Small, Ashy sandmat, oc- casional perennial herb, psc, DAR, DCA, svo, 1800-4450 ft; OCW s.n. (BGWMA); BA2, E TE & B 4934 (BIBE); BA2 5291, 5526, 5705 (SAT); JF 174, 340, 342, 489, 547, 653, 684, 863, 1078, 1087, 1171, 1337, 2133, 1645, AMP & SAP 3612, 5305, AMP SAP MLE &AL2 6101, MT 3,66, 67, 165, 166, BLT 22-146, BHW 10756, 10762, 10840, 18393, BHW & LCH BG-66, BHW & 08, GLW 4442 (SRSC); DSC & HBC 30632, BLT 22-146, 6, 10762, BHW & LCH BG-66 (TEX-LL). Chamaesyce fendleri (Torr. & A. Gray) Small, Fendler sandmat, uncommon perennial dal DSC, SYG, CHP, 3000-5250 ft; JAM 0, 5276, 5619, 5692 (SAT); JF 784, RAS dq did 1239, 1243, 1470; 2197 (SRSO), Cf Small, Rib-seed sandmat, “frequent” annual herb, RIO, DSC, 1800- 3120ft: JF 223, BHW e BLI 8317 SIS)! Së ir HIER LL). T | Cl g Bow illas at but loca!ly f t(G2S2) , DSC, ca. 1800 ft; AL 175, BHW 998( (BIBE); -WRCE PM212199, AMP & SAP 3613, BHW 20900, 20951, 20954, 23035, BHW & MCJ 16854, BHW & BLT 8325a, 8325b, GLW 4484 (SRSC); AMP & I 3613 (TEX- E on al Na J I nual or perennial herb, DSC/DAR, 2400 ft; JF 1422 SC S Bee serpyllifolia (Pers.) Small, Thyme-leaf sand- n annual herb, psc, 3800 ft; BHW 10794 (TEX-LL). Chamaesyce serrula (Engelm.) Wooton & Standl, Sawtooth sandmat, uncommon but locally frequent annual herb, psc, sva, 1800-5100 ft; EGM 165 (BIBE); JF 2196, BHW 10794, 10838 (SRSC); BHW 10838 (TEX-LL). Chamaesyce theriaca (Wheeler) Shinners, Terlingua sandmat, DSC pcA, 1800-3800 ft; CMR 15117 (SAT); JF 2018, 2095, AMP & pers 6139, BHW 13033, 23402, BHW &LCH BG-180, BHW MCJ 16678, 2 BHW & BLT 8325, 8326 (SRSC); BHW & o 8326 (TEX-LL). Chamaesyce rlgulata (L.C. Wheeler) B.L. Turner [Chamae- syce SEET var. KSE TE sele. e Wheeler) Shinners], Bristlecup sandmat, Ge spurge, rare (G5T151) perennial herb, osc, Dea, 1800-2200 ft; AL 164, 165 (BIBE); JF 1695, AMP € SAP 3611, BHW 20899, BHW & MCJ 16960 # pille villifera (Scheele) Small, Hairy sandmat, rare nial herb, sra, 4360-5840 ft; JF 1173, 1305, 1960 Ge Croton bigbendensis B.L. Turner, Big Bend croton, occasional perennial herb, psc, 3000-3880 ft; JF 292, 707, 1463, 1904, MT 158 (SRSC); BLT & e 163 Mar § Croton dioicus Cav, G "frequent and widespread" perennial herb, D DSC, 2150 ft EGM 144 (BIBE); WGD s.n. (TAES); BHW & LCH BG-64 (SRSC S TEK LE), EN) de ,locally ex Torr, Bush croton, encinilla, oc- casional perennial herb, DAR, DCA, 2200-4100 ft; BRM 3137, TR 72, 73, OCW 314 (BGWMA); JF 298 (BIBE); JF 298, 300, 509, 1114, TR 72, 73, MT 40, 98, 256, OCW 340, BHW 10748 (SRSC); BHW 10748 (TEX-LL). Croton incanus Kunth, Torrey croton, rare shrub, pca, 1800 ft; e 2 128, Md 18406 E I C C. Johnst, Tharp croton, rare annual herb, DSC/DAR, 2400 ft; JF 1427 (BIBE & SRSC). 8 Croton pcttsii (Klotzsch) Müll. Arg, Leatherweed, perennial herb, 2200- 2400 ft; 4 (BGWMA & SR C. Johnst) B.L. Turner Golo poi var. thermophilus (MC. pe M.C. Johnst], Leather- weed, occasional (G5T251) perennial herb, psc, DCA, svo, 1850-4400 ft; BRM 3113 (BGWMA); AL 158 (BIBE); JF 175, ^ 431, 457, 682, 1077, 1397, 1541, 1932, 2011, 2058, MT 159, BHW 18401, 20895, BHW & MCJ 16837, 16862, BHW & BLT 8321, GLW 4439 (SRSC). Ditaxis humilis (Engelm. & A. Gray) Pax, Low silverbush, rare EE herb, psc, 2560-2760 ft; JF 152, 1404, BHW & LCH BG-201 (SRSC). Ditaxis Lais cana (Müll. Arg.) A. Hiele New Mexico ditaxis, | R 1800-3500 ft; CMR 5032 (BGWMA); EGM 133, BHW 12889 (BIBE); BA2 SE 5584 (SAT); JF 168, 439, 674, 2051, BLT 22-151, BHW 2889, BHW & LCH BG-199, BHW 8: MCJ 16858 (SRSC); BLT 2 a l, BHW 12889 LES LL). , Candelilla Herb, DSC, DAR, DCA, SYG, 1900-44 400 ft; BA2 5241, 5695 (SAT); JF 259, 573, 778, MT 84, 170, BHW & MCI 16839 (SRSC); AT 2243 (TEX-LL) Euphorbia brachycera Engelm, Horned spurge, uncommon perennial herb, pca, syG/cHp, 3500-5680 ft; JF 160 (BIBE); BHW 8: LCH s.n. (BGWMA); BA2 2513, 5445 (SAT); JF 1273, 1290, 1934, 1940, AMP € SAP 5141, BHW E LCH BG-1/4 (SRSC). abs eriantha Benth, Desert poinsettia, beetle spurge, ommon annual herb, osc, 1800-1850 ft; JF 2010, 2103; 4 (SRSC). rpg Euphorbia exstipulata Engelm, Square-seed spurge, uncom- mon annual herb, psc, sra, 2800-5840 ft; JF 198, 1980, BHW & MCJ 16832 (SRSC). Jatropha dioica E var. graminea Mevaugn, Se de go, leathers 1800-4360 ft; Sr 17230(BIBE); BA2 5479 (SAT); if Geet E dn ak 12902, pue oe DI ay Spreng, Je | | ER er, occasional subshrub, psc, DAR, DCA, 1850- 4360 ft; BHW& LCH s.n. (BGWMA); JF 160 Sdn is 5206, 5292, 5593, 5693 (SAT); JF 160, 205, 297, 526, 825, 1074, 1400, TRV s.n., BHW & LCH BG-146, BHW & MC 1 Ge GLW 4450 (SRSC). 717 Tragia amblyodonta (Müll. Arg.) Pax & K. Hoffm, Dogtooth noseburn, uncommon perennial herb, psc, DAR, DCA, 2680-5680 ft; JF 146, 718, 832, 1297, BLT 22-150, BHW & LCH BG-188 GE GH ae VES d dn herb, D GE 5840 ft; TR 74, CMR 5085 (BGWMA); BA2 5208, pen 5694 (SAT); JF 688, 1966 (SRSC). Fabaceae jt Al Gu Guajillo, rare tree, psc, sva, 2520-4700 ft; JF 669 (BIBE); JF 669, 2187, BHW 21189b (SRSC). Acacia constricta Benth. ex A. Gray, Whitethorn acacia, occa- sional shrub or tree, psc, DCA, 2350 ft; C 1592 (BGWMA); LF 86, PAS 58 (SHST); JF 1385 (SRSC); VB 355 (TEX-LL). Acacia ee (L) Willd. [Acacia smallii Isely], Huisache, on tree, psc, Rio, 1800 ft; BHW 13040 (BIBE); JF 2030, EJL n TCM 2003-66b, PM 2037, AMP 3348, BHW 13040, 18386 (SRSC); AMP 3348, BHW 13040 (TEX-LL). Acacia Ml Gray var. EC Catclaw acacia, uncomm shr rub, DAR, DCA, 2350 ft; SJ 6505 (SAT); JF 1372, aW 21201, GE o Acacia neovernicosa Isely, Viscid acacia, uncommon to occa- sional shrub or tree, DSC, DAR, 3408 ft; EGM 149 (BIBE); + e e od E G BH 20777 ORG) psc/ RIO, y 1000- i fo o 2002, 2027 (SRSO. er acacia, tree, DSC, DAR, DCA, 1 800-41 80 aa BRM 3116, BHW & LCH s.n. (BGWMA); JF 249, 644, 1012, BHW & LCH BG-147 (SRSC). S Acacia vias Torr, Schott acacia, rare shrub or tree, psc; BHW 20 96 (SRSC). II CA I C má + ial herb, DSC, DAR, pca, 3000-3560 ft; ual o p n pn 304 (S RSC). S Astragalus mollissimus Torr, Woolly locoweed, perennial hei b; nd ao 34419 5 (SAT). A. Gray, Wright milkvetch, annual herb; VLC 43 964 (TEX-LL). + Calliandra iselyi B.L. Turner, Falsemesquite, occasional subshrub, psc, svo, 2800-5150 ft; JF 813, 938, 1142, 1324, Bus (SRSC). ic | subsp (R YE Murray, Mexi- c can redbud,'"infrequent'to'common' 'tree, 3500-4600 ft; BA2 5232 (SAT); AMP & SAP 5140 (SRSC), BHW & LCH BG-172 (SRSC & TEX-LL). 8 Dalea aurea Nutt. ex Fraser var. aurea, Golden prairie clover, perennial herb; BRM 3223 (BGWMA); BA2 5318 (SAT). Dalea formosa Torr, Feather dalea, plume dalea, occasional shrub, psc, DAR, DCA, SYG/CHP, 2560-5840 ft; CMR 17225 (BIBE); BA2 315,5191, 5538 (SAT); JF 483, 559, 1014, 1047 1096, 1402, 1914, 1996, MT 136 (SRSC); VB 342 (TEX-LL). Dalea frutescens A. Gray var. frutescens, Black dalea, uncom- on shrub, DSC, DAR, DCA, SYG, 3350—5800 ft; BA? 5484, 5332 (SAT); JF 374a, 1195, MT 230, 245, BHW 10754 (SRSC); BHW 10754 (TEX-LL). Dalea greggii A. Gray, Gregg prairie clover, uncommon sub- shrub, psc, svo, 3800-4600 ft; OCW 9022 (BGWMA); BA2 5277, 5469, 5548 (SAT); JF 1038, OCW 337 (SRSC). 718 S Dalea jamesii (Torr) Torr. & A. Gray, James prairie clover, perennial herb; BA2 5228 (SAT). herb; MT1 50 (SRSC). $ Dalea dee eds SE Foxtail prairie clover, annual , Dsc; OC 21 (TAES). § aes as os Cory, Downy prairie clover, uncom- nial herb, osc, DAR, ca. 2500 ft; AMP & SAP 6140, BLT & GH en (SRSC); BLT & MT2 97-344 (TEX-LL). + Dalea nana Torr. & A. Gray, Dwarf prairie clover, uncom- mon perennial herb, DAR, syG/cHp, 2750-5700 ft; JF 959, 1265 (SRSC). Dalea nec mencana a (A. Ges pue New mexico dalea, “fre- A, 1800-3440 ft; BHW 13032 KO JF 357, BHW GE ER MCJ 16679, BHW & BLT 8319, GLW 4473 (SRSC); BHW 13030 (TAES); BLT & MT2 97- 334, BHW 13032, BHW & BLT 8319 (TEX-LL). Dalea dsc A. Gray var. pogonathera, Bearded dalea, on perennial herb, psc, DAR, 3560 ft; BRM 3119, OC EE (BGWMA); JF 461 (BIBE €: SRSC); BHW & LCH BG- 196 (SRSC); BA2 5377, 5488 (SAT); VLC 43968 (TEX-LL). 8 Dalea terminalis M.E. Jones, [Dalea lanata var. terminalis (M.E. Jones) Barneby], Wooly s clover, uncommon perennial herb, psc, DCA, Nen Ps 2, DG, TE, & PC 4602, BHW 13030 (BIBE); BLT, JDA & s BHW 13030, GLW 4480 (SRSC); BHW 13030 SECH DSC & HBC 30629, BHW 13030 (TEX-LL) Dalea wrightii A. Gray, Wright dalea, uncommon perennial herb, psc, DAR, 2750-3560 ft; BA2 5430 (SAT); JF 282, ode 962, MT 13, 15, 220, BLT & MT2 97-345, 97-349, BHW C30 (SRSC); VLC 43969, BHW C300 (TEX-LL RE Modi Scheele, Velvet bundlefswen uncom- nnial herb, psc, DAR svc, 3360-5840 ft; JF 227, 322, 7130, a 1910, 1978, MT 35 (SRSC). Eysenhardtia texana Scheele, Texas kidneywood, occasional shrub, pca, DAR, 1800-4150 ft; WFM 27, CMR 5125, 11611, OCW s.n., 320a (BGWMA); BA2 5254 (SAT); SCB 223, JF 201, 1112, 1929, 2136, JF & MY 1318, MT 195, OCW 320, BHW & Ko ial herb, par, oca, 1800-2780 ft; JF 961 (BIBE); 188, 940, 961, 1060, ECH 2141 (SRSC); Ke 2090 (TEX-L L). | A. Gray, S rare BT ber SYG, 5300 ft JF 1998 (SRSO). Leucaena retusa Benth, Goldenball leadtree, woohoo tree, occasional tree, psc, DAR, DCA, 2200-5200 ft; WFM s.n. (BGWMA); PVW s.n. (KANU); BA2 5190 (SAT); JF 748, 1020, 1309, AMP & SAP 5144, TR 19, OCW 1, BHW 10849, BHW & LCH BG-135, GLW 4472 (SRSC); RB 1502, BHW € LCH BG-135 8 Lupinus havardii S. Watson, Big bend bluebonnet, annual herb, DAR, 2560 ft; OES 1344 (TEX-LL). "Se GEN sativa L, Alfalfa, annual herb, 1800 ft; EGM 136a (BIBE *§ ipa mei (L) All, Annual yellow sweetclover, "infre- quent "nnus herb, Td ur E nos) AA; Vtonawa AL, ND | Catclaw os common Gre DSC, Se DCA, 3050 ft; WEM 26, 29, 30 (BGWMA); BA2 5238, 5702 (SAT); JF 1150 (SRSC) Mimosa borealis A. Gray, Fragrant mimosa, "infrequent" to common SCH DAR, DCA, 1900-3760 ft; JF 711, 1693, BHW 21166 (SRSC). Mimosa WE Benth, Emory mimosa, occasional shrub, DSC DCA, 2850-3560 ft; JF 848, 970, GLW 4445 (SRSC). + nin texana (A. Gray) Small, Catclaw, occasional shrub, DAR, DCA, SYG, 1800-4600 ft; JF 1015, 1019, 1357, 2145 (SRSC). it Mimosa turneri Barneby, Desert mimosa, Turner mimosa, uncommon shrub, psc, DAR, svo, 3000-4900 ft; JF 1161, ad 1552, eos o | +l , rare tree, RIO, 1800 ft; o pps (SRSC). # Peteria scoparia A. Gray, Rush peteria, rare shrub, psc, 3840 ft; JF 247 (SRSC Pomaria melanosticta S. Schauer [Caesalpinia parryi (E. Fisher) Eifert, Correll & E Caesalpinia melao tica (5. Schauer Fisher; C i (Fisher) Bl al Parry holdback occasional subshrub, DSC, DAR, DCA, 1800— 3500 ft; BRM 3148, M a-c, BHW & LCH s.n. (BGWMA); AL 183; BHW 884, 12891 (BIBE); BA2 5383, E (SAT); JF 148, 343, 655, 1328, 2130, 2182, BLT & MT2 97-332, BHW 10830, 12891, 20952, BHW&LCH BG-141, BHW& MCI 16063, 16835, MT 221 (SRSC); BHW & MCJ 16835 (SWT); BLT& BD 23-168, BLT 8: MT2 97-332, BHW 12891, BHW & LCH BG-141 (TEX-LL). + E glandulosa Torr. var. torreyana (L. D. Benson) M.C. , Mesquite, occasional shrub or tree, DSC, DAR, SYG, SC p ft; JF 807 (SRSC). + Rhynchosia senna Gillies ex Hook. & Arn. var. texana (Torr. & A. Gray) M.C. Johnst, Texas snoutbean, rare perennial herbaceous vine, DCA, 3050 ft; JF 1119 (BIBE €: SRSC) + ges bauhinioides (A. Gray) H.S. Irwin & Barneby, Twinleaf nna, occasional perennial herb, psc, 2400-3000 ft; JF pot 1465 (SRSC). Senna o (Scheele ex Senar a. Irwin & Barneby, b, DSC, DAR, 1800 ft; E 2072, MT 101 (SRSC). Senna orcuttii (Britton & Rose) H.S. Dui & Barneby, Orcutt mon (G252) herb, sva, cue, 4200- ue JF 1030 (BIBE); JF827, 1030, 1218, 1244, 2191, MT 110, BLT 22-153 (SRSC); JF 1030, BLT 22-153 (TEX-LL). Senn pilosior eo ou GE E ies Ge Ge S: a & Barneby arneby ], Trans-Pe- el LL | | , DSC, DCA, -3500 ft; JF 437, 673, 1326, 1603, 2020, 2046, Ge 1263, BHW 20953, GLW 4475 (SRSC); LEB 9640 (TAES); BHW 20727 (TEX-LL). a da . Gray) H.S. Irwin & Barneby, Dwarf senna, armar ) LIC Imanm H Darmalw Ripley senna, rare (G1SH) n herb, sva, 4320-5000 ft; JF 820, 1518, MT 214 (SRSC) S y secundiflora (Gomez-Ortega) Lag. ex DC, Mescal shrub, oca; BRM 3248 (BGWMA). Vicia E Nutt. va lis (Shinners) B.L. Turner, Deer pe A, 3560 ft; BRM 719 3114 (BG eg JF 540, BHW & MCJ 15994 esch Fagaceae Quercus grisea Liebm, Gray oak, uncommon shrub, che, 4880—5680 ft; JF 757 (BIBE); JF 744c, 757, 1246, 1258, 1280, , 1320, MT 149, 225, 240, 250 (SRSC); JF 1282 (TEX-LL). § Direicüs intricata Trel, Dwarf oak, shrub, 5800 ft; PVW s.n. KANU). im TS Small, Lacey Sg rare shrub, oca, 4350 ft; JF C). In Sue Peaks c rue ba Buckl. SE db. Mohr shin-oak, uncom n shrub, cup, 5240-5840 ft; JF 1245, 1972, 2161 (SRSO). Dues pungens Liebm, [Quercus pungens subsp. vaseyana (Buckley; E. Murray], Sandpaper oak, occasional shrub, DSC, DCA, SYG/CHP, 3150-5680 ft; CMR 15828, BHW & LCH s.n. (BGWMA); ER 1223 (BIBE); PVW s.n. (KANU); BA2 5193, 5203, 5214, 5224 5725 5243, 525] 52/5 52/9 5328, 5450 5459, 5485, (SAT); JF 234, 303, 725, 736a, 749a-b, 818, 819, 1 209a, 1220, 1022, 1045, 1223, 1308, 1550, 1709, 1897, JF &MY 13180, AMP & SAP 5139, MT 76,82,83, 111, 112,113, 117,118, 130, 135, 139, 140, 142, 143, 186, 192, 194, 247, BHW 10755, 21184, 23397, BHW & LCH BG-163 (SRSC); BHW 10755, BHW & LCH BG-163 (TEX-LL). Fouquieriaceae decimal Rip di Engelm, Ocotillo, devil's coachwhip, rub, osc, svo, 1800-4880 ft; JM 89 (SAT); JF 2 2 Leo Garryaceae al aa jad "PRATS 4 D CA Il Mal I J ing, Goldman il SYG, CHP, 4040— 5840 ft PVW s.n. (KANU); JF 504, 770, 1202, 1239, 1307, PM 975, MT 114 (SRSC). Gentianaceae Centaurium arizonicum (A. Gray) A. Heller, Arizona centaury, Rosita, uncommon annual herb, psc, DAR, Ro, 1800-3640 T JF 710a, HTF 1232, MT 10 (SRSC). mE calycosum (Buckley) Fernald, Centaury, uncom- o"rare"or"sparse"annual herb, chp, 5150 ft; BA2 5194, Ge (SAT); JF 1323, AMP € SAP 5130, TR 153, MT 178, 196, W & LCH BG-156 (SRSC). Fustoma exaltatum (L.) Salisb. ex G. Don, Catchfly prairie gentian, uncommon annual herb, no, 1800 ft; BA2, DG, TF, & PC 4604, EGM 135, BHW 648, 13048 (BIBE); JF 2254, BHW 13048 (SRSC); BHW 13048 (TEX-LL). drangaceae # ees GE Thornber, Cliff fendlerbush, rare shrub, sva, 4040-5840 ft; JF 721a, 1266, 1277, 1963 (SRSC). $ Tee SE Gray var. microphyllus, Littleleaf mockorange, rare shrub, sya, cHr, 4040-4900 ft; JF 719, 120,433 GROG) Hydrophyllaceae t+ Nama dichotomum (Ruiz & Pav.) Choisy, Wishbone fiddle- ex rare annual aa CHP, 3840 ft; T 1965 (SRSC). herb, DSC, DAR P. 3480 ft; JF 317, 2035 (SRSC); AT 2241 (TEX-LL). DSC, DAR, 1850-3560 ft; AL 176 (BIBE); JF 477, 1006, GLW 4467 (SRSC); BHW C307 (TEX-LL Phacelia congesta Hook, Spike phacelia, bluecurls, uncom- mon annual herb, DAR, DCA, sv, 2560-5080 ft; JF 423, 505a, 744a (SRSC); RRI 556 (TEX-LL). + Phacelia pallida IM. bs [Phacelia O l. e Johnst;], Pale phacelia, ial herb, pca, 2780-2850 ft; JF 185, 954 (SRSC). S ores Ge Torr, 8 A, Sg Pope phacelia, annual herb, O ft; RRI 553 (TEX-LL). MES wá (J.F. TA LM. Johnst, Stout phacelia, rare to"infrequent" annual herb, par, oca, 2000-3150 ft; JF 1587, 1703, BHW 18402 (SRSC). Loermlandaraan § Juglans major (Torr) A. Heller, Ari Inut, tree, DCA; CMR 2602, 2607 (SAT). Juglans ii Berland, Little walnut, occasional tree, DCA, 3050 ft; JF 1151 (BIBE & SRSC); WFM s.n, CMR 5130 (BGWMAJ; BA2 5235 (SAT). Krameriaceae Krameria erecta WER ex Schult, Range ratany, uncommo rub, psc, sra, 3800-4360 ft; BA2 5344 (SAT); JF 569, Pe 0792 (SRSC); "m 10792 (TEX-LL). Krameria grayi Rose & s White ratany, occasional shrub, DSC, DCA, RIO, SYG; 1800-4800 ft; SMU 49, TR 52 (BGWMA); BA2, DG, TE, & B 4920 (BIBE); Ge 5480 (SAT); JF 236, 371, 828, 1056, 1188, 1898, 2102, TR 17, 152, MT 85, BHW & LCH BG-195, BHW & MU 16863 (SRSC); BHW 636 (TEX-LL). Lamiaceae tt Hedeoma costata Hemsl., Ribbed false pennyroyal, rare e herb, DCA, syG/cHP, 4650-5680 ft; JF 1283a, 1951 (SRSC). S Hedeoma drummondii Benth, Drummond pennyroyal, an- MEA or PET WE SH ec? (TEX-LL). sional annual or perennial herb, 2120- 5800 ft; JF ES ECH JF 494, 534, 546, 679, 869, 949, 1042, 1270, 1329, 1702, 1719, BLT 22-152 (SRSC Se Hedeoma plicata Torr, Veiny false pennyroyal, perennial ep BRM 3129, CMR 5071 (BGWMA); BLT 22-152 (TEX- t4 ro serpyllifolia Small, Reverchon false pennyroyal, occasional perennial herb, psc, syc, 3560-5080 ft; JF 740 (BIBE); JF 230, 281, 740, 834 SRSC). § Mentha arvensis L, Wild mint, rare perennial herb, 1800 ft; EGM 158, 167, BHW 832 e BHW 832 (SRSC & TEX- id Salvia greggii A. Gray, Autumn sage, uncommon shrub, D DCA, SYG/CHP, 3500-5680 ^ BRM & BPM 3103 (BGWMA); P 505, 766, 1283, 2171, TR 154, MT 107, BHW € LCH BG-171 (SRSC). Salvii ] Sc] r sage, Roemer sage, uncom- mon perennial herb, : DCA, ae 2000-5840 ft; JF 1284, 1671, 1675, 1975, BHW 18404 (SRSC); BHW 18404 (TEX-LL). 8 Teucrium depressum Small [Teucrium cubense var. densum Je eucrium cubense subsp. depressum (Small) E.M. McClint. €: Epling], Small coastal germander, uncom- sá 720 mon annual herb, 1800-3520 ft; HTF 1234, BHW 8331, GLW 4465 (SRSC). inaceae Linum berlandieri Hook, Berlandier flax, occasional annual or perennial herb, DSC, DAR, DCA, SYG, 600 ft; CMR 5090; n. (BGWMA); i 827 (BIBE); BA2 5338, 5568 W BHW & LCH (SAT): JF SC 295 525, 552, 8 & LCH BG-149 (SRSC). it ped hudsonioides Planch, Texas flax, rare perennial herb, 93 (SRSC). ft; JF & HL 18 Linum Ee (A. Gray) Engelm. ex A. Gray, Rock flax, oc- casional perennial herb, psc, DAR, DCA, 1800-3840 ft; BR 3136, BHW & LCH s.n. (BGWMA); BA2 5218, 5310, 5703 (SAT); JF 241, 276, 847, 1016, 2071, MT 48, 177, TR 282, OCW 509, BHW 10828, BHW & LCH BG-84, BG-164 (SRSC); BHW 10828 (TEX-LL). Loasaceae Cevallia sinuata D Stinging cevallia, occasional perennial herb, p 2 MR 5077 (BGWMA); JF 172, c Se 1/24, MT 218 (SRSC) Eucnide ges ee lew ttle, uncommon to herb, psc, DCA, 1800-3700 ft; BA2& PC 4754 (BIBE); BRM 3104 (BGWMA]; BA2 4754, 5509 (SAT); JF 177, 286, 532, 2089, BHW 18400, 20619 (SRSC). +o Mentzelia mexicana HJ. Thom mpson & Zavortink, Mexican blazingstar, rare perennial herb, DAR, pca, 1800 ft; JF 2031, 2066 (SRSC). +Ao Mentzelia multiflora (Nutt.) A. Gray, Desert mentzelia, or perennial herb, observed at the Boquillas [eStore o site. rennial herb, DSC, DCA, 2350- 3720 JF 23}, 1333 (SRSC). 4. Johnst, mon perennial herb, osc, DCA, 2850-3640 ft JF 312 840, 950, BHW 13071 (SRSC). a taa ldlej ifoli butterfly! , Occasional hrub, DSC, DAR, DCA, 1850- 3900 f CMR 11586 ba 5J 6502 (SAT); JF 150, 436, 835, 1375, 1721, 2012, MT 9, 168, BHW 10759, BHW 8: LCH BG-77 (SRSC); BHW 10759, BHW A LCH BG-77 (TEX-LL). hraceae § Lythrum californicum Torr. & A. Gray, California loosestrife, “infrequent” perennial herb, 1800 ft; BHW 896 (TEX-LL). Malphigiaceae Janusia gracilis A. Gray, Helicopter bush, slender janusia, occa- 1 | ial herl ine, DSC, DAR, DCA, 1800-3720 ft; EGM 122 (BIBE); CMR 15112 (SAT); JF 242, 647, 1332, AMP & SAP 5308, BLT & MT2 97-377, BHW 10793 (SRSC); BLT & MT2 97-337 (TEX-LL). Malvaceae Abutilon malacum S. Watson, Yellow indian mallow, uncom- mon subshrub, osc, DAR, DCA, 1850-2050 ft; EGM 154 (BIBE); E 643, 1082, 1604, A ii a7 e 12886 O A Gra ommon TEA! VES perennial herb, psc, DAR, DCA, 2750-3700 ft; AF 187 (BIBE); JF 841, 969, 1140, 2169 (SRSC). E EI A. Gray, Wright indian mallow, occasional perennial herb, psc, DAR DCA, 1800-3560 ft; TR s.n. (BGWMA); JF Ps 668, 1011, 1648, 2084, BLT & BD 23-171 (SRSC); BLT & BD 23-171, BHW 9158, 10839 (TEX-LL). S Herissantia jd (L) Brizicky, Netvein mallow, “frequent” , DAR, 1800-1900 ft; HCH s.n., BHW 13064 Hibiscus coulteri Harv. ex A. Gray, Desert rosemallow, oc- casional subshrub, 2400-4300 ft; JF 444, 846, 1480, 1582, 2184, MT 51, BHW 10820, BHW & LCH BG-152 (SRSC); BHW 10820, BHW & LCH BG-152 (TEX-LL). psc, DCA, 3200 ft; BRM 3120 (BGWMA); JF 1471 (SRSC). S Hibiscus martianus Zucc.- bin rosemallow, "sparse" subshrub, 2100 ft; OCW 358 (SRSC). *# Malva ROO L, Cheeseweed mallow, rare annual or nial herb, rio, 1800 ft; AL 304, 308 (BIBE €: SRSC). Se ela lepidota (A. Gray) Fryxell, Scurfymallow, perennial MENO, DAR, 2560 ft; OFS 1335 (TAES). la! (Ortega) Krapov., Alkali mallow, rare perennial herb, psc/ban, 2400 ft; JF 1423 (SRSC). 8 Rhynchosida physocalyx (A. Gray) Fryxell, Spearleaf sida, buffpetal, “infrequent” perennial herb, 2250 ft; BRM 3171 (BGWMA); BHW & LCH BG-194 (SRS Sida abutifolia Mill, Spreading sida, occasional perennial herb, psc, DCA, 1800-2850 ft; JF 166, 956, 1325, 1439, 2006, HCH s.n. (SRSC). +0 Sida longipes A. Gray, Stockflower fanpetals, uncommon perennial herb, psc, DAR, svc, 3440-4800 ft; JF 1539, 1920, 2175 (SRSC). Sphaeralcea angustifolia (Cav) G. Don, teure globe- mallow, occasional perennial herb, DSC, DAR, DCA, RIO, SYG, 1800-5300 ft; BRM 3121, T 11593 (BGWMA) CN 499 (BIBE); JF 585, 803, 1415, 1461, 1986, 2104, 2268, BH 641 (SRSC); BHW 247 (TEX-LL). Se Sphaeralcea hastulata A. Gray, Spear globemallow, “infre- quent" perennial herb, pca, 2560 ft; RRI 21431 (TEX-LL). rleliamunarni — Nyctagina a pa ia (Torr) R.A. Levin, Narrowleaf mono- pod, occasional to uncommon subshrub, DSC, DAR, DCA, 1800-3720 ft; JF 243 (BIBE); JF 243, 844, 1352, 2015, 2022, BLT & BD 23-160, BHW 9159, 10790, BHW & MCJ 16846, BHW & BLT 8324 (SRSC); BLT & BD 23-160, BLT & MT2 97-346b, BHW Acleisanthes SE A. Gray, Ang Ge hierba de la rabia, uncomm | her! oca, 1850-4000 ft; BA2 5188 (SAT); p 1070, MT 157, Eeer RSC). Allionia incarnata e var. ea eee d pro pas windmills, DCA ft; EGM 168 (BIBE); BA2 5356 (SAT); JF a 651, En p AMP & SAP 5304, BHW 20879, BHW & MCJ 16848 (SRSC); BHW 702 (TE s Ammocodon chenopodioides (A. as SEH E A.l perennial herb; a s aa iem Fenstermacher et al., Vascular flora of the Dead Horse Mountains en 8 Anul lis eriosol (A. Gray) Standl. [B A. Gray], Big bend ringstem, “infrequent to locally com- mon” annual or perennial herb, psc, DAR, DCA, 1800-2320 ft; EGM 146 (BIBE); AMP & SAP 6141, BLT 8 MT2 97-351, BHW 16676, 20878, 20949 (SRSC); DSC & HBC 30639, OES 1632, BLT -340 (TEX-LL). Boerhavia anisophylla Torr, Wineflower, uncommon perennial herb, DAR, DCA, 1800-3000 ft; JF 1468, 2073, 2134, AMP & SAP 3614, BHW 16690, BHW, BLT, & JDA 8335 (SRSC); BHW 21031, W. BLT, & JDA 8335 (TEX-LL). + Boerhavia coccinea Miller, Red spiderling, rare annual or perennial herb, pca, 3065 ft; JF 7720 (SRSC). 8 eran id M.E. Jones, Spreading spiderling, o “frequent” annual herb, DSC, DAR, DCA, 1800- ne En we SAP MLP & AL? 6102, TRV s.n, BHW 20877, BHW & LCH BG-185BHW € BLT 8315 (SRSC AMP & SAP 3614, BLT & MT2 97-342 (TEX-LL). § Boerhavia linearifolia A. Gray, Narrowleaf spiderling, annual , 4600 ft; BA2 6190, CMR 5060, OCW 5810-1 (BGWMA); BA2 5326 (SAT); AMP & SAP 5120 (SRSC). S Boerhavia spicata Choisy, Creeping spiderling, "infrequent to frequent" annual herb, 1800 ft; BHW 695, 20948, BHW RK Med 16849 oz ^ C4 J] annual herb, 2100 BHW 8315 (TEX- LL). Head a Canyon in 1948. A newer taxon recognized in the Fior Cyphomeris sypsophicices (M. Martens & Garcon) Stand, ruit, DSC DAR, DCA, SYG, 1800-4360 ft: = E (BIBE): BA2 5449 (SAT); JF 348, 545, 951, 1172, 2086 (S 8 Mirabilis albida (Walter) ibi White four oclock, peren- nial herb; BA2 5527 (SAT) + Mirabilis linearis (Pursh) Heimerl, Linearleaf spiderling, rare perennial herb, sva/cHP, 5300-5840 ft; JF 1982, 2159 Mirabilis texensis (J.M. Coult) B.L. Turner, Texas mirabilis, un- common perennial herb, psc, par, DCA, 2750—4200 ft; JF 165, 809, 960, 1502, 2173, BLT & MI2 97-343, BHW 10776, 10844, BHW & MCJ 16829 (SRSC); BLT & MT2 97-343 (TEX-LL). 8 Nyctaginia capitata Choisy, Devil's boquet, perennial herb; IM 137 (SAT). Oleaceae Forestiera angustifolia Torr, Narrowleaf forestiera, common Ub, DSC, DAR, DCA, 1920-4400 ft; BHW & LCH BG-153 (BGWMA); BA2 5256, 5266 (SAT); JF 374, 493, 572a, 2181, JF & HL 1899, AMP & SAP 5826, BHW 18411, 21205, BHW & LCH BG-153, BHW & MCJ 15996, GLW 4459 (SRSC); WBM 2022 (TAES). Fraxinus cuspidata Torr, Fragrant ash, rare to “sparse” tree, DAR/ DCA, 3500-4800 ft; BHW & LCH BG-173 (BGWMA & SRSC); JF 1947 (SRSC) Fraxinus greggii A e Gregg ash, common shrub, DSC, DAR, O ft; TR 243 (BGWMA); BA2 5358, 5570 (SAT); JF 9» he p 1131, TR 296, MT 07b, BHW 20623, BHW & MCJ 15983 (SRSC). Menodora longiflora A. Gray, Showy menodora, twinpod, oc- casional perennial herb, psc, DAR, DCA, 2350-4600 ft; CMR 721 5062, OCW 9027 (BGWMA); JF 334 (BIBE); BA2 5447, 5528 (SAT); JF 226, 334, 517, 946, 1115, 1359, 1360, AMP & SAP 5122, 5145, MT 7 (SRSC). Menodora scabra A. Gray, Rough menodora, common s shrub, psc, DAR DCA, 1800-5080 ft; BHW 1123, ee BA2 5699 (SAT); JF 582, 754, 1482, 1520, BGH 639, MT 56, 180, 22, BHW 10783, 10803, 12887, 13037 (SRSC); BHW 10783, 12887 (TEX-LL). Onagraceae S Calylophus greggii A. Gray [Calylophus hartwegii (Benth) P.H. Raven subsp. pubescens (A. Gray) Towner & PH. Raven], Gregg sundrops, "infrequent" subshrub, 2150 ft; HTF SC). Calylophus hartwegii (Benth) PH. Raven subsp. ed Hartweg sundrops, uncommon perennial herb, D 1800-3050 ft; BA2, DG TE, & PC 4609 (BIBE); Bcc dos a, GR >L). Mar harav niloncic DU Ravan 2 mD Cra egent occasional (6252) subshrub, psc, DCA, RIO, , 1800-4400 ft; JS & AL 301, 304 (BIBE); JF 1363, 1380, 1538, 2091, 2117, 2147, AMP & SAP 3608, BHW 1008, 18397, GLW 4481 (SRSC); BHW 1008 (TAES); ae SAP ada de Sch ee herb; 2200- 2400 ft; TR 75 (SRSO). Gaura coccinea Pursh, Scarlet gaura, occasional perennial herb, psc, par, 2200-3520 ft; TR 75 (BGWMA); BA2 5186, 5248, 5290, 5306, Le 5563, 5576 (SAT); JF 1418, 1460, BLT & BD 23-173, GLW 4449 (SRSC); BLT & BD 23-173 (TEX-LL). 8 SC Ge Lehm. [Gaura mollis Kunth], Velvetweed, per nnial herb, rio, 1800 ft; AL 305 (BIBE). 8 Ludwigia peploides (Kunth) PH. Raven, bd pua willow, ‘infrequent to rare’ : | 1231, ECM bí (SRSC); EGM E dd C297 T iE rare perennial herb, DCA, SYGÍCHP, 3560- 5200 ft IF 521, 1311, 1953 (SRSC). Oenothera EG (Spach) liu. Kunth inea OCCa- Mo 1800- ft; BRM 3170, CMR 5139 (BGWMA); JE 5 10, Se 1229 p & SAP 4300, 5388, BHW 18384, 47055 (SRSC); RRI & BHW 21362 (TEX-LL). Se Oenothera laciniata Hill, Cutleaf evening-primrose, annual Or perennial herb; GE 5574 (SAT) m "requer annua Ee 200 ft BHW & MCI 16088 (SRSC X-LL). ex Aiton, Rose si indrops, rare peren- nial herb, pca, 1800-3000 ft; JF 798, 2076 (SRSC). 8 Oenothera triloba Nutt, Stemless evening-primrose, “infre- quent" annual or biennial herb, 1800-4000 ft; BA2 5564 (SAT); HTF 1228 (SRSC). A Orbanchac Orobanche oc Nutt, Louisiana broomrape, uncom- mon annual herb, DAR, DCA, 1800-2120 ft; BRM 3172 BONAN a 677 (SRSC); DSC & WOD 30714, BHW 836 TEX-L E i Orobanche multicaulis Brandegee, Spiked broomrape, oc- 722 casional annual herb, DAR, nca, nio, 1800-4080 ft; JF 564, 931, BHW 738 (SRSC) Oxa'igareap dii A. Gray | nial herb; MT 228 (SRSC), , peren- Papaveraceae A LS is G.B Enter Chi tain | b, DSC, DAR, 2350-2400 ho ft UE 1414, 1579 SRSC). Aexican pric klepoppy, rare annual herb, rio, 1800 ft; AL 299 (BIBE); AL 300 (SRSC). Argemone sanguinea Greene, Red pricklypoppy, rare annual or perennial herb, osc/Dar, pca, 2800 ft; CMR 2605 (SAT); JF 966 (SRSC). Passifloraceae Passiflora tenuiloba Engelm, Passionflower vine, occasional perennial herbaceous vine, psc, DAR, DCA, 1800—4280 ft; BRM 3150, 3150b; CMR 5082 (BGWMA); AL 142, 143, 318 (BIBE); BA2 5215 (SAT); JF 189, 306, 963, 1076, 1361, 1591, 2129, TR 281, BHW 20606 (SRSC); TR 249 (TEX-LL). Pedaliaceae 8 Proboscidea louisianica Thell. subsp. fragrans (Lindl) Bret- ting, Devil's claw, "infrequent" annual herb; 1800 ft; BHW 12890 (SRSC). ytolaccaceae Rivinia humilis L, Pigeonberry, uncommon perennial herb, DAR, DCA, 2920-3050 ft; CMR 5031 (BGWMA); JF 216a, 1137 (SRSC) Plantagin Plantago ees SCH dig ya locally common annual 300 ft; OCW 584-20 (BGWMA); JF 620, 785, 1250 SE 8 Plantago hookeriana Fisch. & C.A. Mey, California plantain, "infrequent" annual herb, pca, 2560 ft; BHW 21422 (TEX- LL) +0 Plant t j Jacq, Bristlebract plantain, uncom- mon annual herb DSC, DAR, DCA, SYG, 1800—5300 ft; JF 1252, 1609, 2082 (S w taper Decne, Redseed plantain, uncom- ual herb, psc, DAR, DCA, 2350-2560 ft; BRM 3112 BONDA RH 20, 43 (SAT); JF 1335, 1430 (SRSC); RRI 21423 (TEX-LL) Polemoniaceae Gilia incisa o Splitleaf gilia, rare annual or perennial eri 850-4000 ft; BA2 5308, 5573, 5577 (SAT); JF 944 (SR sch § Gilia insignis (Brand) Cory €: H.B. Parks, Marked gilia, “in- frequent" perennial herb, 2250 ft; BHW & LCH BG-200 (SRSC) S i Gilia rigidula Benth. subsp. acerosa (A. Gray) irt Button gilia, blue gilia, uncommon annual herb, psc, 3560-4300 ft; BA2 5372 (SAT); JF 245, E» pa MT 97 (SRS Gilia stewartii |M. Johnst, Stewart giiia, uncommon annua herb, psc, pan, 3400-4800 ft; BRM a (BGWMA); BA2 [| £ al Dos D In LI PET E £T [1 5557 (SAT); JF 498, 814, 1026, 2163, BHW 10761 (SRSC); BHW 10761 (TEX-LL). § SR havardii (A. Gray) VE Grant, Havard ipomopsis, requent" perennial herb; BHW & MU 15907 (SRSC). Se o Nutt, Santa Fe phlox, [Ph/ox mesoleuca Greene], perennial herb; BA2 5371, 5519, 5562 (SAT). Polygalace dei aba Nutt, White E pde perennial erb, psc, cHP, 3500-5150 ft; ACK 826 (BIBE); BA2 5212, 5378 e JF 251, 1227, MT 72, BHW & ICH BG- 158 (SRSC). Polygala barbeyana Chodat, [Polygala longa S.F. Blake], Nar- rowleaf polygala, occasional to uncommon perennial he b DSC, DAR, DCA, SYG, 2200- 3500 ft M 122, BHW & LCH s.n \ 1006 BA? 5286, SJ 6506 (SAT); JF 158, 252, 267, 288, 864, 1099 1160, 1259, 1503, 2174, 2183, TR 122, CMR 5070, MT 28, 244, BHW 10798, 21197, 23401, BHW & LCH BG-169, BHW 8: MCI 16826 (SRSC); BLT & BD 23-1720, BHW 10798 (TEX-LL). ires diii A. Gray var. eri Se bby milkwort, onal perennial herb, D O ft; BA2 5196, E KC Se JF e. 1009 ps SH e 2070, MT , & LCH BG-70, BG-134, BG-182 (SRSC) JF 107 I, oS: ich BG- di Mesta uu he be |, (2003). PIN) ALCA! These ti two varieties as in Brewster Fano bin County, and acters [e.g., JF 964 (SRSC); Powell in prep.]. + jn lindheimeri A. Gray var. parvifolia Wheelock [Po- lygala tweedyi Britton ex Wheelock], Shrubby milkwort, mmon perennial herb, psc, Dea, 1850-2600 ft; JF 929, 1081 (SRSC). Polygala macradenia A. Ge Glandleaf milkwort, common perennial herb, o oca, 1850-4360 ft; BRM 3226 (BGWMA); ACK e BO: BA2 5228, 5547, 5553 (SAT); JF 239, 275, 354, 854, 1529, 2007, BHW 10797, 20955, BHw & BLT 8316, GLW 4446 (SRSC); BLT & BD 23-166, TRV, GLB & WEH 85-46, Ge BLT 83 16 dee D il are perennial "herb psc, DCA, 1800- a BA2, DG, TE, HN AL 312 (BIBE); JF 1670, 1698, 2126 (SRSC) Polygala scoparioides Chodat, Broom milkwort, occasional Geier herb, DSC, DAR, DCA, 2200-4800 ft; CMR 5064, 11590 BGWMA); BA2 5289, SJ 6499 (SAT); JF 425, 506, 580, 1165, 1190, 1365, TR 124, OCW 502, BHW 20624, 21199, GLW 4440, 4452 (SRSC); BLT & BD 23-165, 23-172a (TEX-LL). Polygonaceae # Eriogonum havardii S. Watson, Havard buckwheat, rare perennial herb, osc, 3350 ft; JF 1718 (BIBE €: SRSC); JF ipei platyphyllum Torr. ex Benth, Broad-leaf wild buck- at, occasional perennial herb, par, pca, 1950-4600 ft; i 939, 1381, 1635, AMP & SAP 5123 +0 Eriogonum rotundifolium Benth, Roundleaf buckwheat, rare annual herb, par, 2350 ft; JF 1384 (SRSC § Eriogonum tenellum Torr, Tall wild Duck eds perennial herb; BRM 3206 (BGWMA). Fenstermacher et al., Vascular flora of the Dead Horse Mountains S EE hydropiperoides Michx, Swamp smartweed, 'infrequent" perennial herb, 1800 ft; BHW, BLT & JDA 8336 (SRSC) 8 de dll DER UNTEN L, Pennsylvania smartweed, M. Gómez], annual herb, 1800 A BHW 641 GER ray [Po] ia L.], Spotted jadyethuinb, infeauent*anntlal herb 1800-1850 ft; EGM 161, BHW 642 (BIBE); BHW 21517 (TEX-LL *§ pbs crispus L, Curly Sg “infrequent” serena herb, 800 ft; BHW 18387 (SRSC L, Gold Dod: rare annual or biennial herb, pca, 1800 ft; JF 21 eee m Portulacaceae # Talinum angustissimum (Engelm.) Wooton & Standl [Phe- us aurantiacus (Englem.) Kiger], Narrow-leaf fameflower, rare perennial herb, par, 1900 ft; JF 2040a (SRSC). Many treatments include this species under Premia musa GUIDES the ole El NONO Aerea 23 casional annual herb, psc, rio, 1800-2600 ft; JF 932, 2226 (SRSC Rhamnaceae Ceanothus greggii A. Gray, Desert ceanothus, uncommon shrub, pca, syo, 4040-5840 ft; PVW s.n. (KANU); BA2 5537 (SAT); JF 727, 746a, 771, 1211, 1275, 1976, PM 974, MT 226, Eon (SAS). les (A. Gray) M.C. Johnst, Javelinabush, shrub; E | d SAT. nst, FS Aa A alia | ch DSC, DAR, DCA, SYG, o 4880 ft; JF 161, 744b, 1348, 1386, 1421, TVR s.n. (SRS Condalia warnockii M.C. Johnst, Warnock condalia, uncom- on shrub, psc, DAR, DCA, svc, 1800-4650 ft; JF 1185, 2085, BHW 10814 (SRSC); BHW 10814 (TEX-LL). Karwinskia humboldtiana (Willd. ex Roem. & Schult.) cc, Coyotillo, locally uncommon shrub, DCA, RIO /DSC, 1800-3150 ft; BHW 834 (BIBE, SRSC, €: TEX-LL); JF 1595, 200 1, E 135 (SRSC) k.ex Torr. & A Gray) A Gray, Lotebush, (FNA 2 ttl ing separate species break down under a continuum of intergradation, but collections e n Brewster Cou nty UF did di SRSC) demonstrate deg Individ uals ] aut the same site IK OI) with n no intergradation or characters (Powell in prep.). [Talii IMG Fon pp har] J de UC 1,0 £. £] “all DSC, 3880 ft: JF 293, 2040a (SRSC). # Phemeranthus brevicaulis (S. Watson) Kiger, Dwarf fame- flo ower, [Talinum brevicaule S. Watson], rare, locally ommon perennial herb, svc/cup, 5320-5840 ft; JF 1967, pu (SRSC). Portulaca See L, Purslane, verdolaga, rare annual herb, bar, 2500-2880 ft; TR s.n. (BGWMA); JE di (SRSC). tul losa L, Kiss me quick, | eren- nial herb, DSC, DAR, DCA, SYG/CHP, 1800-5840 ft; BHW 12901 (BIBE); BA2 5287, pus (SAT); JF 176, 240, 272, 1154, 1979, 2 3, BHW 12901, JF & MY 1316, BHW & MCJ 16843 (SRSC); Bu 12901 (TEX-LL). Primulaceae S Anagallis arvensis L, Scarlet pimpernel, annual or biennial herb, rio, 1800 ft; AL 303 (BIBE). S lus ebracteat bs} I) R. Knuth, Lime- water brookweed, “infrequent” perennial herb, RIO, 1800 ft; EGM 130 (BIBE); HTF 1223, BHW 13049 (SRSC). Ranunculaceae Ane Edna SUDO var. texana Enquist & Ge Desert j , DSC, DCA, 2560-4440 ft JF 562, BHW & MU 15986 (SRSC). Clematis drummondii Torr. & A. Gray, Virgin’s bower, old man's b mmon perennial herbaceous vine, DAR, DCA, RIO, Na 4080 ft a a Ge MA ui 295 Klee, BHW & RLU s.n. Dm Resedaceae +0 Oligomeris linifolia (Vahl) J.F. Macbr, Desert spikes, oc- ocasional PS DSC, DAR, DCA, 2200- ca. 4 000 ft: BA 25226, 5261, 5706 (SAT); JF 1312, 1419, 1426, TR 117 (SRSC); AG 380 (TEX-LL). osaceae Cercocarpus breviflorus A. Gray [Cercocarpus montanus Raf. var. paucidentatus (S. Wats) FL. Martin], Small-leaf mountain ma cus rare shrub, cup, 5800-5840 ft; PVW s.n. (KANU); JF 1267, 1977 (SRSC) Cercocarpus ga R.A. Denham var, glaber [Cercocarpus glaber (S. Wats) Fl Martin], S n on shrub, DCA, syG/cHP, 3500-5680 ft; OCW 9036, BHW & on (BGWMA); PVW s.n. (KANU); BA2 5439, 5452 (SAT); JF 716, 1021, 1197, 1274, 1278, 1281, MT 64, 109, 115, OCW 338, BHW &LCH BG-150 (SRSC). Trans-Pecos mountain mahoga- nies were previously treated as varieties of Cercocarpus montanus Ge 1998) and are now recognized by some as differe (Turner et al. 2003, Powell in prep.). Fallugia pial (D. Don) Endi. E Tos Mu qi OC- A, 2350-4040 ft; BRM 3108, TR s.n. (BGWMA); BA2 Rn p (SAT); JF 157, 1111, 1378, PM 978, TR 114, MT 201, BHW & LCH BG-168 (SRSC); BHW & LCH BG-88 (TEX-LL). Malacomeles denticulata (Kunth) G.N. Jones, [Amelanchier clita lata (Kunth) K ee Serviceberry, uncommon DCA/sYG, CHP, 3800-5080 ft; PVW s.n. (KANU); BA? s en JF 511; ZLAZ288, = 1184, 1954, BHW 10826 RSC); DHR 2298, add ee 6 ( | Bt) Mat rockspirea, tufted rockmat rare ssubshrüb. SYG, 4750- 5840 ft; JF 786, 1269, Mie (SRSC). lii (W. Wight) S. C. Mason, Havard plum, uncom- mon to occasional shrub DAR, pc, svc, 3050-5840 ft; JF 1 109 (BIBE); JF 742, 1109, 1266a, 1973, PM 973 (SRSC); BHW & LCH d 140 DIRCE & TEX- LL). | + ] icl , Heat th cliffrose, com- mon -— | DCA, SYG, CHP, » 2050- 5150 ft; BA2 6192, CMR 724 5067, 5133, 15832, TR s.n. (BGWMAJ; PVW s.n. (KANU); BA2 5366 (SAT); JF 751, 948, 1098, 1601, 1672, 1902, MT 25, 94, 171, TR 115, BHW 10836, 16682, 20622, BHW & LCH BG-136, BHW & MCJ 16831 (SRSC); BHW 10836, BHW & LCH BG-138 TERA Vauguelinia copmibosa Humb. & Bonpl. subsp. angustifolia (Rydb.) W.J. Hess & Henrickson, Narrow-leaf vauquelinia, occasional shrub or tree, DCA, svc, 4200-4800 ft; BA2 555 (SAD); JF 496a, 577, 862, PM 972, BHW 10778, 10784, BHW & MCJ 16683 (SRSC); BHW 10778 (TEX-LL). dadas LL A erb, DAR DCA, SYG, 3500- Ge ft; JF 428, 745 (SRSQ. Hedyotis acerosa A. Gray var. acerosa, Needleleaf bluet, com- mon perennial herb, o R, DCA, svG/cHP, 3500-5840 ft; - BRM 3229 (BGWMA); ee (BIBE); JF 250, 287, 330, 480, 491,824, 1051, 1101, 1717, MT 27a, 57, OCW 323, BHW 10825 (SRSC); BHW 10825 (TEX-LL). Hedyotis intricata Fosberg, Tangled starviolet, fascicled bluet, occasional shrub, psc, pca, svc/cHP, 3800-5800 ft; PVW s.n. (KANU); JF 294, 481, 749, 866, 1487, MT 91, 169, BHW & MU TOODA PRECI B | nigricans, Diamondflow- ers, c common perennial herb, DAR Dx DCA, 1800- 4360 ft; BRM 3139 (BGWMA); JF 500, AL 196 (BIBE); BA2 5342 (SAT); JF 156, 186, 424, 500, 541, 801, 823, 1129, 1530, 1924, 1925, 1926, 1931, JF & MY 1354 (SRSC); BLT & MT2 97-335 (TEX-LL). Hedyotis pooleana B.L. Turner, [Stenaria mullerae var. pooleana (B.L. Turner) Terrell]. Jackies bluet, occasional (G1T1S1) perennial herb, psc, svc, 2700-5680 ft; JF 237, 713, 775, 1168, 1288, 1553, 1605, 1715, 1716, 1916, 1961, 1962, 2192, 2195, JF & ME 553, JMP 2527 (RC; id MS l L). TI ist Pp J H | | ! Er + mu in addon to the previously mentioned EE over two specimens of Hedyotis angulata seem to approach vegetative characters of Hedyotis pooleana (Tharp 270, Warnock 23716 SRSC) which is i restricted to h sms e mountains slightly morphological past with | a Mew specimens showing a habit, even approac Hedyotis pl as Ens to the sc densely-matted growth. Rutaceae: folia (Benth) V.L. Bailey, Common hoptree, skunkbush, rare shrub, SYG, CHP, 4800— 5680 ft BA2 327 (SAT); JF 1213, 1294, 2162, MT 185 (SRSC). Thamnosma texanum (A. Gray) Torr, Dutchman's breeches, Texas desertrue, occasional to uncommon perennial herb, psc, DAR, DCA, SYG, 1800-5840 ft; BRM 3122, CMR 5081 (BGWMA); JF 488, 747, 1583, 1995, BHW 18394, BHW & LCH BG-69, BHW & MCI 15906, GLW 4441 (SRSC). f Salicaceae § Populus fremontii S. Wats, Fremont cottonwood, locally occasional tree, nio, 1800 ft; EW 19709 (SRSC). | gal rn a a Em L | de" dau ndo f Texas 2(1) § Salix exigua Nutt, Narrowleaf willow, coyote willow, “in- frequent to frequent" tree; BHW 12911 (SRSC); BHW s.n. TEX-L 8 Salix exilifolia Dorn [Salix taxifolia Kunth], Yewleaf willow, Salix nigra Marshall, Black willow, uncommon tree, DCa, 1800-2350 ft; BHW 647 (BIBE); JF 1344, BHW 18389 fee EGM 71, BHW 21505 (TEX-LL). Sapindac + ae saponario L. id aei p (Hook. & Arn.) L.D. Benson oncillo, rare tree, DAR, 3050 ft; JF 1116 (SRSC). AI AA H kh I 4- (SAT); AASA Er BPM 3109 TRB4(BGWMA); ro T DCA, 18 JP 817, 1144 2075, 2127, TR 84 (SRSC). Sapotaceae § Sideroxylon lanuginosum Michx. ssp. rigidum (Gray) T.D. nnington, Gum bully, ironwood, rare tree, DCA, 2560 ft; AL 240 (BIBE); BHW & MC] s.n. (SRSC). Scrophulariaceae 8 Castilleja integra ^. Gray var. integra, Wholeleaf indian paint- brush, perennial herb; BA2 5227, 5345, 5390, 5448, 5460, 5542, 5588 (SAT); BLT 8: BD 23-169 ( TEX-LL). Castilleja rigida Eastw, Rigid indian paintbrush, broadbract p occasional perennial herb, psc, syG/cHP, 2150-5200 ft; TR 111, OCW 9035 (BGWMA); OES 1332 (BIBE); $)6497 (SAT; JF IST, 2 14, 331, 485, 774, 830, 833, 1210, 1580, 1720, 1908, AMP 8: SAP 5121, TR 111, MT 14, 126, 133, 148, on 154, BHW 10851, BHW & LCH BG-71 (SRSC); BHW 10851, W & LCH BG-71 (TEX-LL). Wee candidum IM. Johnst, Boquillas pee OC- casional to b, DSC, DAR, DCA, 180 EGM 153, BHW 13070 (BIBE); JF 329, 567, 836, 837, Se Es 2092, 2124, PM 238, BHW 13070, BHW & LCH BG-615HW & MCI 16857 (SRSC); BHW s.n. (TEX-LL). + Leucophyllum frutescens (Berland.) LM. Johnst, Barometer bush, ceniza, purple sage, rare shrub, pca, 1800 ft; JF 2083 (SRSC). Leucophyilum minus A. Gray, Big Bend silverleaf, common shrub, DSG, DAR, DCA, 2600-4300 ft; CMR 5066 (BGWMA); BA 107 (BIBE); DB s.n. (SAT); JF 244, 266, 470, 831,842, 1162, MT 23, 24, 125, BLT 22-148, BHW 10804, BHW & LCH BG-161, GLW 4471 (SRSC); BLT 22-148 (TEX-LL). Maurandya antirrhiniflora Humb. & Bonpl. ex Willd. [Mauran- della antirrhiniflora (Humb. & Bonpl. ex Willd.) Rothm.], Snapdragon vine, common perennial herbaceous vine, DSC, DAR, DCA, 1800-4080 ft; BRM 3105, TR 110 (BGWMA); JF 438, 971, TR 110, 113, MT 100, BHW 10807 (SRSC). 8 Maurandya wislizeni Engelm. ex A. Gray, Balloonbush, rare perennial herbaceous vine, rio, 1800 ft; AL 307 (BIBE); AL 302 (SRSC). Penstemon baccharifolius Hook, Baccharisleaf beardtongue, ccasional perennial herb, pca, 1800-4600 ft; CMR 5061, MU 39 (BGWMA); AL 145 (BIBE); JAS 3450 (NYBG); JLB 866 (SAT); JF 180, 838, 974, 1313, 1681, 2106, AMP & SAP 5124, TR 112, MT 95, 183, BHW & LCH BG-162 (SRSC). Penstemon barbatus (Cav) Roth, Beardlip penstemon, rare mm Fenstermacher et al., Vascular flora of the Dead Horse Mountains perennial herb, Dar, DCA, 3160-4000 ft; JF 222, MT 216, BHW e (SRSC), LCH & BHW BG-175 ÉS LL). ly Gray, C | rare pes herb ABER 4120 ft; JF 724 (SRSC). Penstemon lanceolatus Benth., Lanceleaf beardtounge, mmon to "rare" perennial! herb, 3500-5840 ft; JF 1306, BHW & LCH BG-175 (SRSC); JMP 2725, BHW 10752 (TEX-LL) # Seymeria faicata B.L. Turner var. falcata, Falcate blacksenna, locally common annual herb, par/syc, 4000-5500 ft; JF 1943 (BIBE, BRIT, SRSC, & TEX-LL); JF 571, 1187, 1548, 1952, 1959; MT 137 (SRSC). New to the U.S. (Fenstermacher 2006). GE scabra A. Gray, Limpia blacksenna, uncomm uai herb, svc, 4880-5840 ft; BA2 5330 (SAT); JF 2 M MT 255 (SRSC). $ Stemodia coahuilensis B.L. Turner, Coahuila twintip, rare annual herb, pan, 2200 ft. Photo documented by Roy Morey in February 2007: at the first mini-side drainage encountered when walking up the wash towards Ernst Tinaja from Tue parking area; « one e clump" of plant(s) was found. 4 , à Rare in the U.S.; mostly a Mexican species from Cuatro Cienegas area (Henrickson 2005). Simaroubac an stewartii C.H. Mull, Crucifixion thorn, rare shrub, E ~3700 ft; JF 1145, JF & HL 1900, BHW 11760 (SRSC). Solanaceae 8 Calibrachoa parviflora Juss.) D'Arcy, Wild petunia, annual herb, rio, 1800 ft; AL 309 (BIBE). Chamaesaracha a Averett, Ge five E false night- de, uncommon perennial herb, psc, DCA, svG/CHP, 1800—4800 ft; Jr 2204, mm 16846, pus BHW & MCJ 16685 (SRSC). Chamaesaracha sordida (Dunal) A. Gray, Hairy false night- shade, occasional Po herb, psc, DCA, 1900-3640 ft; BHW A & MWB 519, 520, CMR 17157, TXDOT 1324 GE p 290, 449 629, 654, 965, 2047, BLT & BD 23-175, BHW 20898 (SRSC); BLT & BD 23-175 (TEX-LL). Chamaesaracha villosa Rydb, Trans-Pecos five eyes, wooly false nightshade, uncommon perennial herb, psc, DAR, DCA, 1900-2560 ft; JF 1442, BE NES BHW & LCH BG-74, BHW & MCI GC BHW & BLT 8327 (SRSC); JEA Edid ii Ge X-LL). | red datura, thorn apple, annual or perennial herb, R RIO, SE ft; JF 2262 (SRSC). Lycium berlandieri Dunal, Berlandier wolfberry, occasional shrub, DSC, DAR, DCA, 1800-3480 ft; EGM 10, BHW 13025 (BIBE); JF 1090, 1387, 1390, 1391, 1451, 2045, 2110, MT 205, BHW 13025, 13067, A (SRSC). +o Lycium puberulum A. Gray var. berberioides (Correll) F. Chiang, Downy wolfberry, rare shrub, osc, 2350 ft; JF 1392 (SRSC). ! Nicotiana glauca Graham, Tree tobacco, uncommon shrub, DCA, RIo, 185C ft; CN 11739 (BIBE); CMR s.n. (SAT); JF 1063, 2260 (SRSC) 725 Nicotiana trigonophylla Dunal, Desert tobacco, uncommon ual herb, psc, DAR, DCA, 2400-3500 ft; CMR 16411 (SAT); 8 ps 440, 1446 (SRSC). Physalis cinerascens (Dunal) Hitchc. var. cinerascens, Small- flower groundcherry, uncommon perennial herb, syc/ CHP, Ste p ft; BA2 5231, 5311 (SAT); ie 1981, MT 128, OMe dd ) pi Gray, | icl ncommon to occasional perennial herb, t DSC, DAR, DCA, d: 4650 ft; OES 1334 (BIBE); JF 159, 216, 441, 1086, 1906, 2069, 2074, MT 36, 37, o. BHW & MCJ 16825 (SRSC); TRV & GLB 85-36, BHW 10779 (TE Quincula lobata (Torr.) - £ [Physalis lobata Torr.], Chinese lantern, purple groundcherry, "infrequent/sparse" peren- nial herb, psc, nio, 1800-3800 ft; BHW 1227, 10747, JF 2269 (SRSC); BHW 10747 (TEX-LL). Solanum e Cav, Silverleaf nightshade, uncom- on perennial herb, pca, nio, 1800-1850 ft; EGM 160, BHW 645 (Eo. 7 1064 (SRSC). § pon ptychanthum Dunal [Solanum americanum Mill], st Indian nightshade, annual herb; EGM 137 (BIBE). SE triquetrum Cav, Texas SÉIER uncommon rennial woody vine, DSC, DAR, DCA, svo, 2780-4800 ft; JF 162, 249a, 1035, 2176, 2178, 2185, PM 976, BHW & MU 16838 (SRSC) Sterculiaceae Ayenia filiformis S. Watson, Trans-Pecos or narrowleaf ayenia, shrub, osc, DCA, RIO, 1800-2250 ft; CMR 17160 (BIBE); EE BHW 18398, 20950, BHW & LCH BG- 187 (SR SC). y hylla A Gr ray, Dense ayenia, casional subshrub, DSC, DAR, DCA, 1800-4500 JF 1124, 1957 2038, 2172, BGH 632, AMP & SAP 5307, BHW 20620 (SRSC); DSC & HBC 30627, BHW 10791 (TEX-LL). 8 Ayenia pilosa Cristóbal, Hairy or dwarf ayenia, subshrub, 3000 ft; BHW & LCH s.n. (BGWMA); BHW & LCH BG-139 (SRSC). + Tamaricacea 118 Tamarix cm” (L) H. Karst, Athel tamarisk, saltcedar, locally occasional tree, psc, no, 1800-1900 ft; BHW 13052 (BIBE); dici dii IM Ee GL SN 3349 dee — J Al V +l Bis I + J J removed by NPS exotic plant eradication teams, as are the Tamarix chinensis, but they are firmly established in upstream areas and so will likely continue to reestablish along the river corridor into the future. I! Tamarix chinensis Lour, Five-stamen tamarisk, saltcedar, locally abundant tree, oca, nio, 1800-ca, 4000 ft; JF 1346, 2233, 2253, MT 179 (SRSC). Ulmaceae Celtis pallida Torr, Spiny hackberry, common shrub or tree, DAR, 00 ft; CMR 5078 (BGWMA); JF 2112, TR 80, MT 204 BHW 14048 (SRSC) SA Celtis reticulata Torr, Netleaf hackberry, palo blanco, tree. Listed by Amos and Giles (1992) as occurring in Brushy Canyon, Known from BBNP and commonly through the rest of Brewster County. 726 Urticaceae +0 Parietaria pensylvanica Muhl. ex Willd., Pennsylvania pellitory, rare annual herb, psc, ca, 3150-3560 ft; JF 538, 1701 (SRS Verbenaceae o Aloysia dere (Gillies & Hook.) Tronc, Beebrush, white- sh, occasional shrub, psc, DAR, DCA, 1880-3480 ft; JF 67 1, 1362, TR ge SE +o Aloysia wrightii A. Heller ex Abrams, Wright beebrush, oreganillo, b, DSC, DCA, SYG/CHP, 3560-5080 ft; JF 476, 746 (SRSC). Bouchea spathulata Torr, Spoonleaf, occasional shrub, DSC, DAR, DCA, SYG, 1800-5000 ft; BA2, DG, TE, & B 4927 (BIBE); PVW s.n. (KANU); BA2 4927, SJ & GJ 6503 (SAT); JF 170, 857, 1511, MT 1 (SRSC); BHW 10749, 10766 (TEX-LL). 8 Glandularia pe (Nutt) i vat. kee Ce B.L. Tur a vervain, uncommon perennial herb, DSC, - DAR RIO, 1800- 3520ft; OES E (BIBE); HTF 1226, GLW 4466 USE mul | a ids on this genus (Turner and Poweil 2005). o ali (A. Heller) Umber [Glandularia unda Um eaked verbena, uncommon annual he a DSC, DCA, RIO, e A 40 (BGWMA); JF 618 (BIBE); CMR 16409 (SAT); JF 618, us o (SRSC); BHW & RRI 21430 (TEX-LL). The individuals found in BBNP may be more appropriately placed under Glandularia vere- f La | a +l | ¡EA | ñ | A (Pow iell in n prep, Mannicksan 2003) 8 Glandularia wrightii (A. Gray) Umber, Wright verbena, pinky, perenni sis: Cu e (S ATA i | , desert lantana, uncommon to occasional shrub, DSC, DAR, DCA, 1800-2780 ft; TR 103 (BGWMA); BA2 4636, 4936 (SAT); JF 163, 947, 1364, TR 103 (SRSC); BHW 831a (TEX-LL) Lippia graveolens Kunth, Mexican oregano, scented lippia, uncommon shrub, osc, DAR, pca, 1800-2350 ft; DG & TE 4613, EGM 148, BHW 13065 (BIBE); DG 4613 (SAT); JF 1080, 1341, 1691, 2032, 2142, AMP s.n., 5306, 5823, BHW 13065, 21202 (SRSC); BHW 831 (TEX-LL). Phyla nodiflora (L.) Greene, bs pu ee “frequent erennial herb, RIO, OCA, 0-230 1, BHW 833 (BIBE); JF 2243, OCW 510, DE GE 8 Tetraclea coulteri A. Stay, EE wrinklefruit, stinkweed, “infrequent” | i , 2250 ft; BRM 3209, CMR 5073 (BGWMA); BA2 5717 (SAT); BHW & LCH BG-184 (SRSC). Verbena neomexicana (A. Gray) Small, Hillside vervain, New " A || E al Dos = im LI det de nho e TR f Texas 2(1) Mexico verbena, occasional to “abundant” perennial herb, nio, DCA, 1800-? ft; BRM 3047, CMR 11635 (BGWMA); JF 2265 (SRSC). Violac Pini verticillatus (Ortega) Baill, Babyslippers, uncom- mon to rare perennial herb, psc, sva, 3000-4900 ft; JF 220, 1322, 2170, BHW 10895 (SRSC). Viscaceae # Phoradendron hawksworthii (Wiens) Wiens , Hawksworth mistletoe, rare subshrub, psc/sva, 4160 ft; JF 1896 (SRSC). Phoradendron tomentosum (DC) Engelm. ex A. Gray, Christmas mistletoe, uncommon subshrub, psc, oca, 2780 ft; CMR Sg (BGWMA); JF 182 (SRSC); AMP & SAP 4301 (SRSC € X-LL). "mon villosum Nutt, Oak or Cory mistletoe, uncom- mon subshrub, svc/cue, 2300-5000 ft; PVW s.n. (KANU); JF 769, 1209, BHW 10845, 10846, OCW 505, Ger & LCH BG-145 (SRSC). Vitaceae «a A E ^ P» |, Sorrelvine, ivy lr av Tarr 8 reebine, rare perennial woody vine, DSC, DAR, 2050-3740 A JF 232, 1682 (SRSC). 8 Vitis arizonica Engelm, Canyon grape,"abundant" perennial ody vine, rio, 1800 ft; EGM 156, BHW 13038 (BIBE); BHW 13038, GLW 4489 (SRSC). Zygophyllac Guaiacum men o Guayacan, Texas lignum- , comm DAR, DCA, RIO, 1800-3280 ft; an 5033 BEN. y 2 690, 2111, TR 96, BHW 18413 SR SC). 5 baena grandiflora Torr. ex A. Gray, Arizona poppy, inf t" annual herb, psc, oca, 1800-2320 ft; BLT & ee 97-336, BHW & MCJ 16867 (SRSC); EGM 53, BLT & M12 97-336 (TEX-LL o Kallstroemia hirsutissima Vail, Hairy caltrop, uncommon ual herb, osc, 1900-3000 ft; JF 199, 219, BHW 13066 (SRSC). 8 Kallstroemia parviflora Norton, Warty caltrop, annual herb; BH BG-197 (SRSC) Larrea tridentata (Sessé € Moc. ex DC) Coville, Creosote, adora, abundant shrub, psc, par, 1800-3560 ft; ACK 829 (BIBE); JJ 40, JT 30 (SAT); JF 606 (SRSC); DS 1091 (TEX-LL). ! Tribulus SE L, Goathead, puncturevine, "infrequent to fi nual herb, rio, 1800 ft; BAZ, DG, TE, & PC 4607, EGM SE Ge BA2 4607 (SAT); JF 2227, BHW 23036, BHW & MCJ 16850 (SRSC). ACKNOWLEDGMENTS D . à 1 1.1. 0 PEE De S Billie Turner (TEX-LL) is recognized and anes for EES y EEN Appreciation for (nancial support goes to SRSU, to this project, in addition to assisting with L BBNP, Friends of Big Bend National Park, Big Bend Natural History Association, and the Texas Academy of Science. Many thanks go to BBNP staff, especially Betty Alex for GIS support, Tom Alex for cultural his- tory information, John Forsythe for weather data, Don Corrick for geologic insight, Don Sharlow, the trails program, and the Rio District rangers for field support, and Allison Leavitt for her field observations that Fenstermacher et al., Vascular flora of the Dead Horse Mountains 727 made significant additions to this flora. Lynn Loomis (USDA, Marfa, TX) provided an excellent primer on local soil dynamics. Thanks go to the SRSU biology department for providing overall support and continu- ing to support an active herbarium. Herbarium techs Fay Burton and Molly Klein mounted the majority of specimens. The SRSU GIS lab also provided important assistance. Enthusiastic and valuable field support was supplied by Mark Yuhas, Mark Darrach, and Hilary Loring. Tom Vanzant, Mike Pittman, and Todd Montandon (TPWD) were helpful in providing access to and data for the BGWMA. Bill Dodson and the Shackelford family are thanked for facilitating and allowing access to the eastern part of the study area. Richard Worthington (UTEP) and James Henrickson (TEX-LL) provided important floristic data, while Jackie Poole (TPWD) was a valuable source of information about rare plants and regional floristic publica- tions. Final appreciation, after thanking the three reviewers for helping to improve the manuscript, goes to all of the herbaria and curators that provided information and assistance to this project especially Caleb Morse (KANU), Bonnie Amos (SAT), Monique Reed (TAES), and Tom Wendt (TEX-LL). This research was conducted in part under National Park Service Task Agreement Number J717003001A and Cooperative Agreement Number H7170030001 with Sul Ross State University. REFERENCES ALEX, BL, A. LeAvrrT, J. Timer, and J. SIROTNAK. 2006. Final report on the sensitive plant project, Big Bend National Park, Texas, Field Seasons: June 2003-April 2006. Science & Resource Management, Big Bend National Park, Texas. Unpublished report to the National Biological Infrastructure Inventory, U.S. Geological Survey, Wash- ington, D.C. Amos, B. and D. Gites. 1992. Survey of the vascular plants of Brushy Canyon. Unpublished report to the Nature Conservancy of Texas, Fort Davis. AntHony, M. 1956. 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YARBOROUGH, S.C. and A.M. PoweLL. 2002. Fern and fern allies of the Trans-Pecos and adjacent areas. Texas Tech WorLD WILDLIFE FUND | _J E University Press, Lubbock. DISTRIBUTION AND TAXONOMY OF SYMPHYOTRICHUM SERICEUM AND S. PRATENSE (ASTERACEAE: ASTEREAE) Ronald L. Jones C. Theo Witsel! Dept. H Mcd Sciences Arkansas Natural Heritage Commission 21 er Avenue 1500 Tower Building B dep) University 323 Center Street Richmond, Kentucky 40475, U.S.A. Little Rock, Arkansas 72201, U.S.A. Guy L. Nesom 2925 Hartwood Drive Fort Worth, Texas 76109, U.S.A ABSTRACT " 1 mm I 3 e l2 Bao gk E hoe era 1. a : -1 f +1 tralTT CG A Hrs ] in Texas E , Pi 2 +1 J J Taga 4 4 ae r I i L a e listributi fS pratense is in Texas, southern Arkansas , and Louisiana, with disjuncts to Arkansas, Oklahoma, and Texas. TI en to aii nue ee dentur Missisippi, Tennessee, and a Wes an Ge record from nou ene p = of which are still | ericeum here treated as S. pratense. Sy pas tric} i larg la f diploid populations, while both tetraploid TEE C DES Id : A | AS : "Em and diploid sovulatigns occur in 5. pratense. Count; p pped. Symph; listed in 4 states and S. pratense in 7 states. RESUMEN Symphyotrichum sericeum y S. pratense son especies distintas, que no se intergradan, primariamente del centro de U.S.A., simpátricas sólo en Texas. La distribución más densa de S. sericeum está en los estados del centro-norte, con poa disyuntas y sistemas de pobl a el sur hasta Arkansas, Ge y Texas. La dist de S pratense está en Texas sur de Han y Louisiana, con hasta Alabama, Florida, Georgia, Kentucky, Mississippi, cen y Virginia, con una cita histórica de North Carolina. Especimenes de estas disyunciones en el sudeste de U.S., algunos de los cuales aún se listan como S. sericeum en listados estatales y citas de especies raras, se tratan aquí como S. pratense. Symphyotrichum sericeum consiste mayormente en poblaciones diploides, mientras que en S. pratense se encuentran tanto poblaciones tetraploides y diploides. C t A zs ain zx oe dud | 1 i C I dere c nux] ler tata) q + A T3 1/8 eh Eu ES L + estados y S. pratense en 7. Symphyotrichum sericeum (Vent.) Nesom and S. pratense (Raf.) Nesom are two closely related taxa of central and eastern United States. In most treat ts they have been maintained as distinct species (e.g., Correll & Johnston 1970, Cronquist 1980, Jones 1990, Nesom 1994, Brouillet et al. 2006). When the taxa are considered to be conspecific (see Wunderlin & Hansen 2004), S. pratense is treated as S. sericeum var. microphyllum (DC) Wunderlin & B.F. Hansen. Symphyotrichum pratense was known for many years as Aster phyllolepis Torr. & A. Gray. Symphyotrichum sericeum is generally EE in the EE states, from Texas to North Dakota and southern Canada, to western Michigan. Sym; trated in eastern Texas and western Louisiana. kopu anonsi of piante in his corales that are disjunct further eastward—to Florida, Virginia, and Kentucky te in phyllary morphology and have been variously treated as S. sericeum, S. pratense, or S. sericeum var microphyllus, and a number of states treat these often widely scattered populations as state-rarities. Chromosome counts also need clarification, as both ploidy levels @n = 10 and 2n = 20) have been published for both taxa (Brouillet et al. 2006). The objective of this paper is to update available information on this complex, clarify geographical EE point out distinguishing features, identify typical habitats and associates, review availabl tion, and summarize the state-raritv status of the taxa. J. Bot. Res. Inst. Texas 2(1): 731 — 739. 2008 732 J d tanical h Inst Texas 2(1) The present study provides a detailed view of the geographic distribution of Symphyotrichum sericeum and S. pratense, based primarily on a study of herbarium specimens from BRIT-SMU, EKY, GA, LSU, MO, NCU, NLU, TENN, TEX-LL, UNC, VDB, and VPI. Maps are augmented by records from various published sources. The available literature on habitats, associates, and chromosome numbers was also reviewed. A new chromosome report from a Kentucky population is based on mitotic counts from root tip squashes from germinating achenes. The key is based on a detailed examination of morphological characteristics of herbarium specimens from across the ranges of the two taxa. SYMPHYOTRICHUM SERICEUM Distribution and Habitat.—The primary distribution of Symphyotrichum sericeum is from Arkansas and Oklahoma to southern Canada to western Michigan and Indiana, with disjunct populations in central Texas (Fig. 1). Over the larger part of its range, S. sericeum occurs in a variety of habitats, e.g., dunes, sandy woods, glades, sandstone glades, over granite outcrop— but typically associated with seasonally dry, open to semi-open habitat. Symphyotrichum sericeum in Texas mostly occurs on the Edwards Plateau, where it grows on limestone hills, commonly on open, rocky slopes and roadbanks. An early collection supposedly made far to the east (Harris Co.: Lindheimer 78, BRIT) is similar in every respect to Edwards Plateau plants and may be mislabeled. Swink and Wilhelm (1994) list a number of associates of this species in Illinois, including Amorpha canescens, Schizachyrium scoparium, and Bouteloua curtipendula in dry hill prairies, and Anemone cylindrica, Liatris aspera, Liatris cylindracea, and Spiranthes cernua in sand barrens sites near Lake Michigan. In Wisconsin it occurs at dry prairie sites on hillsides, outcrops, and bluffs, sometimes under oaks, jack pine, or red cedar, but usually in the open in habitats dominated by Schizachyrium scoparium, with Asclepias viridiflora, Dalea purpurea, and Brickellia eupatorioides (Cochrane & Iltis SEL In Manitoba the species occurs in remnant tall-grass prairie sites and in openings in Q pa/Populus tremuloides woodlands (Foster & Hamel 2006). The species is primarily associated with the Central Lowlands Physiographic Province in the m S. punt it also occurs in the Ozark Plateau Province of Arl d Missouri. A recent collection sericeum from open shale barrens on a southwest-facing slope in central Arkansas, is pandy the first record from the Ouachita Mountains Province. This site is disjunct from the nearest populations about 155 km to the northeast on the Ozark Plateau. Voucher: Garland Co.: Ouachita Mountains, Ouachita National Forest, steep S-facing shale barrens (Mazarn Formation) on upper slope, east of and above Walnut Creek near point at which it flows into Lake Ouachita, Crystal Springs 7.5” quadrangle, steep glades, elev. 750 ft, hundreds of plants in high quality shale barrens, 29 Sep 2006, Witsell 06-471 (ANHC,BRIT, UARK). A second population was subsequently found in similar habitat 1.9 km northwest of the above site Chromosome Numbers.—There are about a dozen published diploid counts for Symphyotrichum seri- ceum, these mostly from the northern portions of its range (Jones 1980; Semple & Brouillet 1980; Semple & Chmielewski 1987; Semple et al. 1989; poms etal. 20921, with the most Se count from Izard Co., Ar- kansas (Sherif et al. 1983). There is apy only le report of count (Love 1982) for S. seri- ceum, this count from Canada; t | diploid counts from this same cay (J. Semple, pers. comm.). Taxonomy.—Brouillet et al. (2006) treated Symphyotrichum lucayanum (Britt.) Nesom as a synonym of S. sericeum, but S. lucayanum is here id to be more closely related to S. concolor (L.) Nesom, based on their similarities in capitulescence and phyllary morphology and their closer geographical proximity. The latter is widespread in the southeastern U.S.A. Symphyotrichum lucayanum was treated by Nesom (1994) as a distinct species, based on its very loosely racemiform capitulescence with interspersed leaves only slightly reduced or not at all from the proximal cauline, different from the more densely congested, bracteate capit- ulescence of S. concolor. Turner (1982) treated the taxon as a synonym of Aster concolor L., and the type of Aster lucayanum Britt. See E EE 2448, NY!) was annotated be W.T. Gillis in 1974 as A. concolor. State-Rarity. | sericeum is currently ranked as S2 (Threatened or Rare) in Arkansas, Indiana, Michigan, See North Dakota (NatureServe 2008). It is listed as Nationally Threatened in Canada dS pratense 733 Jones et al Dictributi LS E SE — e pm 2 we Wi S a. E "el iu wae TE LH SINE o A jac dub Br HE EE as TIT e Te ECC HE Bp ORE dë Rep E lo. Lr EINER A I-A LEES a aos $ E EAE N | e T DA SL o? EE Kate, bet ST EE e ee E EHE EE OE ae HT] SC) RA zd -HH NE Jp. e a EEN E Ito E É doa FEAS Bh e [t Prt uM Ee E RE SE E KINN, COH ! X e E Ed 2 i URDU mU HUI | SS UE E iss Ke Se fecht E Fic. 1. Distributi 734 (Foster & Hamel 2006). We could find no substantiation of the occurrence of S. sericeum in Georgia, Ohio, or Tennessee, where indicated to occur by Brouillet et al. (2006) and NatureServe (2008). SYMPHYOTRICHUM PRATENSE Distribution and Habitat.—The densest distribution of 9 phyotrichum pratense isi ce E and southern iind ace disjunct Ge of populati l over and in a variety of physi E. 2). 5 wish pratense in Texas charsetetistisálly occurs in sandy soil m) € lesa Bands ge silty clay, silty clay-loam), being largely restricted to the follow- ing natural regions: Piney Woods, Oak Woods and Prairie, Gulf Coast Prairie and Marshes, and Blackland Prairie. Within these regions, it is less frequent in blackland prairie: e.g., Texas. Dallas Co.: Lundell 12032 (LL, SMU). Shinners (1950), who recognized the legitimacy of the Rafinesque name ahead of the Torrey & Gray name, described this species as "a frequent and showy plant of open oak woods and transition-belt prairies thoughout east Texas ..." In Texas, S. pratense also occurs in longleaf and loblolly pine areas and in coastal prae EE and it as Reco a GER fields, roadsides, clearings, and fencerows. In 1 d sites , blackland prairie, chalk hills, and cal la as well as along dry EE A blaciland prairie locality for Symphyotrichum pratense is on the “Kieffer Prairie:” Winn Par.: Brown 8220 (SMU) and five other collections from the same locality (LSU Herbarium Online Database 2008). At the nid Prairie E Ene grows with associates such as Schizachyrium sco- parium, Andropogon gerardii, Sorgl , Echi pallida, Oligoneuron rigidum, and Vernonia missurica (USDA MA Sopas 2008). In the Gulf Coastal Plain of me typical habitat for Symphyotrichum pratense is open saline barrens—treeless openings in Pinus taed imilis flatwoods with clay soils containing high levels of sodium and/or magnesium. Eric Sundell, viis first discovered the species in Arkansas at the Warren Prairie (Sundell 1983), noted that the pane occurred with prickly pear and ER palmetto and that a number of other rare species were jated with these sites, including G and Schoenolirion wrightii. The USDA Plants website (USDA, NRCS 2008) lists Izard and Neon counties in Arkansas for S. pratense, but these records from northern Arkansas are based on collections of S. sericeum. Eastward (to Alabama, Florida, Georgia, Kentucky, Mississippi, Tennessee, and Virginia), Symphyotrichum pratense occurs mostly in calcareous habitats, e.g., chalk barrens, cedar glades, and limestone outcrops. Kral (1981) reported on the species at Coastal Plain sites in Alabama (Sumter Co.) and Florida (Gadsden Co.) and noted that the plants in Alabama were found in blackland prairie patches and that those in Florida were found at rocky calcareous outcrops in association with Schoenus nigricans, a plant more normally found near the coast. The species subsequently was discovered in Decatur Co., Georgia, just across the state line from Gadsden Co., in calcareous glades on a bluff: Decatur Co.: Godfrey & Gholson 80099 (GA). In Mississippi the species occurs in association SR Ge chalky outcrops (L. McCook, pers. comm.). The central Alabama and Mississippi popul ted in the Black Belt prairie lands (see Barone 2005). The most northerly populations in Alabama occur in o prairie Caen within the Highland Rim province, along with many other rare species, including Dalea gattingeri, Leavenworthia alabamica, Cypripedium can- didum, Eriogonum longifolium var. harperi, Schoenolirion croceum, and Spiranthes magnicamporum (Webb et al. 1997). At its most northerly range limits in Kentucky, Symphyotrichum pratense is found in glades or barrens habitats of the Interior Low Plateaus (chiefly Highland Rim sites) with Juniperus virginiana and Quercus marilandica as the woody dominants and with Andropogon gerardii, Schizachyrium scoparium, Sorghastrum nutans, Sporobolus compositus, and S. vaginiflorus as the conspicuous grasses. These habitats often contain a variety of other rare or infrequent plants, such as Cypripedium candidum, Echinacea pallida, Liatris aspera, Liatris cylindracea, Lithospermum canescens, Silphium pinnatifidum, Dalea candida, Dalea purpurea, and Phys- ostegia virginiana (Cranfill 1991; Kentucky State Nature Preserves Commission 2008). The barrens sites in Tennessee (Highland Rim and Appalachian Plateau sites) and in southwestern Virginia (Valley and Ridge) fS hvotricl j d$ pratense 735 Jones et al.. Dictributi Tele TEE RA ER MEC AR ONES p E use | SI eese EAR LEA ES N Séi + E pure FO LA p Se a | [V Q ei ¡9 E A Y Ve S TE Pao Bis At c ‘An, E DAS al Së a aS e SC MAS de - y A i n Yon Ca NH RUE M a Ge [P e XC p SE mis MODE Sas AS Pe > CREE ORAL oy ERGO Dp ces PAES e eeh E ae e A Tr $ LAS es AT e LO EI j AN AN E MON = BUS LE] de E e. / NO muB: LIIS US (| | reos ú e o Symphyotrichum pratense E De 2 Diet AE f 5j pl asf sl are very similar to the Kentucky sites and also contain a number of very rare species (DeSelm 1990; Ludwig 1999; Tennessee Natural Heritage Program 2008). In northern Georgia the plants occur at both Appalachian Plateau and at Valley and Ridge sites (T. Patrick, pers. comm.). In the Coosa river watershed of Georgia, S. pratense occurs in limestone glades; other species associated with this habitat include Asclepias hirtella, Baptisia australis, Buchnera americana, Dalea gattingeri, Echinacea pallida, Hypericum dolabriforme, Hypericum sphaerocarpum, Liatris squarrosa, and Spiranthes magnicamporum (Georgia Department of Natural Resources Recent investigations of dolomite and limestone barrens in southwestern Virginia have discovered several populations of Symphyotrichum pratense -- initially reported by Ludwig (1999), subsequently treated by Brouillet et al. (2006), Weakley (2007), and Virginia Botanical Associates (2008). We have studied the following vouchers: Russell Co.: 1 km E of Carterton, one-acre barren at top of SW-facing slope on W end of a short ridge, dense grass cover with dolomitic limestone cobbles, 20 May 1996 [not yet in flower], Wie- boldt 9415 (VPI); river knob 1 km E of Carterton, top of SW-facing hardwood slope, substrate cobbly with dolomitic (?) limestone, 2100 ft, dominant species with Liatris aspera and Andropogon gerardii, scattered over most of one-acre barren, 17 Sep 1996, Wieboldt 9605 (VDB, VPI); E side of Mill Creek where it cuts through ridge between Clinch River and Reeds Valley, ca. 1.9 km N of Jessees Mill, SW-facing limestone hillside in open grassland dominated by Schizachyrium scoparium, 2040 ft, ca. 20 plants, 11 Sep 1996, Ludwig 3082 (VPI). Wise Co.: Fleming 14122 (GMUF). Ludwig (1999) noted that this species has | found at six Russell County sites. | | | E ul np Li MA L | | P» ru 736 t t f Texas 2(1) Chromosome Numbers.—The first report of the tetraploid level of 2n=20 for Symphyotrichum pratense was from Gonzales Co., Texas, by Semple and Brouillet (1980), who noted that a diploid count had been made earlier for the species, but that report was unvouchered and part of a 1945 embryological study by German anatomists. Semple and Chmielewski (1987) published diploid counts for plants from Grimes and Walker counties in Texas and a tetraploid count for plants in Milam Co., Texas. Two years later Semple et. al. (1989), reported a diploid count of 2n=10 for plants from Gadsden Co., Florida. In addition, we report a previously unpublished 2n=20 count for S. pratense: Kentucky. Hardin Co.: cedar glade off KY 2762, 6 Nov 1987, R.L. Jones 5457 (EKY). Taxonomy.—in the main part of the range of Symphyotrichum pratense, the plants typically have phyl- lary surfaces mostly AMAA ang ge Ge with the spreading-ciliate margins. Phyllaries of the Virginia plants are faces, with or without strongly ciliate margins, but resemble typical S. marae of ee and Lona: in the proximal indurate portion of the phyllary being relatively short, the distal leafy portion being relatively broad and long, and the overall phyllary length about 1.5 cm. Their geographic position also obviously suggests that they are part of S. pratense. Some plants from Tennessee are similar in their phyllary features (e.g., Roane Co.: Morton 1307, SMU) The description of Aster montanus Nutt. seems to refer to Symphyotrichum pratense; Nuttall noted that it was “nearly allied to the preceding [Aster sericeus], but distinct." Torrey and Gray (in Fl. N. Amer.) treated it as A. sericeus var. p. They also noted that the species was known from "Tennessee and North Carolina near the mountains." On Nuttall's 1816 trip, he crossed from Tennessee to Asheville, N.C., and went on to Rutherfordton and Morganton (Graustein 1967). Gray (in Synop. Fl. N. Amer. ) formalized the name at varietal rank, as Aster sericeus var. montanus (Nutt.) Gray, noting that the involucre is “sometimes glabrate and villose-ciliate; approaching the next species [A. phyllolepis].” An image of the holotype of A. montanus was obtained from PH, and an examination indicates that the involucral widths are about 2 cm wide, much larger than typical S. sericeum, and we therefore place this name in the synonomy of S. pratense. Today neither S. sericeum nor S. pratense is known to occur in North Carolina. Gray (in Fl. N. Amer.) listed North Carolina as included in the range of A. sericeus var. B, and later (in Synop. Fl. N. Amer), he specified Buncombe County as the site of the collection. The specimen upon which Gray apparently based his comment was located at NY, and an image was obtained. The sheet bears three plants from two dif- ferent areas—two are from Minnesota and identified as Aster sericeus, while the other was collected from “Buncombe, North Carolina, Gibbes” and identified as “A. sericeus B. (montanus Nutt.).” This latter specimen exhibits the large heads of S. pratense, and is here assigned to that taxon; it represents the only known record of the species from North Carolina. S. pratense has not been subsequently rediscovered in the state and can be assumed to be extirpated there. Aster montanus Nutt., Gen. N. Ámer. Pl. 2:156. 1818 (non ge montanus All. 1785). Aster sericeus var. montanus (Nutt.) A. Mia goen Fl. N. dina 1, part 2:179. 1884. Lasall dpa nov.], Leafl. Bot. Observ. Crit. E E USA. Tennessee. N data, [T. Nuttall s.n., S pt t ly Oct r 1816, fide Graustein 1967], tall’s anta as “Aster montanus" (HOLOTYPE: PH, digital image!) Nuttall: prot ] g “HAB. On the mountains of Tennessee and North Carolina, rare. (Near Asheville and Morganton).” When these disj lati f Symphyotricl i were first discovered in eastern U.S., especially at the more northerly bes in Albane. eessen and Kentucky, they were often identified as S. sericeum. They are still listed as such in several state natural heritage databases. We now confirm all examined speci- mens from Alabama, Florida, Georgia, Kentucky, Mississippi, North Carolina, Tennessee, and Virginia as S. pratense. State-Rarity.—S icl lly rare outside its main range in Texas and Louisiana — itis currently nnn Te (S2) in as, Special Concern (S3) in Kentucky, and $1 (critically imperiled ) in Alabama, Georgia, Mississippi, Tennessee, and Virginia (NatureServe 2008). Although there is only one known county of occurrence in Florida (the only state Where it still is treated as a variety of S. seri- ceum, S. pratense is currently not listed among that state’ E (Florida Natural Areas Inventory 2008). Jones et al., Distributi fS hyotrict i i S. pratense 737 c E T PE E "9m ishina he Tasa =O through- out mic of ilie: ranges of the two taxa lena: that they can be reliably diemgar ied as follows: 1. Phyllaries densely sericeous, vestiture of margins similar to surfaces, not spreading ciliate; terminal, green, pubescent portion of largest phyllaries 1/4—1/3(-1/2) the total phyllary length, usually distinctly shorter than the proximal, indurate and glabrate portion; middle phyllaries mostly « 7 mm long an wide, averaging 6(-8) mm long and 2(-3) mm wide; involucra! height mostly « 10 mm (7-11 mm) high mo um sericeum 1. Phyllaries with abaxial faces ually glabrous to glabrate, less commonly sparsely st 0.5—1.0 mm lona EUER longer than ek on the surfaces; terminal, green, pubescent portion of largest phyllaries 2/3-3/4 the total phyllary GE longer than the proximal, indurate and durae portan E EE dnd > mm long and > 2.5 mm wide, averaging 9-14) mm long, and 3(-5) m (8-15 mm) high Symphyotrichum pratense CONCLUSION Symphyotrichum pratense and S. sericeum are distinct species of central and eastern U.S. They are largely al- lopatric in distribution, and occur sympatrically only in Texas, where S. pratense is common and S. sericeum rare. Symphyotrichum pratense is associated with a range of physiographic provinces—it is chiefly Coastal Plain in distribution, but its range extends to the Interior Low Plateaus, the Appalachian Plateau, and the Valley and Ridge. There is an historical record from the Blue Ridge of North Carolina. Symphyotrichum seri- ceum is largely associated with the Central Lowlands Province and the Ozark Plateau, and only sporadically into adjacent provinces to the south, north, and west. Disjunct populations and population systems are a prominent feature of the biology of both taxa. There is some EE in phyllary pubescence in the easternmost disjunct populations, but tl f lting from local genetic drift in populations far removed from the main range of the species, and there is no ev ideme of any significant gene interchange between the taxa. The available evidence indicates that S. sericeum consists of diploid populations (with the exception of one lous report) and that S. pratense contains populations at both ploidy levels. Symphyotrichum sericeum is generally common throughout much of its a considered rare only in the states at the eastern and southern periphery of its range (Michigan, Indiana, and Arkansas), and is Nationally Threatened in Canada. Outside of Texas and Louisiana, S. pratense is a rare plant and is con- sidered critically imperiled in several states (Alabama, ee amd Tennessee, and Virginia). The identity of these disjunct eastern populations has been the soi ome confusion in the past — they have been variously treated by the duret states as 5. pratense or S. sericeum, and this latter name continues to be used by some eastern states for their rare species lists. Our study concludes that all specimens examined from populations in Alabama, Florida, Georgia, Kentucky, Mississippi, North Carolina, Tennessee, and Virginia, are S. pratense. ACKNOWLEDGMENTS We are grateful to staff at MO, TENN, and TEX-LL for help during visits there and to the curators and staff of all herbaria that provided loans. In particular, we thank Tom Wieboldt for a loan of specimens from VPI, Johnny Townsend (Natural Heritage Program, Virginia Dept. of Conservation and Recreation) for information on Virginia collections, Andrea Weeks at GMUF for an image of the Wise County (Virginia) collection of Symphyotrichum pratense, Alina Freire-Fierro of PH for an image of the holotype of Aster montanus, Jacquelyn Kallunki, Thomas Zanoni, and Lucy Klebieko of NY for providing a virtual herbarium loan of the speci- men from North Carolina, and John Pruski (MO) for help with obtaining literature. Brent Baker (UARK) provided distribution data for specimens compiled by the Arkansas Vascular Flora Project, Deborah White provided data from the Kentucky State Nature Preserves Commission database, Lucile McCook provided information on Mississippi populations, Emily Wood and Walter Kittredge for specimen searches at GH, and Wendy Zomlefer and David Giannasi for information on Georgia records. In addition, we thank John Semple for his assistance in the gathering of information on chromosome counts. 738 | | tanical h Insti Texas 2( REFERENCES BARONE, J.A. 2005. Historical presence and distribution of prairies in the Black Belt of Mississippi and Alabama. Castanea 70:1/0-183. BrouiLLET, L., J.C. SEMPLE, G.A. ALLEN, K.L. Chambers, and S.D. SUNDBERG. 2006. Symphyotrict In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. Oxford Univ. Press, New York and Oxford. Vol. 20:465-539. COCHRANE, T.S. and H.H. Iris, 2000. Atlas of the Wisconsin prairie and savanna flora. Tech. Bull. No. 191. Wisconsin Department of Natural Resources and the University of Wisconsin-Madison Herbarium, Madison CorreLL, D.S. and M.C. JoHNsTON. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner, Texas. CRANFILL, R. 1991. Flora of Hardin County, Kentucky. Castanea 6:228-26/. Cronauist, A. 1980. Flora of the Southeastern United States. Vol. 1. Asteraceae. Univ. of North Carolina Press, Chapel Hill. Desem, H.R. 1990. Flora and vegetation of some barrens of the eastern Highland Rim of Tennessee. Castanea 55:187-206 FLORIDA NATURAL AREAS INVENTORY. 2008. Field guide to the rare plants and animals of Florida. «ttp//www.fnai.org/ fieldguide/> Accessed March 2008. Foster, C. & C. HAMEL. 2006. Rare species surveys of the Mamas FERA pala Centre, SCC MS Report 06-01. Manitoba Conservation Data Centre, Winnepeg. Accessed March 2008, GEORGIA DEPARTMENT OF NATURAL Resources. 2007. Wildlife R Division. Known occurrences of Special Concern plants, animals, and natural communities of the Coosa River Watershed. «georgiawildlife.dnr.state.ga.us/as- sets/documents/HUC8_03150105.rtf> Accessed March 2008. GRAUSTEIN, J.E. 1967. Thomas Nuttall, naturalist: Explorations in America, 1808-1841. Harvard University Press, Cambridge. sts Te Al 4 4 | | es pot Hips öf certain North American species. Brittonia 32:240-261. | Jones, R.L. 1990. The taxonomy and distribution of Aster pratensis Raf. ASB Bull. 37:99 (Abstract). KraL, R. 1981. Further additions to some notes on the flora of the southern states, particularly Alabama and middle Tennessee. Rhodora 83:301-315. KENTUCKY STATE NATURE Preserves COMMISSION. 2008. Kentucky Rare Plant Database. Accessed March 2008. LOUISIANA STATE UNIVERSITY HERBARIUM ON-LINE DATABASE, 2008. Dept. of Biological Sciences, LSU, Baton Rouge. < http:// www.herbarium.Isu.edu» Accessed January 2008. Love, A. (ed). 1982. IOPB chromosome number reports LXXV. Taxon 31:342-368 (count on page 359 under Virgulus sericeus). Lupwi, J.C. 1999. we dore of EE and limestone barrens in southwestern Virginia. Castanea 64:209—230, NATURESERVE 2008. ^ 'er:an online encyclopedia of life (web application). Version 7.0. NatureServe, Arlington, Virginia. htt "Ünaturesere: org/explorer. (Accessed April 2008). Nesom, G.L. 1994. EN overview of Aster sensu lato (Asteraceae: Astereae), emphasizing the New World species. Phytologia 77:141-297. SEMPLE, J.C. and L. BeouiLer. 1980. Chromosome numbers and satellite chromosome morphology in Aster and Lasallea. Amer. J. Bot. 67:1027-1039. SEMPLE, J.C. and J.G. CHmieewski. 1987. Chromosome numbers in Fam. Compositae, Tribe Astereae. Il. Additional Counts. Rhodora 89:319-325. SEMPLE, J.C., J.G. CHMIELEWsKI, and M. Lane. 1989. Chromosome numbers in Fam. Compositae, Tribe Astereae. IIl. Ad- ditional counts and comments on some generic limits and ancestral base numbers. Rhodora 91:296-314. Jones et al., Distribution of Symphyotrict i d S. pratense 739 d É d SEMPLE, J.C., S.B. HEARD and L. BroutLLer. 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. No. 41:1-134 SHERIF, A.S., ER SMITH, and K.L. HORNBERGER. 1983. In IOPB Chromosome Number Reports LXXX, Taxon 32:508. SHINNERS, L.H. 1950. Notes on Texas Compositae-V. Field & Lab. 18:32-42. SUNDELL, E. 1983. Two additions to the Arkansas flora from Warren Prairie. Sida 10:188-189. Swink, F. and G. WILHELM. 1994. Plants of the Chicago Region. Indiana Academy of Science, Indianapolis. TENNESSEE NATURAL HERITAGE PROGRAM. 2008a. Natural Heritage Inventory Program. Accessed March 2008 VIRGINIA BOTANICAL ASSOCIATES. 2008. Digital Atlas of the Virginia Flora. Accessed January 2008. WEakLEY, A.S. 2007. Flora of the Carolinas, Virginia, Georgia, and surrounding areas. Working Draft of 11 January 2007. Univ. of North Carolina Herbarium (NCU). Ac- cessed January 2008. Wess, D.H., H.R. Dese. M, and W.M. Dennis. 1997. Studies of prairie barrens of northwestern Alabama. Castanea 62:173-184. WUNDERLIN, R.P. and B.F. Hansen. 2004. Atlas of Florida Vascular Plants [S.M. Landry and KN Campbell (applica- tion development), Florida Center for Community Design and Research.] Institute for Systematic Botany, University of South Florida, Tampa. Last modified 11 July 2007. Accessed January 2008. EE I H Hi JIL] f / Ll HA IE, 740 BOOK NOTICE RACHEL WiLson (ed.). 2007. Sweeteners Handbook, Third Edition. (ISBN 978-1-905224-42-5, hbk.). Blackwell Publishing Professional, 2121 State Avenue, Ames, Iowa 50014-8500, U.S.A. (Orders: www. blackwellfood.com, 1-800-862-6657, 1-515-292-3348 fax). $169.99, 304 pp., b/w figures and tables, n x 9". Contents: Foreword (by the author, with acknowledgments) Contributors Introduction Pat 1. Intense Sweeteners Acesulfame K, Alitame, Aspartame, Brazzeine, Cyclamate, Glycyrrhizin, Neohesperidine DC, Neotame, Saccharin, Stevioside, Su- cralose, Thaumatin Part 2. Bulk Sweeteners Crystalline F , Erythritol, Isomalt, Isomaltulose, Lactitol, Malitol & Malitol Syrup, Mannitol, Sorbitol & Sorbitol Syrup, Tre- halose, Xyitol Part 3. Legislation European Community Legislation, UK Legislation, Gulf States Legislation, USA Legislation, Other Countries, References Suppliers Index From the publisher —“Sweet titut jor add | ingredients group for the food industry, with their appli ane 11 ] 4 4 "r1 1 1 r eT Lr + "m 1 ] 1 LI +1 t r zz 1 + r r Iu or “1 + +1 | ; PS PA Er Asus 1 T] L x £ 1 11 J e 11 ug ee f Lei Som pP 5 ep: e de E. | 1 = 1 dicus EE E 1 J ] +l f:1 H A” r r E E L “First published in 1996, The Sweet Handbook has | an ial ref tool for the food industry, providing detailed ti he properti d applicati fel ly permitted f in food The third edition of this book " = le441 na lal ] £,. 4] = A -l a x 1 11 ] x " MAL LALA id LEN YT CALLA LAI LY | 1 1 Cs ar +1 E AA 1 ] 1 A » ELE Ee ee? D Lu J. Bot. Res. Inst. Texas 2(1): 740. 2008 RANUNCULUS FICARIA (RANUNCULACEAE), NATURALIZED IN TEXAS Guy L. Nesom 2925 Hartwood Drive Fort Worth, Texas 76109, U.S.A. guynesom@sbcglobal.net ABSTRACT R lus ficaria i lized al creek in Tarrant Co, Texas. The plants Gate iai bulbils ges infertile E and per- hap best identified ubsp ubiera, although other morphological featur ly accord p f that subspecies. Key Worbs: Ranunculus ficaria, naturalized, Texas RESUMEN Ranunculus Jemig está BEE a Se Targo de un ee en Tarrant Co, Texas. Las ieee pa aulbiige a axilares y aquenios ins ertl tile y bulbifera, ju E y de acuerdo L E I I i A population of Ranunculus ficaria L. (lesser celandine, fig buttercup) was discovered in flower immediately adjacent to Overton Creek in Fort Worth's Overton Park, near the center of the city. The creek at this point runs through a broad but relatively steep-sided ravine and is about three miles above confluence with the Trinity River. Tributaries of Overton Creek pass through several miles of urban neighborhood upstream from this area. The creek commonly rises beyond its banks in heavy rain. Thirty discrete plants, clearly Ruan. ey in an area of about 30 square feet—the largest produced lants occurs on the stream terrace about 300 yards at least 20 flowers. A second p downstream. Voucher Specimen: TEXAS. Tarrant Co.: Fort Worth, Overton Park between the ae I d Hulen Street, Ge Gees pe lel to Shady Creek Drive, immediately adjacent to the creek on the wet, sandy terra i 2 Mar axillary bulbils abundantly in production by 2 Apr, Nesom FW08-1 (BRIT, MO, NCSC, NCU, TEX). The Overton Creek plants arise from tuber-producing fibrous roots and produce dark green, glabrous, ovate and deeply cordate but otherwise entire (to faintly crenate) basal leaves and EE Jeu flowers P mm on erect to ascending leafless peduncles. The habitat is typical for the species in North America, the plants occur on shaded stream banks, floodplains, low woods, and moist disturbed areas. “Flowering usually occurs from late winter through mid-spring (March through May), depending on conditions. Afterwards, the above-ground portions die back" (Swearingen 2005). Ranunculus ficaria is a beautiful horticultural species, and many photos are available on the internet. Sell (1994) NR DAE five EE based primarily on leaf and bd size, achene fertility, and whether or not axillary | I ft flowering Among t the fi ,only subsp. bul bifera (Marsden-Jones) Lawalree (as Saken bulbilifer Lambinon”) and subsp. carlos (F.W. Schultz) Rouy & Fouc. were described by Sell as producing bulbils, and of these two, only subsp. bulbifera has infertile, glabrous achenes. The Overton Creek plants produce axillary bulbils and infertile achenes and probably are best identified as subsp. bulbifera, but otherwise EM are not a clear bu ge UN match for any of the subspecific taxa in Sell's key. The | ly crowded at the base on short stems. Most leaves are ca. 6—7 cm long and 5-5.5 cm wide, ranging smaller to 3 x 3 cm, on petioles 8-10 cm long. The flowers are ca. 2.5—4 cm in diameter with 7—9 petals 15-19 mm long, 6-8 mm wide. Pollen (2 plants) is ca 7096 viable, as judged by uniform size and full-staining in lactophenol-cotton blue; the remaining 3096 J. Bot. Res. Inst. Texas 2(1): 741 — 742. 2008 742 of the grains are non-staining and distinctly smaller (and highly variable in size). All achenes apparently are glabrous and infertile. Axillary bulbils were not present at the time of collection (8 March) but were in abundant production by 2 April—they are whitish, conic-cylindric to short-cylindric, and essentially without surface features. Superficially, at least, the axillary bulbils are very similar to the root tubers. Many stems and leaves had at least begun to wither by 5 May, with the axillary bulbils loosened and easily detached. Cn in Renan Caen of bovis ME taxa is further tempered by Whittemore's observations (1997): * ( ially in leaf size and stem posture), and many att ts have been made to dide it into varieties or sbspeciés Ges Sell 1994). The different forms, however, intergrade extensively and the varieties are often impossible to distinguish." Sell himself noted (p. 47) that *a large proportion of herbarium specimens cannot be determined with certainty" and relied on field experience in the British Isles and on published literature to summarize distributions of the infraspecific taxa. Sell (1994) noted that “all the subspecies can spread by tubers (Clones of all the subspecies can be produced by division of the root tubers.), two spread by axillary bulbils, and all except subsp. [bulbifera] can spread by seed." Both the tubers and bulbils are easily carried downstream during flood events (Bargeron et al. 2003). Bulbils are produced near the end of flowering or in fruit maturation, and Sell noted that the five taxa can be recognized “if cultivated, or if examined at intervals through their flowering and fruit- ing periods." Detailed observations of this nature, however, apparently have not been recorded in North America—Magee and Ahles (1999) and Rhoads and Block (2007) unequivocally specify that the plants produce bulbils, but other U.S.A. floras note that bulbils are produced "sometimes" or "often." Only Magee and id HAE E E pae to infraspecific rank (as subsp. bulbifera). , northern Africa, and western Asia. In North America it is natural- ized in Oregon, Washington, and British Columbia and from Missouri, Tennessee, and Virginia northward to Ontario, Quebec, and Newfoundland (PLANTS Database 2008). The Texas plants are the southernmost known for the species as naturalized in the U.S.A. It is reported to be invasive in Connecticut, Delaware, Maryland, New Jersey, Oregon, Pennsylvania, Virginia, Wisconsin, West Virginia, and the District of Co- lumbia (Plant Conservation Alliance 2007). REFERENCES BARGERON, C.T, DJ. MoorHEAD, G.K. Douce, R.C. ReARDON, and AE Mier. 2003. The invasive Spa of ne Faster United States: Identification and Control. Invasive and Exotic Plants. Accessed April 2008. Macer, 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. PLANT CONSERVATION ALLIANCE. 2007. GEN pia nt invaders "i natural areas (WeedUS Database). PCA Alien Plant Work- ing Group. Accessed April 2008 PLANTS DATABASE. 2008. Ranunculus fiai: USDA Natural Roues Consemation Service. AT + 1 - o = — Be -J 5 At 1 4 + lada: en atrae maíces in dican t + I L E il 1 1 d 1 = 1 J Ja E re "n x L J F Kate Pp. Por lo tanto, las 1 ] 1 1 T3 H A E f 1 A 1 1 J J ] È 1 T. a e i + del le | lales de | lel le California. La T ia de P. vaginatum, probablemente no reconocida en el Is J : 1: 4 " Lë 11 1 4 SR ] ry 2 1 z H 4 A | J 4 I L + LA + L 1 r1 J 1 A m m 1 15:1 1 1 £ J 1 1 ial | California. INTRODUCTION Plant invasions are a major issue for ecosystems around the world (Williamson 1996; Gurevitch & Padilla 2004). The EEN e SES ey Eyer urbanization, intensive agriculture and horticultural practices, i of natural habitats, biog hical factors, and climate change are some olt the driving forces hari increase the movement of species around the world (Mooney & Hobbs 2000; Vila & Pujadas 2001; Myers & Bazely 2003; Py3ek € Richardson 2006). As a result, the flora in many regions of the United States, including California, is now rich in nonnative (alien, exotic) plants (Westbrooks 1998; Bossard & Randall 2007). Invasive plant species reproduce in very large numbers, are superior competitors, and have the poten- tial to spread across large areas and at considerable distances from parent plants (Richardson et al. 2000). Environmental plant pests (species that affect the loss of native biodiversity) and ecosystem transformers J. Bot. Res. Inst, Texas 2(1): 743 — 759. 2008 744 J tani it Texas 2(1) (species that alter the character, condition, form, or function of ecosystems) pose a serious threat to wild- lands in California and elsewhere (Randall 1997; Vitousek et al. 1997; Richardson et al. 2000; Bossard & Randall 2007). In California's estuarine wetlands, some of the most problematic and well-known ecosystem transformers are the introduced Spartina species, especially S. alterniflora Loisel. and its hybrids (Callaway 2005; California Conservancy 2007). Historically, the coastal plains of California have been 11 ltiple introductions for long periods of time, and consequently they support more nonnative ESCH species ihan many other parts of Je state (Randall et al. 1998; Bossard & Randall 2007) E are respec any ee to DAN invasions, Da ticularly in coastal or urban areas that et al. 1999; Ruiz et al. 2000; Callaway & Zedler 2004: ie & Kercher 2004). Even salt KM which HIDE have not experienced significant i i D to ES E in levels and anaerobic t inhibit establisl t of most ti ts, are now highly ystems (Callaway 2005; Grewell et al. d Acorn the pomum al wetlands to large urban centers, which meron as | sinks Iterations to hydrological regimes, and of ers har promote the establishment of nonnative facultative wetland plants (Galatowitsch et al. 1999: Cronk & Fennessy 2001; Zedler & Kercher 2004). In southern California, expanding urbanization provides year-round sources of water from runoff and discharge of wastewaters into natural drainages, which provide direct pathways of dispersal for nonnative species spreading from uplands to wetlands at the wildland-urban interface (White & Greer 2006; Riefner & Boyd 2007). In this paper, we provide the first documented records of Paspalum vaginatum Sw. (Poaceae, Panicoideae, Paniceae) for California. Paspalum vaginatum (seashore paspalum, saltwater couch, saltwater paspalum, silt- grass) is widely recognized as a serious invasive pest and an ecosystem transformer that has oM in atrasa tira many pubiropica! and tropical coastal gene environments, often with detrimen and function of indi ( & uae A00 Shaw n 2003; Weber 2003; Dana et al. e 2004; Siemens 2006). In M. to understand tl an t of thi E * T | a RI D e n 2 ld EI “and + LEM 1 D 1 tl | traits e California, we review that promote invasiveness and ithe formation of monocultures in estuarine wetlands. Rapidly expanding populations and the formation of dense monocultures pose a serious threat to the structure, function, and biological composition of tidal marshes, sloughs, and mudflats in southern California. thern California is depicted on Figure 1. Individual populations documented bs Seen. The known ane of P. vaginat e dat CALIFORNIA. Los Angeles Co.: City of Malibu, Zuma Lagoon at Westward Beach Rd., UTM gie 83) us 0331858E 3765305N, elev. 3 m (10 ft), rare, € flat with Jaumea, 4 Nov 2005, Riefner 05-744 (RSA); City of Malibu, Malibu Lagoon, W lagoon, S of Pacific Coast Hwy., UTM (NAD a 115 iino a elev. 1 m (3 ft), common on mud flats, channel banks, and in salt marsh with Frankenia and eee highly I t ies, 8 Jul 2006, Riefner 06-290 (RSA); City of Malibu, Malibu Lagoon, E lag Beach, S of Pacific Coast Hw: UTM (NAD 83) 115 0344854E 3767212N, elev. sea level, l | inlet l, on mud flats, d in salt h with Sali ia, 8 Jul 2006, Riefner 06-293 (RSA), ae] hli 4 1, 29 Jul 2006, Riefner 06-327 (RSA); City of Carson, Dominguez Channel, S Sol oe s Blvd. near on St., UTM oe 29) 115 0392702E dd 99N, idi 7 m (23 ft), widespread and ver Salicornia, Distichlis, and Atrif nel, 29 Jul 2006, Riefner 06-330 y " Fui near Victoria Park, Domi GE at Del e nw pene (NAD 83) 115 0392422E 3746119N, elev. 7 m (23 ft), well established, (RSA). Orange Co.: City of San Juan Capistrano, dibus Creek, N side of Del Obispo St. at Paseo Adelento St., UTM (NAD 83) 11S 0438127E 3706763N, elev. 27 m (89 ft), uncommon, ii streambed in shallow fresh water with Paspalum distichum, 14 Oct 2003, Riefner 03-455 (RSA); City of San Juan Capistrano, Trabuco Creek, S side of Del Obispo St. at Paseo Adelento St., UTM (NAD 83) 115 0438127E 3706763N, elev. 26 m (85 ft), uncommon, E streambed in shallow fresh water, 14 Oct 2003, Riefner 03-456 (RSA); City of Mission Viejo, Oso Creek, ca. 0.5 mi N of Paseo ii Comas on oe es UTM (NAD = 115 0437527E 3714753N, elev 86 m (281 ft), well ee and highly i i gI : [e] E growing with Bacchari hefnha Puli ludsoa, and Salix lasiolepis, 9 Jul ment plant, and soil in 2005, defer 05-543 (RSA); City of fission Viejo, Oso Creek, ca. 1.4 mi N of Paseo De Colinas on Comes Capistrano, UTM (NAD 83) s 0437526E 3715062N, elev. 88 m Ge e Wes aman a di invasive i pones E in aa a shallow water icifoli , Paspalum Wal I D distichum, and Scl | i 9 Jul 2 2005, Riefner 05-545 (RSA); City of I Niguel "n Mn Alica Pwky. at Skate Park O with 1 11 f. 1 Way, UTM (NAD 83) 11s 0433416E 3713654N, elev. 46 m (151 fo, locally established in wet ith shallow fresh water, i T i X | VENTURA COUNTY 1 LOS ANGELES COUNT j j E i Y | i Part H | i e ueneme p d j ` kun E Ba T ML ¿ E e ge E we e T es US A if * Riverside X eLos Angeles [TT Sa E \Carsone Ze: H S 4 " Ve VENT er B ar UMEDE SLUM, Indian Wellse* El Tor $ “Mission vis Newport Beach* Ls op eg, , E gt GC 3eSan pistrano NNNM “sea ’ CN pon s [^ — Sarka Catalina DN | = Jsland' H S | m ^ = Carlsbad * * Rancho Santa Fe A) KL AS \ LS | ` 2] LN 4 SE xi d = po Diego Les m pu be 1 M D A | +: £n Ê H A a al FANE EFI dl A | LER | | ns! J A IL L inc Ai L E i ELS d | À J in red. AF7 2M (DC A); Cit [WT D 1 E of Jamboree Rd. at growing with SEN n in Polypogon, a DIS. ici 2007, Riefner 07-304 (RS/ (NAD 83) 115 0419719E 3723895N, elev. 6m (20 ft), tidal wetland, ayview Way, widespread and well established on mud flats and in salt "e Di with Salicornia, Limonium, and Jaumea, highly invasive and excluding native species, 23 Jul 2007, Riefner 07-314 (RSA); City o a Point, Doheny State Beach, E side of San Juan Creek SS UTM (NAD 83) 115 0436583E 3702748N, elev. 4 m (13 ft), E d but invasive, many individual plants scattered on m flats and wet sand, 17 Oct 2007, Riefner 07-434 (RSA); City of D Point, Doheny State nem W side a an de icd did POM (NAD 83) 11S idis 3702895N, elev. 6 m e ID, y common on dry mo with Jaumea, gulatus, B , Distichlis, and Typha, 17 Oct 2007, Riefner 07- 436 GER E of Dana Point, Dana Point Sege Dana Point Harbor pi near Ss Lantern I Dr., SC o ES ge 0436040E 3702905N, elev. 4 m (13 ft), locally hrharta erecta, Paspalum dilatatum and near Puerto Pl, UTM (NAD Typha, 17 Oct 2007, kigi 07-437 (RSA); Cay of Dam Point, E SE - Beach Dan 83) 115 0435964E 3702721N, elev. 10 m (33 ft), local, sand dune swale with Distichlis " Juncus acutus, 17 Oct 2007, Riefner 07-439 (RSA); City of Laguna Niguel, Laguna Niue in e oa de La Paz Rd. at iis Creek, UTM (NAD 83) 115 0434146E 3712609N, elev. 64 m (210 ft), locally est Typha, 26 Oct p ee EE SAN City of Laguna wA d of La geg Rd. at aS de eg , UTM (NAD 83) 1150434619E 3711316N, in concrete ch , growing with Atriplex prostrata and Typha, 26 Oct 2007, Rioja 07-459 (RSA); City of Newport Beach, North s Se Bench, EH shore of Newport Bay near noe Star Ln., UTM (NAD e 11S 0417162E 3721028N, elev. 9 m ft), t h, g growing near p lvlanti I marit imus, 26 Oct 2007, Riefner 07-461 s NA of Laguna Beach, Ali com at Pacific Coast Buy: UTM (NAD 83) 115 0430150E 3708320N, elev. ca. 9 m (30 ft), j s with Pulicaria and Jaumea, 13 Jan 2008, Riefner 08-4 (RSA); City of Ga Se San ud rd at Ld 73,E in bd i [M ndi 85) 115 0419906E 3723854N, elev. ca. 6 m (20 ft), locally g g n 2008, Riefner 08-9 (RSA). Riverside Co.: Salton Si D ] O The Nature Conservancy preserve, 33°30, 04 » 115°49.9°W, elev. -37 m DA palu m disti tated Nov. 2007]; In- ft), wet soil and shallow water, 4 Dec 1994, Sanders et al. 15885 (RSA) [originally det Whi River, Hwy. 111 near Happy Point, UTM (NAD 82) 11S 0564872E 3730987N, elev. ca. 20 m (66 ft), rors dian Wells, tributary to 746 | | tani titute of Texas 2(1) 1 al f urb reek, growing with Scirpus robustus, I hl bsp. uninervia, and Eclipta, ` 7 Aug 2005, Riefner 05-596 (RSA). San Diego Co.: City of o Santa Fe, = o St. at H Canine eni SA Elio EE UTM (NAD 83) 11S 477707E 3652459N, elev. 12 m (39 eh ll est water subsp. peploides, 8 Sep 2002, Riefner 02-224 (RSA); City ho Santa Fe, La Orilla St. ca. 0.4 mi E of El Camino Real UTM (NAD 83) 11S 0478231E 3652819N, elev. 21 m (69 ft), well blished but local rsh, growing with Aster subulatus var. ioi Distichlis, and Salicornia, 8 Sep 2002, Riefner 02-224 (RSA); City of ses Loma CH Sbugh b Buccaneer Park, Pacific St. at Morse St., UTM (NAD 83) 11S 0465660E 3671043N, elev. 1 m (3 ft), well li on creek banks growing ER podus Se ee en and Atriplex BECH 8 Sep 2002, Riefner 02-221 (RSA) loriginally det. as aspalu widespread, highly invasive and excluding native species, 5 Oct 2007, Riefner 07-420 (RSA); GC di Oceanside, Loma Alt ba at Pacific Coast Hwy., UTM (NAD 83) 115 0465836E 3671307N, elev. 13 m blis! water, growing with e Schoenoplectus ig. and Typha, 5 Oct 2007, 7. Riefner 07-420 (RSA); City of M ocu Buena Vista ku Rue e Ge at Ocean St., UTM (NAD 83) 115 0466593E 3669614N, elev. 1 m (3 ft), locally | , Juncus acutus, and Tamarix, 6 15 Oct 2007, Riefner 07-429 (RSA); City of Carlsbad, Batiquitos Lagoon, vicinity of i Rd. at Columbine Dr., UTM (NAD 83) 11S 0474831E 3661679N, elev. 5 m (16 ft), rare, invading Jaumea marsh, 15 Oct 2007, Riefner 07-432 (RSA). Ventura Co.: City of Point Hueneme, Bubbling Springs Ge E of See Dr. dod Port Beer Dr., UTM (NAD 83) 115 0297999E 3780473N, elev. 3 m (10 ft), locally common on creek banks, Į with Ludwigia, Typha, and Schoenoplectus, 8 Jul 2006, Riefner 06-280 (RSA). PREVIOUS KNOWLEDGE Distribution. eim vaginatum is a perennial grass documented from warm, coastal regions around the world, incl t and southeast Europe, tropical Africa, temperate and tropical Ásia (Arabia, China, India, Malaysia), Australia, New Zealand, French Polynesia, the Galápagos Islands, Hawaii, and other Pacific Islands. the Caribbean, Central and South America, and the Atlantic and Gulf Coasts of the United States and Mexico in North America (Waterl 1997; Allen & Hall 2003; Zuloaga et al. 2003; Clayton et al. 2006). Its exact native range, which is centered in the tropical and subtropical regions of the New World, remains uncertain (Chase 1929; Wagner et al. 1999; Edgar & Connor 2000; Weber 2003). In the United States, P. vaginatum occurs from North Carolina to Florida, west in the Gulf States to Texas, and in New Mexico, but was not previously reported for California (Webster 1993; Allen & Hall 2003; DiTomaso & SECH 200r, USDA id cam Flora Project 2008). ge,an t of cultivation ; Pr agin atum ] tal salt and brackish water marshes, shallow-water l d tidal channels, mangroves, coastal ET EECH , dunes and beaches, summer-moist saltpans, wet pastures, and freshwater ripari 1 floodplain habitats in many warm temperate, subtropical, and tropica l ts (SI & Riveros 1990; Stutzenbaker 1999; Wagner et al. 1999; Edgar & Con- nor 2000: Filigheddu et al. 2001; Allen & Hall 2003; Shaw & Allen 2003; Dana et al. 2004; Siemens 2006). Anthropogenic Uses and Worldwide Introductions.—Paspalum vaginatum has numerous anthro- pogenic uses that have transported it to many countries around the world, and eventually to southern California. Historically, it was used as bedding in trade ships that sailed between Africa, North and South America, Central America, and the Caribbean Islands during the 1700s and 1800s (Duncan & Carrow 1999). Paspalum vaginatum has also been used widely for rehabilitation of salt-affected lands, stabilization of sand dunes and dredge materials, restoration of coastal wetlands, and forage for livestock (Leithead et al. 1971; OIC 1990; Vargas 1995; NRCS 1999; Loch & Lees 2001; Barrett-Lennard et al. 2003; Fontenot 2007). As water shortages continue to escalate worldwide, increased use of saline water or poor-quality re- claimed effluent waters for turf landscape irrigation is necessary (Snow 2001). The systematic development and cultivation of varieties (cultivars) of turf grasses with improved tolerance to saline soils and low-quality irrigation waters, including P. vaginatum, are playing a central role in the turf grass industry (Henry 1981; Hall 1994; Brown et al. 1998; Carrow & Duncan 1998; Duncan 2003; Marcum 2004). To meet the demands of water conservation and expanding recreational needs, P. vaginatum is now grown on golf courses in Ásia (including Malaysia, Indonesia, and the Philippines), South Africa, the Middle East, Sardinia, New Zealand, South America, Hawaii, the Caribbean Islands, and across the southern United States (Duncan & Carrow 1999; Beard 2002). Paspalum vaginatum is also used as a ground cover in gardens, and new cultivars are Riefner and Columbus, Paspalum vaginatum in southern Californi 747 highly promoted for lawns in the southern United States (USDA Zones 7-10), Australia, Hawaii, and else- where (Duncan & Carrow 1999; Rushing 2006; Christians et al. 2007; Tukey 2007). Environmental Tolerances Create a Competitive Advantage.—Paspalum vaginatum has broad ecological tolerances and life history traits that aid its widespread establishment and utility to land and resource managers worldwide. It is an EE aggressive, SC and stoloniferous mat-forming perennial grass that often f ies (Hitchcock 1951; Weber 2003). It roots well in sand, clay, silt or mucky soils in acidi Ikaline conditions, is tolerant ih waterlogged and highly saline soils, prospers in low fertility soils and poor-quality effluent waters used for irrigation, is highly tolerant of mechanical Done and heavy livestock grazing, has high shoot density for effective soil ao develops a thick f tolerant of foot traffic and wear, and is not readily tible to drought or pathogens ica 1999; Duncan & Carrow 1999; Loch & Lees 2001; BxirettLeunard e et al. 2003; Lee et al. 2004a,b; Fontenot 2007). Therefore, its rapid growth, abundant propagules (in this case rhizome and stolon frag- ments), and broad environmental tolerances are important characteristics typical of an “ideal weed" (Sakai et al. 2001). In a given environment, these characteristics often confer an advantage to a nonnative species relative to indigenous species (Barney & DiTomaso 2008). Invasive Status Worldwide.—Paspalum vaginatum is well known for its aggressive behavior and its ability to dee nes ecosystem renun (Weber 2003). It is an bean) and eee STEE of disturbed, bare o T deposit ta and once established it ] indi | | for many years Shaw & Allen 2003). "eiim XE DONUM is, however, selfincompatible rarely produces viable seed t quantity, and therefore, must | ted vegetatively (I Iall 1994- Duncan & Carrow 1999). Hone it would be careless to conclude that Se lacking sexual reproduction cannot become serious invaders; giant reed (Arundo donax L.) is a classic example of a sterile but highly invasive plant of southern California's riparian habitats (Bossard et al. 2000; Barney & DiTomaso 2008). Outside of cultivation, vegetative reproduction of P. vaginatum would be possible through animal graz- ing and other natural physical regimes that would disturb and transport rhizomes and stolons (Graeme & Kendal 2001). Paspalum vaginatum is apparently dispersed primarily by water when pieces of stolon or rhizome are carried often long distances by streams. ZE ye fragments root easily, forming new plants that start new infestations (Weber 2003). Est ts, shallow brackish lagoons or tidal creeks, and gaps in disturbed wetlands or scoured streambeds that are not adi colonized by indigenous species are highly vulnerable to invasion (Filigheddu et al. 2001; Shaw & Allen 2003; Siemens 2006). Importantly, P. vaginatum and its impacts on the function of native estuarine ecosystems are significant and comparable to the well-documented invasions by nonnative Spartina species (Graeme & genii 2001; Allen & Shaw 2003; Dana et al. 2004; Siemens 2006). In New Zealand estuaries, and much like the ecological impacts effected by introduced Spartina spe- cies, P. vaginatum transforms the composition and structure of indigenous vegetation by growing over and displacing low-stature species, endangering populations of threatened species, altering fish spawning and feeding grounds, prohibiting burrowing fauna and reducing access to feeding sites and roosting by shore birds due to its high shoot and root density, and altering estuarine hydrology by accumulating sediments (Graeme & Kendal 2001; Shaw & Allen 2003). Similar negative effects in brackish wetlands have been documented in Spain, where P. vaginatum dis- places halophytes and alters the composition and structure of native communities, promotes accumulation of sediments and organic detritus, and alters hydrological and nutrient regimes (Dana et al. 2004). Accord- ingly, P. vaginatum has been classified as *dangerous" (a species causing ecological damage or alteration to natural ecosystems) (Dana et al. 2007). Paspalum vaginatum has also been classified as invasive in Portugal (Domingues de Almeida & Freitas 2006). In the Galapagos Islands, EEN P. agi atum stands out as a special case. It has aggressively invaded disturbed and natural wetlands where ive plants, and affects wai t and soil moisture content (Vargas 1995; Gravez et al. 2004; Guézou et al. 2007). Paspalum vaginatum has also successfully 748 Journal of the Botanical Research Institute of Texas 2(1) invaded many lagoons, threatening these ecosystems and their globally significant avifaunal biodiversity. Quantitative investigations have GE that P. Laid eus are EE for E fiddler crab habitat, and for alter ing T bitat of | f water availability during feeding (Siemens 2006). Ov eval. these iio lacilitaie e a shift kom an aquatic to a more ter- restrial inverte brate community, thereby affecting food availability for native fauna foraging in and around lagoons (Siemens 2006). Accordingly, its control and eradication are necessary in order to maintain the status of the Galápagos Islands lagoons as a Ramsar Internationally Important Wetland (Siemens 2006). In South Africa, Héfliger & Scholz (1980) documented similar negative affects to native community functions, including competition for space, light, water, and nutrients, replacement of native vegetation, en of water flow, iu contaminant seed loading. Further, the invasion of P. vaginatum and other nonna 1 the sand and mudflats of the Wilderness Lakes Wetland (designated a Ramsar site) unstable for wading birds (Randall & Russell 1995). Paspalum vaginatum is also invasive and has been classified as a "significant environmental weed" in Australia (Randall 2007). It is also invasive in the Hawaiian Islands and the Marshall Islands (Wagner et al. 1999; Whistler & Steele 1999; Vander Velde 2003). Accordingly, PIER (2007) has ranked P. vaginatum as a "high risk" invasive species. 3 SIGNIFICANCE IN SOUTHERN CALIFORNIA Introduction, Dispersal, and Naturalization.—These naturalized populations represent the first docu- mented records of P. vaginatum for California (Webster 1993; Allen & Hall 2003; USDA 2007). Paspalum vaginatum has also not been included in major publications on nonnative species established in California (Bossard et al. 2000; Hrusa et al. 2002; DiTomaso & Healy 2003; Bossard & Randall 2007; DiTomaso & Healy 2007; Grewell et al. 2007). Thus far, P. vaginatum has not been of concern in southern California (Roberts 1998; Roberts et al. 2004; Rebman & Simpson 2006; Clarke et al. 2007; Riefner & Boyd 2007) Biologists often have difficulty tracing the source of nonnative organisms and verifying their acciden- tal versus intentional human transport across natural boundaries. However, the arrival of P. vaginatum in California is most likely the result of purposeful introductions, presumably for soil stabilization and habitat enhancement projects or more likely for cultivation of turf grass for lawns, parks, and golt courses. Native in the southeastern United States, P. vaginatum has been used for restoration of coastal wetlands, stabilization of dunes, banks, and dredge materials in the Gulf States (Duncan & Carrow 1999; NRCS 1999; Fontenot 2007). Accordingly, it may have been introduced for enhancement or restoration of salt marsh habitats in southern California, but we have not been able to confirm this. However, P. vaginatum was introduced to southern California for cultivation as a turf grass during the 1970s and 1980s for research at the University of California South Coast Field Station in El Toro, Orange County, and for commercial production in Indio, Riverside County. It was also planted for turf grass in Laguna Niguel, Orange County, and at Rancho Santa Fe, San Diego County (Henry et al. 1979; Duncan & Carrow 1999). As documented by herbarium records (Sanders et al. 15885, RSA; originally identified as P. distichum), the earliest escaped and naturalized plants were collected in 1994 near the Salton Sea, Riverside County (Fig. 1). The senior author also made observations of P. vaginatum in 1994 (originally identified as P. distichum and not collected) at Doheny State Beach, City of Dana Point, Orange County, during surveys conducted for California State Parks (Fig. 1). Species beginning to escape cultivation receive little attention, largely because of the difficulty in identifying and locating them before they become established in native communities. Time lags in plant invasions, a phase when plants are difficult to document during early dis- persal, then exponential growth, naturalization in native communities, and subsequent spread describes a typical invasion scenario (Sakai et al. 2001; Schierenbeck et al. 2007). In the case of P. vaginatum, plants escaping cultivation from sites in El Toro and Laguna Niguel, Orange County, to native habitats on the coast at Dana Point, Orange County, and from inland sites at Indio to near the Salton Sea in Riverside County, perhaps during the 1990s, appears to be a logical scenario for southern California (Fig. 1). nH TË a | L ,P | a A H al CRUE a 749 For this reason, we believe turf plantings may be the most likely source of propagules that dispersed from irrigated urban landscapes downstream to native plant wetland communities. Although its reproduc- tion outside of cultivation and dispersal mechanisms are not well understood, P. vaginatum may have been spread vegetatively by animals, carried by mechanized landscape equipment to new sites, or rhizome and stolon fragments may have been dispersed by water during storm events from turf plantings in the uplands into urban drainages and then downstream to native wetland communities. Paspalum vaginatum is now widely naturalized in southern California's coastal wetlands and at inland habitats near the Salton Sea (Fig. 1). Itis most common in estuarine wetlands where it can form monocultures on mudflats, tidal marshes, and shores of estuary lagoons (Fig. 2), or along tidal creeks and sloughs (Fig. 3). Bare soils of tidal mudflats (Fig. 4) and wet sand of disturbed streambeds (Fig. 5) appear to be highly vulnerable to colonization. Paspalum vaginatum also grows in many non-saline wetland habitats, including disturbed riparian scrub and slow-moving waters of urban creeks in coastal and desert regions. However, P. vaginatum is unlikely to form extensive monocultures or survive long-term competition from large, fast- growing plants and shrubs Bus pi a environments. Urbanization a Wetl Paspalum vaginatum is one of many subtropical and tropical non- hed and ding in southern California. Expanding urbanization has fragmented natu habitats, encouraging nonnative planis cultivated in cities to move easily from human-maintained landscapes to native communities (Vitousek et al. 1996; Cronk & Fennessy 2001). In urban environments, wetlands often accumulate debris and sediments, and receive floodwaters that supply resources (nutrient enrichment, elevated levels of light following flood scour) that may accelerate the growth of opportunistic nonnative plants and facilitate invasion of sunny paps in ES cann (Davis et al. 2000; nd & os 2004). In addition, hydrological and hyd ons, accelerated erosion, and genic disturbances to wetland and "s habitats have created numerous —á for invasion by nonnative hydrophytes (Newman et al. 1996; Tickner at al. 2001; Werner & Zedler 2002). Historically, summer drought and the seasonal nature of stream flows in coastal southern California have acted as a barrier to colonization by nonnative facultative wetland plants in the wildland-urban inter- face (Brigham 2007; Riefner & Boyd 2007). Increased moisture availability in urban environments, anda oa a ephemeral to perennial stream flows owing to waste water E into a M e inan er-dry climate] ilitated the dispersal of tl I g California (Greer & re 2003; NE S Boyd 2007). Biological Traits P in Estuarine Wetlands.—Many wetland plant invasions can be explained by the ecological requirements and attributes of the potentially invasive plant species and the local environmental opportunities afforded by anthropogenic disturbances (flooding, nutrients, propagules, sediments) that may promote invasion and the formation of monocultures in wetland sinks (Werner € Zedler 2002; Kercher & Zedler 2004; Zedler & Kercher 2004). In addition, the selection and breeding for horticultural or agronomic purposes of an individual species, its a in eine naive em mxTosT os mrt with Acirimantal environments, followed by escape and naturalization, often describes a scenario typical of many invasive plants introduced and now established in the United States (Reichard & White 2001). Paspalum vaginatum cultivars with improved salt-tolerant characteristics have been developed and are now used widely in the turf grass industry (Lee et al. 2004a,b; Duncan & Carrow 1999, Duncan 2003). Paspalum vaginatum is one of the most saline-tolerant turf species available, with ecotypes exhibiting high- vigor root and shoot growth at high salinity levels (Carrow & Duncan 1998; Beard 2002; Lee et al. 2005). Additionally, cultivars have also been developed and evaluated for improved growth rate and transplant survival for restoration and soil stabilization of brackish shorelines, dunes, mudflats, and dredge materials in the Gulf Coast region of the southern United States (Duncan & Carrow 1999; NRCS 1999; Fontenot 2007). ur" RA Ex 23M LL L Stot J E. £144 i L RR NZ .L [| J | L ] A la cr z de al b EL | PR L [| phytes Phot h tal tl Alta SI gh, Oceanside San Diego County i e f 8 T NERO ão — "T I ch EN n mL: | P | E be T1 A va - Q Ta Ta: Fr ibi 752 a ) £l n.a H In Ly ZA ol ET 2(1) lexas 2(1) TABLE 1. Attributes of P bn aai tet promote! Invasive PENMON in sd Wetanas and its SH to form monocultures in estuarine wetlands in southern C MT 1234. Opportunity or Wetland Niche Open to Invasion A ru E Agito! A = A. nm A I H B aL Formation of Monocultures Unvegetated mudflats Hin cun ena anoxic c soils Alteration |, flucti lating wa des and shallow water Saline soils Inundation tolerance, aggressive colonization, lack of competition from native hydrophytes erenchyma in roots Broad ecological tolerance, rhizom | stolons that float and grow out into water, adventitious roots, and rapid growth J { pi Fäi 5 Ie [t " Pus High salt tolerance, sustained vigo! Dhi | nt] E n Ir It y water, aggressive colonization t rhizome mats, and lack of competition Flooding, nutrient influx, and sediment deposition anri acta Lal el rM | E from native hydrophytes Warm-season o with C, pathway, wide pH tolerance and adaptation to diverse soil t Canopy gaps High light requirements, aggressi from fast-growing native hydrophytes Summer-wet urban drainages I . Lt , and lack of competition D D ds D (fe A AA A ás 141 , and migh 1 m tal aff ATATIA sf a 14 le Accordingly, P. to native estuarine ecosystems if they po e urban environments or if they are used in restorations United States. The biological adaptations outside the native d of the species, i and attributes of P. 1 atum that promote its invasive SES vior associated with anthropogenic disturbance in ann wetlands, and its ability to form monocultures in native estuarine wetlands are summarized in Table SS to the Environment.—Our field observations indicate P. vaginatum is a threat to the ecological function of estuarine wetlands in southern California. We currently lack experimental data to substanti- ate the changes to wetland composition and structure that we have observed. However, strong anecdotal evidence and quantitative data gathered in other countries provide sufficient baseline information indicat- Ing its impacts to native ecosystems may be significant and comparable to the well-documented invasions by introduced Spartina species (Graeme & Kendal 2001; Shaw & Allen 2003; Dana et al. 2004; Siemens 2006). ie of Lar dde species Spec S. alterniflora and its hybrids, have engineered the alteration of calto ftid al mudflats to , changes to creek morphology and ead E failure of local wetland restoration projects, and local extinctions of the native California cord grass (Spartina foliosa Trin.) in the San Francisco Estuary (Ayres et al. 2004; Callaway 2005; Strong & Ayres 2005; California Conservancy 2007; Grewell et al. 2007). Comparisons of the environmental impacts of introduced Spartina species versus P. vaginatum are summarized in Table 2 CETA TEE P211 DISCUSSION Wetlands are uniquely important one resources EE are highly vulnerable and threatened in southern California (Ferren & Fiedler 1993). P , rapidly expanding urbanization, habitat fragmentation, and poorly coordinated conservation programs have resulted in the long-term, unmitigated loss and degradation of wetland resources, which are proportionally higher in California than in other parts of the United SEN (Dahl 1990; E et al. a) To compound historic losses, wetland invasions by threatei l ecosystem function, which may soon surpass habitat loss as diem main cause of E on (Chapin et al. 2000; Grewell et al. 2007). Hann 5 Fx 76 liforni 753 (fe H " J ic TABLE 2. C i fi ts to estuarine wetl in the San F j Est northern California, f P. vaginatum in southern California's af estuarine wetlands. Impacts of Spartina Species: San Francisco Estuary Impacts of Paspalum vaginatum: Southern California Estuarine Wetlands | anum gen, en ete A P4 with the federally | isted Cordylanthus mollis subsp. mollis chokes channels used by the endangered California clapper rail to forage, and displaces pickle weed marsh of the endangered salt marsh harvest mous fe A P | || Fa] a L VI TIGE oad, r coalesces t Cord grass meadows ~ | xl Ir tamong dense shoots, which increases the elevation of the mud flat for further colonization | | E L 1 € I £f Li p Bo "a4 a fll aL the alterina ! 3 open mud of cl marsh hydrology | £ SAS J L BL | ti m i la |] hydrology and channel habitat Local extinction of the native California cord grass (S. roosa): conversion of Diae California rp = wit! LI th mh federally and state-listed Cordylanthus maritimus DEN e insufficient inf available for potential impacts to listed animal species potentially competes with, grows over, or inhibits germination of ar am I + ci b A i | pl m | PEU RN ES = GUICI coulteri, and S | F Li -t ' L OF Mails, L E 2l + [| J Wi A SÉ : : SII IVY I tabili | ib! A m € | : | I ki y ng dense shoots, to stolons, and rhi further colonization gh shoot, rhizo ciue d E EE SA EIA Y CAVIS S VI Se alter ELS n of burrowing falta I f cl bird f. ina | itat flats tidal creek KAN channels, and shallow lagoon waters, which could become unsuitable for wading birds h ^ and smothers Batis, m Distichlis, Frankeni ++ H ZE Atriml low | E tO S. alterniflora Isolepis, jaumea, Limonium, M hloe, Salicornia, UA hybrids and Triglochin Failure of wetland restoration project Failure of wetland restoration project obj | ly ia uds acd MOM MAE d tof displaces and exclud f native species native species in aed i rid. isa E gg 1 D ogical AAA CI fiada that jous threat to wetland diversity JA ue California. Beine the use of P. vaginatum will probably increase in turf grass] SH marshes and H irr na ME southern California, it is unlikely that the species will decline. In fact, P. vaginatum will most likely: spread to new DEE which points to an immediate need to control or eradicate expanding infestations. tand research programs have been developed to monitor and restore P. vaginatum-affected wetlands in € countries (Gree & Kendal 2001). Potential strategies and resource management topics relevant for urbanized southern California are: 1) Create awareness and educate biologists, field researchers, and resource agencies regarding the invasive behavior of P vaginatum, [| £ n a Inm LI PEE cd ET 4 754 Jo tl t lexas 2(1) 2) Add P vaginatum to the California Invasive Plant Council list of plants of greatest ecological concern (severe ecological impacts on physical processes, plant and animal communities, and vegetation structure); 3) Expand field surveys to determine the complete distribution and abundance in native habitats, quantify expanding use in urban landscapes, and determine if it is used in restorations; 4) Assign P vaginatum an obligate wetland indicator status (OBL, found >99% of the time in wetlands) for California (Region 0) on the National List of Plant Species that Occur in Wetlands to aid wetland delineations; 5) Identify and monitor adverse ecological changes to wetland ecosystems and potential impacts to sensitive floral and faunal elements; 6) Develop habitat- and region-specific control programs; 7) Conduct ecological risk assessments of potential control techniques to avoid secondary impacts to sensi- tive resources. Perroi | naturalization of P. vaginatum d tes t gent need to implement detailed floristic monitoring of wetland outhern California's rapidly url g ities. Early detection of new invasive plants would likely minimize their impacts on biodiversity and ecosystem func- tions, and reduce the high cost of control and eradication. Declines in biodiversity of wetland ecosystems in urbanized environments are becoming a major concern to resource managers, especially in California, which has the highest number of wildland-urban interface housing units (and associated parks, lawns, recreational landscapes planted with turf grass) in the United States (McKinney 2002; Radeloff et al. 2005; Schwartz et al. 2006). Accordingly, the widespread utility of P. vaginatum for turf landscapes, and potential use in wetland restorations, may require the participation of agronomists, biologists, conservation groups, and local and federal government ce ies where wetland biodiversity and conservation objectives a potentially conflict with urban landscape management practices in southern California. ACKNOWLEDGMENTS We are deeply indebted to Catherine Beard (Environment Waikato, Hamilton East, New Zealand), Hamish Kendal (Natural Solutions, Coromandel, New Zealand), Rossella Filigheddu (University of Sassari, Italy), and Tania J. Siemens (Cornell University and Charles Darwin Research Station, Ecuador), who kindly shared their knowledge, experiences, and documentation regarding the invasive behavior of Paspalum vaginatum. We also thank Ruben Ramirez, Cadre Environmental, who prepared the distribution map. Barney Lipscomb (Botanical Research Institute of Texas), Roy J. Shlemon (RJ. 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KercHer 2004. Causes and consequences of plant invasions in wetlands: opportunities, op- portunists, and outcomes. Crit. Rev. PI. Sci. 23:431-452. ZuLoaGa, F.O., O. Morrone, G. Davipse, T.S. Figueiras, P.M. Peterson, R.J. Soreng, and E.J. Juoziewicz. 2003. Catalogue of New World grasses (Poaceae): Ill. Subfamilies Panicoideae, Aristidoideae, Arundinoideae, and Danthonioideae. Contr. U.S. Natl. Herb. 46:1-662. 760 L I A£«kAD SC hi dese £T. TEN Kë BOOK NOTICE CHARLES Kress. 2008. The Ecological World View. (ISBN 978-0-520-25479-4, pbk.). University of California Press, Berkeley and Los Angeles, California, U.S.A. (Orders: www.ucpress.edu, orders@cpfsinc.com, The University of California Press, c/o California/Princeton Fulfillment Services, 1445 Lower Ferry Road Ewing, NJ 08618, U.S.A., 1-800-777-4726, 1-800-999-1958 fax). $59.95, 574 pp., 8 1/4" x 10 3/4". Contents: , Preface 1). An Introduction to Ecology 2). Geographic Ecology 3). What Limits Geographic ain m 7). 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