VOLUME 1, NUMBER 2, 12 DECEMBER 2007 "AT ES Y x SY 9. x A IN > M ‘ x Ni JOURNAL OF THE \ i BOTANICAL RESEARCH TEXAS ( JOURNAL OF THE BOTANICAL RESEARCH INSTITUTE OF TEXAS HISTORY AND DEDICATION 1962—Lloyd H. Shinners (left), a member of the Southern Methodist University (SMU) faculty and a prolific researcher and writer, published the first issues of Sida, Contributions to Botany (now J. Bot. Res. Inst. Texas) 1971—William F. Mahler (right), professor of botany at SMU and director emeritus of BRIT, inherited editorship nd iaht nvri FED 1993—BRIT becomes publisher/copyright holder. 2007 —First issue of J. Bot. Res. Inst. Texas. MISSION AND GOALS The BRIT Press seeks innovation and excellence in preparation, manufacture, and distribution of botanical research and scientific discoveries for the twenty-first century. The BRIT Press—bringing out the best in botanical science for plant conservation and education. DIRECTION AND COVERAGE The BRIT Press considers original research papers concerned with classical and modern systematic botany, sensu lato, for publication in J. Bot. Res. Inst. Texas. All submissions are peer-reviewed. Guidelines for submissions are available from the BRIT Press website, http://www.britpress.org. BIBLIOGRAPHICAL Citation abbreviation for the Journal of the Botanical Research Institute of Texas is J. Bot. Res. Inst. Texas following the principles of B.P.H. (informally JBRIT). International Standard Serial No. (ISSN 1934-5259) FREQUENCY OF PUBLICATION J. Bot. Res. Inst. Texas is published semiannually (summer/fall) n volume by the Botanical Research Institute of Texas. J. Bot. Res. Inst. Texas VOLUME 1 NUMBER 2 ISSN 1934-5259 12 DECEMBER 2007 COPYRIGHT 2007 Botanical Research Institute of Texas (BRIT) 509 Pecan Street Fort Worth, Texas 76102-4060, USA EDITOR Barney L Lipscomb Botanical Research Institute of Texas 509 Pecan Street Fort Worth, Texas 76102-4060, USA 817-332-7432; 817-332-4112 fax Electronic brit.org; jbrit@brit.org CONTRIBUTING SPANISH EDITOR Félix Llamas Dpto. de Botanica, Facultad de Biologia Universidad de Léon E-2471 Léon, SPAIN SUBSCRIPTION PRICES (2008) $41. Personal (Individual/Family) $90. USA (Institutional) $100. Outside USA (Institutional —= SUBSCRIPTIONS/BACK VOLUMES J. Bot. Res. Inst. Texas and Sida, Contr. Bot. Judy MacKenzie, Publications Assistant Botanical Research Institute of Texas 509 Pecan Street Fort Worth, Texas 76102-4060, USA Electronic mail: jmackenzie@brit.org COMPOSITION rhorngraphics, Plano, Texas; rlrc! PRINTING Prepress production and printing in the United States of America by Millet the Printer, Dallas, Texas www.millettheprinter.com COVER ILLUSTRATION Electronically tinted botanical illustration of Liatris aestivalis originally used on BRIT's anniversary poster 2001. Summer gayfeather flowers mid Jul-Aug(-Sep) and is endemic to Oklahoma and Texas. Sida 19:768. 2001. Botanical illustration by Linny Heagy ©2001. TABLE OF CONTENTS SYSTEMATICS Revision of Lobelia sect. Galeatella (Campanulaceae: Lobelioideae) Thomas G. LAMMERS A new species and two new combinations in Potentilla sect. Niveae (Rosaceae) David F. MURRAY AND REIDAR ELVEN A new species of Alocasia (Araceae: Colocasieae) from Panay Island, Philippines MELANIE P. MEDECILO, GEORGE C. YAO, AND DOMINGO A. MADULID Folia taxonomica 2. New species of Passiflora subgenus Passiflora (Passifloraceae) from the Guianas CHRISTIAN FEUILLET Contributions to the genus Festuca (Poaceae: Poeae) in Mexico and a key to the Central American species STEPHEN J. DARBYSHIRE Aphanelytrum peruvianum (Poaceae: Poinae): a new species from Peru IsIDORO SÁNCHEZ VEGA, PAUL M. PETERSON, ROBERT J. SORENG, AND SIMON L44GAARD A new species of Chusquea sect. Verticillatae (Poaceae: Bambusoideae) from Ecuador LYNN G. CLARK, CHRISTOPHER D. TYRRELL, Jimmy K. TRIPLETT, AND AMANDA E. FISHER Estudios en las Apocynaceae Neotropicales XXX: tres nuevas especies andinas de Mandevilla (Apocynoideae: Mesechiteae) J. FRANCISCO MORALES Estudios en las Apocynaceae Neotropicales XXXI: el complejo de Mandevilla hirsuta y cuatro nuevas especies J. FRANCISCO MORALES Boerhavia triquetra var. intermedia (Nyctaginaceae); a new combination and varietal status for the widespread southwestern North American B. intermedia RICHARD SPELLENBERG e be se a 1 fel 1 > 4 1 yperaceae) L Davip J. ROSEN, STEPHAN L. HATCH, AND RICHARD CARTER Fourteen new combinations in Sedum (Crassulaceae) HIDEAKI OHBA A new gypsophilous species of Erigeron (Asteraceae: Astereae) from northeastern Mexico Guy L. Nesom Folia taxonomica 3. Passiflora davidii (Passifloraceae), a new species in sub- genus Passiflora and a key to the sections of supersection Stipulata CHRISTIAN FEUILLET Four new combinations in Mosiera (Myrtaceae) from the Caribbean ANDREW SALYWON New combinations in Philadelphus (Philadelphaceae) JAMES HENRICKSON Five new varieties of Agapetes (Ericaceae) from India D. BANIK AND M. SANJAPPA 789 815 819 Phylogenetic analyses of the genus Baptistonia (Orchidaceae: Oncidiinae) sensu lato based on morphological characters Guy R. CHIRON Muhlenbergiinae (Poaceae: Chloridoideae: Cynodonteae): from northeastern México PauL M. PETERSON, Jesus VALDÉS-REYNA, AND YOLANDA HERRERA ARRIETA A second species of Cyclopogon s.s. (Orchidaceae: Spiranthinae) Eric A. CHRISTENSON Lectotypification of Maxillaria heteroclita (Orchidaceae) Eric A. CHRISTENSON Miscellaneous typifications, new combinations and one new variety in North American Crataegus (Rosaceae) J.B. PHIPPS Kruschke names in North American Crataegus (Rosaceae) J.B. PHIPPS New combinations in Dudleya (Crassulaceae) and Ribes (Grossulariaceae) Nancy R. Morin New combinations in Limnanthes (Limnanthaceae) Nancy R. Morin New combinations in Micranthes (a segregate of Saxifraga, Saxifragaceae) in North America Luc BROUILLET AND RICHARD GORNALL es New | | new iesin the genus Spermacoce (Rubiaceae) for the Flora of Goiás and Tocantins (Brazil) and the Flora of the Guianas PIERO G. DELPRETE Hawthorns (Crataegus: Rosaceae) of the Cypress Hills, Alberta and Saskatchewan J.B. Puipps AND R.J. O'KENNON Thomas Walter Typification Project, IV: neotypes and epitypes for 43 Walter names, of genera A through C DanteL B. WARD ANATOMY AND MORPHOLOGY Cotyledon areoles in the Fabaceae subtribe Clitoriinae JAMES A. LACKEY FLORISTICS, ECOLOGY, AND CONSERVATION Rediscovery of Eleocharis kleinii (Cyperaceae), an overlooked species from the highlands of South Brazil R. Trevisan, D.J. ROSEN, AND I.I. BOLDRINI Distribution of Gamochaeta (Asteraceae: Gnaphalieae) in Texas, Oklahoma, Arkansas, and Louisiana Guy L. Nesom Flow diagrams for plant succession in the Middle Tennessee cedar glades J.M. BASKIN, E. QUARTERMAN, AND C.C. BASKIN 913 1005 1011 1015 1017 1019 1023 1031 1091 1101 1119 1125 1131 Observation on vegetation changes in Cajun Prairie, a coastal prairie flora in southwest Louisiana CHARLES ÁLLEN AND SARA THAMES Phytogeography of the Big Thicket, East Texas MicHAEL H. MACROBERTS AND BARBARA R. MACROBERTS A floristic survey of National Park Service areas of Timucuan Ecological and Historic Preserve (including Fort Caroline National Memorial), Duval County, Florida Wenny B. ZOMLEFER, DaviD E. GIANNASI, AND WALTER S. JUDD The vascular flora of the Chunky River (Mississippi) Lucas C. MAJURE New plant discoveries for Sonoran Islands, Gulf of California, Mexico BENJAMIN T. WILDER, RICHARD S. FELGER, HUMBERTO ROMERO-MORALES, AND ADRIAN QUIJADA-MASCAREÑAS Checklist of the vascular plants of Washington County, Pennsylvania CYNTHIA M. MORTON AND LOREE SPEEDY Saxifraga texana (Saxifragaceae) new to Louisiana CHRISTOPHER S. REID, PATRICIA L. FAULKNER, BARBARA R. MACROBERTS, AND MICHAEL H. MACROBERTS Vicia lathyroides (Fabaceae): new to the flora of Texas SONNIA HILL, RUTH LOPER, JASON R. SINGHURST, AND WALTER C. HOLMES Notes on the flora of Texas with additions and other significant records. III Larry E. Brown, Eric L. Kern, Davin J. ROSEN, AND JOE LIGGIO Notes on Texas orchids (Orchidaceae) PauL MARTIN BROWN Galium anglicum (Rubiaceae) new for Texas and notes on the taxonomy of the G. parisiense/divaricatum complex BARNEY L. LIPSCOMB AND Guy L. Nesom IN MEMORIAM James Hinton (1915-2006) GEORGE S. HINTON AND BILLIE L. TURNER Book Reviews AND Notices 826, 846, 852, 858, 870, 902, 932, 1018, 1148, 1156, 1228, 1250 Announcements 1281 1141 1149 1157 1179 1203 1229 1251 1253 1255 1265 1269 1277 INDEX to new names and new combinations in J. Bot. Res. Inst. Texas 1(2), 2007 Agapetes acuminata var. tipiensis Banik € Sanjappa, var. nov.—903 Agapetes flava var. nagensis Banik € Sanjappa, var. nov.—905 Agapetes megacarpa var. lohitensis Banik € Sanjappa, var. nov.—905 Agapetes odontocera var. mizoramensis Banik € Sanjappa, var. nov.—908 Agapetes salicifolia var. glanduliflora Banik € Sanjappa, var. nov.—910 Alocasia nycteris Medecilo, Yao & Madulid, sp. nov.—817 Aphanelytrum peruvianum Sánchez Vega, PM. Peterson, Soreng & Leegaard, sp. nov.—842 Boerhavia triquetra S. Wats. var. intermedia (M.E. Jones) Spellenb., comb. nov.—874 Chusquea annagardneriae L.G. Clark, C.D. Tyrrell, Triplett & A.E. Fisher, sp. nov.—847 Crataegus aquacervensis J.B. Phipps & O'Kennon, sp. nov.—1061 Crataegus brazoria var. viburnifolia (Sarg.) J.B. Phipps, comb. nov.—1007 Crataegus chrysocarpa var. blanchardii (Sarg.) J.B. Phipps, comb. et stat. nov.—1009 Crataegus chrysocarpa var. praecox (Sarg.) J.B. Phipps, comb. et stat. nov.—1009 Crataegus chrysocarpa var. vigintistamina J.B. Phipps, var. nov.—1008 Crataegus coccinea var. pringlei (Sarg.) J.A. Macklin & J.B. Phipps, comb. et stat. nov.—1005 Crataegus cupressocollina J.B. Phipps & O'Kennon, sp. nov.—1056 Crataegus series Cupressocollinae J.B. Phipps & O'Kennon, ser. nov.—1056 Crataegus series Montaninsulae J.B. Phipps & O'Kennon, ser. nov.—1065 Crataegus pruinosa var. dissona (Sarg.) J.B. Phipps, comb. et stat. nov.—1007 Crataegus purpurella J.B. Phipps & O'Kennon, sp. nov.—1066 Crataegus reverchonii var. mohrii (Beadle) J.B. Phipps, comb. et stat. nov.—1006 Crataegus rivuloadamensis J.B. Phipps & O'Kennon, sp. nov.—1065 Crataegus rivulopugnensis J.B. Phipps & O'Kennon, sp. nov.—1070 Crataegus rubribracteolata J.B. Phipps & O'Kennon, sp. nov.—1073 Crataegus sheila-phippsiae var. saskatchewanensis J.B. Phipps & O'Kennon, var. nov.—1079 Crataegus ursopedensis J.B.Phipps & O'Kennon, sp. nov.—1081 Crataegus viridis var. velutina (Sarg.) J.B. Phipps, comb. et stat. nov.—1009 Cyclopogon secundum Christenson, sp. nov.—1001 Dudleya abramsii subsp. costifolia (Bartel & Shevock) Moran, comb. nov.—1015 Dudleya abramsii subsp. setchellii (Jeps.) Moran, comb. nov.—1015 Eleocharis acutangula subsp. breviseta D.J. Rosen, subsp. nov.—885 Eleocharis acutangula subsp. neotropica D.J. Rosen, subsp. nov.—886 Erigeron heleniae Nesom, sp. nov.—891 Festuca xgonzalez-ledesmae S.J. Darbyshire, nothosp. nov.—832 Festuca longiligula S.J. Darbyshire, sp. nov.—827 Festuca xmiscella S.J. Darbyshire, nothosp. nov.—835 Limnanthes alba subsp. gracilis (Howell) Morin, comb. et stat. nov.—1017 Limnanthes alba subsp. parishii (Jeps.) Morin, comb. et stat. nov.—1017 Limnanthes douglasii subsp. striata (Jeps.) Morin, comb. et stat. nov.—1017 Lobelia gloria-montis subsp. longibracteata (Rock) Lammers, stat. nov.—806 Lobelia xkauaiensis (A. Gray) A. Heller, stat. nov.—801 Lobelia koolauensis (Hosaka & Fosberg) Lammers, comb. et stat. nov.—803 Lobelia wahiawa Lammers, sp. nov.—802 Mandevilla aequatorialis J.F Morales, sp. nov.—853 Mandevilla frigida J.E Morales, sp. nov.—855 Mandevilla horrida J.E Morales, sp. nov.—863 Mandevilla inexperata J.F Morales, sp. nov.—865 Mandevilla lojana J.E Morales, sp. nov.—855 Mandevilla megabracteata J.E Morales, sp. nov.—866 Mandevilla similaris J.E Morales, sp. nov.—867 Micranthes bryophora (A. Gray) Brouillet & Gornall, comb. nov.—1020 Micranthes calycina (Sternb.) Gornall & H. Ohba, comb. nov.—1020 Micranthes ferruginea (Graham) Brouillet & Gornall, comb. nov.—1020 Micranthes foliolosa (R. Br.) Gornall, comb. nov.—1020 Micranthes gormanii (Suksd.) Brouillet & Gornall, comb. nov.—1020 Micranthes idahoensis (Piper) Brouillet & Gornall, comb. nov.—1020 Micranthes mexicana (Engl. & Irmsch.) Brouillet & Gornall, comb. nov.—1020 Micranthes nelsoniana var. aestivalis (Fisch. & C.A. Mey.) Gornall & H. Ohba, comb. et stat. nov.—1020 Micranthes nelsoniana var. carlottae (Calder & Savile) Gornall & H. Ohba, comb. et stat. nov.—1020 Micranthes nelsoniana var. cascadensis (Calder & Savile) Gornall & H. Ohba, comb. et stat. nov.—1020 Micranthes nelsoniana var. insularis (Hultén) Gornall & H. Ohba, comb. et stat. nov.—1020 Micranthes nelsoniana var. pacifica (Hultén) Gornall & H. Ohba, comb. et stat. nov.—1020 Micranthes nudicaulis (D. Don) Gornall & H. Ohba, comb. nov.—1021 Micranthes petiolaris (Raf.) Brouillet & Gornall, comb. nov.—1021 Micranthes razshivinii (P. Yu. Zhmylev) Brouillet & Gornall, comb. nov.—1021 Micranthes tempestiva (Elvander & Denton) Brouillet & Gornall, comb. nov.—1021 Micranthes tischii (Skelly) Brouillet & Gornall, comb. nov.—1021 Micranthes tolmiei (Torrey & A. Gray) Brouillet & Gornall, comb. nov.—1022 Micranthes unalaschcensis (Sternb.) Gornall & H. Ohba, comb. nov.—1022 Mosiera androsiana (Urban) Salywon, comb. nov.—899 Mosiera cuspidata (Alain) Salywon, comb. nov.—899 Mosiera gracilipes (Alain) Salywon, comb. nov.—899 Mosiera xerophytica (Britton) Salywon, comb. nov.—900 Passiflora arta Feuillet, sp. nov.—819 Passiflora compar Feuillet, sp. nov.—821 Passiflora davidii Feuillet, sp. nov.—896 Philadelphus microphyllus var. argyrocalyx (Wooton) Henrickson, comb. & stat. nov.—901 Philadelphus microphyllus var. madrensis (Hemsley) Henrickson, comb. & stat. nov.—901 Philadelphus microphyllus var. stramineus (Rydb.) Henrickson, comb. & stat. nov.—901 Potentilla arenosa subsp. chamissonis (Hultén) Elven & D.E Murray, comb. nov.—813 Potentilla holmgrenii D.E Murray & Elven, sp. nov. —811 Potentilla hyparctica Malte subsp. elatior (Abrom.) Elven & D.E Murray, comb. nov.—813 Ribes oxyacanthoides var. cognatum (Greene) Morin, comb. et stat. nov.—1015 Sedum lanceolatum var. subalpinum (Fród.) H. Ohba, comb. & stat. nov.—890 Sedum laxum var. eastwoodiae (Britton) H. Ohba, comb. & stat. nov.—889 Sedum laxum var. flavidum (Denton) H. Ohba, comb. & stat. nov.—889 Sedum laxum var. heckneri (M. Peck) H. Ohba, comb. & stat. nov.—889 Sedum laxum var. latifolium (R.T. Clausen) H. Ohba, comb. & stat. nov.—890 Sedum obtusatum var. boreale (R.T. Clausen) H. Ohba, comb. & stat. nov.—889 Sedum obtusatum var. paradisum (Denton) H. Ohba, comb. & stat. nov.—889 Sedum obtusatum var. retusum (Rose) H. Ohba, comb. & stat. nov.—889 Sedum oreganum var. tenue (R.T. Clausen) H. Ohba, comb. & stat. nov.—890 Sedum radiatum var. ciliosum (Howell) H. Ohba, comb. & stat. nov.—889 Sedum radiatum var. depauperatum (R.T. Clausen) H. Ohba, comb. & stat. nov.—889 Sedum stenopetalum var. monanthum (Suksd.) H. Ohba, comb. & stat. nov.—890 Sedum wrightii var. densiflorum (R.T. Clausen) H. Ohba, comb. & stat. nov.—889 Sedum wrightii var. priscum (R.T. Clausen) H. Ohba, comb. & stat. nov.—889 Spermacoce burchellii (E.L. Cabral & Bacigalupo) Delprete, comb. nov.—1025 Spermacoce crispata (K. Schum.) Delprete, comb. nov.—1025 Spermacoce delicatula (E.L. Cabral) Delprete, comb. nov.—1025 Spermacoce dimorpha (J.H. Kirkbr.) Delprete, comb. nov.—1025 Spermacoce incognita (E.L. Cabral) Delprete, comb. nov.—1025 Spermacoce irwiniana (E.L. Cabral), Delprete, comb. nov.—1025 Spermacoce multiflora (DC.) Delprete, comb. nov.—1026 Spermacoce paraensis (E.L. Cabral & Bacigalupo) Delprete, comb. nov.—1027 Spermacoce pulchristipula (Bremek.) Delprete, comb. nov.—1027 Spermacoce semiamplexicale (E.L. Cabral) Delprete, comb. nov.—1028 Spermacoce spicata (Miq.) Delprete, comb. nov.—1028 Spermacoce tocantinsiana (E.L. Cabral € Bacigalupo) Delprete, comb. nov. 1028 REVISION OF LOBELIA SECT. GALEATEELA (CAMPANULACEAE: LOBELIOIDEAE) Thomas G. Lammers dolia of Biology and eee University of Wisconsin Osh E Wisconsin 54901, po ABSTRACT Pew: 1 T: 1 Lobelia sect. Galeatella is a group of pachycaul treelets distinguished by a solitary pliestesial | stock; a dense apical rosette of sessile coriaceous leaves leaving a helical pattern of raised ee cicatrices on the sida stem; a on. or d MR short- pet teat f nie rnithophilous flowers on o let pedicels; calyx hli hic lo! hed l d horizontally about as n as a stout ‘ube to 3 times d the ventral connate for 7 their rM or more, e, forming a ilaa inrolled trifid lip ee th rted, all five anthers with RUN HN Rd niu E L 1 à 1 1 [xe 1! 11 2 s 21 0] ue Hawaiian Islands, with two pn each on Kauai i. villosa D L. wahiawa, sp. nov.) and Oʻahu (E; al and L. koolauensis, comb. et stat. nov.), an g L. gloria-montis) on Maui and Molokai, divided into two subspecies: subsp. gloria-montis (West Maui and Moloka‘i) and bus longibracte ata stat. nov. (West Maui and East Maui). The two species on Kauai hybridize, producing L. xkauaiensis, stat. n RESUMEN : “J T2211 : 1 A E 4 1; A qus 4 : Lobelia sect. Galeatella es un grup paq I g p vástago solitari p pico qu J > J a Tuo s pe 1 A 1 a s i pia N 3 1 1 o dE J É J E T TT s o 1 5) 4 T J 1 J pa Ti A | 1 SES ii ; Į g Į ebracteolados; lóbulos del cáliz cad el fruto joven; corola no azul con lóbulos lineares monomórficos, el par dorsal extendido o recurvado pM dr tan d como el tubo rígido s tres veces más cd el n e en ’/, de su nud o más, pu un labi j q p p A 5 t ES RATE 1 1 Tae 1 JA] 1 bl 1 l l l d b la madure pol y semillas oblongas alad testa débil te estriada. La ión ndémica de las Islas ns con dos especies en ds una de las Kauai (L. villosa y L. wahiawa, sp. nov) y O'ahu (L. gaudichaudii y L. koolauensis, comb. et stat. nov.), y una sola especie (L. gloria-montis) en Maui y Molokai, dividida en dos subespecies: subsp. gloria-montis (West Maui and HS y subsp. longibracteata, stat. nov. (West Maui y East Maui). Las d pecies de Kauai hibridan produciendo L. xkauaiensis, stat. nov. The finest of our lobelias is Lobelia gloria-montis, a truly royal and superb plant ... To see this species growing is one of the delights of the botanist.—Rock (1919: 35, 117) The Hawaiian archipelago is renowned among botanists for the many compositional peculiarities of its flora (MacCaughey 1917; Hitchcock 1919; Carlquist 1974, 1980; Wagner et al. 1990). Among the most peculiar is the fact that the family with the greatest number of native species is not Gramineae Juss. or Compositae Giseke, but Campanulaceae Juss.; at no other spot on the globe does this family predominate statistically. In a flora of not quite 1000 indigenous species (Wagner et al. 1990), 134 of them (all endemic) belong to Campanulaceae (Lammers 20072). Furthermore, these 134 species represent nearly 696 of all species in this cosmopolitan family, although the Hawaiian Islands only comprise 0.00496 of Earth's land surface. Six genera are present, all members of subfamily Lobelioideae Burnett. Five are endemic and assigned to the tribe Delisseeae C. Presl: Brighamia A. Gray, Clermontia Gaudich., Cyanea Gaudich., Delissea Gaudich., and Trematolobelia Zahlbr. ex Rock. The only one of the six not endemic to the archipelago is Lobelia L., a member of tribe Lobelieae Rchb., which is virtually cosmopolitan in its distribution. Lobelia is the largest of the 29 genera in the Lobelioideae, comprising 405 species of annual and peren- nial herbs, shrubs, and trees (Lammers 20072). The genus is characterized by solitary or racemose flowers on usually bibracteolate pedicels; corollas with the tube dorsally cleft to the base and the lobes monomorphic, J. Bot. Res. Inst. Texas 1(2): 789 — 810. 2007 790 Journal of the Botanical R h Institute of Texas 1(2) docs or dimorphic and the ventral larger; and apically capsules or rarely berries (Murata 1995; Lammers 2004a, 2007b). Because of its large number of species, Lobelia has been divided into a number of sections and other infrageneric taxa over the years. The first species described from the Hawaiian Islands, L. gaudichaudii, was assigned to sect. Homochilus A. DC. by Candolle (1839). As additional Hawaiian species of Lobelia were discovered during the Nineteenth Century, they all were assumed to be related to L. gaudichaudii. Bentham (1876) removed these Hawaiian species from sect. Homochilus to sect. Rhynchopetalum (Fresen.) Benth. (cf., Lammers 20045); the latter subsequently was subsumed into sect. Tylomium (C. Presl) Benth. by Schonland (1889). In contrast to this single-taxon view of Hawaiian Lobelia, Rock (1919) believed that the Hawaiian species of Lobelia actually formed two taxa that were not each other's closest relatives: (1) L. gaudichaudii and allied species, characterized by a large white, yellow, red, or purple-striped corolla with stout tube and spreading or horizontally recurved dorsal lobes, and all five anthers apically tufted; and (2) L. yuccoides Hillebr. and allied species, characterized by a small blue or magenta corolla with slender tube and recoiled dorsal lobes, and only the ventral pair of anthers apically tufted. This two-taxon hypothesis supported by Skottsberg (1928) and Carlquist (1980), and formalized by Wimmer (1948), who erected two sections within Lobelia subg. Tupa (G. Don) E. Wimm. to accomodate the Hawaiian endemics: sect. Galeatella E. Wimm. for L. gaudichaudii and its allies, and sect. Revolutella E. Wimm. for L. yuccoides Hillebr. and its allies (Wimmer 1953, 1968). Although Mabberley (1974, 1975) suggested returning both groups to different parts of sect. Rhynchopetalum, he did not effect these changes and Wimmer's arrangement is the classification in current use (Lammers 1990, 20072, 2007b; Murata 1995). Over the past 15 years, the summary of Hawaiian Campanulaceae that I prepared (Lammers 1990) for the Manual of the flowering plants of Hawai'i (Wagner et al. 1990) has been expanded into a series of monographs. Previous entries covered Brighamia (Lammers 1989), Clermontia (Lammers 1991), and Delissea (Lammers 2005). The present effort covers Lobelia sect. Galeatella. This treatment will also form part of a series of monographs of the sections of Lobelia subg. Tupa; previous entries in that series accounted for sect. Tupa (G. Don) Benth. (Lammers 2000) and sect. Homochilus (Lammers 2004b). TAXONOMIC HISTORY As might be expected from their predominance in the Hawaiian flora, Lobelioideae were among the first botanical specimens collected in the islands at the time of their discovery by Europeans in 1778 (St. John 19772, 197 Tb, 1979). However, it was another half century before botanists visiting the archipelago obtained specimens referable to Lobelia sect. Galeatella. During an October 1836 visit to O'ahu by the French corvette La Bonite (cf., Lasegue 1845; Kuykendall 1947), ship's naturalist Charles Gaudichaud-Beaupré reached the summit of the Ko'olau Range above Honolulu harbor (St. John & Titcomb 1983). Here he collected a pachy- caul species of Lobelia a meter tall with a thick pithy stem densely covered in transversely rhombic leaf scars, a dense apical rosette of coriaceous leaves, a terminal raceme of large red flowers, and capsules filled with flat winged seeds. Not long after, Swiss botanist Alphonse de Candolle saw the specimen in the herbarium at Paris and named it in Gaudichaud's honor (Candolle 1839). Although Gaudichaud never published a description of Lobelia gaudichaudii himself, the atlas of La Bonite's voyage (Gaudichaud s.d.) did include an illustration, labeled with Candolle's binomial; the fascicle of the book containing this plate is believed to have appeared in 1842 (Stafleu & Cowan 1976). The United States Exploring Expedition (cf., Wilkes 1845; Jackson 1985) called at the Hawaiian Islands during 1840-41 and collected specimens of Lobelia on Kauai that were very similar to L. gaudichaudii of O'ahu. Harvard botanist Asa Gray (cf., Dupree 1959; St. John 1985) segregated these plants from the species as L. gaudichaudii var. kauaiensis (Gray 1861; Mann 1867). However, Austrian botanist Heinrich Wawra, who visited the archipelago aboard the frigate Donau during 1869—70, did not distinguish the specimens he had gathered on O'ahu and Kauai, calling them all L. gaudichaudii (Wawra 1873). This single-taxon approach Lammers, Revision of Lobelia sect. Galeatella 791 was also adopted by the islands' first resident botanist, Wilhelm Hillebrand (cf., Degener 1957; Lammers 1994), who broadened the ci iption of the species even further by including plants he had discovered on West Maui (Hillebrand 1888). American Amos Arthur Heller, who botanized on O'ahu and Kauai in 1895 (cf., Wagner & Shannon 1999), not only disagreed with this single-taxon view, but further disagreed with their original treatment as conspecific varieties. He segregated the Kauai populations from L. gaudichaudii as L. kauaiensis, commenting (Heller 1897: 911), “Had Dr. Gray seen [plants of O'ahu and Kauai] in the living state, he certainly would not have considered the one a mere form of the other, worthy of varietal rank only." The next botanist to deal with these plants was Austrian expatriate Joseph Rock, who resided in the islands from 1907 to 1920 and again from 1953 until his death in 1962 (cf., Chock 1963; Sutton 1974). At first, he accepted Heller's two-species treatment, but modified it slightly by describing three new varieties. Thinking that the white-flowered plants on West Maui were the original nomenclatural type of L. gaudi- chaudii, he segregated the red-flowered O'ahu populations as L. gaudichaudii var. coccinea (Rock 1917); only later did he realize that this was an error and that the type of the species was actually a red-flowered plant from O'ahu. Tall plants with very long floral bracts and calyx lobes that he discovered on West Maui were distinguished as L. gaudichaudii var. longibracteata (Rock 1913a, 1913b), while plants with densely pubes- cent inflorescences that he discovered on Kauai were segregated as L. kauaiensis var. villosa (Rock 1917). However, in his subsequent monograph of the Hawaiian Lobelioideae (Rock 1919), he treated the plants of each island as a discrete species: L. kauaiensis and its var. villosa on Kauai, L. gaudichaudii (with var. coccinea in synonymy) on O'ahu; and the newly described L. gloria-montis and its var. longibracteata on West Maui. When plants referable to sect. Galeatella were found on Molokai, Degener (19382) named them L. glo- ria-montis var. molokaiensis, though a paucity of good material precluded an adequate description. Similarly, when white-flowered plants with a branched inflorescence was discovered in the northwestern Ko'olau Range, Fosberg and Hosaka (1938) described them as a variety of the O'ahu species, L. gaudichaudii var. koolauensis. The first dedicated taxonomic study of these plants was by University of Hawaii professor Harold St. John (cf., Anonymous 1980, 1991) and his student Edward Hosaka. Their revision (St. John & Hosaka 1938) resurrected Hillebrand's (1888) broad circumscription of L. gaudichaudii, including in it plants of O'ahu, Kauai, and West Maui. Within this broad circumscription, they distinguished five geographic varieties: var. gaudichaudii (as *var. typica" and including var. coccinea as a synonym), var. gloria-montis (with three heterotypic formae), var. kauaiensis (with one heterotypic forma), var. koolauensis, and var. longibracteata; var. molokaiensis was excluded for want of adequate material. However, not all Kauai plants were included in L. gaudichaudii var. kauaiensis; those recognized previously as L. kauaiensis var. villosa were accorded specific rank as L. villosa. Years later, a population discovered on East Maui was described (St. John & Medeiros 1987) as a sixth geographical variety, L. gaudichaudii var. albiflora. When Viennese botanist Franz Elfried Wimmer (cf., Rechinger 1961; Degener & Degener 1962a) prepared his monograph of Lobelia for Das Pflanzenreich (Wimmer 1953), his treatment of sect. Galeatella was identical to that of St. John and Hosaka (1938). In subsequent years, however, his view of the group changed. In a letter to Otto Degener dated 20 November 1958 (quoted by Degener & Degener 1974), he indicated his intention to accord the group generic status under the name “Schiexelia,” and to recognize five species therein. His posthumously published supplement to the Pflanzenreich monograph (Wimmer 1968) did recognize five species: L. kauaiensis and L. villosa on Kauai, L. gaudichaudii on O'ahu; and L. gloria-montis and L. longibracteata (newly elevated from varietal rank) on West Maui. However, no mention was made of elevating the section to generic rank and it remained a section of Lobelia subg. Tupa. Degener and Degener (1962b, 1974, 1983) carried through on the elevation to generic rank, though they disregarded Wimmer's provisional name and took up the sectional epithet at the new rank. Their synopsis of the group (Degener & Degener 1974) maintained Wimmer's (1968) five-species classification. My treatment in the Manual (Lam- mers 1990) was essentially the same, except for including L. longibracteata within L. gloria-montis. 792 Journal of the Botanical R h Institute of Texas 1(2) MATERIALS AND METHODS Revision of the classification of Lobelia sect. Galeatella was based upon morphological data; definitions of qualitative character states follow Harris and Harris (1994). These data were gathered from almost 300 specimens (including the types of all heterotypic names) deposited in 31 herbaria (see Acknowledgments for a complete list). The data obtained from these specimens were analyzed via traditional taxonomic methodology (Leen- houts 1968; Qualls 1986; Vogel 1987; Maxted 1992; Watson 1997; Winston 1999) within a geographic framework. Populations referable to sect. Galeatella are known from just six discrete areas on four islands: (1) the Wahiawa drainage of south-central Kauai at 570-800 m; (2) the Alaka‘i Swamp of north-central Kauai, from Pihea to Wai'aleale, at 1100-1590 m; (3) the Ko'olau Range of O'ahu, from Pu'u Ka'inapuaa to Mt. Olympus, at 670-960 m; (4) the mountains of West Maui at 915-1760 m; (5) the northwestern slopes of Haleakala on East Maui at 1280—1700 m; and (6) eastern Moloka'i from Papa‘ala to Kalua'aha at 1165-1350 m. Because of the extreme precinctiveness of the Hawaiian flora (MacCaughey 1917; Hitchcock 1919; Carlquist 1980; Wagner et al. 1990), I hypothesized that each of these six areas would harbor a single taxon unique to that locale, or at least to that island. Therefore, when inspecting the data, I asked two questions. First, were all specimens from a given area relatively homogeneous, with most characters evincing a continuous pattern of variation? If so, the hypothesis would be supported; if instead, several characters consistently showed correlated gaps in their patterns of variation within an area, the hypothesis would be refuted. Second, are the plants that evince a given correlated suite of morphological features restricted to a single area or island? If so, the hypothesis would be supported; if not, the hypothesis would be refuted. Once taxa had been discerned in this fashion, they were compared to nomenclatural type specimens to determine the correct name under the International Code of Botanical Nomenclature (ICBN; McNeill et al. 2006). Decisions on rank for the taxa were made in light of the definitions of species and subspecies that I have employed previously (Lammers 1991, 20073). RESULTS AND DISCUSSION In revising Lobelia sect. Galeatella, three aspects of its classification were evaluated in light of the assembled data: (1) the circumscription of the group, (2) its position and rank, and (3) its division into component taxa. Circumscription.—The circumscription and characterization of Lobelia sect. Galeatella have been invari- ant since its publication by Wimmer (1948). Overall similarity among populations is quite high; recall that the entire section has been treated as just one or two species until recently, with most variation expressed by distinguishing geographic varieties. Furthermore, much of this overall similarity involves characters that are quite unusual among Lobelioideae. Most species of Lobelia are hemicryptophytes or therophytes (Lammers 20072); woodiness is less com- mon, confined to certain groups within subg. Tupa (Carlquist 1969; Murata 1995). Even among those species that are woody, the growth form of sect. Galeatella is unusual. In sect. Tylomium (Wilbur 1991), sect. Tupa (Lammers 2000), and sect. Homochilus (Lammers 2004b), the woody species are iteroparous (polycarpic) shrubs with several leptocaul stems that persist, branch, and flower each growing season. In sect. Galeatella, in contrast, the stem is pachycaul, unbranched, pliestesial (i.e., requiring several to many years of growth before it achieves sufficient mass to flower), and hapaxanthic (i.e., dying after flowering). Although Truman and Augspurger (1991) described these species as semelparous (monocarpic), this is incorrect. An indi- vidual shoot does indeed die by the time the capsules dehisce and disperse ripe seed. However, a new shoot emerges around the base of the plant (presumably from a root bud) to replace it. This sucker, which is far more pubescent than a mature shoot (cf., the “juvenile plant” of Degener 19382), will eventually grow into a new flowering shoot (Hillebrand 1888; pers. observ.). As such, the individual plant is in fact iteroparous (polycarpic), conforming to the *Tomlinson" architectural model (cf., Hallé & Oldeman 1970; Hallé et al. Lammers, Revision of Lobelia sect. Galeatella 793 Similarly, the winged seeds of sect. Galeatella (Fig. 1) are unusual in Lobelia, occurring elsewhere only in some species of sect. Rhynchopetalum and sect. Trimeris (C. Presl) A. DC. [the earliest available name for sect. Colensoa (Hook. f.) J. Murata] (Mabberley 1975; Murata 1995; Buss et al. 2001). The possession of apical tufts of trichomes on all five anthers rather than just the ventral pair is otherwise unknown in subg. Tupa, though it does characterize sect. Cryptostemon (E. Wimm.) J. Murata and sect. Delostemon (E. Wimm.) J. Murata in herbaceous subg. Lobelia (Murata 1995). In summary, the assemblage of morphological features that characterize these Hawaiian species is found in no other species of Lobelioideae and the circumscription adopted here is confidently believed to create a natural group. Position and rank.—In contrast to circumscription, there has been a diversity of opinion regarding the relationships of Lobelia sect. Galeatella to other taxa of Lobelioideae and the appropriate rank at which to recognize the taxon. Most authors who have commented on relationships agree in identifying as the group's closest relatives certain species assigned to another section in Lobelia subg. Tupa, sect. Trimeris. However, they disagree on specifics. Rock (1919) allied the group to tropical Asian members of the group, such as L. boninensis Koidz. and L. pyramidalis Wall., while Mabberley (1974, 1975) saw a relationship to Brazilian species such as L. hassleri Zahlbr. and L. organensis Gardner. The endemic Hawaiian genus Trematolobelia has also been suggested as a close relative by some (Candolle 1839; Stapf 1893; Mabberley 1975). As for rank, the group has been treated as either a section within Lobelia subg. Tupa (Wimmer 1948, 1953, 1968; Lammers 1990, 2007b; Murata 1995) or as a genus distinct from Lobelia (Degener & Degener 1962b, 1974, 1983). Because of the geological youth and de novo volcanic origin of the Hawaiian Islands, it is assumed that its flora developed from continental species that in some way were able to colonize the archipelago via long-distance dispersal (Guppy 1906; Brown 1921; Carlquist 1974; Funk & Wagner 1995). For many years, it was hypothesized that multiple colonizations were required to explain the 134 endemic species of Lobelioideae: a common one for Clermontia, Cyanea, and Delissea, and separate ones for each of the remain- ing taxa, including Lobelia sect. Galeatella (Rock 1919; Stone 1967; Carlquist 1980; Mabberley 1974, 1975; Lammers 1990). However, recent analyses of molecular data (Knox et al. 1993; Givnish et al. 1995; E. Knox, Indiana University, pers. comm.; A. Antonelli, Góteborg University, pers. comm.) have dealt a devastating blow to this polyphyletic hypothesis. All these studies indicate that the endemic Hawaiian Lobelioideae are mono- phyletic, i.e., that all 134 species in six genera evolved from a single ancestral colonization of the archipelago. The Hawaiian species (as well as Apetahia Baill. and Sclerotheca A. DC., genera endemic to islands of the southwestern Pacific) form a well supported clade in these analyses. Its sister-group is a clade comprising the African and Brazilian species of sect. Rhynchopetalum and sect. Trimeris, and the outgroup of this larger Hawaiian-African-Brazilian clade is a group of tropical Asian species of sect. Trimeris. Consequently, the species that Rock (1919) and Mabberley (1974, 1975) believed were the immediate ancestors of sect. Galeatella are more distantly related, and the groups closest relatives actually are to be found among the Hawaiian endemics. The molecular analyses cited above indicate that the Hawaiian clade comprises two subordinate clades: one comprising the genera with axillary inflorescences (Brighamia, Clermontia, Cyanea, Delissea) or solitary axillary flowers (Apetahia, Sclerotheca) and the other comprising the taxa with terminal inflorescences (Lo- belia sect. Galeatella, Lobelia sect. Revolutella, and Trematolobelia). Within the latter subclade, sect. Galeatella is sister to Trematolobelia while sect. Revolutella is their outgroup. The sister-group relationship between sect. Galeatella andTrematolobelia is supported by morphology. Overall, the two are very similar in structure and appearance. Both are pachycaul treelets with a solitary pliestesial hapaxanthic shoot; a dense apical rosette of coriaceous leaves leaving a helical pattern of raised corky cicatrices on the unbranched stem; a terminal bracteate raceme of large ornithophilous flowers; calyx lobes caducous from the young fruit; non-blue corolla with linear monomorphic lobes; and exserted staminal 794 Journal of the Botanical R h Institute of Texas 1(2) Is of Lobeli t. Galeatella prey 1 following B t al. (2001). Scale b 625 um. A. L villosa (Flynn 685, PTBG). B. L. xkauaiensis (Fost 14224, K). E. 1 lichaudii (Gart Fic. 1. S g (Hitchcock 15437, US). C. L. wahiawa (Forbes 294, K, US). D. L. kool j subsp. gloria-montis (Hitchcock 14735, US). ( ( EL gloria-montis column. Both groups also have relatively large winged seeds and an irregular network of sclerenchymatous bundles in the pericarp, character states unusual among Lobelioideae, which may serve as synapomorphies uniting them. In attempting to distinguish Lobelia sect. Galeatella from Trematolobelia, 1 could identify just five morpho- logical characters that were consistently useful. Using the horizontal posture of the inflorescence branches as a distinguishing character for Trematolobelia is compromised by the erect unbranched inflorescence of T. singularis H. St. John; cf., Carlquist 1980, pg. 212D [top]. Similarly, use of corolla color is precluded by overlap; red and white flowers are found in species of both groups. Lammers, Revision of Lobelia sect. Galeatella 795 Although both taxa are characterized by a woody stem that is pachycaul, unbranched, pliestesial, and hapaxanthic, there is a fundamental difference in longevity. While individual plants of sect. Galeatella are iteroparous (polycarpic) via basal offshoots (the *Tomlinson" architectural model), individuals of Trema- tolobelia are semelparous (monocarpic), dying by the time the capsules dehisce and disperse ripe seed (the ^Holttum" architectural model; Hallé & Oldeman 1970; Hallé et al. 1978). The latter plan may have evolved from the former through the simple suppression of the basal offshoots. Lobelia sect. Galeatella further differs from Trematolobelia, and is unusual among species of Lobelia subg. Tupa, in having ebracteolate (vs. bibracteolate) pedicels and all five anthers (vs. just the ventral pair) bearded at the apex with tufts of stiff white trichomes; these two characters are potential synapomorphies for the species of sect. Galeatella. Conversely, Trematolobelia differs from sect. Galetella and is unique among Lobelioideae in its strongly secund (vs. ample) inflorescences and laterally poricidal (vs. apically loculicidal) capsule; these characters are potential synapomorphies for the species of Trematolobelia. Although the fruit of Trematolobelia (described in detail by Carlquist 1962) is unique in the subfamily, there is some similarity to the fruit of sect. Galeatella, as noted by Hillebrand (1888) and Stapf (1893). In all Lobelioideae, the ovary is adnate to a hypanthium of appendicular origin (Kaplan 1967). As a result, the pericarp of the fruit is actually a combination of hypanthial and gynoecial tissue. In both Trematolobelia and sect. Galeatella, the calyx lobes are caducous early on; the exocarp and the parenchymatous portions of the mesocarp deliquesce and are washed away by the frequent rains, leaving behind a series of sclerenchymatous bundles surrounding the endocarp. In sect. Galeatella, this mesocarpal network is open distally, forming an open basket or cup. The endocarp it loosely encloses is thick and woody, and dehisces apically through a pair of triangular loculicidal valves. In Trematolobelia in contrast, this mesocarpal network t distally, forming a laterally f i sphere. The endocarp is tightly enclosed and adnate to the sclerenchymatous mesocarp at its summit, precluding the formation of dehiscent valves. Instead, the wall is thin and papery, rupturing irregularly at its base. This releases the seeds, which accumulate in the bottom of the mesocarpal network. When the plant is flailed by the strong winds characteristic of its montane habitat, the winged seeds are shaken out through the lateral foramina in the sclerenchymatous mesocarp. These similarities suggest that the fruit of sect. Galeatella represents an intermediate step in the evo- lution of the unique poricidal capsule of Trematolobelia from the loculicidal capsules with intact exocarp and mesocarp that are more typical of the Lobelioideae. All of the necessary components are in place: the caducous calyx lobes, the mesocarpal bundles, the deliquescent exocarp and mesocarp parenchyma, and the winged seeds. All that was required was (1) the apical closure and contraction of the mesocarpal network, (2) cessation of apical dehiscence via adnation of the endocarp to the mesocarpal sclerenchyma, and (3) the thinning of the endocarp. In light of the well supported phylogenetic relationship between sect. Galeatella and Trematolobelia, the current classification at these ranks appears suboptimal. From a cladistic perspective, sister-groups should have the same rank. This could be achieved in the present case by elevating sect. Galeatella to generic rank, following Degener and Degener (1962b, 1974, 1983); by including Trematolobelia in Lobelia as another sec- tion, following Post and Kuntze (1903); or by expanding Trematolobelia to encompass sect. Galeatella. However, I have elected to do none of these at present but rather to maintain the status quo and con- tinue pro tempore to treat this taxon as a section within Lobelia subg. Tupa. First, it seems best to complete projected studies of sect. Revolutella and Trematolobelia before making changes in the position and rank of sect. Galeatella. These three taxa are intimately connected and any necessary changes would best be made in concert. Second, the molecular analyses cited above raise issues that extend beyond far beyond the Hawai- ian Islands. The sampled species of Lobelia in these studies fall throughout the phylogeny of the subfamily, from the very first branch to the last; all other genera sampled thus far are embedded among them. Thus, as currently circumscribed, Lobelia is one of the most unnatural genera imaginable. It not only violates key principles of cladistic classification, but of traditional taxonomy as well. If we are to avoid treating the entire subfamily as a single genus comprising 1192 species (Lammers 2007a), it will be necessary to segregate 796 Journal of the Botanical R h Institute of Texas 1(2) a number of genera from Lobelia. However, the studies cited above have not yet reached the point where itis feasible to do so. Although the overall problem is obvious, the best resolution is not yet evident. Any changes in the position and rank of the Hawaiian taxa of Lobelia would best be done as part of the overall realignment of the genus. Component taxa.—My original hypothesis, that each of the six areas in which sect. Galeatella occurs would possess a taxon unique to that locale, was not supported. Analyses of the data revealed a total of seven taxa. Half the sites harbored two or even three taxa, and only four of the seven taxa were confined to a single site. On the other hand, all taxa but one were confined to a single island, and even that one exception may prove to be spurious once additional specimens have been studied. The specimens examined from the Alaka‘i Swamp of north-central Kauai formed three groups on the basis of morphology. The most widely distributed, occurring throughout the length of the region from Pihea to Wai'aleale, is also the most distinctive member of the entire section, differing from all others by its densely pubescent inflorescence on a conspicuously bracteate peduncle, pale yellow flowers that are the smallest in the section, floral bracts rounded or subcordate at base, and calyx lobes rounded or obtuse at apex. The type of L. kauaiensis var. villosa is a representative of this taxon. At the southeastern terminus of the Alakai Swamp, on the summit of Wai'aleale, are found plants that differ considerably from the last. This taxon is characterized by depressed obtrullate or depressed obovate cicatrices on the stems, linear or oblanceolate leaves, a tall (1.5—2.5 m) glabrous branched inflorescence with blackish purple axes and a largely naked peduncle, relatively narrow floral bracts, relatively large flowers, calyx lobes acute or acuminate at apex, a pale purple corolla with dark purple longitudinal veins, and large (1.7-2.5 mm long) seeds. This glabrous taxon corresponds to St. John and Hosaka's (1938) concept of L. gaudichaudii var. kauaiensis; however, as pointed out by Skottsberg (1944), it does not match the type of that name: the inflorescence in the type specimen is somewhat pubescent, while the apex of the calyx lobes is rounded. In fact, none of the type specimens referable to sect. Galeatella match these plants, meaning that none of the available names apply to the taxon; though known for well over a century, it has no name. The third taxon is likewise found on Wai'aleale, but also has been collected at the opposite end of Alakai Swamp, at Pihea. It resembles the unnamed glabrous taxon in its depressed obtrullate or depressed obovate cicatrices on the stems, linear or oblanceolate leaves, branched inflorescence, relatively large flowers, longitudinally striped pale purple corolla, and relatively large (2-3 mm long) seeds. However, it differs in its somewhat pubescent inflorescence with green axes, broader floral bracts, rounded calyx lobes, and pink corolla venation; the inflorescence is also shorter on average (1-2 m). The types of L. gaudichaudii f. hirsuta and L. gaudichaudii var. kauaiensis represent this taxon. As noted above, the densely pubescent taxon that corresponds to the type of L. kauaiensis var. villosa is found throughout the Alaka'i Swamp and is the most distinctive member of the section. The totally glabrous taxon occurs at just one place within its range, and at that place, the somewhat pubescent taxon is also found. The densely pubescent taxon and the glabrous taxon exhibit pronounced differences in flower color and size. Especially noteworthy is the gap in the lengths of their staminal columns: 45—57 mm long in the densely pubescent plants vs. 70-80 mm long in the glabrous ones. All this suggests that they are isolated reproductively under normal circumstances and merit distinction at specific rank. The name L. villosa is available for the densely pubescent species; the glabrous species is christened L. wahiawa below. The somewhat pubescent plants at Pihea and Wai'ale'ale resemble L. wahiawa in several characters, as noted above; the characters by which they differ may be interpreted as due to the genetic influence of L. villosa. The inflorescence is shorter, more pubescent, and not darkly pigmented; the floral bracts are wider and the calyx lobes more rounded; and the pigmentation of the corolla veins is diluted. For these reasons, osa and L. wahiawa. It is this taxon == this taxon is here interpreted as an interspecific hybrid between L. vil to which the name L. kauaiensis properly applies. I generally have not endorsed the practice of christening occasional interpecific hybrids with binomials. In this case, however, retaining a well known name in a new status may result in | nfusion than if it simply disappeared. Therefore, L. villosa x L. wahiawa, which has Lammers, Revision of Lobelia sect. Galeatella 797 been called L. gaudichaudii var. kauaiensis f. hirsuta (St. John & Hosaka 1938; Wimmer 1953) or Galeatella kauaiensis var. hirsuta (Degener & Degener 1962b, 1974), now will be called L. xkauaiensis. In the Wahiawa drainage of south-central Kauai, only a single taxon is found and it is L. wahiawa. Thus, the range of this species is largely allopatric from L. villosa. The latter is confined to the high elevations of the Alakai Swamp while L. wahiawa occurs in the low elevations of Wahiawa Swamp, except for the single population in the Alaka'i Swamp at Wai‘ale‘ale. It is possible that the sympatry of the two on Wai'aleale is a relatively recent phenomenon: the winged seeds of lowland L. wahiawa may have been carried the 10 km to the mountaintop and successfully established there. Hybridization then occurred from time to time. The presence of the hybrid without L. wahiawa at Pihea (15 km northwest of Wai'aleale) could be explained by the dispersal of pollen of the absent parent via nectarivorous bird (cf., Lammers & Freeman 1986; Lammers et al. 1988) or by dispersal of hybrid seed from Wai‘ale‘ale. It could also be evidence of a second montane colonization, as yet undocumented or now extirpated, by L. wahiawa. The specimens examined from O'ahu differ from all others in the section by their relatively short (0.4—1 m) cylindrical inflorescences, distinctly revolute leaves typically pubescent on the abaxial midrib, and small (1-1.8 mm long) seeds. Other characters, however, vary among these specimens; furthermore, this variation is correlated with geography. Specimens from the northwestern portion of the Ko'olau Range, between Pu'u Ka'inapua'a and ‘Eleao at elevations of 670—860 m, are characterized by a 2-6-branched inflorescence, white corolla, dorsal anthers 14-16 mm long, capsules 18-21 mm long and 9-10 mm in diameter, and seeds 1.5-1.8 mm long; the type of L. gaudichaudii var. koolauensis represents this taxon. Specimens from the southeastern Ko'olau Range, between Pu'u Pauao and Mt. Olympus at 825-960 m, differ by their usually unbranched inflorescence, red corolla, shorter (10-13 mm) dorsal anthers, larger (23-30 x 11-10 mm) capsules, and smaller (1-1.5 mm) seeds; the types of L. gaudichaudii and L. gaudichaudii var. coccinea are representatives of this taxon. Note that the ranges of these two taxa overlap between Pu'u Pauao and ‘Eleao. These two taxa were treated previously as conspecific and distinguished at either varietal (St. John & Hosaka 1938; Wimmer 1953, 1968; Degener & Degener 1962b, 1974) or subspecific (Lammers 1988, 1990) rank. However, as noted by St. John and Hosaka (1938), the northwestern Ko'olau plants seem almost as closely related to the plants of Kauai here called L. wahiawa as they do to the southeastern Ko'olau plants. Characters shared by that species and the northwestern Ko'olau plants (but not the southeastern) include the branched (vs. unbranched) inflorescence, light-colored (vs. red) corolla, dorsal anthers 14-16 (vs. 10-13) mm long, and seeds 1.5-2.5 (vs. 1-1.5) mm long. While the unique characters shared by the O'ahu plants may suggest their derivation from a common ancestor, the characters that differ, particularly corolla color and anther length, are the sort that might be indicative of reproductive isolation. This hypothesis is supported by the absence of morphologically inter- mediate specimens in the zone of sympatry between Pu'u Pauao and 'Eleao. Because the evidence suggests that these two taxa are unable to interbreed in nature, they are accorded specific rank. The name L. gaudi- chaudii applies to the southeastern taxon, and the new combination L. koolauensis will be effected below for the northwestern one. Specimens seen from West Maui share a distinctive suite of features that at once distinguishes them from the plants of Kauai and O'ahu: a pyramidal outline to the inflorescence, created by a progressive shortening of the pedicels from base to apex; lamina with a thickened or subrevolute margin typically ciliate toward base; the largest flowers and fruits in the group; and seeds of moderate size (2-2.3 mm long). Certain other characters, however, evince considerable variation. In the majority of plants from West Maui (including the type specimens of L. gaudichaudii f. kukuiensis and L. gloria-montis), the inflorescence is glabrous and the corolla white. However, occasional specimens (such as the type of L. gaudichaudii f. bryanii) have a densely pubescent inflorescence, while in others (including the type of L. gaudichaudii f. sanguinea) the corolla is red. In neither case could I detect any correlation of the unusual character state with other morphological characters, or with geography, elevation, or habitat; the red-flowered variants at least are definitely reported to grow side-by-side with white-flowered plants 798 Journal of the Botanical R h Institute of Texas 1(2) (Skottsberg 1944; Selling 1947). As such, it appears that each character—pubescent inflorescence and red corolla—is a simple genetic variant occurring sporadically within a population. There are other specimens from West Maui that resemble the dominant morphotype in their glabrous inflorescence and white corolla, but that differ in other characters; furthermore, they are restricted to a dif- ferent habitat. The plants mentioned in the preceding paragraph occur in open montane bogs on Pu'u Kukui, 'Eke, and other summits. All have a stem less than 1.5 m tall; an unbranched inflorescence; and relatively short floral bracts, pedicels, and calyx lobes. Specimens collected in forests at the head of Tao Valley (cf., Rock 1913a, 1919) differ in their much taller (2.5—4 m) stems, 5-7-branched inflorescence, floral bracts 60-80 (vs. 20-47) mm long, pedicels 50-60 (vs. 15-45) mm long, and calyx lobes 30-35 (vs. 7.5225) mm long. The type of L. gaudichaudii var. longibracteata is an example of these plants. In my previous treatment (Lammers 1990), I did not distinguish these larger forest plants from the smaller bog plants at any rank, primarily because of the close proximity of their distributions. I certainly do not favor according them each specific rank as Wimmer (1968) and Degener and Degener (1974) have done. The characters that distinguish the two taxa are not the sort that might be expected to confer reproductive isolation, and in any case, evince some overlap in their ranges of values. As such, it is deemed best to treat them as conspecific ecological subspecies. The name for the species is L. gloria-montis. The small bog plants constitute autonymic subsp. gloria-montis, while the new combination subsp. longibracteata is effected below for the large forest plants. Specimens from East Maui (including the type of L. gaudichaudii var. albiflora) are very similar to the tall forest plants of West Maui, with stem 2-4 m tall, floral bracts 62-65 mm long, pedicels 38-55 mm long, and calyx lobes 25-30 mm long; their habitat, described as forested margins of bogs, also seems quite similar. The only substantive difference is that the inflorescence of the East Maui plants, so far as known, does not branch. Given the small number of specimens representing the tall forest plants of East and West Maui, it seems best at present to assign them all to a single taxon. Consequently, the specimens from East Maui are here assigned to L. gloria-montis subsp. longibracteata. The optimal disposition of the populations on Moloka'i is uncertain. All known specimens (including the type of L. gloria-montis var. molokaiensis) are sterile. Although one would predict that a geographically isolated population might at least merit recognition at subspecific rank, the specimens at hand are indistin- guishable from the typical morphotype of L. gloria-montis subsp. gloria-montis. Until more adequate material is available for study, these plants are assigned to that taxon. In summary, sect. Galeatella is treated here as comprising five species. Kauai and O'ahu each have two endemic species: L. villosa and L. wahiawa on the former, L. koolauensis and L. gaudichaudii on the latter. The Kauai pair hybridize where sympatric (L. xkauaiensis), while the O'ahu pair do not. The fifth species (L. gloria- montis) is represented by two ecological races on West Maui: subsp. gloria-montis and subsp. longibracteata. The populations on East Maui and Molokai are rather poorly known, but are assigned to L. gloria-montis subsp. longibracteata and L. gloria-montis subsp. gloria-montis, respectively. TAXONOMIC TREATMENT Toc sect. Galeatella E. Wimm., Ann. Naturhist. Mus. Wien 56:369. 1948. Galeatella (E. Wimm.) O. Deg. € I. ., Fl. Hawaiiensis, Fam. 339. 1962. Tre (designated by Degener & Degener 1962b): L. kauaiensis var. villosa Rock Pachycaul treelets of the *Tomlinson" architectural model (cf., Hallé & Oldeman 1970; Hallé et al. 1978), 0.7—5 m tall; plant polycarpic but the shoot pliestesial, hapaxanthic, and replaced by a sucker from the rootstock after fruit maturation. Stem 0.3-4 m tall, 0.8-10 cm diam., solitary and unbranched, erect or ascending, leaty only at apex; cicatrices raised, corky, transversely rhombic, depressed obtrullate, or depressed obovate in outline with a single central vein scar, closely packed in a helical pattern and long persistent; pith soft, broader than the lignified cortex toward apex but narrowing toward base as the cortex increases in thickness, becoming hollow with age; latex white, viscous. Leaves alternate, simple, exstipulate, sessile, glabrous or pubescent, forming a dense spherical or oblate apical rosette; lamina oblong, elliptic, lanceolate, oblanceo- late, or linear, coriaceous, stiff, flat; margin entire, commonly revolute, with few to several intramarginal Lammers, Revision of Lobelia sect. Galeatella 799 callosities distally; apex acuminate, acute, or rarely obtuse; base attenuate or cuneate. Flowers tetracyclic, perfect, proterandrous, zygomorphic, epigynous, pedicellate, resupinate, ornithophilous, large; inflorescence an erect terminal pedunculate bracteate raceme, 0.4—2.5 m tall, unbranched or 2-7-branched from base, up to 100-flowered over its life, but seldom more than 4—8 open at any one time, glabrous or pubescent; peduncle usually shorter than rachis, densely to sparsely covered with reduced leaves (sterile bracts) that rapidly or gradually decrease in size acropetally; floral bracts conspicuous, equaling or shorter than the pedicels; pedicels ascending, stiff, ebracteolate. Calyx synsepalous, glabrous or pubescent; tube adnate to the basal '/,—/, of the ovary, forming a broadly obconic (rarely campanulate or hemispherical) appendicular hypanthium //,—,, as long as the corolla; lobes 5, valvate, triangular, lanceolate, ovate, or oblong, shorter than the corolla tube, a little shorter than the hypanthium to over 4 times longer, the dorsal slightly longer than the ventral, caducous from the young fruit, the margin entire, the apex acuminate, acute, obtuse, or rounded and sometimes apiculate. Corolla sympetalous, zygomorphic, subbilabiate, of various colors (white, pale yel- low, red, or pale purple with darker longitudinal veins) but never blue, glabrous (the lobes rarely pubescent at apex); tube straight or slightly curved, cylindric or expanding slightly towards the mouth, 1.3—5 times longer than broad, dorsally cleft nearly to base but never fenestrate; lobes 5, valvate, linear, monomorphic, acuminate or acute at apex, the dorsal pair spreading or recurved horizontally, about as long as the tube to 3 times longer, the three ventral lobes connate for ?/, their length or more, forming a deflexed inrolled trifid lip shorter than the dorsal lobes. Stamens 5, antisepalous, connate distally for most of their length, strongly exserted, emerging from the dorsal corolla slit above or between the dorsal lobes; filament tube suberect or slightly deflexed, 2.8—6.3 times longer than anther tube, glabrous (rarely short pubescent ventrally); dorsal anthers longer than the ventral, overhanging the orifice of the tube and somewhat occluding it; all 5 anthers with tufts of white trichomes at apex; pollen 52-54 um polar diam. x 35-41 pm equatorial diam., prolate, tricolporate, 3-celled at anthesis, the exine shallowly and minutely reticulate (Selling 1947; Erdtman 1952; Pandey et al. 1993). Ovary 2-loculed, inferior, adnate to the hypanthium for /,—/, its length, the conic apex free, forming a pronounced beak on the young fruit; placentae axile; ovules numerous; style 1, slender, terete, with a ring of stiff white hairs near the apex; stigma 2-lobed, the lobes appressed and non-receptive as the style grows through the anther tube, pushing out pollen, after which the stigmas spread and become receptive. Fruit an ovoid or ellipsoid capsule; exocarp and parenchymatous portions of the mesocarp deli- quescent, leaving a loosely enshrouding series of sclerenchymatous bundles, open at top; endocarp rigid and durable, its conical apex splitting loculicidally into a pair of triangular valves; seeds (Fig. 1) winged, oblong, compressed, honey- or chestnut-brown, the narrow wing paler, the testa faintly striate (Type C of Murata 1992). Chromosome number unknown (Lammers 1993). These species are restricted to wet habitats at elevations of 570-1760 m on four of the Hawaiian Islands. On Kauai and Maui, they are found in level saturated bogs, while on O'ahu and Molokai, they occur on windswept ridge crests and summit slopes. The plant communities that develop in all these areas (cf., Gagné & Cuddihy 1990) typically are dominated by dwarfed shrubs and various graminoids, and the species of Lobelia sect. Galeatella are typically among the tallest and most conspicuous elements in the vegetation (cf., Carlquist 1980, pp. 346, 350, 352 [bottom]). Flowering begins in mid-summer and is generally finished by October, with most seed maturing by January. The flowers of these species, like most Hawaiian Lobelioideae, are presumed to be ornithophilous (Lammers & Freeman 1986; Lammers et al. 1988; Smith et al. 1995). Peale's (1848: 152) illustration of the endemic nectarivorous bird Vestiaria coccinea (Forster) of the Drepanidini (Aves: Fringillidae) includes an unequivocal depiction of Lobelia gaudichaudii; he commented that the bird's bill was *admirably adapted" for extracting nectar from its corolla. The flat winged seeds (Fig. 1) are presumably suited for a limited degree of anemochory. Although none of the species has been examined cytologically, the group is hypothesized to be tetra- ploid with 2n = 28, in light of the fact that all counted species of its sister-group (Trematolobelia) and their outgroup (Lobelia sect. Revolutella) have that number (Lammers 1993). 800 Journal of the Botanical R h Institute of Texas 1(2) KEY TO THE SPECIES AND SUBSPECIES 1. Inflorescense densely pul the peduncle d ly covered with sterile bract lually d g l H | JI AA P | | | 13 s | d ; l | Ii y 24mm long; fi filament tube EE 45 mm long; capsule 1 13- 18 mm "d de Swamp ofKaual) — — 1. Lobelia villosa 1. Inflorescense glabrous (rarely pubescent), th covered with sterile bracts rapidly decreas- ing in size acropetaly, the floral Pine cuneate at base; I ee ithium SE. e) en eax lobes acuminate or acute (rar ly rouna obtuse) a | with darker lor \gitudinal veins, red, or white, 53-103 mm long, the Abe 2 5 times uid than broad, the ventral lip 24-50 mm long; filament tube 57-80 mm long; capsule 19-37 mm lon 2. Cicatrices depressed obtrullate or depressed obovate, the corners rounded; lamina 17-37 cm long, com- monly linear; inflorescence 1-2.5 m tall; corolla pale purple with dark purple or pink longitudinal veins; seeds (1.7-)2-3 mm long (Kaua'i). 3. Inflorescence 1-2 m tall, ee 35-flowered, its axi | | | | urled trichomes; floral bracts elliptic, oblong, or obovate; calyx lobes eed ae COS yth DOE (Alakaʻi Swamp) 2.L jen xkauaiensis 3. Inflorescence 1.5-2.5 m tall, 20-85-flowered, its axis blackish purple and glabrous; floral bract narrowly elliptic, narrowly oblong, or lanceolate; calyx lobes acuminate or acute (rarely obtuse) a apex; corolla with dark purple veins (Wahiawa & Alaka'i Swamps) 3. Lobelia wahiawa 2. Cicatrices transversely rhombic, the corners acute; lamina 9-24 cm long, commonly oblanceolate; inflo- rescence 0.4-1.5 m tall; corolla red or white; seeds 1-2(-2.3) mm long. 4. Inflorescence 0.4-1 m tall, cylindrical in outline; lamina di tly revolute at margin, pubescence (if any confined to abaxial midrib; corolla 58-78 mm long, the tube 19-31 mm long, the dorsal lobes 27-45 a mm long; capsule 18-30 mm long; seeds 1-1.8 mm long (O'ahu). . Inflorescence 2-7- E corolla ane dorsal anthers 14-16 mm long; capsule 2 mm long x 9-10 mm diameter; seeds 1.5-1.8 mm long (NW Ko'olau Range) Lobelia koolauensis . Inflorescence unbranched; corolla red; dorsal anthers 10-13 mm long; capsule 2: 33430. "y mm lon 11-19 mm diameter; id 15 mm TOO GE Galan Range 5. Lobelia T — 4. mc 15mtall, tel in, pubescence , EN SOLES D E long; capsule 27-37 mm long; seeds 2-2. s mm eng LOU MED 6. Lobelia ions montis 6. Stems0.4-1.5 mtallifloralbracts 20-47 lat licels 15-45mmlong;cal lobes 7.5-25 mm long (Moloka'i, West Maui) 6a. Lobelia gloria-montis subsp. gloria-montis 6. Stems2-4m tall floral bracts 60-90 mm long, oblong or linear; pedicels 38-60 mm long; calyx lobes 25-35 mm long (West and East Maui) 6b. Lobelia gloria-montis subsp. longibracteata 1. Lobelia villosa (Rock) H. St. John & Hosaka, Occas. Pap. Bernice Pauahi Bishop Mus. 14:124. 1938. Lobelia kauaiensis var. villosa Rock, Bull. Torrey Bot. Club 44:237. 1917. Galeatella villosa (Rock) O. Deg. & I. Deg., Fl. Hawaiiensis, Fam. 339. 1962. Type: HAWAIIAN ISLANDS. Kauas: Wai‘ale‘ale, 21 Oct 1916, Rock 12844 (HoLoTYPE: BISH!; isotypes: BISH! K!). The collection number is given as “12741” in the protologue and subsequent monograph (Rock 1919), but the figured specimen in the latter (pl. 61; called “Type” in the caption) shows this sheet with this number. Plant 1.2-2.2 m tall. Stem 0.6-2 m tall x1-6 cm diam.; cicatrices 3-3.5 mm tall x 6-7 mm wide, transversely rhombic, the corners acute. Lamina 14-22 cm long x 1.73 cm wide, oblong or oblanceolate, densely pubescent on margin, sometimes also pubescent on the dorsal and/or ventral midrib; margin subrevolute; apex obtuse or acute; base cuneate or attenuate. Inflorescence 0.5-1 m tall, 2-5-branched (rarely unbranched in small plants), densely pubescent, each branch 13-45-flowered and cylindrical in outline, its axis green; peduncle densely covered with sterile bracts that gradually decrease in size acropetally; sterile bracts 25-120 mm long x 10-30 mm wide, elliptic, oblong, lanceolate, ovate, or obovate, the apex acute or obtuse, the base rounded; floral bracts 17-30 mm long x 8-18 mm wide, elliptic, ovate or obovate, the apex acute or obtuse, the base rounded or subcordate; pedicels 15-35 mm long, of more-or-less equal length throughout. Hypan- thium 5-9 mm long x 7-11 mm diam., broadly obconic, '/,—'/,, as long as the corolla, densely pubescent; calyx lobes 8.5-12 mm long x 4-5 mm wide, triangular or oblong, equaling the hypanthium to 2.2 times as long, the apex obtuse or rounded. Corolla 44-57 mm long, pale yellow, glabrous; tube 15-18 mm long x 8-13 mm diam. at middle, straight, cylindric or expanding slightly towards the mouth, 1.3-2 times longer Lammers, Revision of Lobelia sect. Galeatella 801 than broad; dorsal lobes 29-39 mm long x 3.5-5 mm wide, 1.6-2.2 times longer than the tube; ventral lip 22-24 mm long x 12-17 mm wide. Filament tube 36-45 mm long, slightly deflexed, 2.8—4.5 times longer than anther tube, glabrous; anther tube 4 mm diam.; dorsal anthers 9-13 mm long; ventral anthers 8-9.5 mm long. Capsule 13-18 mm long x 10-12 mm diam.; seeds (Fig. 14; Wimmer 1953, fig. 104c) 1.5-2.5 mm long x 0.6-0.8 mm wide. Icon.—Rock (1919), pl. 43. Figured specimen.—Rock (1919), pl. 61 [holotype]. Distribution, habitat, and phenology.—Endemic to north-central Kauai, known only from Montane Wet Mixed Communities (Gagné & Cuddihy 1990) of the Alaka‘i Swamp (cf., Carlquist 1980, p. 350) from Pihea to Wai'ale'ale, at 1100-1560 m. Flowering from August to October. Considered “rare” by Wagner et al. (1999). Representative specimens: KAUAI. Waimea: Alakai, bogs between Lel | d Kilohana, Cranwell et al. 2984 (BISH, GB); Alaka‘i Swamp, in 2"™ large bog on trail to old USGS cabin on way to Wai‘ale‘ale, Davis & Kores 131 (BISH); Alaka'i Swamp, Flynn 685 (PTBG), Flynn 686 (PTBG); Alaka'i Swamp, Forbes 1122.K (BISH); Wai‘ale‘ale, Hitchcock 15450 (US); Alaka'i Swamp, along trail between Pihea and Kilohana, Lammers et al. 5376 (DUKE, F, OS); Wai‘ale‘ale, Rock 5823 pro parte (GH), Rock 8856 (BISH); Wai'ale'ale, 23 Sep 1909, Rock s.n. (BISH), Oct 1911, Rock s.n. (BISH); Alaka'i Swamp, Yuncker 3524 (DPU, US). 2. Lobelia xkauaiensis (A. Gray) A. Heller, pM. Bot. Stud. 1:911. 1897 (as ‘kauaensis’, pro sp.) stat. NOV. Lobelia gaudichaudii var. kauaiensis A. Gray, Proc. Amer. Acad. Arts 5:150. 1861 (as “kauaensis’). Galeatella kauaiensis (A. Gray) O. Deg. & I. Deg., Fl. Hawaiiensis, Fam. 339. 1962 d p Type: HAWAIIAN ISLANDS. Kauai: [Alaka‘i Swamp, 30 Oct 1840, leg. W. | & C. Pickering (fide Wilkes 1845),] U.S. Exploring p s.n. is OTYPE: US!; ISOTYPE: PU i: d is Es on m fable Hawaiian proper noun “Kaua'i;” Grays (1861 error to be corrected, following Skottsberg (1944), Wimmer (1968), and Degen and egerat (1974) du Lobelia gaudichaudii f. hirsuta H. St. John & Hosaka, Occas. Pap. Bernice Pauahi Bishop Mus. 14:121. 1938. Galeatella kauaiensis var. hirsuta (H. St. John & Hosaka) O. Deg. & I. Deg., Fl. Hawaiiensis, Fam. 339. 1962. Type: HAWAIIAN ISLANDS. Kaua‘t: Wai‘ale‘ale, Oct 1916, Rock 12845a (HOLOTYPE: BISH!; IsoTYPE: BISH!). Plant 2.4—4 m tall. Stem 1-2 m tall x1-6 cm diam.; cicatrices 3-4.5 mm tall x 4.5-8 mm wide, depressed obtrullate or depressed obovate, the corners rounded. Lamina 24-32 cm long x 2.2-3.3 cm wide, linear, oblanceolate, or narrowly elliptic, glabrous or pubescent on ventral midrib; margin subrevolute; apex acu- minate; base attenuate or cuneate. Inflorescence 1-2 m tall, 5-branched, sparsely pubescent with relatively short curled trichomes, each branch 13-35-flowered and cylindrical in outline, its axis green; peduncle sparsely covered with sterile bracts that rapidly decrease in size acropetally; sterile bracts 30-105 mm long x 4-14 mm wide, linear or narrowly oblong, the apex acuminate, the base cuneate; floral bracts1 2-23 mm long x 4-10 mm wide, elliptic, oblong, or obovate, the apex obtuse or rounded, the base cuneate; pedicels 25-45 mm long, of more-or-less equal length throughout. Hypanthium 6-10 mm long x 13-15 mm diam., broadly obconic, 1/7-1/11 as long as the corolla, glabrous or pubescent; calyx lobes 8-13 mm long x 3-6 mm wide, triangular or oblong, equaling to a bit longer than the hypanthium, the margin entire, the apex rounded. Corolla 64-66 mm long, pale purple with pink longitudinal veins, glabrous; tube 20-22 mm long x 10-12 mm diam. at middle, slightly curved, expanding slightly towards the mouth, twice as long as broad; dorsal lobes 42-46 mm long x 4—5 mm wide, about twice as long as the tube; ventral lip 37-39 mm long x 15-17 mm wide. Filament tube 64—66 mm long, suberect, 4 times longer than anther tube, glabrous; anther tube 4 mm diam.; dorsal anthers 14-16 mm long; ventral anthers 10-11 mm long. Capsule 19-30 mm long x 8-20 mm diam.; seeds (Fig. 1B) 2-3 mm long x 0.7-0.9 mm wide, chestnut-brown. Icones.—Rock (1919), pl. 42 [right]; Wimmer (1968), fig. 29. Distribution, habitat, and phenology.—A putative hybrid of L. villosa and the next species, known only from the Montane Wet Mixed C ities (Gagné & Cuddihy 1990) at the southeastern and northwestern extremities of the Alakai Swamp on north-central Kaua‘i. At the former site (Wai'aleale, 1370-1570 m), it occurs with both parents, as evinced by Roch 5823; the sheet at BISH represents the hybrid, while that at GH is a mixed gathering of the two parents. At the latter site (Pihea, 1220-1350 m), L. villosa co-occurs but not the other parent. Flowering in August, fruit ripening in October. This taxon is listed as “rare” by Wagner et al. (1999), an assessment that also encompasses L. wahiawa. 802 Journal of the Botanical R h Institute of Texas 1(2) Discussion.—Plants on Kauai that resemble the following species except for their puberulous raceme have been treated as a minor variant of it by most authors (Rock 1919; St. John & Hosaka 1938; Wimmer 1953; Degener & Degener 1962b, 1974). In my earlier treatment (Lammer 1990), I suggested that they were more likely interspecific hybrids, the pubescence indicating gene-flow from L. villosa. Unfortunately, I missed clues (Gray 1861; Skottsberg 1944) that the type specimen of L. kauaiensis was an example of this same hybrid. As a result, not only does the name L. gaudichaudii f. hirsuta apply to the hybrid, but L. kauaiensis as well. The latter name is used here to denote the hybrid, with the addition of the multiplication sign to denote its status as a nothotaxon. R KAUA'I. Hanalei: ridge E of Pil | 1s Kilol Flynn & DeLappe 3198 (BISH, F, K, MO, PTBG, US); Norn Bog one Wainiha pali, Wood & Perlman 3672 E PTBG). Waimea: Napali-Kona Forest AUR Wai'ale'ale trail, Alaka'i Swamp Keáku River, Chock 994 (BISH, NSW); Wai‘ale‘ale, along trail from “Kauku” [Keakü?], Cranwell et al. 3047 pro parte (GB); Wai'aleale, Hitchcock rd a (US); Wai'ale'ale, Rock 5109 (BISH); Wai'ale'ale, Sep 1909, Roch s.n. (BISH); Wai'ale'ale, Oct 1911, Rock s.n. (BISH); Wai'ale'ale, R I BISH); ridge SW of Pihea, on east rim of Kalalau Valley, St. John et al. 22945 (DUKE). E 3. Lobelia wahiawa Lammers, Sp. nov. Type: HAWAIIAN ISLANDS. Kauai: Koloa District, Wahiawa Swamp near headwaters of Wahiawa Stream, 630 m, open swamp with some shrubby areas, frequent, 24 Apr 1964, Stauffer & Dehler 5910 (HoLotyeE: BISH!; isotypes: A! DUKE! K! US). leatellae | i ] i ] i imile L. xkauaiensi, cuj ,sed o [2] E E in E a} L^ is vel acuminatis, et corollae venatione atro-purpurea differt. Species Lobeliae sect. Tai dead inflorescentia glabra calvcis Plant 2-4 m tall. Stem 1-2.5 m tall x 1-10 cm diam.; cicatrices 3-5 mm tall x 4.5-9 mm wide, depressed obtrullate or depressed obovate, the corners rounded. Lamina 17-37 cm long x 13.3 cm wide, linear, oblanceolate, or narrowly elliptic, glabrous; margin subrevolute; apex acuminate; base attenuate. Inflores- cence 1.5-2.5 m tall, 2-5-branched (rarely unbranched in small plants), each branch 20-85-flowered and cylindrical in outline, its axis blackish purple, glabrous; peduncle sparsely covered with sterile bracts that rapidly decrease in size acropetally; sterile bracts 12-160 mm long x 4-22 mm wide, linear, narrowly el- liptic, or lanceolate, the apex acuminate, the base cuneate; floral bracts 16-27 mm long x 3-7.5 mm wide, narrowly elliptic, narrowly oblong, or lanceolate, the apex acute or obtuse and sometimes apiculate, the base cuneate; pedicels 17-35 mm long, of more-or-less equal length throughout. Hypanthium 5-9 mm long x 8-14 mm diam., broadly obconic or rarely almost hemispherical, 1/8-1/12 as long as the corolla, glabrous; calyx lobes 9-15 mm long x 3-6.5 mm wide, triangular, narrowly oblong, or lanceolate, equaling to 3 times longer than the hypanthium, the margin entire, the apex acuminate or acute (rarely obtuse) and sometimes apiculate. Corolla 53-75 mm long, pale purple with dark purple longitudinal stripes, glabrous; tube 18-28 mm long x 6-10 mm diam. at middle, suberect or slightly curved, expanding slightly towards the mouth, 2—4.5 times longer than broad; dorsal lobes 30-48 mm long x 3-5 mm wide, 1.3-2.2 times as long as the tube; ventral lip 25-38 mm long x 11-18 mm wide. Filament tube 57-65 mm long, decurved slightly, 4—5 times longer than anther tube, glabrous; anther tube 4-5 mm diam.; dorsal anthers 12-16 mm long; ventral anthers 9-12 mm long. Capsule 20-28 mm long x 10-14 mm diam.; seeds (Fig. 1C) 1.7-2.5 mm long x 0.6-1.4 mm wide, honey- or chestnut-brown. Icones.—Rock (1919), pl. 16; Carlquist (1980), p. 245 [left], p. 348 [top left]; Sohmer and Gustafson (1987), fig. 114; Lammers (1990), pl. 57 [lower left]. Figured specimen.—Rock (1919), pl. 60 [Heller 2888, GH]. Distribution, habitat, and phenology.—Endemic to central Kauai, known primarily from the Lowland Wet Mixed Communities (Gagné & Cuddihy 1990) at 570-800 m in the watershed of Wahiawa Stream west of Kahili (cf., Carlquist 1980, p. 346), but also found in the unique Montane Wet Mixed Community (Gagné € Cuddihy 1990) of the summit bog on Wai'aleale at 1100-1590 m. Flowering July to March, fruit ripening December to April. Regarded as “rare” by Wagner et al. (1999), an assessment that includes its hybrid with L. villosa (see previous taxon). Etymology.—The specific epithet is the name of the type locality, used as a noun in apposition; it is Hawaiian for “milkfish place" (Pukui et al. 1974), a reference to widespread Chanos chanos (Forsskál) of the Chanidae (Actinopterygii). Lammers, Revision of Lobelia sect. Galeatella 803 Vernacular name.—Pu'e (Rock 1919; Pukui & Elbert 1971). Discussion.—This is the glabrous species that has been known for many years (Heller 1897; Rock 1919; Wimmer 1968; Lammers 1990) as Lobelia kauaiensis. However, as noted by Skottsberg (1944), the original description of the basionym of that name (Gray 1861) called for a plant *racemo puberulo; calycis viscosi" and its typ indeed conforms to that diagnosis. As a result, this glabrous species has no name and r so is named here as a new species. Representative specimens: KAUAT. [Kawaihau: Pohakupili (fide Wawra 1873), l Wawra 2044 (W). Koloa: Wahiawa Bog, Carlquist 1980 (RSA); Wahiawa Bog, 26 Nov 1972, Carlquist s.n. (RSA); Wahiawa Bog, Fagerlind & Skottsberg 6480 (GB, UPS, S); Hanapepe, in turfosis Faurie 552 (A, BISH, P, W); Wahiawa Bog, Flynn 371 (PTBG); Wahiawa Bog, Flynn 668 (OS); Wahiawa Bog, Flynn 699 (OS); Wahiawa Swamp, Forbes 294.K (BISH, US, W); Wahiawa Swamp, Aug 1909, Forbes s.n. (BISH); Wahiawa Bog, Gustafson 1049 (BISH, RSA); in and near a bog at the head of the Wahiawa, 19 Oct 1895, Heller 2888 a GH, MO, NY, US); Wahiawa Bog, Kato LK-2 (BISH); Wahiawa Bog, Lammers et al. 5861 (OS); Wahiawa Bog, 28 Dec 1980, Lucas s.n. (RS Wahiawa, MacDaniels 636 (BH, BISH); Wahiawa Swamp, Sparre H.75 (S); Wahiawa Bog area west of Mt. Kahili, Stern & Carlquist 1331 (RSA, US); Wahiawa Bog, Stern & Herbst 2942 (PTBG, RSA); kahili Swamp, Wahiawa, St. John et al. 10849 (A, K, BISH, NSW); Kahili Bog, Wahiawa, St. John & Fosberg 13561 (BISH, POM); west side of Wahiawa drainage below ridge between Hulua and Pu'u'au'uka, Wood et al. 1059 (MO, PTBG). Waimea: Wai'aleale, along trail from “Kauku” [Ke'akú?], Cranwell et al. 3047 pro parte (BISH); Wai'ale'ale summit, Davis & Kores 113 (A, BISH) & 113a (BISH); Wai'aleale, Hitch- cock 15437 pro parte (US); Wai'ale'ale, Hitchcock 15499 (US); Wai‘ale‘ale, Rock 5115 (BISH); Wai‘ale‘ale, Rock 5823 pro parte (GH); Wai'ale'ale, Rock 12845 (BISH, K); Wai‘ale‘ale, Sep 1909, Roch s.n. (NY, US, W); along Alaka'i Swamp Trail, Stone 1536 (BISH). 4. Lobelia koolauensis (Hosaka & Fosberg) Lammers, comb. et stat. nov. Lobelia gaudichaudii var. koolauensis Hosaka € dde Occas. Pap. Bernice Pauahi Bishop Mus. 14:4. 1938. e NH CM var. aa auensis dd: & Fosberg) O. Deg. Y I. Fl. Hawaiiensis, Fam. 339. 1962. Lobeli g Syst. Bot. 13:506. 1988 Deg., Type: HAWAIIAN ISLANDS. O'aHu: Ko'olau Range, divide | lof} i Nui and Kai lc] sloping bog, 2600 ft, 31 May 1937, Hosaka & gn 1915 (uoLotyre: BISH [mounted on 5 pm ; ISOTYPES: BISH[2]! US! WD. Plant 1-2.5 m tall. Stem 1-1.5 m tall x 1.8-3 cm diam.; cicatrices 3-4 mm tall x 6.5-10 mm wide, transversely rhombic, the corners acute. Lamina 11.8-20 cm long x 0.9-2.8 cm wide, linear, oblanceolate, or narrowly elliptic, glabrous or the abaxial midrib pubescent; margin distinctly revolute; apex acute or acuminate; base attenuate. Inflorescence 0.5-1 m tall, 2-7-branched (rarely unbranched in small plants), each branch 12-25-flowered and cylindrical in outline, its axis green, glabrous; peduncle sparsely covered with sterile bracts that rapidly decrease in size acropetally; sterile bracts 20-95 mm long x 4-13 mm wide, narrowly oblong, narrowly elliptic, or lanceolate, the apex acute, the base cuneate; floral bracts 20-48 mm long x 5-10 mm wide, narrowly elliptic or lanceolate, the apex acuminate, the base cuneate; pedicels 22-60 mm long, of more-or-less equal length throughout. Hypanthium 5-10 mm long x 7-11 mm diam., broadly obconic, 1/6-1/8 as long as the corolla, glabrous; calyx lobes 10-22 mm long x 3.5-6 mm wide, narrowly d or lanceolate, equaling to 4.4 times longer than the hypanthium, the margin entire, the apex and sometimes apiculate. Corolla 58-78 mm long, white, glabrous; tube 19-31 mm long x 8-12.5 mm diam. at middle, suberect, expanding slightly towards the mouth, 2-3.5 times longer than broad; dorsal lobes 27-45 mm long x 4.5-6.5 mm wide, 0.9-2.4 times as long as the tube; ventral lip 24-40 mm long x 17-24 mm wide. Filament tube 61-78 mm long, suberect, 4-5 times longer than anther tube, glabrous; anther tube 4—4.5 mm diam.; dorsal anthers 14-16 mm long; ventral anthers 9-10 mm long. Capsule 18-21 mm long x 9-10 mm diam.; seeds (Fig. 1D) 1.5-1.8 mm long x 0.8-1 mm wide, honey-brown. Icones.—Hosaka and Fosberg (1938), fig. 2; Wimmer (1953), fig. 104a. Distribution, habitat, and phenology.—Endemic to O'ahu, known only from Montane Wet Shrublands (Gagné € Cuddihy 1990) on the summit of the northwestern Ko'olau Range, between Puu Ka'inapua'a and 'Eleao, at 670—860 m. Flowering from late May to October, fruits ripening November to January. Considered “endangered” by Wagner et al. (1999). Representative specimens: O'AHU. Ko'oLau Mrs.: Poamoho Trail, Paaloa-Wahiawa, Cowan 252A (WAB); Poamoho Trail, Laie, Degener et al. 11497 (MO, W); top of Poamoho Trail crest at head of Punalu'u Valley, Fosberg 13330 (BISH); main divide above Kaipapa'u Gulch, Fosberg 14224 (BISH, K); Manana Trail, NE of Pacific Palisades, 12 Jul 1985, Obata s.n. (F, OS); Mánana Trail 1 km from summit, 12 Aug 1996, Obata et al. s.n. (BISH, OSH); Manana Trail 0.8 km from summit, Obata et al. 569 (BISH), Obata et al. 570 (BISH); Manana Trail, below summit, Perlman & Obata 6484 (BISH, PTBG, US); Manana Valley, north-facing slope, Perlman et al. 17171 (PTBG); Manana Trail to summit, next ridge north of Mànana, Wood et al. 8513 (PTBG). 804 Journal of the Botanical R h Institute of Texas 1(2) 5. Lobelia gaudichaudii A. DC. in DC., Prodr. 7:384. 1839. Dortmanna gaudichaudii (A. DC.) Kuntze, Revis. Gen. Pl 2:972. 1891. Galeatella gaudichaudii (A. DC.) O. Deg. & I. Deg., Fl. Hawaiiensis, Fam. 339. 1962. Type: HAWAIIAN ISLANDS. O'aHU: sommet de la montagnes à gauche de là vallée de Onoruru [i.e., Honolulu], [8-24 Oct 1836, fide Kuykendall (1947)] Gaudichaud s.n. (HOLOTYPE: P-00520806!; isotypes: GH! G-DC! P-00520807!) Lobelia gaudichaudii var. coccinea Rock, Bull. Torrey Bot. Club 44:238. 1917. Type: HAWAIIAN ISLANDS. O‘anu: summit of Mt. Konahuanui, 1912, leg. G.W. Shaw, Rock 12742 (HoLotyPE: BISH!; isotypes: BISH[3]! GH! US). Plant 1-3 m tall. Stem 0.8-2 m tall x 1.3-5 cm diam.; cicatrices 2.5—4.5 mm tall x 7-11 mm wide, trans- versely rhombic, the corners acute. Lamina 9.5-19.5 cm long x 1.1-2.8 cm wide, oblanceolate, densely long pubescent on the abaxial midrib (rarely glabrous); margin distinctly revolute; apex acute or acuminate; base attenuate. Inflorescence 0.4-0.8 m tall, unbranched, 18-36-flowered and cylindrical in outline, its axis green, glabrous; peduncle sparsely covered with sterile bracts that rapidly decrease in size acropetally; sterile bracts 25-110 mm long x 10-18 mm wide, linear, narrowly elliptic, oblong, or ovate, the apex acuminate or acute, the base cuneate; floral bracts 18-32 mm long x 6-8 mm wide, ovate or lanceolate, the apex acute or acuminate, the base cuneate; pedicels 20-40 mm long, of more-or-less equal length throughout. Hypan- thium 6-10 mm long x 7.5-12 mm diam., broadly obconic or campanulate, 1/8-1/9 as long as the corolla glabrous; calyx lobes 13-18 mm long x 4.5-7 mm wide, ovate or lanceolate, 1.6-2.5 times longer than the hypanthium, the margin entire, the apex acuminate or acute. Corolla 60-65 mm long, red, glabrous; tube 20-30 mm long x 6-11 mm diam. at middle, slightly curved, expanding slightly towards the mouth 2—4.5 times longer than broad; dorsal lobes 30-40 mm long x 4-5 mm wide, equaling to twice as long as the tube; ventral lip 24-30 mm long x 12-20 mm wide. Filament tube 58-66 mm long, decurved slightly, 4—6 times longer than anther tube, glabrous; anther tube 4-6 mm diam.; dorsal anthers 10-13 mm long; ventral anthers 7.5-10 mm long. Capsule 23-30 mm long x 11-19 mm diam.; seeds (Fig. 1E) 1-1.5 mm long x 0.5-0.7 mm wide, chestnut-brown. Icones.—Gaudichaud (s.d.), pl. 45; Peale (1848), pg. 152; Rock (1919), pl. 56; Degener (1938b); Murata (1995), figs. 56-57 [seed only]; Smith et al. (1995), fig. 1. Figured specimen.—Rock (1919), pl. 57 [holotype of L. gaudichaudii var. coccinea]. Distribution, habitat, and phenology. —Endemic to O'ahu, known only from Montane Wet Shrublands (Gagné & Cuddihy 1990) on the summit of the southeastern Ko'olau Range, between Pu'u Pauao and Mt. Olympus, at 805-960 m. Flowering from July to October, fruits ripening November to January. Considered “vulnerable” by Wagner et al. (1999) Representative specimens: O'AHU. Ko'oLau Mrs.: windward pali of Puu Kónáhuanui, Bishop 1188 (BISH); Konahuanui, 10 Jan 1904, Bryan s.n. (BISH); near summit Konahuanui (42), 30 Jul 1922, Bryan s.n. (BISH); main ridge between Konahuanui and Mt. Olympus, Degener 7965 (NY); Bowman Trail, at divide, leg. M. Kerr, i ud 19218 (NY); near summit ur im Trail, leg. M. Kerr, Degener 21674 (B, W); mauka of Tripler Hospital, Degener 30174 (N Konahuanui 0.0 (B MO, NY); Forbes 2182.0 (BISH); between Punalu'u and Kaipapa'u, 8-13 May 1909, Forbes & Thompson s.n. (BISH); ae of Mt. umm Garber 82 (BISH); Mt. Lanihuli peak Garber 261 (BISH); Lanihuli, Garber 262 (BISH); Kipapa-Waiahole crest, Grant 7217 (BISH); summit of Konahuanui, Hillebrand 72 (K), Kipapa M on main ie Hosaka 686 (BISH, K); Konahuanui, 100 yd E of East Peak, 13 Oct 1974, Obata s.n. (B, BISH); summit of ‘Aiea Ridge tr t, 3 Nov 1974, Obata s.n. (BISH); Kipapa Trail, right ridge of Kipapa Gulch, Ozaki 561 (BISH); Punalu‘u, Rock 64 (BISH, GH), Rock 65 (BISH); Mt. Lanihuli, leg. Nelson & Stone, Rock 10003 (BISH); Konahuanui, Rock 10003 (BISH); Lanihuli, leg. Daingerfiel ¡[añ B, S); about Y2 hr Poamoho side of Castle Trail Summit along the Ko'olau Crest Trail, Stemmermann & i pid 1207 (BISH); Waipi o-Waiahole divide, Kipapa Gulch, St. John 12077 (BISH, RSA); S of Pu'u Ka'aumakua, St. John 20252 (BISH); u Konàhuanui, E. brink, St. John 20315 ee Konahuanui, Topping 3223 (NY); [above Honolulu, 25 Sep 1840 — 5 Apr 1841 (fide cm 1845),] U.S. Exploring Expedition s.n. (GH 6. Lobelia gloria-montis Rock, Monogr. Stud. Haw. Lobelioid.: 17. 1919. Lobelia gaudichaudii var. gloria-montis (Rock) H. St. John & Hosaka, Occas. Pap. Bernice Pauahi Bishop Mus. 14:121. 1938. Galeatella gloria-montis (Rock) O. Deg. & I. Deg., Fl. Hawaiiensis, Fam. 339. 1962. Type: HAWAIIAN ISLANDS. West Maur: Pu'u Kukui Mt., 5700 i aed 1910, Rock & Hammond 8209 (HOLOTYPE: BISH!; isotypes: A[4]! BISH[4]! GH! K! NY! OSH! US! W!). Thi Hammond is not mentioned as co-collector on many of the isotypes. I f pl. 58 as “Type;” Plant 1.5-5 m tall. Stem 0.4—4 m tall x 1.5—10 cm diam.; cicatrices 3-5 mm tall x 6-13 mm wide, transversely rhombic, the corners acute. Lamina 10-24 cm long x 1.3-3.8 cm wide, oblanceolate or narrowly oblong, Lammers, Revision of Lobelia sect. Galeatella 805 glabrous or margin ciliate toward base; margin subrevolute or merely thickened; apex acute or acuminate; base attenuate, cuneate, or obtuse. Inflorescence 1-1.5 m tall, unbranched or 5-7-branched, each branch 18-100-flowered and pyramidal in outline, its axis green, glabrous or pubescent; peduncle sparsely covered with sterile bracts that rapidly decrease in size acropetally; sterile bracts 30-120 mm long x 10-25 mm wide, oblanceolate, narrowly oblong, oblong, or elliptic, the apex acuminate or acute, the base cuneate; floral bracts 20-90 mm long x 6-16 mm wide, obovate, elliptic, lanceolate, oblong, or linear, the apex acute or acumi- nate, the base cuneate; pedicels 15-60 mm long. Hypanthium 8-15 mm long x 6-17 mm diam., broadly obconic or campanulate, '/.—'/, as long as the corolla, glabrous or pubescent; calyx lobes 7.535 mm long x 3.5-9 mm wide, narrowly triangular or lanceolate, ?4 as long to 31⁄2 times longer than the hypanthium, the margin entire and sometimes ciliate, the apex acuminate or acute and sometimes apiculate. Corolla 67-103 mm long, white or rarely red, glabrous (rarely sparsely pubescent at apex of lobes); tube 26-45 mm long x 7-15 mm diam. at middle, suberect or slightly curved, expanding slightly towards the mouth, 2.2-5 times longer than broad; dorsal lobes 35-70 mm long x 4-6 mm wide, 0.9-3.1 times as long as the tube; ventral lip 30-52 mm long x 12-28 mm wide. Filament tube 58-82 mm long, decurved slightly, 415—614 times longer than anther tube, glabrous; anther tube 4—6.5 mm diam.; dorsal anthers 11-16 mm long; ventral anthers 8-13 mm long. Capsule 27-37 mm long x 10-18 mm diam.; seeds [Fig. 1F] 2-2.3 mm long x 1-1.2 mm wide, honey- or chestnut-brown. Distribution, habitat, and phenology.—Known from Montane Wet Mixed Communities and Montane Wet Forest (Gagné & Cuddihy 1990) on Maui and Moloka‘i, where two subspecies occur; see below for details. 6a. Lobelia gloria-montis subsp. gloria-montis Lobelia gloria-montis var. molokaiensis O. Deg., Fl. Hawaiiensis, Fam. 339. 1938. Galeatella i ea montis var. ee. ud p (2: Deg. & I. Deg., Fl. Hawaiiensis, Fam. 339. 1962. Type: HAWAIIAN ISLANDS. ^ g Par pali, g g g- and windswept pali, 6 Jun 1928, Degener 7778 (HoLotyPE: NY!). Lobelia gaudichaudii f. bryanii H. St. John & Hosaka, Occas. Pap. Bernice Pauahi Bishop Mus. 14:123. 1938. Galeatella gloria-montis var. bryanii (H. St. John & Hosaka) O. Deg. & I. Deg., Fl. Hawaiiensis, Fam. 339. 1962. Tre: HAWAIIAN ISLANDS. West Maur: Puu Kukui, May 1910, Forbes 68.M (HOLoTYPE: BISH!; isotypes: BISH! NSW! P! US! WI). Lobelia ed f. kukuiensis H. St. John & Hosaka, Occas. Pap. Bernice Pauahi Bishop Mus. 14:123. 1938. Galeatella gloria-montis kuiensis (H. St. John & Hosaka) O. Deg. & ., Fl. Hawaiiensis, Fam. 339. 1962. Type: HAWAIIAN ISLANDS. West Maul: em au-Pu'u Kukui trail, brushy opening in rain a 4400 ft, 19 Dec 1928, Ewart 89 (HoLoTYPE: BISH!; isotype: BISH!). Lobelia gaudichaudii f. sanguinea H. St. John & Hosaka, Occas. Pap. Bernice Pauahi Bishop Mus. 14:123. 1938. Galeatel sanguinea (H. St. John & Hosaka) O. Deg. & I. Deg., Fl. Hawaiiensis, Fam. 339. 1962. Type: HAWAIIAN ISLANDS. West Maut: Pu‘u Kukui, in swamp, 4500 ft, 16 Aug 1933, Hartt s.n. (HoLoTYPE: BISH!; isotypes: BISH[4]! US!). Plant 1.5-2.4 m tall. Stem 0.4-1.5 m tall x 1.5-4 cm diam. Lamina 10-24 cm long x 1.3-3.8 cm wide, oblanceolate or narrowly oblong. Inflorescence 1-1.5 m tall, unbranched (rarely with 1 or two clearly sub- a gloria-montis f. ordinate basal branches), 18-100-flowered; floral bracts 20-47 mm long x 6-16 mm wide, obovate, elliptic, or lanceolate; pedicels 15-45 mm long. Hypanthium 8-15 mm long x 6-17 mm diam., broadly obconic or campanulate, '/.—'/,, as long as the corolla, glabrous or rarely pubescent; calyx lobes 7.5-25 mm long x 3.5-9 mm wide, narrowly triangular or lanceolate, ?4 as long to 342 times longer than the hypanthium. Corolla 67-96 mm long, white or rarely red, glabrous (rarely sparsely pubescent at apex of lobes); tube 26-45 mm long x 7-13 mm diam. at middle, suberect, 2.2—5 times longer than broad; dorsal lobes 35-70 mm long x 4—6 mm wide, 0.9-3.1 times as long as the tube; ventral lip 30-48 mm long x 12-28 mm wide. Filament tube 58-80 mm long, 415-64 times longer than anther tube; anther tube 4—6.5 mm diam.; dorsal anthers 11-16 mm long; ventral anthers 8-13 mm long. Capsule 27-37 mm long x 10-15 mm diam.; seeds [Fig. 1F] 222.3 mm long x 1 mm wide, honey-brown. Icones.—Hitchcock (1919), pl. 25-1; Rock (1919), pl. 15 [right], 17, 18; Degener (193823); Wimmer (1943), fig. 11d; Wimmer (1953), fig. 104b [seed only]; Degener and Degener (1974), p. 2; Carlquist (1980), pp. 212C [bottom], 245 [right]; Degener and Degener (1983), p. 456; Kepler (1983), p. 80 [upper right]; Sohmer and Gustafson (1987), fig. 121; Lammers (1990), pl. 57 [upper right]. Figured specimen.—Rock (1919), pl. 58 [holotype of L. gloria-montis]. 806 Journal of the Botanical R h Institute of Texas 1(2) Distribution, habitat, and phenology.—Known primarily from Montane Wet Mixed Communities (Gagné & Cuddihy 1990) on Pu'u Kukui, West Maui, at 915-1760 m (cf., Carlquist 1980, p. 352); sterile plants from Montane Wet Shrublands (Gagné & Cuddihy 1990) on eastern Molokai, at 1165-1350 m are placed here as well. Flowering July through December, fruit ripening August through May. Considered “apparently secure” by Wagner et al. (1999). Representative specimens: Moloka‘i: Papa‘ala pali, Anderson 513 (BISH); Kalua'aha Ahu., ridge E of Manawai Gulch, Warshauer & McEl- downey 2390 (BISH). West Maui: Mountains, 3 Aug 1961, Bonsey & Suthers s.n. (MSC); Pu'u Kukui, Bryan 632 (BISH); Pu'u Kukui, Dec 1928, Bryan s.n. (BISED; below summit, Pu'u exi Carlquist 553 (RSA); Pu'u Kukui, Carlquist 555 (RSA); Pu'u Kukui, Carlquist 2142 (BISH, MICH, RSA); S of Puu Kukui summit, Char & Arakawa PKWM-55-76 (PTBG); Puu Kukui, Cranwell et al. 2667 (BISH, GB, K, S), Pu'u Kukui, on road to Nakalaloa, Cranwell 2668 (BISH, GB, K, S); just below summit of Mt. ‘Eke on northwest side, Degener 7937 (B, n CAS, GH, K, MICH, MO, NY, WIS), Degener 7938a (B, CU, K, MASS, MO, NSW, NY, US, WELO, Degener 7938b (BISH, NY); Puu Kukui summit, Degener et al. 25023 pun ‘Eke, Forbes 456.M (BISH); Puu Kukui, Gillett 1793 (BISH); trail up to Puu Kukui, Gillett et al. 2444 (BISH); Keahikauo Bog, NE of ‘Eke Crater ifson 2020 (OS, RSA); Pu'u Kukui, Higashino 76-29 (BISH); summit of Mt. ‘Eke, 1870, Hillebrand s.n. (B); ‘Eke, s.d., Hillebrand s.n. (C, GH, K, NSW, S, US); Putu Kukui, Hitchcock 14735 (US); Pu'u Kukui, near summit, Hitch- cock 14836 (US); Pu'u Kukui, Hitchcock 14861 (US); from Violet Lake to near the summit, Hobdy 665 (BISH); top of mt., Mann & Brigham 462 (BISH, GH, K); top of mts., Mann & Brigham s.n. (US); ‘Eke trail, base of ‘Eke Crater, Medeiros 341 (BISH); Puu Kukui, McAlpin 635 (DUKE); Mt. Kukui, Munro 418 (BISH); Pu‘u Kukui, Munro 419 (BISH); Pu'u Kukui, Munro 615 (BISH, K); Pu'u Kukui, 16 Aug 1933, Neal & Hartt s.n. (BISH); western headwaters of Kahakuloa Str. N of ‘Eke, Oppenheimer & Kunna H119910 (BISH); Violet Lake, Perlman et al. 6884 (MO, PTBG); Pu'u Kukui, Aug 1910, Roch s.n. (A, NSW, NY); summit of Pu'u Kukui, Aug 1910, Rock s.n. (NSW, NY); summit of Mt ‘Eke, Sep 1918, Rock & Hashimoto s.n. (BISH); Puu Kukui, summit bog, Skottsberg 773 (BISH, GB, S); Pu'u Kukui, St. John 10274 (BISH); Pu'u Kukui, Tessene 1786 (WIS); Pu'u Kukui, Wilbur & Webster 834 (DUKE); Pu'u Kukui summit bogs, Wood 11505 (OSH, PTBG). 6b. Lobelia gloria-montis subsp. longibracteata (Rock) Lammers, stat. nov. Lobelia gaudichaudii var. longibracteata Rock, Indig. Trees Haw. Isl.: 78. 1913. Lobelia gloria-montis var. longibracteata (Rock) Rock, Monogr. Stud. Haw. Lobelioid.: 119. 1919. Lobelia longibracteata (Rock) E. Wimm., Pflanzenr. IV.276c: 889. 1968. Galeatella gloria-montis var. longibracteata (Rock) O. Deg. & I. Deg., Fl. Hawaiiensis, Fam. 339. 1962. Galeatella longibracteata (Rock) O. Deg. & I. Deg., Prodr. Galeatella & Neowim- meria: 6. 1974. Tyre (designated by Rock po HAWAIIAN ISLANDS. West Maur: Mt. Pu'u Kukui, 5780 ft, Aug 1910, Rock 8818 (LECTOTYPE: BISH!; ISOLECTOTYPES: A! BISH! GH se ee var. albiflora H. St. John & A.C. Medeiros, Phytologia 63:366. 1987. Type: HAWAIIAN ISLANDS. East Maur: Haleakala itt on bog margin above mid amp bog , 4200 m" [ft], 13 Jun 1982, Medeiros 4 (HOLOTYPE: BISH! ). Plant 2.4—5 m tall. Stem 2-4 m tall x 2-10 cm diam. Lamina 18-23 cm long x 1.8-4.3 cm wide, oblanceo- late. Inflorescence 1-1.5 m tall, unbranched or 5-7-branched, each branch 40-80-flowered; floral bracts 60-90 mm long x 6-10 mm wide, oblong or linear; pedicels 38-60 mm long. Hypanthium 10-13 mm long x 11-18 mm diam., broadly campanulate, 1/8 as long as the corolla, pubescent; calyx lobes 25-35 mm long x 5-9 mm wide, narrowly triangular, 2-342 times longer than the hypanthium. Corolla 78-103 mm long, white, glabrous; tube 28-44 mm long x 12-15 mm diam. at middle, slightly curved, 2.5-3 times longer than broad; dorsal lobes 45-70 mm long x 5-6 mm wide, half again as long as the tube; ventral lip 40-52 mm long x 22-26 mm wide. Filament tube 68-82 mm long, 4.5—5 times longer than anther tube; anther tube 4.5-6 mm diam .; dorsal anthers 15-17 mm long; ventral anthers 11-13 mm long. Capsule 33-36 mm long x 14-18 mm diam.; seeds 2 mm long x 1.2 mm wide, chestnut-brown. Figured specimen.—Rock (1919), pl. 59 [lectotype]. Distribution, habitat, and phenology.—Known only from Montane Wet Forest (Gagné & Cuddihy 1990) on the rim of Tao Valley, West Maui, at 1740-1760 m; and on the northwestern slopes of Haleakala, East Maui, at 1280-1700 m. Flowering from June to August. Representative specimens: West Maui: Pu'u Kukui summit and extending to east bog and Helu Peak along the ridge line, Hobdy 664 BISH). East Maui: NE outer rift on margin of Mid Camp bog, Medeiros 2076 (BISH); S rim of Kipahulu Valley, Kuiki, Haleakala, St. John & Mitchell 21237 (BISH). ACKNOWLEDGMENTS The curators and staffs of the following institutions are gratefully acknowledged for allowing study of speci- mens in their care, either via loans or during personal visits: A, B, BH, BISH, C, CAS, CU, DUKE, F, GB, GH, K, MASS, MICH, MO, MSC, NSW, NY, OS, OSH, P, POM, PTBG, RSA, S, UPS, US, W, WAB, WELC, Lammers, Revision of Lobelia sect. Galeatella 807 WIS. Figure 1 was expertly prepared by Dr. Todd Kostman, Director of the Electron Microscopy Facility at the University of Wisconsin Oshkosh. The helpful suggestions of manuscript reviewers Eric Knox and David Lorence were much appreciated. REFERENCES ANONYMOUS. 1980. Harold St. John— career synopsis and bibliography. Pacific Sci. 33:435-447. ANONYMOUS. 1991. In memoriam: Harold St. John dies at age 99. Newslett. Haw. Bot. Soc. 30(1):3-4. BENTHAM, G. 1876. Campanulaceae. In: G. Bentham and J. D. Hooker, Genera plantarum, vol. 2. Reeve and Co, London. Pp. 541—564. Brown, F.B.H. 1921. Origin of the Hawaiian flora. 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Murray Reidar Elven University of Alaska Museum of the North Natural History M |B ical / 7 Yukon Driv University of Oslo Fairbanks, Alaska 99775-6960, U.S.A. PO. Box 1172 Blindern ffdfm@uafedu NO-0318 Oslo, Norway reidar.elven@nhm.uio.no ABSTRACT Potentilla holmgrenii D.F. Murray & Elven, sp. nov. is described and illustrated. Two new combinations are made: Potentilla arenosa (Turcz.) Juz. subsp. chamissonis (Hultén) Elven € D.F. Murray and P. hyparctica Malte subsp. elatior (Abrom.) Elven € D.F. Murray. RESUMEN Se describe e ilustra porcus uns dns D.F. Murray & Elven, sp. nov.. Se hacen nue mbinaciones: Potentilla arenosa (Turcz.) Juz. subsp. Elven & D.F. Murray y P. hyparctica Malte subsp. elatior (Abrom.) Elven & D.F. Murray. While preparing a treatment of boreal and arctic Potentilla for Flora of North America it became necessary to assess how well the names of widespread arctic plants have been applied to Rocky Mountain species. A case in point is Potentilla nivea L., in the sense of Eriksen et al. (1999) not Soják (1989). Whereas P. nivea is indeed found in the Rocky Mountain region, some plants so named are morphologi- cally well beyond the limits accepted for the species throughout its circumpolar and boreal-alpine ranges. A densely tufted plant of mountain summits in Nevada and Utah is distinct by its small, compact stature, its ternate leaves with densely floccose petioles and laminae, and leaflets with fewer teeth directed forward and having rounded apices. This new species presented below is named P. holmgrenii. It can be distinguished from P. nivea by the key provided. Basal leaves densely hairy on both surfaces; central leaflets with teeth 2-4 per side, apices obtuse to rounded, directed forward, terminal tooth often orbicular and frequently overlapped by ad- jacent pair of lateral teeth; epicalyx bractlets elliptic, shorter and narrower than sepals; styles 1 - mm or more in OR without eue basal prs sags holmgrenii ely hairy on upper itl h 3-5 per side, apices acute, not direct forward, ten tooth not orbicular, not overlapped by adjacent pair of teeth; epicalyx bractlets eee or lanceolate, as wide as sepals at the base; styles less than 1.2 mm long, evenly tubular, with distinct basal papillae Potentilla nivea Potentilla holmgrenii D.E Murray & Elven, sp. nov. (Figs. 1-2). Tyee: NEVADA. Wurre PINE Co.: Mt. Moriah, 6 Jul 1966, Holmgren and Reveal 2818 (HoLoTYPE: NY; isotypes BRY, UTC). Planta pulvinata. C ] il f. d v f. I : £A EET LE vel idi apicibus obtusis vel qu: Ad Potentillae niveae L. E sed "armis foliorum supra adpressi-tomentosis, Plants perennial, forming dense cushions to 10 cm in diameter (Fig. 1) and occasionally small mats gray- ish-white or white, eglandular; caudex woody, stout, strongly ramified with branches 0.4-0.6 cm thick, densely sheathed with persistent, blackish-brown, indurate remains of stipules, petioles and some whole leaves, giving the branches a compact, columnar structure; stems ascending, 2-8 cm, densely white-floccose; T: leaves basal and cauline (0-1); stipules broadly ovate, obtuse, light brown, proximally glabrous, distally and abaxially with sparse, long, soft hairs, becoming subglabrous; petioles 0.5-2 cm, densely floccose; leaflets three, margins slightly revolute; central leaflet broadly elliptic or obovate, 0.7-1.3 x 0.4-0.6 cm, short-stipitate, base cuneate, incised half way to midvein, teeth 2-4 per side, broadly elliptic to obovate, J. Bot. Res. Inst. Texas 1(2): 811 — 814. 2007 812 Journal of the Botanical R h Institute of Texas 1(2) Fe. 1.C NT £ D, Ha hal Fic. 2. Shoot with and leaf cluster. Note shape and orientation of the teeth (Tiehm 9862, BRY 305045). Scale bar = d Reveal 2819, UTC 115752 isotype). Scale bar = 2 cm. apices obtuse to rounded Fig.2), abaxial surface with very dense tomentum, white or grayish-white floccose with some straight, soft, verrucose, subappressed hairs ca. 1 mm long on veins, adaxial surface gray, densely hairy with straight, silky, subappressed hairs ca. 1-1.5 mm long; inflorescence simple or open, 1-2) flowers; bracts lanceolate or ovate; pedicels straight, 0.4—0.7 cm in flower, 1.5-2 cm in fruit, densely floccose and mixed with a few straight, subappressed to subpatent hairs; hypanthium 1.8-2.2 mm. with | dense, straight, 1 hairs and shorter stiff hairs ca. 0.7 mm long; epicalyx bractlets narrowly elliptic, obtuse, margins revolute, gray-green or reddish, 2-2.5 x 0.5-0.9 mm, shorter and narrower than the sepals; sepals broadly lanceolate or triangular, subacute or acute, gray-green, 3-3.5 mm, petals pale yellow, broadly obovate or obcordate, retuse, + overlapping, 4-5 mm, longer than sepals; stamens ca. 20; filaments 1-2 mm; anthers ca. 0.5-0.4 mm; carpels ca. 30-40; styles tubular, 1.2-1.4 mm, thickened at base and with a few, indistinct papillae, distinctly dilated beneath stigma; achenes pale olive-green, glabrous, 1.1-1.4 x 1 mm. Habitat.—On wind-swept ridges, fellfields, rocky slopes; 3300-4500 m. Distribution.—Potentilla holmgrenii is known presently from the Schell Creek and Snake ranges of White Pine County in eastern Nevada, and the Deep Creek Range in adjacent Juab County in western Utah. Etymology.—The name was chosen to honor Noel H. Holmgren, who has long been a student of the intermountain flora. In his treat- AA y IEI „A . J he 4° Dat PRTI 813 ment of Potentilla for the Intermountain Flora (Holmgren 1997), Holmgren called attention to the compact nature of the populations in the Snake and Schell Creek ranges but refrained from treating them as distinct from P. nivea. Specim xamined: NEVADA. White Pine Co.: North Schell Peak, 31 Jul 1989, Mosely 1592A (BYU, NY, UC); North Schell Peak, 3 Ju al 1982, Lavin 4208 (NY); Shell [sic.], Creek Mts., 13 Jul 1938, Pennell and Schaeffer, Jr. 22959 (NY); Taft Peak area, 30 Jun 1985, Tiehm 9862 (BRY, COLO, NY, RM, UTC,); Mt. Wheeler, 20 Jun 1928, Cottam 3303 (BRY); Wheeler Peak, 20 Jul 1941, McVaugh 6032 (UTC); Mt. Wheeler, 19 Jul 1954, Langenheim s.n. (UC); Mt. Wheeler, 5 Aug 1985, Holmgren and Vincent 10970 (NY, UTC, UC); Mt. Wheeler, 3 Jul 1980, White and Neese 323 (BYU); NW shoulder Wheeler Peak, 9 Jul 1969, Lewis 1698 (RM); Wheeler Peak, 29 Jul 1976, Albee 3319 (NY); Mt. Moriah, 2 Aug 1999, Niles et al. 5817 (RENO). UTAH. Juab Co.: Deep Cr. Mts., 13 Jul 1983, Goodrich 19024 (BRY, NY). Additionally, our treatment of arctic Potentilla requires two new combinations that are presented below. otentilla arenosa (Turcz.) Juz. is the common, widespread arctic and boreal-alpine species to which the name P. hookeriana has been largely misapplied, with P. hookeriana sensu str. now restricted to the mountains that differ only, but significantly, in one character: the petiole hairs. Subspecies arenosa is characterized by petioles with two of Colorado, Nevada, and Utah. Potentilla arenosa comprises two largely allopatric sul r layers of hairs: long and straight and short and stiff or curly. Subspecies chamissonis has petioles with only long, straight, verrucose hairs. Potentilla arenosa (Turcz.) Juz. subsp. chamissonis (Hultén) Elven & D.E Murray, comb. nov. Basioxvw. Po- a chamissonis Hultén, Bot. Not. ad 140. 1945; P hooheriana Lehmann subsp. chami is (Hultén) Hultén; P nivea sensu tenti Soják (1989), non L.; P nivea L. subsp. chamissonis (Hultén) Hiitonen. Habitat.—On cliffs, dry heath tundra, ridge tops. Distribution.—southern Greenland, Man., N.W.T., Nunavut, Que., n. Europe. We have not been able to confirm reports by Hultén (1968) of this sul É from Alaska or the Yukon Territory. Southern arctic plants of Potentilla hyparctica Malte differ from the high arctic ones in several presum- ably independent features. The two forms are sympatric in Greenland, Baffin Island, northern Alaska, and Yukon Territory and appear there to remain distinct. We propose for the southern taxon the combination P hyparctica subsp elatior. Potentilla hyparctica Malte subsp. elatior (Abrom.) Elven & D.E Murray, comb. nov. Basionwm. Potentilla emarginata Pursh var. elatior Abrom., Biblioth. Bot. 8(Heft 42, part 2):8. 1899; P hyparctica var. elatior (Abrom.) Fernald Plants mostly 4—20 cm, sparsely hairy; leaflets broadly obovate, lobes broadly oblong, rounded or obtuse, central leaflet long-stipitate (2-3 mm), base broadly cuneate or rounded; inflorescence 1-3(-5)-flowered; epicalyx bractlets broadly oblong-ovate, 2-5 mm wide; sepals rounded or obtuse. Habitat. —In moist ericaceous heath, snowpatch meadows, moist arctic and alpine tundra, rocky out- crops, solifluction slopes, and talus. Distribution.—Greenland; Alta., B.C., Nfld. and Labr., N.W.T., Nunavut, Que, Yukon; Alaska; ne. Asia. ACKNOWLEDGMENTS Authors wish to thank the curators at BRY, COLO, NY, RENO, RM, UTC, UC for loans of specimens and Barbara Ertter and James Reveal for their careful reading of the manuscript and suggestions for improve- ment. Arnold Tiehm and an anonymous reviewer provided helpful comments. Thanks to Guy Nesom for the Latin diagnosis and to Barbara Murray for creating the figures. REFERENCES ERIKSEN, B, B. JonseLL, and O. NiLssow. 1999. Proposal to conserve the name Potentilla nivea (Rosaceae) with a con- served type. Taxon 48:165-166. HOLMGREN, N.H. 1997. Rosaceae. In: A. Cronquist, N.H. Holmgren, and PK. Holmgren. Intermountain flora: vascular 814 Journal of the Botanical R h Institute of Texas 1(2) plants ofthe Intermountain West, U.S.A. Vol. 3, Part A: Subclass Rosidae (except Fabales). The New York Botani- cal Garden, Bronx. Pp. 64-158. HuLTÉN, E. 1968. Flora of Alaska and neighboring territories. Stanford Univ. Press, Stanford. SoJAk, J. 1989. Notes on Potentilla (Rosaceae). VIII. P nivea L. aggr. Candollea 44:741-762. A NEW SPECIES OF ALOCASIA (ARACEAE: COLOCASIEAR FROM PANAY ISLAND, PHILIPPINES Melanie P. Medecilo George C. Yao n Sciences Department Philippine Horticultural Society LIE Las lle University- asmarinas ila S edli g Bank pom Cavite 4115 Quezon City PHILIPPINES PHILIPPINES Domingo A. Madulid you Division nal Museum PHILIPPINES ABSTRACT Alocasia nycteris Medecilo, Y & Madulid is d ibed ies f Aklan and Antique Panay Island Philippines The spe- NAM A Te ts bat-wing shaped blade and deeply undulated t bpi o E E E d leaf margins. The new species is most closely similar to A. sanderiana W. Bull. Key WoRps: Araceae, Philippines, Panay Island, endemic plant, threatened plant RESUMEN Se describe un isa ia Yao & PE bos nueva v nS y e ne. dn oa La especie se La a nueva especie eM Y O es Eon más AR a A. sanderiana W. Bull. There are 20 genera and about 130 species belonging to the family Araceae in the Philippines. The largest genus in the Philippines is Alocasia (Schott) G. Don with 14 species recorded so far (Hay 1999) but based on == recent field studies, the list increases to at least 18 species (Medecilo, in prep.). All species of Alocasia in the Philippines are endemic, except A. macrorrhizos (L.) G. Don which is widespread throughout the Asian tropics and never found away from human disturbance anywhere in its range. Alocasia has more than 75 species in the Malesian region, Sri Lanka, Australia, southern China and the southernmost parts of Japan. The most recent taxonomic revision on the genus in the country was done by Hay (1999). Twelve species enumerated by Merrill (1925) was reduced to ten and four new species were added, namely: A. boyceana A. Hay, A. clypeolata A. Hay, A. ramosii A. Hay, and A. scalprum A. Hay. The center of diversity of the genus is Borneo with at least 21 species (Hay 1999, 2000) and the country is the second most species-rich sub region, next to Borneo in the Malesian region. Some of the Philippine species have West Malesian and continental Asian affinity. The conservation status and local endemicity was discussed in the revisionary work of Hay (1999). Six Philippine species, namely: A. atropurpurea Engl., A. sinuata N.E. Br., A. portei Becc. & Engl., A. scalprum A. Hay, A. sanderiana Hort. ex. Bull, and A. clypeolata A. Hay were known from two or three herbarium collections taken in the wild but most of the species are known from cultivation. Most species are known in horticulture except A. atropurpurea Engl., and systematic measures for protection are desired. Several species are of ornamental interest because of the appearance of the foliage leaves. A. sanderiana Hort. ex. Bullis being used for hybrid utilization and reported as extinct in the wild. Other species include A. micholitziana Sander, A. scalprum A. Hay, A. zebrina K. Koch & Veitch, A. sinuata N.E. Br., A. heterophylla Merr. and A. ramosii A. Hay. Some of these species are being tissue cultured for commercial purposes. J. Bot. Res. Inst. Texas 1(2): 815 — 818. 2007 816 J Infl ¿VU Mal OF Das P | te of Texas 1(2) Fic. 1. Al Jecilo, Yao & Madulid Sp. Nov. A. Habit: B. venation: C. Inflorescence: D. Pistils: E. N F. Sy ice); G. Infructescence. Scale: A,B,C and G = 2 cm; D-F = 3 mm. Medecilo et al., A pecies of Al ia f the Philippines 817 Wen £1 £cl r Lal LA Iul J L 162. Close-uj i idely sagi l i Fic 3. Close-up view of infructescence. natifid margins. Alocasia nycteris Medecilo, Yao & Madulid, sp. nov. (Figs. 1-3). Tw: PHILIPPINES: Nabas, Aklan, Panay Island, 11°54'18"N and 121°59'43"E, 3 Nov 2005, Medecilo 397 (HoLotype: PNH; isotypes: DLSU, DLSU-D) Alocasia sanderiana arte similis in foliis late sagittatis profunde undulatis, sed spadicis appendice longa, interstitiis sterilibus irregulariter factis differt. | Terrestrial robust herb to 1.5 m tall; stem erect to decumbent, up to 12 cm x 2 cm; Leaves 2-4 together, subtended by cataphylls; cataphylls papery, up to 12 cm long by 2 cm wide at base, narrowly ovate, acumi- nate; petiole 45-100 cm x 1.5-2.0 cm wide, green with blackish green streaks, sheathed from one-third to one-quarter its length, marginally marcescent; blade hastate to sagittate, 20-37 cm x 35 cm wide at the middle, coriaceous, dark glossy green adaxially, paler green abaxially, membranous when dry, margin deeply undulated to subpinnatifid along margin; anterior lobe widely triangular or deltoid, acuminate at apex, 3-4 pairs of lateral veins nearly opposite diverging at 60—65? angle on each side of the anterior costa; axillary glands conspicuous, brownish green, 0.1 mm diam.; secondary veins very fine, 3 mm apart arising from the anterior costa and primary lateral veins then abruptly deflected to the margin not forming interprimary collective veins; posterior lobes diverging at 85?—95? angle, up to 24 cm long; posterior costae naked in the sinus for 3—4 cm, tip rounded, in some leaves forked, with 3 primary veins arising from the posterior costa; inner side of the posterior lobe up to 4 cm wide at the middle, lanceolate. Inflorescence 1—2 together, sub- tended by cataphylls; cataphyll whitish green, narrowly ovate, 10-12cm x 3-4 cm at the middle, mucronate to 2 mm long, thick when fresh; peduncle up to 24 cm long x 1 cm wide at base, light green with grooves at the base; spathe up to 10 cm; lower spathe tube ovoid, light green, 1.5cm x lcm diam.; limb narrowly lanceolate, cucullate, pale yellow, tip inate, separated from the limb by a short deep constriction; spadix shorter than the spathe, partly adnate to the lower spathe at 1.5 mm; female zone 10 mm long, 7 mm wide at the middle; ovaries light green, subglobose, 3 mm in diam.; stigma 3-lobed; lobes, erect, 1 mm diam.; style 0.5 mm long, brownish green; pistillodes reduced, irregular shape with holes in the center, 1-2mm wide, whitish; sterile interstice 5 mm long x 4 mm wide, cylindric; whorl comprised of diamond-shaped to rhombo-hexagonal synandrodes, 2 mm wide; male 14 x 6 mm, cylindric, whitish green; synandria irregular to rhombo-hexagonal, slightly wavy at the edges, 1.5-2 mm diam., apical pores capped by synconnective; appendix 40 mm long x 5mm wide, yellow, tapering. Infructescence oblong. Fruit a berry, globose, 10 mm diam, orange when mature, 1—3 seeded. Seeds, greenish-black, globular, 2 mm in diam. Distribution. —PHILIPPINES: Panay Island, Nabas to Ibajay, Aklan and Antique, 0-20 masl. Endemic. 818 Journal of the Botanical R h Institute of Texas 1(2) Habitat. —Remnant lowland forests and secondary forests. Common on rocky areas. Prefers shaded places along roadsides. It also grows in disturbed secondary forest near ricefields. Specimens examined: PHILIPPINES. Aklan Prov.: Nabas, living collections of Ray Ong in San Juan, Metro Manila, 3 Nov 2005, Medecilo 397; Medecilo 398 (DLSU-DH 701-702 Notes.—Alocasia nycteris resembles A. sanderiana W. Bull and A. portei Schott in having deeply undulated to subpinnatifid blade margins. It differs from A. sanderiana by its larger, taller habit, not having peltate leaves, the broadly triangular, hastate-sagittate blade, longer posterior lobes (in some individuals) and the absence of a silver white midrib and primary lateral veins. The color of the leaves of A. nycteris is green on both lower and upper surfaces while A. sanderiana is shiny, deep blackish green in the adaxial side and purple in the abaxial surface. The species described here was first introduced to the public by an aroid enthusiast, Antonio Advin- cula during a garden show of the Philippine Horticultural Society, Inc. (PHSI) in 2003. George Yao, former President of the PHSI featured the species in the International Aroid Society Newsletter Vol.5, No. 3 (Aug 2003) and named it Alocasia advincula. The species name, however, could not be considered as validly published in accordance with the International Code of Botanical Nomenclature (Greuter et al. 2000) as it lacking a Latin diagnosis, and there is no rank indicated and no citation of the holotype. Etymology.—The specific epithet ‘nycteris’ alludes to the bat-shaped leaves of this plant. Nycteris is a genus of African and Asiatic bats comprising the hollow-faced bats. The species is known by its horticultural name “Bat Alocasia.” It was recently introduced in the horticultural trade and is now becoming popular as an indoor or pot plant. It can be easily propagated vegetatively by separation of the suckers and division of the rhizomes. It is best grown under shade with adequate water supply. The plant is not only known in local markets but also in the international markets. Conservation status.—Plants are still commonly found along roadsides and in private lands near second- ary forests. At present the populations along accessible places are under threat because of over collection for horticultural purposes and from habitat destruction. No legal protection is given to the plant at present. Thus, it is highly recommended that appropriate conservation measures are imposed to prevent rapid decline of population. Mass propagation of the plant by tissue culture is an alternative to collection of the plants in the wild. No populational studies have been made and information is lacking about the biology, phenology, pollination and reproductive biology of the species. ACKNOWLEDGMENTS The authors would like to thank the Philippine Horticultural Society, Inc. (PHSD for funding the collection trip of Melanie Medecilo in Aklan. Thanks to Ray Ong for providing valuable information on the species in Panay. The Abellar family provided accommodation and warm hospitality to Medecilo in Aklan. Peter Boyce is acknowledged for his suggestion on the specific epithet and for commenting on the early draft of the manuscript. Jeff Veldkamp is acknowledged for the Latin diagnosis. Nemesio Diego, Jr., artist from the Botany Division, National Museum rendered the illustration. We are very grateful for the comments of Tom Croat and Peter Boyce. We also wish to thank Marlon C. Pareja, Chairman of the Biological Sciences Depart- ment, De La Salle University-Dasmariñas, Dasmariñas, Cavite and Director Corazon S. Alvina, National Museum for their constant support and encouragement. REFERENCES GREUTER, W, J. McNeiLL, F.R. Barre, H.M. Bunper, V. DEMOULIN, T.S. FituEiRAs, D.H. Nicotson, P.C. Sit vA J.E. Skoc, P. TREHANE, N.J. TurLAND, and D.L. HAwKswortH (eds). 2000. International code of botanical nomenclature (St. Louis Code). Regnum Veg. 131. Koeltz Scientific Books, Konegstein. http://www.bgbm.fuberlin.de/iapt/nomenclature/ code/SaintLouis/0000St. Luistile.htm Hay, A. 1999. The genus Alocasia (Araceae-Colocasieae) in the Philippines. Gard. Bull. Singapore 51:1-41. Hay, A. 2000. Alocasia nebula. Bot. Mag., n.s. 17(1):14—18, pl. 381. Mebecito, M.P in prep. Monograph of the Genus Alocasia in the Philippines. FOLIA TAXONOMICA 2. NEW SPECIES OF PASSIFLORA SUBGENUS PASSIFLORA (PASSIFLORACEAE) FROM THE GUIANAS Christian Feuillet Department of Botany, MRC-166 Smithsonian Institution, PO. Box 3712 Washington, D.C. 20013-7012, U.S.A. feuillec@si.edu ABSTRACT Two new species of uoc re d with ps flowers are T Vids the Guianas. Du new an poe in subgenus d fro om P. coccinea Į row leaves and bracts. P compen from Guyana and French Guiana | j leaves wi ntinuoush its glandula dL ith P. coccinea an 1 1 1 1 E o B A key to p persecti tone is edad: RÉSUMÉ D 11 peces de Passiflora (Passifl d à fl dé Les nouvelles espéces e au sous-genre Passifl Coccinea. Passiflora arta de Guyana est facile à distinguer de P. coccinea grâce à ses feuilles et ses bractées étroites. Passiflora compar de Guyana et de o a t feuilles p i in la marge n'est pas sandals de eo continue. La couronne et operculum des d P. coccinea. U lé E E la supersection C Passiflora L. is the largest genus of the Passifloraceae with around 530 described species (Feuillet & Mac- Dougal 2007). Although members of the genus Passiflora L. are usually very visible and well collected, more than 27 species of passionflowers have been added to the flora of the Guianas since Killip (1937 & 1938) who mentioned the presence in the Guianas of 33 species. Including the two new species described below, this is an 8296 increase. Although they have bright red flowers, the two species described herein have been overlooked for a long time because they are uncommon and close enough to the most common species in the Guianas, P. coccinea Aubl., to have been confused with it and rarely collected. The two new species belong in subg. Passiflora supersect. Coccinea Feuillet & J.M. MacDougal. Passiflora supersection Coccinea includes 13 South American species, two of them expanding into Central America. Traditionally these species have been included in subg. Distephana (Juss. ex DC.) Killip (now a supersection). They are here separated from Distephana on the base of a series of morphological characters. The leaf blades are hairy, glandular at margin, dentate or serrate or crenate. The bracts are foliaceous, usu- ally red. The erect and compact corona is much smaller than in supersect. Passiflora, Stipulata Feuillet € J.M. MacDougal and Laurifolia (Cervi) Feuillet & J. M. MacDougal, but larger than the reduced corona of supersect. Distephana (DC.) Feuillet & J. M. MacDougal and Tacsonia (Juss.) Feuillet & J[.M. MacDougal. Passiflora arta Feuillet, sp. nov. (Fig. LA-C). Wee: GUYANA. U. Takutu - U. EsstQUIBO: Acarai M of unnamed peak, 6 km S of Sipu River, 1?22'N, 58°56'W, 700 m, fl., 30 Aug 1998, H.D. Clarke 7230 roue US). LES T T4 1 Passiflora arta in subg. xa ora supersect. i P. llipticis, corona uniseriata, fer operculo longiore dif Liana probably reaching the canopy. Vegetative parts with short rufous trichomes. Young stems terete, stri- ated. Tendrils long, base strong and straight. Vegetative bud although part of the axillary complex (leaves, stipules, pedicel, tendril), clearly in apical position on the stem, 5-7 mm away from the petiole, prophylls acute at apex. Stipule linear, 6 x 0.2 mm, soon deciduous. Leaves: petiole 5-13 mm long, with 2 lateral glands at the base; glands sessile, round, ca. 1 mm in diam., glabrous; blade narrow-elliptic, 7-7.5 x 1.2-1.5 cm, acuminate, mucronate at the apex, rounded to acute at the base, the margin shortly glandular serrate J. Bot. Res. Inst. Texas 1(2): 819 — 825. 2007 1(2) AA £T. Lat +h 1@Xd5 Journal of 820 PTE” IO Üj : S h Il E : EN CIAT M jf ff pium a J A-C from Clarke 7230 (US); D-G I. € rl J iole, stipules, tendril, ls: F. Bracts: G. Fl D. Leaf: E. A m Fic. 1. A-C. Passiflora arta: A. S from Feuillet 1423 (US). section; D-G. P Feuillet, N e £D Ora | è 4l Cart 821 to bi-serrate, drying olivaceous adaxially and yellow-brown abaxially, 4-6 main lateral veins on each side of the midrib, tertiary venation reticulate. Inflorescence sessile, 1-flowered. Pedicels 11 cm long, with short pubescence. Bracts three, verticillate at 2.5 mm below the flower, pink red, narrow-elliptic, 5-5.5 x 1-1.3 cm, free to base, acuminate at the apex, glandular crenate to loosely serrate at the margin, with short scat- tered trichomes on both surfaces. Flowers pink red: hypanthium 9 mm long, round at the base, then short- cylindric, short pubescent outside, glabrous inside; sepals narrow-oblong, 3-3.5 cm long plus a 9 mm long, subapical awn, 3-7 mm broad, short pubescent in the parts exposed in bud, otherwise glabrous; petals similar to the sepals, thinner, not awned, glabrous; corona in 1 row of filaments 8-12 mm long, subulate, erect, slightly leaning against the androgynophore, linked ina 1.5 mm long membrane at the base, glabrous; operculum borne at the top of the hypanthium, membranous, shortly laciniate at the margin, 9-10 mm long, dependent to the base of the hypanthium, then shortly recurved inward, glabrous; disc annular at the bottom of the hypanthium; limen tubular membranous, dentate at the margin, 2 mm long, surrounding the base of the androgynophore, glabrous; androgynophore cylindric, ca. 2 cm long, glabrous; stamen glabrous, filaments flat, 13-15 x 2 mm, joint for 2-3 mm in a membrane at base, anthers dorsifixed, rectangular, 7 x 2 mm; gynophore shorter than the membrane at the base of the stamens; ovary 3-3.5 mm long, 2-2.5 mm in diam., subglobose, densely pubescent; styles 10-12 mm long, glabrous; stigmas ca. 2.5 x 3.5 mm, glabrous. Fruit not seen. Distribution.—Passiflora arta is known only from the type locality, Acarai Mts., Upper Takutu - Upper Essequibo region in Guyana. It inhabits a dense forest on brown sand at 700 m. The forest was dominated by Clusia L., Oenocarpus Mart., and Palicourea Aubl. species. Because of leaves with serrate margin, bracts with glandular-serrate margin, red bracts and perianth, short corona filaments, straight, and forming a cone appressed to the androgynophore, Passiflora arta sp. nov. clearly belong in P. supersect. Coccinea. In this group it is close to P. coccinea (figure 1 D-G; plate 105a in Feuillet 2002b) which has a large Amazonian and Guianan distribution. The Guianan P. arta has nar- row-elliptic rather than ovate leaves and bracts, a uniseriate rather than triseriate corona, and an operculum dependent nearly to the bottom rather than to the middle of the hypanthium. Differences with other species of Passiflora supersect. Coccinea are summed up in the key below. The flower measurements have been done on the dried flower of the holotype. Etymology.—The Latin specific epithet means *narrow" (street) or “thin” (person). It refers to the narrow leaves and bracts of the new species as compared to the sympatric P. coccinea. Passiflora compar Feuillet, Sp. nov. (Figs. 2, 3). Tee: GUYANA. PoTARO-SIPARUNI: 1-3 km NNE of Kato along trail to Para- makatoi, 4?0'N, 59°49’W, 675 m, 19 Mar 1989, fl. & fr., LJ. Gillespie 873 (HoLotypE: US (f1.); isotypes: BRIT, MO, K, NY, US (fr.)). Passiflora compar in subg. do dud EE Coccinea P P. coccineae affinis; foliis subtiliter biserratis, tubo floris breviore, corona biseriata, prima serie | ti, operculo vix breviore quam tubum differt Liana 8 m tall (de Granville 12498), m reaching the canopy. Vegetative parts with short white or rufous trichomes. Young stems terete to subangular, striated. Tendrils long, base thin and straight. Vegetative bud although part of the axillary complex (leaves, stipules, pedicel, tendril), clearly in apical position on the stem, 5-7 mm away from the petiole, with 6 long acuminate prophylls. Stipule filiform, 8-12 x 0.2 mm, dentate-glandular at margin, soon deciduous. Leaves: petiole 5-13 mm long, with 2 lateral glands at the base, one petiole (of de Granville 12498) with a pair of smaller glands at the apex; glands sessile, round, ca. 1 mm in diam., glabrous; blade oblong, 3-7 x 1.5-3 cm, acuminate, cuspidate at the apex, rounded at the base, the margin shortly glandular serrate to bi-serrate, drying olivaceous adaxially and yellow-brown ab- axially, 3-5 main lateral veins on each side of the midrib, tertiary venation reticulate. Inflorescence sessile, 1-flowered. Pedicels 5-9 cm long, with short pubescence. Bracts three, verticillate at 2-3 mm below the flower, pink red or orange red, oval, 4.5—5.5 x 2-3 cm, free to base, rounded at the apex, loosely glandular crenate at the margin to glandular serrate at the apex, with short scattered trichomes on both surfaces. Flow- ers: hypanthium 7-8 mm long, round at the base, then short-cylindric, short pubescent outside, glabrous inside; sepals narrow-oblong, 2.5 cm long plus a 7 mm long, subapical awn, 8 mm broad, “pale red” (sic!) 822 Journal of the Botanical R h Institute of Texas 1(2) Cathy faaqale- 2003 Fic. 2 P, 77) AC uhi 14 stipule, tendril, pedicel, and vegetative | ain: C. Node with I tiole, a ion. A-B from Jansen-Jacobs et al. 1262 (US); C: from de Granville 12498 (US); D-E from Jansen- P bud: D. Flower, face view: E. F Jacobs et al. 1262 (US) & photograph by C. Feuillet. Feuillet, N e £D Ora | è 4l Cart 823 Fic. 3. Passifl par Feuillet, Gillespie 873 (photo L. Gillespie). adaxially, red abaxially, short pubescent in the parts exposed in bud, otherwise glabrous; petals similar to the sepals, thinner, not awned, red, glabrous; corona in 2 rows of filaments, filaments 5-7 mm long, outer subulate, inner filiform, red or pink, erect, slightly leaning against the androgynophore, free, glabrous; operculum membranous, shortly laciniate at the margin, 8-9 mm long, borne at the top of the hypanthium, dependent to the base of the hypanthium, then long recurved inward, glabrous; disc annular at the bottom of the hypanthium; limen tubular membranous, upper third laciniate, 5 mm long, surrounding the base of the androgynophore, glabrous; androgynophore cylindric, ca. 1.-2 cm long, glabrous; stamen glabrous, filaments flat, 12-15 x 1 mm, joint for 3-4 mm in a membrane at base, anthers dorsifixed, rectangular, 5-6 x 2.5 mm, green, pollen yellow; gynophore shorter than the membrane at the base of the stamens; ovary 5 mm long, 3 mm in diam., subglobose, densely pubescent; styles 10-12 mm long, thin, red or rose with red spots, with scattered trichomes; stigmas ca. 2.5 x 2 mm, yellow, glabrous. Fruit not seen. Distribution.—Passiflora compar is known in Guyana from the Kanuku Mts. (Upper Takutu-Upper Es- sequibo) and near Kato and in French Guiana from the upper Maroni basin, the Saúl area, and the hills SW of Cayenne. It inhabits moist forest below 400 m of elevation, near rivers or on slopes. Because of serrate leaf margins, bracts with glandular-serrate margin, red bracts and perianth, short corona filaments, straight, and forming a cone appressed to the androgynophore, Passiflora compar sp. nov. clearly belong in P. supersect. Coccinea. In this group it is close to P. coccinea (Fig. 1 D-G) which has a large Amazonian and Guianan distribution. The Guianan P. compar has thin rather than leathery or rubbery leaves and bracts, a corona in two rows rather than three, and an operculum dependent nearly to the bottom rather than to the middle of the hypanthium. Differences with other species of Passiflora supersect. Coccinea are summed up in the key below. Etymology.—The Latin noun in apposition, “compar” means “equal, companion” and refers to the second row of corona filaments nearly as long as the first row, and to the fact that this new species stands close to P. coccinea. 824 £L Dag *..1D Journal UI h Institute of Texas 1(2) PARATYPES: GUYANA. U. Takutu - U. Essequibo: Kanuku Mts., 3°08’N, 59°23’W, 280—400 m, fl. & fr., 20 Feb 1985, MJ. Jansen-Jacobs, C. Feuillet, P. Hiepko, L.E. Skog & B.J.H. ter Welle 330 (B, BBS, BRG, CAY, K, P, NY, U, US); 3°21’N, 59?29"W, 140-320 m, 26 Nov 1987 Jansen-Jacobs et al. 1262 (BRG, U, US). FRENCH GUIANA: Bassin de la Litani (= upper Maroni), Saut Lavaud, 2?45'N, 54?14"W, 140m 15 Sep. 1994, de Granville 12498 (CAY, US). KEY LOCTHE:SPEGIES-OFP.PASSTEELORA-SUBERSECTz ‘COCGINEA 1. Bracts linear, less than 5 mm wide. 2. Peduncle 10-15 cm long; petals white, corona 3 ranked, the 2 outmost of filaments free to ba se ea Wurdack 2. Peduncle ii to 5 cm long; petals pink or red, corona 2 ranked, both fused in a membrane half t third of the w . Bracts ues de cong lanceolate, mostly more than 5 mm wide. 3. Leaves always unlobe HR Hu ER Rodschied 4. S nd crenate 5. Operculum erect P. aimae Annonay & Feuillet 5. Operculum decumbent P. longicuspis Vanderplank 4. Leaf margin viable or serrate 6. Stipules fimbria P. callimorpha Harms 6. E dus serrate or linear. 7. Operculum filamentose. 8. Corona with 4 rows of filaments P. quadrifaria Vanderplank 8. Corona with 1 or 2 rows of filaments. 9. Corona with two rows of subulate filaments P. araujoi Sacco 9. Corona with one row of pad filaments P. tholozanii Sacco 7. Operculum membranous, fimbria 10. Corona with one row of filam P. arta Feuillet 10. Corona with two or three rows M ues 11. Corona with two rows of filaments subequal in length P. compar Feuillet 11. Corona with three rows of filaments. 12. Outermost corona dee erect, slightly diverging, dark red to violet-black miniata Vanderplank 12. Outermost corona filaments erect, converging around the androgynophore, white red. 13. Outermost corona filaments red P. sp. (Guyana) 13. Outermost corona filaments white P. coccinea Aubl. 3. Leaves 3-5-lobed. 14. Bracts fimbriate P. involucrata (Mast) Gentry 14. Bracts entire, dentate, or serrate. 15. Bracts more than 10 mm wide. 16. Stipules 3-5 mm long; third row of the corona tubular 16. Stipules 6-15 mm long; third row of the corona filamentose . P vitifolia var. bracteosa (Karsten) Killip , margaritae Sacco 1:5: ae less than 10 mm wide. 7. Stipules incised-serrate P. buchtienii Killip if p. cl serrate. Outer rank of the corona red; operculum fimbriate P. vitifolia Kunth RM . P. speciosa Gardner e Outer a corona white; operculum crenate or denticulate ACKNOWLEDGMENTS I would like to thank the curators of the herbaria CAY and U for lending the material in their care. I am grateful to Cathy Pasquale-Johnson for the fine illustrations and to three reviewers who made this paper better through their corrections and suggestions. As always, discussions with John MacDougal have been stimulating. This is number 83 in the Smithsonian's Biological Diversity of the Guiana Shield Program publication series. REFERENCES FeuiLLeT C. 2002b. Passifloraceae. In: S.A. Mori, G. Cremers, C. Gracie, J.-J. de Granville, S.V. Heald, M. Hoff, and J.D Feuillet, New species of Passiflora from the Gui 825 Mitchell, eds. Guide to the vascular plants of central French Guiana. Part 2. Dicotyledons. Mem. New York Bot. Gard. 76:566-570. FeuiLLET, C. and J.M. MacDoucaL. 2003 [May 2004]. A new infrageneric classification of Passiflora L. (Passifloraceae). Passiflora 13(2):34-38. Feuittet, C. and J.M. MacDoucat. 2007 [Dec. 2006]. Passifloraceae. In: K. Kubitzki, ed. The families and genera of vascular plants IX:270-281. Springer, Berlin. HANSEN, A.K., L.E. GILBERT, B.B. Simpson, S.R. Downie, A.C. Cervi, and R.K. Jansen. 2006. Phylogenetic relashionships and chromosome number evolution in Passiflora. Syst. Bot. 31:138-150. Kiur, E.P. 1937. Passifloraceae. In A.A. Pulle, ed. Flora of Suriname 3(1):306-327. KiLuP, E.P. 1938. The American species of Passifloraceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 19:1-613. MacDouca. J.M. and C. Feuittet. 2004. Systematics. In: T. Ulmer and J.M. MacDougal, Passiflora: Passionflowers of the World. Timber Press. Pp. 27-31. 826 Journal of the Botanical R h Institute of Texas 1(2) BOOK REVIEWS Thomas G. Lammers. 2007. World Checklist and Bibliography of Campanulaceae. (ISBN 978-1-84246- 186-0, pbk.). Royal Botanic Gardens, Kew, Richmond, Surrey, TWO 3AB, UK. (Orders: www.kew.org, +44 (0)20-8332-5646 fax). $185.55, 675 pp., 8 1/4" x 11 5/8". From the 67 species of Adenophora to the two species of Zeugandra, Thomas Lammers seems to cover all the known genera, species, nde: and nee iu in the pu daa nu As science goes, no sooner s the oun is UE there are taxa montis subsp. longibracteata, Lobeli 1 Lobelia wahiawa) E in this issue of J. Bot. Res. Inst. Texas d aE 5 789- d 1 n easily con. fth Campanulaceae “Eighty -four g d encompassing e e 2319 2 391 subspecies and 27 pL Mad another 8135 names are ies as Mid j D. checklist is alphabetical by the “checklist” d SESS for the pm Not sure e what subfamily a eons bela dne t pe that m each genus ais n the number o I g n known, and additional notes. The bibliography in and of teal is quite a contribution The author says it is selective rather than ee a it’s pretty exhausting just to read through the 11 pages of references under the eae Campanula or the 12 pages provided for the B s ia. The checklist includes every validly | Baa Tense PIRR | at or below specific rank, as well as all validly published g place and date of EM Nomenclatural types are provided for each generic name. The format of tl tire checklist i ht to follow and find. Thank goodness! Lord I didn't know there were so many genera in the Campanulaceae; I'll bet the Good Lord was surprised to see so many names on the list as well. A useful work on an important worldwide family.— Barney Lipscomb, Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. Rick Darke. 2007. The Encyclopedia of Grasses For Livable Landscapes. (ISBN 978-0-88192-817-4, hbk.). Timber Press Inc, 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $59.95, 487 pp., color photographs, 2 color Hardiness Zone maps, USDA and European, 10" x 11". Gra & the Global Garden Grass Families (Poaceae, Cyperaceae, Juncaceae, Restionaceae, Typhaceae) The Beauty of Grasses Design with Grasses Cultivation & Maintenance Grass Names & Nomenclatu PADS S I MN ure Encyclopedia of Grasses, Sedges, Rushes, Restios, & Cattails Grasses are beautiful and are used more and more in garden landscapes. This book has excellent introductory discussions of PR gardens, including plant im cultivation, propagation, and maintenance (Chapters 1—6), but the 'encyclopedia' is the heart, covering hundreds of species (in alphabetical or y genus and species, intermixing gea of the five different families), all with DM: color photos and comments on nativity, Pre and cultivation. Sixty-seven species of Carex! A great bargain, even at list price, and find reductions of nearly 40% on the internet.—Guy L. Nesom, Botanical m Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. J. Bot. Res. Inst. Texas 1(2): 826. 2007 CONTRIBUTIONS TO THE GENUS FESTUCA (POACEAE: POEAE) IN MEXICO ANI A KEY TO THE CENTRAL AMERICAN SPECIES Stephen J. Darbyshire Agriculture and Argi-Food Canada Eastern Cereal and Oilseed Research Centre Central a Farm, Wm. Saunders Bldg. #49 wa, Ontario, KIA 0C6, CANADA. ABSTRACT new species, Festuca longiligula (Poaceae: Poeae), is described from the state of Coahuila, Mexico. Names and descriptions are aA for two nothospecies from central Mexico, Festuca meonzaler: Lanne and F. crece putative RA OLE: hil ] F. livida x F. tolucensis, respectively. O Hel leria) TE are presented. A key to the | ee x F. pecies of Festuca in Central America T —— Key Wonps: Festuca, new species, Mexico, Central America, hybridization, nothospecies, subgenus Helleria, keys RESUMEN Sed il ie, F P P delestado de Coahuila, México. Se d 1 1 p dos nothospecies de exito central, A «gonzalez- Eme F. xmiscella, híbridos 1 de F. livida x F. hepl phila y F. livida x F. tolucensi S t TUE. Pu ; i i g Helleria) livida. Se aport | de | peci idas de Festuca en América Central. The pioneering work of E.B. Alexeev on the taxonomy of Festuca in Mexico and Central America (Alexeev 1980, 1984) set the ground work for further detailed investigations on the genus. Not all of his described taxa have been widely accepted (cf. Soreng et al. 2003) and some await further study. A working list of species is given in Table 1. This paper describes new species (including nothospecies), contributes some additional observations and presents an updated key to the species known from Mexico and Central America. Festuca longiligula S.J. Darbyshire, sp. nov. (Figs. 1, 2A, 2B, 3A) Tre: MEXICO. Coanuia: SO de Saltillo, ceja y ladera S de Sierra La Viga, subalpina, 3700 msnm, 22 Aug 1986, [A.] McDonald 2102 (HoLotypE: TEX: isotype: WIS, XAL (non vidi) Plantae perennes caespitosae, innovationibus intravaginalibus praeditae. Culmi 70-90 cm alti, leves. Vaginae foliorum ima basi tan- tum integrae, leves vel scabrae; vaginae foliorum emortuorum in fibras longitudinales non solutae; ligulae scariosae, (2-)5-9.5 mm longae, laceratae; laminae foliorum innovationum conduplicatae, 0.8-1.5 mm diametri, abaxialiter scabrae, adaxialiter hirsutae, cum 9-11 nervis, 7-11-costatae, ug qon de abaxialiter has continuo i d ao I fasciculis cum venas g BOGE j KO Paniculae 10-22 cm longae, S laxae, axe distali et ramulis er Spiculae (6-)7.5-13 mm longae, ovatae ad ellipsiodeae, cum 4-7 flosculis; glumae ovatae ad lanceolatae, scariosae vel margine late scariosae, glabrae vel pleurumque scabrosae ad hispidulae dimidi ti; glumae inferiores 3.4—5.5(—5.7) mm longae, cu nervis; glumae superiores 4.5-6.5 mm longae, cum (123(-5) nervis; ice usque 1.5 mm longae, hispidulae unilateraliter; lemmatae (5-)6-7 mm longae, lanceolatae ad anguste ovatae, be vel apice scabra ad hispidula, cum = a nervis Ms apie late scariosa, obtusa ad abrupte acuta, interdum emarginata 5m ad plus minusve bidentata, longa I 1ga; paleae 5 m longae, apice bidentatae, inter venas et saepe in marginalibus distalibus hispidulae, venae EN ciliatae prope apicem ad 1/2; T. 2.5-3.3 mm longae; ovarium apex sparsim o hispidulus. Caryopses 3.5-4.5 x Plants perennial, caespitose, shoots intravaginal. Culms erect or slightly decumbent at the base, 70-90 cm tall, smooth and glabrous. Leaf sheaths with the margins closed only at the base, smooth and glabrous or sometimes scabrous, not decaying into longitudinal fibres; auricles absent; ligules scarious, (2-)5-9.5 mm long, lacerate; leaf blades of vegetative shoots conduplicate, 0.8-1.5 mm in diameter, abaxially scabrous, adaxially hirsute, with 9-11 veins and 7-11 adaxial ribs; sclerenchyma in a continuous abaxial band, fas- cicles joining the major veins with abaxial epidermis and with adaxial fascicles opposite veins sometimes joining veins to form girders (Fig. 2); bulliform cells absent or inconspicuous. Inflorescence paniculate, J. Bot. Res. Inst. Texas 1(2): 827 — 840. 2007 £L Dag *..1D hi titit f Texas 1(2) 828 Journal of TABLE 1. Sy f the genus A M | [A ll | ( ly from | Species Mexico (States) Central America (other countries) F. aequipaleata E. Fourn., Biol. Centr-Amer. Veracruz 3(20):581. 1885 F. aguana E.B. Alexeev, Bot. Zhurn. (Moscow & | d) 66:1493, f. 1. L ningrad) 1981. Rupr., Bull. Acad. Roy. Sci. Bruxelles 9236. 1842. F. arundinacea Schreb, Spicil. Fl. Lips.:57, 1771 E aperea El p Alexeev, Bot. Zhurn. (Moscow & 1409 | PCA F. bajacalforniana Gonz.-Led. & S.D. Koch, Novon 4:28-30, f. 3. 1 F. beamanii 3 a Dec Bot. Zhurn. (Moscow & 0-1501, f. 2. 1981 12V, ft. E.B. e Bot. Zhurn. (Moscow & => Leningrad) 66:1496, f. 1.1981. F. breviglumis Swallen, Contr. U.S. Natl. Herb. 29:398. 1 F. callosa (Piper) St. Yves, Candollea 2:291. 1925. F. cartagana E.B. Alexeev, Bot. Zhurn. (Moscow & Leningrad) 67:1291-1292, f. 1. 1982 F. a eat Swallen, Ann. Missouri Bot. Gard. 30:116. 1943. F. is Gonz.-Led. & S.D. Koch, Novon 4: 27-27,£.2.1 F. diclina SEN Novon 5:129, f. 1, 2. 1995. F. filiformis Pourret, Hist. & Mém. Rea Roy. Sci. Toulouse 3:319. 1788. k dab d Darbys E.B. Alexeev, Bot Ler :1498 B [= EEG ?] F. hephaestophila Steud., Syn. Pl. Glumac. 1:310. 1854 F. herrerae Davidse, Novon 2:322-324, f. 1. 1992. F. hintoniana E.B. Alexeev, Bot. Zhurn. (Moscow & Leningrad) 67:1292, f. 1. 1982. F. jaliscana E.B. Alexeev, Bot. Zhurn. (Moscow & Leningrad) 66:1493-1495, f. 1. 1981. F. ligulata Swallen, Amer. J. Bot. 19:436, f. 1. 1932. F. livida (Kunth) puta Syst. Veg. 1:353. 1825. F. longiligula Darbys F. ads (E. Fourn.) JA ol., Bol. Soc. Bot. Mexico 23:165. 1958 a (Moscow & J | 1 d) Hl F. xmiscella Darbysh. is eps E.B. Alexeev, Bot. Zhur. 66: —-1498, f. 2. 1981. E ovina L., Sp. Pl 1:73-74. 1 F. panamica E.B. Alexeev, " Zhurn. (Moscow & Leningrad) 67:1290-1291, f. 1. 1982. Chiapas Distrito Federal Durango Guerrero, Hidalgo, Jalisco, México, Michoacán, Morelos, Nuevo León, Puebla, Tlaxcala, Veracruz México, Veracruz México Baja California Veracruz Jalisco, Michoacán, Morelos, Oaxaca Oaxaca, Puebla Coahuila Chihuahua México, Tlaxcala, Veracruz México México, Nuevo León, Puebla, Tamaulipas, Tlaxcala, Veracruz Coahuila, Nuevo León Jalisco oahuila México, Puebla, Tlaxcala, Veracruz Coahuila Chiapas, Chihuahua, Distrito Federal, Hidalgo, Jalisco, México, Michoacán, Morelos, Oaxaca, Puebla, San Luis Potosí, Tamaulipas, Veracruz México Distrito Federal, Durango, Jalisco, México Puebla, Veracruz Guatemala, Panama Costa Rica, Guatemala, Costa Rica, Guatemala Guatemala Costa Rica, Guatemala, Panama Costa Rica, Panama Costa Rica, Panama Costa Rica Guatemala Costa Rica, Guatemala Costa Rica, Panama Honduras Costa Rica Panama Darbyshire, F t in Mavi Val to tl PS Y | . k 829 Taste 1. continued. Species Mexico (States) Central America (other countries) F. pringlei St. Yves, Candollea 2:305-307, f. 53. 1925. Chihuahua, Durango, Jalisco, San Luis Potosí F. roblensis Gonz.-Led., Novon 8:147-149, f. 1. 1998. Guanajuato, Zacatecas F. rosei Piper, Contr. U.S. Natl. Herb. 10:45. 1906. Distrito pa ee Jalisco, México, Guatemala Michoacan, Morelos, Tlaxcala, Veracruz E rubra L, Sp. Pl. 1 753. Chihuahua, México, Nuevo León Costa Rica zedowskiana E E oim Bot. Zhurn. (Moscow & México pros 66:1500, f. 2. 1981 F. subverticillata (Pers) E.B. Alexeev, Novosti Sist. based on a probable label error cae t. 17:52, f. 2. 1980. FUE Pr Alexeev, Bot. ps (Moscow & Panama Leningrad) 66:1495-1496, f. 1. 1981. F. talamancensis Davidse, oo 2:324, f. 1. 1992. Costa Rica F. E Gonz.-Led. & S.D. Koch, Novon Michoacán 4:25-27, f. 1 F. nd qus Nov. Gen. Sp. 1:153. 1815. Chiapas, Distrito Federal, Jalisco, México, Costa Rica, Guatemala Michoacán, Morelos, Oaxaca, Puebla, San Luis Potosí, Tlaxcala, Veracru F. valdesii Gonz.-Led. €: S.D. Koch, Novon 8: Coahuila, Nuevo León, Tamaulipas 149-151, f. 2.1998. F. willdenowidia Schult. & Schult. f, Add. ad Mant. Distrito Federal, México, Michoacan Guatemala 3:650. 1827. Puebla, Tlaxcala, Veracruz 10-22 cm long, pyramidal, lax, rachis glabrous below and scabrous above; branches lax, 1(-2) per lower node, scabrous. Spikelets (6-)7.5-13 mm long, ovate to ellipsoid, more or less distal on the branches, with 4-7 florets; glumes ovate to lanceolate, scarious or with broad scarious margins, glabrous or frequently scabrous to hispidulous in the upper half; lower glumes 3.4—5.5(-5.7) mm long, with 1(-3) veins; upper glumes 4.5-6.5 mm long, with (1-)3(-5) veins; rachilla up to 1.5 mm long, unilaterally hispidulous op- posite to the floret; lemmas (56-7 mm long, lanceolate to narrowly ovate, glabrous or apically scabrous or hispidulous, with 5(-7) obscure veins, the apex broadly scarious, obtuse to abruptly acute, sometimes emarginate to more or less bidentate, awnless or with a short mucro arising between the lobes to 0.5 mm long; paleas 5-7 mm long, bidentate, apically hispidulous between the veins and sometimes on the up- per margins, veins scabrous-ciliate in the upper 1/2-2/3; anthers 2.5-3.3 mm long; ovary apex sparsely hispidulous; styles 2. Caryopses 3.5-4.5 mm long, 0.8-1.5 mm wide. Additional collection examined: MEXICO. Coahuila: Mpio. de Arteaga, Sierra la Viga 6 km al E de Jame, Puerto Maravillas, 25?22'N 100°34'W, bosque de Pinus rudis, P. ayacahuite, Pseudotsuga, Abies, Quercus, con arbustos de Ceanothus y Pinus culminicola, 3000-3150 m, 16 Sep 1989, J.A. Villarreal & et al. VR-1983 (TEX). Fig. 2B Festuca longiligulata belongs to subgenus Festuca section Breviaristatae and is similar to F. thurberi Vasey of the southern Rocky Mountains in the dense caespitose habit, glumes thinner in texture than the lemmas and long ligules. It differs from the latter in the more ovate lemmas which are obtuse (sometimes abruptly acute) to somewhat truncate or emarginate with a wide scarious margin, while in F. thurberi the lemmas are more narrowly ovate to lanceolate and tapering to an entire acute apex with narrow scarious margins (Fig. 3). The anthers of F. longiligulata are usually shorter, 2.5-3.3 mm, versus 3-4.5 mm, and the ovary apex is sparsely hispidulous, versus densely hispidulous. These characters and the isolated distribution in the Sierra Madre Oriental suggest that F. longiligula is a distinct species. It differs from F. ligulata in the more densely caespitose habit, usually longer ligules ((2-)5-9.5 mm, versus 2.5-3.5 mm), lemmas (5-7 mm, versus 4.5-6 mm) and anthers (2.5-3.3 mm, versus 1.5-2.6 mm): £L Dat Journal of 830 HERBARIUM Fig. 1. Fi Darbyshire, F t in Meyi A Rote A : 3 T Fic. 2.1 tion shoot leaf blad ions of Fe longiligula. A — Holotype, McDonald 2102; B — Paratype, Villarreal et al. VR-1983. Fic. 3. Lower lemmas. A — A longiligula, McDonald 2102 (hol ); B - Festuca thurberi, Swallen 6608 (DAO). Festuca subgenus Helleria In hi ; f Festuca Alexeev (1980) established F. subgenus Helleria (E. Fourn.) E.B. Alexeev (Novost. Sist. Vyssh. Rast. 17:51. 1980; = Helleria E. Fourn., Mex. Pl. 2:128. 1886, non Nees & Martius, 1824; Hellerochloa Rauschert, Taxon 31:561. 1982) in which he included F. livida as the sole species. The long glumes (longer than the lemmas and Do uec the whole spikelet) and the unusual strongly drooping panicles were J; within the genus Festuca. Alexeev interpreted these characteristics as a highly e. PUDE o dead to the alpine habitats in which the species occurs. Other cited as uniquely characteristics of F. livi in these rig climatic conditions. The small anthers (0.8-1.5 mm long) and usually non- E ced stamens and NM are typical of a cleistogamous habit. The anthers dehisce at an early stage of inflorescence development with pollen being released into the cavity 832 Journal of the Botanical R h Institute of Texas 1(2) between the lemma and palea. The anthers dehisce at the level of the stigmas and remain entangled in the stigmatic branches as the caryopsis matures. The leaf sheaths of shoots and culms are loose (inflated), trap- ping air and providing insulation for the developing tissues within. Other Central American alpine species of Festuca have loose flag leaf sheaths, but this characteristic is pronounced in F. livida. The pendulous form of the panicle in F. livida is unusual in that the rachis is lax right to the lowest inflorescence node. The lowest rachis internode is scarcely wider than the single branch at the lower node, giving the inflorescence the appearance of consisting of two branches at the lower node. As in many spe- cies of Festuca there is a ring of tissue associated with the lowest inflorescence node. In F. livida it is usually well developed bearing a pointed tip and sometimes extending into short bract up to 25 mm long. In other Central America species of Festuca, the ring of tissue may bear a pointed tip, but it is scarcely more than 1 mm long. Another unusual characteristic of F. livida is the distinctly swollen and slightly flattened upper culm internode with a large inner cavity (Fig. 4). Two chromosome counts have been reported for F. livida, n 2 14 (Nevado de Toluca; Beaman et al. 1962) and n = 21 (Cofre de Perote; González-Ledesma 1991). Meiosis of plants from Cofre de Perote was, however, disrupted (González-Ledesma 1991), indicating pollen fertility problems. Pollen from anthers of 9 herbarium specimens (DAO) showed a considerable range of development and stainability in lactophenol cotton blue preparations. Even though most anthers seemed to be normally dehisced, conservative estimates of pollen 1]- F viability ranged from 0-20%. When slightly less stained and/or non-spherical (but not d) grains were deemed to be possibly fertile, calculations suggested that 2-48% of pollen grains might be viable. However, low pollen viability does not necessarily present a fertility handicap for cleistogamous grasses and may in fact be an adaptive conservation of resources. Field observations in 1997 at Cofre de Perote (19°29'42"N, 97°08'55"W) and La Malinche indicated that large numbers of plants bore inflorescences with abortive caryopses, although most florets usually contained dehisced anthers. It is possible that unusual climatic conditions in that season caused caryopsis abortion, although given the low proportion of stainable pollen, lack of caryopsis development may have been the result of failed pollination. The many specialized morphological and anatomical features of F. livida, unique or highly unusual in Festuca, provide support for the recognition of the separate genus Hellerochloa. However the taxon is clearly related to the members of Festuca subgenus Festuca, as indicated by the similarity of the vegetative shoot structure, leaf blade anatomy (both in cross section and the abaxial epidermis) and ovary/caryopsis mor- phology, as well as the spontaneous hybridization with at least F. hephaestophila and F. tolucensis (see below). In spite of the fact that intergeneric hybrids have been reported between most genera in Loliinae (except Leucopoa) and many genera in Poeae, and that F. livida exhibits several unique or unusual characters, the recognition of Hellerochloa seems premature Two spontaneous hybrids, F. livida x F. hephaestophila and F. livida x F. tolucensis are sterile, intermedi- ate in many morphological characters and not uncommon in areas where the parental species come into contact. How the largely cleistogamous F. livida is able to hybridize with sympatric species of Festuca is yet to be determined. Festuca xgonzalez-ledesmae S.J. Darbyshire, nothosp. nov. (Festuca livida (Kunth) Willd. ex Spreng. x Festuca hephaestophila Nees ex Steudel). (Fig. 5). Type: MEXICO. Veracruz: Mpio. de Perote, Cotre de Perote, lado N en la base del macizo rocoso que esta en la cima, pastizal de Festuca, Calamagrostis, Trisetum, Agrostis y Deschampsia, alt. 4050 m, 15 Sep 1986, Gonzál desma & P Vera C. 196 (HoLoTYPE: DAO 683891; Isotype: CHAPA). Inter Festuca livida et Festuca hephaestophila intermedia est. Plantae perennes caespitosae, interdum rhizomatibus brevibus praeditae, innovationibus intravaginalibus praeditae. Culmi 4.5-22 cm alti, erecti, glabri, internodia superne interdum leviter tumida. Vaginae foliorum ima basi tantum integrae, leves, plus minusve inflatae; vaginae foliorum emortuorum in fibras longitudinales plus minusve solutae; ligulae 0.2-1(-1.5) mm longae; laminae foliorum conduplicatae, (0. a E —0.9 mm diametri, apice acuto ad obtuso, abaxialiter glabrae, adaxialiter hirsutae, cum 5-7 nervis, 5-7-costatae, sclerenchyma li libus, cum venas majore abaxialiter, raro adaxialiter ee cellulis Dor. destitutae. Paniculae 223-6 cm longae, ene minusve erectae, axe distali et ramulis scabris. Spiculae 7-11(-13) mm longae, ellipsiodeae, cum 2-)3-4(-5) flosculis; glumae glabrae ad scabrae vel hispidulae, iru ad lanceolatae ne tantem ellipsoideae, inermes vel arista usque ad 0.3 mm longa praeditae; glumae inferiores (5.8-)6.5-8.5-9) mm longae, lanceolatae vel ovatae, cum 1-3 nervis; glumae superiores (6.5)7-9.5(-10) mm longae, cum 3-5 nervis; rachillae usque ad 1.5 Darbyshire, F t in Mavi Val to tl PS Y | . k 833 L J . £ Luis J L 4 f Fic. 4. Festuca livida, S -1 eera] 5-8.5(-9.5) mm longae, | ] d ellipsoid scabrae ad hirsutae, cum 5 nervis, m longae apice interdum eee inermia vel plerumque praedita arista (0.3-)0.8-2.5(-3.4) mm us paleae 5.5-7.5 mm longae, inter venas et saepe in marginalibus distalibus hispidulae, venae prope apicem ad 2/3(-3/4) cum trichomatibus; antherae (1-)1.2-2 mm longae, plerumque indehiscentes; ovarium apex glabrus vel sparsim hispidulus. Caryopses non praeditae Intermediate between F. livida and F. hephaestophila. Plants perennial, densely caespitose, short rhizomes sometimes present, shoots intravaginal. Culms 4.5-22 cm tall, erect, glabrous throughout, upper internode sometimes slightly swollen. Leaf sheaths closed only at the base, glabrous, more or less loose and disin- tegrating into fibres with age; ligules 0.2-1(-1.5) mm long; leaf blades conduplicate, (0.3—)0.4-0.9 mm in diameter, apex abruptly acute to obtuse, abaxially glabrous (rarely sparsely scabrous), adaxially hirsute, with 5-7 veins and 5-7 adaxial ribs, sclerenchyma in fascicles at the margins and abaxially at the veins, rarely small adaxial fascicles, bulliform cells absent. Inflorescence paniculate, (2-2)3-6 cm long, erect, stiffly open, axis distally scabrous, branches scabrous, a single branch at lower node. Spikelets 7-11(-13) mm long, ellipsoid, with (22)3—4(—5) florets; glumes glabrous to scabrous or hispidulous (especially above), subulate to lanceolate or sometimes narrowly ellipsoid, unawned or with an awn up to 0.3 mm long; lower glumes (5.8—)6.5-8.5(-9) mm long, lanceolate or ovate, with 1-3 veins; upper glumes (6.5-)7-9.5-10) mm long, with 3-5 veins; rachilla up to 1.5 mm long, unilaterally hispidulous opposite to the floret; lem- mas 6.5-8.5-9.5) mm long, lanceolate to ellipsoid, scabrous to hirsute, with 5 veins, bidentate or entire, unawned or usually with an awn (0.3-)0.8-2.5(-3.4) mm long; paleas 5.5-7.5 mm long, hispidulous between the veins and sometimes on the upper margins, veins with trichomes from the apex to about 2/3(-3/4) to the base; anthers (1-)1.2-2 mm long, mostly indehiscent; ovary apex glabrous or sparsely hispidulous. Caryopses not formed. Additional coll ined. MEXICO. México: Nevado de Toluca, talus slopes and rocky S. rim inside crater, 3 Jul 1964, G. Mick & K. Roe 198. Tlaxcala: vertiente NO del volcán La Malinche, zacatonal de Festuca y Calamagrostis, suelo arenoso, alt. 4080 m, 16 Sep 1986, M. González-Ledesma & P. Vera C. 205; La Malinche volcano, SE of Tlaxcala, NE of Puebla, 19°14'02"N 98?01'56"W, alpine talus t to F. livida and F. hephaestophila, mostly on windward side of ridges and peak, 4290 m, 22 Sep 1997, SJ. DI & M. Ganas ez-Ledesma 4824. Veracruz: summit of Cofre de Perote, 19°29'42"N 97?08'55"W, alpine talus and solifluction sites, 4140 m, 20 Sep 1997, S.J. Darbyshire & M. González-Ledesma 47944. The inflorescence of the F. xgonzalez-ledesmae has the erect appearance of F. hephaestophila. The branches may be stiffly spreading, but the lower branch and lower rachis internode are not drooping to pendulous as in F. livida. The glumes are broadly to narrowly lanceolate and almost as long as or longer than the adjacent 834 I Lnaft+h,D ¿VU Mal OF HERBARIO-HORTORIO DEL COLEGIO DE POSTGRADUADOS (CHAPA) CHAPINGO, MEXICO Festuca hephaestoph le x [vida MEXICO. VERACRUZ: Mpio. de Perote. Cofre en la base del i t de Perote, lado N g la cima, Pastizal Calamagrostis, is Trisetum, Agrost Alt. 4 050 a. 15-11-1986 Planta glauca, frecuente. Fic. 5. Fest M. Gonzalez-Ledesma 196 y P. Vera £. 40z INAN\ . Scale bar: left = 2 inches; right = 5 cm. Darbyshire, F t in Mavi Val to tl PS Y | . P 835 lemmas, whereas they are broadly lanceolate to ellipsoid and distinctly shorter than the adjacent lemmas in F. hephaestophila. The lemmas of the hybrid are scabrous to hispid at least in the upper third and sometimes throughout, whereas they are glabrous or only slightly scabrous to hispidulous at the very apex in F. hepha- estophila. A comparison of additional characters is given in Table 2. In lactophenol cotton blue preparations from the type collection, 0-2% of pollen grains were evenly stained and/or regularly formed. This hybrid was first recognized by Manuel González-Ledesma who described it in some detail (González-Ledesma 1991). Festuca xmiscella S.J. Darbyshire, nothosp. nov. (Festuca livida (Kunth) Willd. ex Spreng. x Festuca tolucensis . (Fi g. 6). Type: MEXICO. Mexico: E slope of Nevado de Toluca, near gate to crater, 19°15'N 99°45'W, alpine solifluction sites, about ed m, 26 Sep 1997, SJ. Darbyshire & M. González-Ledesma 4851 (HoLoTYPE: DAO 811264). Inter Festuca livida et Festuca tolucensis cue est. Plantae p Bu: aeditae. Culmi 0—60 cm alti, erecti, scabri ad hispiduli eee tumida. V. leve vel Mn plus minusve inflatae; vaginae foliorum emortuorum in fibras longitudinales non solutae; ligulae (0.5-)1-2.5 mm Vir laminae foliorum conduplicatae, (0.530.7-1.522.1) mm diametri apice attenuato, abaxialiter glabrae vel scabrae, adaxialiter hirsutae, cum 7-9 nervis, 7—9-costatae, d abaxialiter annulo continuo subepidermali disposito, fasciculis adaxialiter interdum cum venas, cellulis bulliformibu utae. Paniculae 5-15(- n cm a ns et nutantes, axe dia: et d M sud 9-14(-16) mm longae, obovatae vel late ellipsiodae, cum 4-6( inermae vel arista usque ad 1 mm longa praeditae; glumae inferiores (6.3 a 10€ i mm a cum 1-3 nervis; glumae superiores (7.5-)8-11(-13.8) mm longae, cum 3-5 nervis; e usque ad 1.5 mm longae, hist lemmatae (7.5-)8-10(-12) mm longae, lanceolatae, scabrae, papill cum 5 nervis, pe nut bt inermia vel plerumque praedita arista (021-2(-2.9) mm longa; paleae "E En mm longae inter hi pi ulae, venae prope apicem a trichomatibus; antherae 1.5-4.5 mm longae, plerumque leed ovarium apex glabrus. Caryposes non praeditae. Intermediate between F. livida and F. tolucensis. Plants perennial, densely caespitose, shoots intravaginal. Culms 20—60 cm tall, erect, scabrous to hispidulous, upper internode sometimes slightly swollen. Leaf sheaths closed only at the base, glabrous or scabrous, somewhat loose, not splitting into fibres with age; ligules (0.5-)1-2.5 mm long; leaf blades conduplicate, (0.5—)0.7-1.5(-2.1) mm in diameter, apex attenu- ate to a sharp point, abaxially glabrous or scabrous, adaxially hirsute, with 7-9 veins and 7-9 adaxial ribs, sclerenchyma in a continuous abaxial band and sometimes with adaxial fascicles at the veins, bulliform cells abscent. Inflorescence paniculate, 5-15(-20) cm long, lax and nodding, axis glabrous below and sca- brous distally, branches scabrous, a single branch at lowest node. Spikelets 9—14(-16) mm long, obovate or broadly ellipsoid, with 4—6(—7) florets; glumes glabrous, scabrous or hispidulous, subulate to lanceolate, unawned or with an awn up to 1 mm long; lower glumes (6.3—)7—10(-11.5) mm long, with 13 veins; upper glumes (7.5—)8—11(-13.8) mm long, with 3—5 veins; rachilla up to 1.5 mm long, unilaterally hispidulous opposite to the floret; lemmas (7.5—8—-10(-12) mm long, lanceolate, scabrous, papillose or hirsute, with 5 veins, apex usually bidentate, unawned or usually with a short awn (O—)1—2(-2.9) mm long; paleas 6-7.5 mm long, hispidulous between the veins and sometimes on the upper margins, veins with trichomes from the apex to about 3/4 to the base; anthers 1.5-4.5 mm long, mostly indehiscent; ovary apex glabrous. puis not formed. A dditi ined. MEXICO. México: E slope of Nevado de Toluca tet ter, 19°15'N 99°45'W, alpine solifluc- tion sites, ca. 4000 m, 26 Sep 1997, S.J. Darbyshire & M. Gonzdlez-Ledesma 4849. The open paniculate inflorescence of the F. xmiscella is similar in appearance to that of F. tolucensis, but with fewer spikelets (usually less than 20 versus usually more than 20). The lower rachis internode is stiff or somewhat lax, but not pendulous as in F. livida. The spikelets are obovate to broadly ellipsoid in F. xmiscella with a length to width ratio usually 2 or less. In F. tolucensis the spikelets are usually lanceolate or narrowly ellipsoid (occasionally broadly ellipsoid) with a length to width ratio usually 2.5 or greater (occasionally as little as 1.5). The glumes are 3/4 as long as or about as long as the adjacent lemmas, whereas they are usually 1/2 to 3/4 as long as the adjacent lemmas in F. tolucensis. A comparison of additional characters is given in Table 2. In lactophenol cotton blue preparations from the type collection, 2-27% of pollen grains were evenly stained and/or regularly formed. This hybrid was first identified and characterized by González- Ledesma (1991). £L Dag PRA D hi PEA 836 Journal of f Texas 1(2) TABLE 2. A compari fd istics of A hepl hila, F livida, E tol character F. hephaestophila F. xgonzalez- F. livida F. xmiscella F. tolucensis ledesmae spikelet length 45-84 7-11(-13) (9.4)10-17(21) | 9-14(-16) (6.5-)8-14(-18) lower glume length — 2.8-4.5(-5.5) (5.8-)655-8.5(-9) — 9.2-15(-20) (6.3-)7-10(-11.5) Q7-)35-7.5(-8.6) upper glume length 3.5-6.3 (6.5-)7-9.5(-10) 94-17(-21) (/5-)8-11(-13.8) | (3.6294.5-8.5(-9.3) lemma length 4-5.5(-6.2) 6.5-8.5(-9.5) 6.5-12.5 (7.5-)8-10(-12) (4.6-)5.5-8.5(-10) anther length 2.2-3.5 (1-)1.2-2 0.8-1.5 1.5-4.5 (2.6-)3-4.5(-5.2) The following key to species of Festuca in Mexico and Central America is largely based on that by Alexeev (1984), but has been updated with additional information and recently described taxa. Although frequently treated in a different genus, Festuca arundinacea Schreb. [Lolium arundinaceum (Schreb.) Darbysh.; Schedonorus arundinaceus (Schreb.) Dumort.] is included in the key and Table 1, because of its similarity to and continued placement in Festuca by some authors. The inclusion of F. ovina L. is based on Godfrey 66649 report by Pohl (1980) for Costa Rica and accepted by Alexeev (1984). The specimen has not been examined and its identity remains unclear. KEY TO THE SPECIES OF FESTUCA IN MEXICO AND CENTRAL AMERICA . Leaf blades of shoots usually more than 2 mm wide, flat in cross section, sometimes round in the upper part (or becoming involute or convolute when dry), p ing distinct (wide) bulliform cells. 2. Margins of the blade-sheath junction with falcate auricles, more or less provided with cilia_______ F. arundinacea 2. Margins of the blade-sheath junction lacking auricles, without cilia. 3. Ligules of culm leaves (1.3-)2-10(-11) mm lon 4. Plants loosely caespitose; ligules of culm leaves less than 3.5 mm long; lemmas less than 7 mm long, apex entire, awnless 5. Ligules of culm leaves 2.5-3.5 mm long; leaf blades up to 2 mm wide; lemmas 4-6 mm long; plants of Mexico and Texas F. ligulata 5. Ligules of culm leaves less than 2.5 mm long; leaf blades 1-4 mm wide; lemmas 5.7-6.8 mm long; BRE of e Rica ang E F. herrerae 4. Plant Im | lly 3-8( ue cuc e a EUM lemmas greater than 7 mm lon bi ¿with an awn or a mucro (sometimes absent) ea from NS the teeth. 6. Apex of ovaries densely hisp F. asperella 6. ign iu ies Pun E at most with a short mucro less d 0. 5; mm jos ng F. tancitaroensis 7. Ligules of culm leaves (1.3-)3-8(-11) mm long; lemmas with an awn (0.5-)1-4 mm long. 8. Leaf blades stiff, 3-5 mm wide; upper glume (5.5-)6-8.5 mm long F. lugens 8. Leaf blades lax, 1.5-3.6(-6) mm wide; upper glume (3.8-)4-5.6(-6) mm long____________ Kroblensis 3. Ligules of culm leaves not more than 1.5 mm long 9. Lemmas 12-17.5 mm long. 10. Lower glumes (1-)2-6(-7.5) mm long; lemmas with 3-5 veins, awns 10-14(-16) mm long F. breviglumis 10. Lower glumes (7.5-)8-9 mm long; lemmas with 5 veins, awns 1.5-3 mm long . .. F chiriquensis 9. Lemmas 3-9.5(-10) mm lon 11. Apex of lemmas bidentate awns (0.3-)0.5-1.8 mm long arising between the teeth. 12. Lemmas 5.5-6.2 mm long, scabrous throughout the back; paleas scabrous along the keels to the base; anthers 2.4-2.7 mm long; apex of ovaries densely hispid; plants of Panama Ligu 15 F. swallenii 12. Lemmas 6.5-7 mm long, scabrous on the upper half; paleas scabrous along the keels on the upper third; anthers 3.5-4 mm long; apex of ovaries with a few hairs (1-4); plants of F. bidenticulata Mexico 11. Apex of lemmas without teeth or awns. 13. Lemmas 3-4.5 mm long, apex obtuse; anthers 1-1.3 mm long F. subverticillata 837 A y PRO J Darbyshire, pé AND AIF ¿E e GS N^ = 3 81126 Y, o > awa, CASES. 2006 | LLLI [ITI] Plants of Mexico Festuca livida x Festuca tolucensis Mexico, Mexico, east slope of Nevado de Toluca; near gate to crater; about 4000 m elev. 19?15'N, 99°45'W alpine solifluction sites J Darbvshi LM c+} Fak 4851 Coll.: S. 26 Sept. 1997 appears to be a hybrid swam with plants intermediate between F. livida and F. tolucensis; 7 replicates 4851 (DAO). Scale bar: left — 2 inches; right — 5 cm. Fic. 6. Fe 838 f the Bot Journal UI titute of Texas 1(2) 13. Lemmas 5.5-9.5(-10) mm "E apex acute or mucronate; anthers 1.6-4 mm long. 14. sag of ovaries glabrou 15. Leaf blades flat or T folded, 1-4 mm wide; spikelets 6.5-9 mm long _______ R herrerae 15: " blades i Du Or E 3 10 mm MuR du ie 17 mm long. ]le leaf blades A or with shallow adaxial ribs; IE branches scabrous or ini pilos .amplissima 16. Dead leaf sheaths not breaking up into longitudinal fibres; leaf blades ae ong adaxial ribs; A branches mostly smooth F. jaliscana 14. Apex T E densely hisp 17. Anthers 2.8-4.5(-5) mm F plants of northeast Mexico F. valdesii 17. Anthers up to 2.5 mm long; plants not of Mexico (further south). 18. a branches smooth or almost smooth; spikelets 6.5-7.5 mm long, with -3 florets; anthers 1.7-2 mm Heng) Sel s of Guatemala F. aguana 18. men brancl | long, with 4-5 florets; anthers 2-2.5 mm long; plants of Panama F. panamica . Leaf blades of shoots usually less than 2 mm in diameter, folded (conduplicate) in cross section, lacking or with indistinct narrow bulliform cells. 9. almost as long or longer than the florets (excluding awns) and longer than adjacent lemmas, sometimes with a mucro up to 1.3 mm long; inflorescence nodding, lower rachis internode lax and more or less pendulous F. livida 19. Glumes distinctly shorter than the florets and shorter than adjacent lemmas (except in sterile hybrids with F. livida keyed below), without a mucro or with a mucro less than 0.3 mm long; panicles erect or nodding, lower rachis internode stiff or lax but not pendulous 20. Plants sterile (anthers mostly indehiscent, caryopses not | ] usually more than 3/4 as long as spikelets and about as long as lemmas; plants alpine, o above tree-line; growing with F livida (Central Mexico). 21. Plants 4.5-22 cm high, leaf blades with 5-7 veins, sclerenchyma fascicles about as wide as v anthers 1-2 mm F- xgonzalez: -ledesmae 21. Plants 20-60 cm high, leaf blades with 7-11 veins, sclerenchyma fascicles much wider th Si rs 1.5-4.5 mm lon Plants usually fertile (anthers usually dehiscent, caryopses produced); glumes usually less than 3/4 as rae as spikelets and distinctly shorter than lemmas; plants montane or alpine; growing with F livida or not. 22. Der blades of shoots with one adaxial rib. 23. Leaf blades of vegetative shoots with 3 veins, sclerenchyma in 7 small abaxial fascicles; apex of ovaries hispid 23. Leaf blades of vegetative shoots with 5-7 veins, scl | ti abaxial band; apex of ovaries glabrous. 24. Upper e usually 2.6-4 mm long; lemmas Ed more than 3.3 mm long, with short awns 0.5-2 mm long; anthers (1.4-)2-2.6 m © =) E xmiscella N e ajacaliforniana interrupted F. ovina 24. aa Fae usually 2-3 mm long; lemmas EV less than 3.3 mm long, without awns (rarely mucronate); anthers (1-)1.5-2 mm long F. filiformis 22. Leaf blades of shoots with 3-21 adaxial ribs. 25. Anthers (0.8-)1-1.4(-1.6) mm long. 26. Spikelets 8-10 mm long; lemmas 6-8.5 mm long, puberulent throughout —— F. willdenowiana 26. Spikelets 6.4-7 mm long; lemmas 5-6 mm long, glabrous or apically scaberulous F. talamancensis 25. Anthers greater than 1.5 mm long. 27. Lemma aj bid i i | or not at all; examine multiple lemmas), awn (0.3-)0.7-3(-3.7) mm long, arising between the teeth or apex rarely emarginate with the awn arising a little below the apex 28. Leaf sheaths usually glabrous; ligules of culm leaves (1-)1.5-3.2(-4.6) mm long; abaxial surface of leaf blades Selects pal cule) ule oil eles ina eon ELON or interrupted abaxial ~ usually g adaxial fascicles often p 28. Leaf sheaths pen ionem scabrous; ligules of culm leaves 0.4-1 mm long; abaxial surface of leaf blades glabrous or slightly scabrous; sclerenchyma of leaf F. tolucensis Darbyshire, r + 27, I y to tl f. PS ee | i. | M 839 hlag i : | re | | E B HT not or girders, adaxial fascicles absent F. aequipaleata Lemma apex entire, without an awn or awned from the apex. 29, ligules o of culm is -)2.5 mm long or more. aves 2.5-3.5 mm long; lemmas 4.5-6 mm long, apex narrowly . Lig scarious; sete 1.5-2.6 mm long F. ligulata 30. Ligules of culm leaves (2-)5-9.5 mm long; lemmas 5-7 mm long, apex broadly carious; anthers 2.5-3.3 mm long F. longiligula 29. aules fcum leaves ess than or equal 2 mm lon sub 31 eaf blades of vegetative shoots (the widest) 1-2 mm in diameter, with 11-23 veins, with 9-21 adaxial ribs. 32. Panicle branches smooth; paleas scabrous along the keels on the upper 2/3 — 3/4; apex of ovaries glabrous; plants of Costa Rica and Panama F. cartagana 32. Panicle! | | pal | | he keel the upp 1/3 — 1/2; apex of ovaries hispid; plants of Mexico F. orizabensis Leaf blades of vegetative shoots always 0.3-1(-1.2) mm in diameter (some- times wider in F. rubra), with 5-9 veins, with 3-7 adaxial ribs 33. Leaf blades Du se: shoots with abaxial sclerenchyma in a continu- ous band, rarely ir Ens 34. Leafblades of g | 5 in di t ith 5(-7) veins, adaxial surfaces with hairs 50- 80 um long; ies of blades and sheaths swollen (callose auricles) F. callosa 34. Leaf blades of vegetative shoots 0.6-0.8 mm in diameter, with 7-9 veins, adaxial surfaces with hairs 20-30 um long; junction of blades and sheaths not swollen (auricles not callose). 35. Margins of leaf sheaths of vegetative shoots closed up to 2/3; ligules of culm leaves 0.2-0 ong; leaf blades usually uco-pruinose; lemmas 6.5-7 mm long, not awned; apex of ovaries densely hispid F. hintoniana 35. Margins of leaf sheaths of vegetative shoots closed up to 1/3; ligules of culm leaves 0.5-1 mm long; leaf blades green, not glaucous or pruinose; lemmas 5-5.2 mm long, awns 2-2.2 mm long; apex of ovaries glabrous F. beamanii 33. Leaf blades of vegetative shoots with abaxial sclerenchyma in fascicles. 36. Leaf blades of vegetative shoots always with 5 veins, with 3 adaxial ribs 37. Margins of leaf sheaths of vegetative shoots closed from the ear the top; dead leaf sheaths not breaking up into longitudinal fibres; leaf blades abaxially more or less scabrous or rarely gata e E dense Meu Sel eee el fascicles the keels on the upper 1 V/;a apex KO CES with a few hairs, rae glabrous; plants alpine F. hephaestophila 27. Margins of leaf sheatt is of tati | ts cl | fi the base o 1/3; dead leaf sheaths more or less breaking up into longi- tudinal fibres; leaf blades abaxially smooth (glabrous), adaxially smooth or almost so, adaxial fascicles of sclerenchyma absent; montane F. rzedowskiana 36. Widest E blades of vegetative shoots with 7-9 veins, with 3-7 adaxial ribs. 38. Plants laxly caespitose, often with short rhizomes; margins of leaf sheaths of vegetative shoots closed from the base to at least 1/2; lemmas with an awn greater than 1 mmlong________ F.rubra 38. Plants more or less densely caespitose, without rhizomes; mar- gins of leaf sheaths of vegetative shoots closed from the base 840 Journal of the Botanical R Institute of Texas 1(2) to not more than 1/3; lemmas awnless or sometimes with an apical mucro up to 1 mm lon 39. Anthersinl r floret in tl ikel lly ak of upper florets well-developed and 3-4.2 mm long_ 39. Anthers in lower florets of the spikelet not abortive, all anthers more or less the same length, 1.6-6 mm long. , those _F.diclina 40. Some ot g f f tose); adaxial surface of leaf blades of vegetative shoots with a few isolated trichomes 20-40 um long; anthers 1.6-2.3 mm lon 41. Ligules 0.1-0.2 mm long; spikelets 10.5-11.5 mm long, with 5-6 florets; lemmas 6.5-7.5 mm long F. guatemalica 41. Ligules 0.3-2 mm long; spikelets 6.5-9 mm long, with 3-5 florets; lemmas 5.7-6.8 mm long F. herrerae 40. All shoots intravaginal; adaxial surface of leaf blades of vegetative shoots with dense trichomes 50-100 um long; anthers greater than 2.5 mm long. 42. Blades of leaves always folded (conduplicate or convolute), 0.3-0.6 mm in diameter, with 5-7(-9) veins; panicles more or less dense, branches erect; apex of ovaries glabrous (rarely with a few hairs) F. rosei 42. Blades of culm leaves flat to loosely folded (con- duplicate), (0.4-)0.6-1.2 mm in diameter, with 9-11 veins; panicles open, branches lax; apex of ovaries more or less hispid (rarely glabrous) F. pringlei ACKNOWLEDGMENTS I am grateful to Jacques Cayouette, Manuel González-Ledesma, Jochen Müller and Barbara Wilson for their useful comments and discussions, to Jacques Cayouette and Jochen Müller for assistance with the Latin descriptions, and to herbarium curators at ARIZ, DAO, TEX, US and WIS for making specimens available for study. Manuel González-Ledesma was of invaluable assistance with field work. REFERENCES ALEXEEV, E.B. 1980. Festuca L. subgenera et sectiones novae ex America boreali et Mexica. Novosti Sist. Vyssh. Rast. 17:42-53. [In Russian]. ALEXEEV, E.B. 1984. Genus Festuca L. (Poaceae) in Mexico et America Centrali. Novosti Sist. Vyssh. Rast. 21:25-59. [In Russian]. Beaman, J.H., D.C.D. De Jone, and W.P. Stoutamire. 1962. Chromosome studies in the alpine and subalpine floras of Mexico and Guatemala. Amer. J. Bot. 49:41—50. GONZALEZ-LEDESMA, M. 1991. Révision del genera Festuca L. (Gramineae) en el Eje Volcánico Transversal. Ph.D. dis- sertation. Colegio de Postgraduados, Chapingo, Mexico. PoHL, RW, 1980. Family 15. Gramineae. In: W.C. Burger, ed. Flora Costaricensis. Fieldiana Bot. New Ser. 4:1-608. SORENG, R.J., P.M. PETERSON, G. DAvipse, E.J. Jupziewicz, F.O. ZULOAGA, T.S. FILGUEIRAS, and O. Marone (eds.). 2003. Catalogue of New World grasses (Poaceae): IV. Subfamily Pooideae. Contr. U.S. Natl. Herb. 48:1—730. APHANELYTRUM PERUVIANUM (POACEAE: POINAE): A NEW SPECIES FROM PERÚ Isidoro Sánchez Vega Paul M. Peterson and Robert J. Soreng Muss de Biología Department of Botany Herbario CPUN National Museum of e eo Universidad Nacional de Cajamarca Smithsonian Institu Apartado 55, ACH PERÜ Washington, DC Ha v U.S.A. svisidoro@yahoo.co peterson@si.edu sorengr@si.edu Simon Leegaard Department of Systematic Botany DK-8000 Aarhus C, DENM, simon.laegaard@biology.au. aa ABSTRACT Aphanelytrum peruvianum Sánchez Vega, P.M. Peterson, Soreng & Leegaard, sp. nov., is described and illustrated. The new species occurs on rocky sites near Cerro Akumullca in Cajamarca, Perú and represents the second known species ascribed to Aphanelytrum. Aphanelytrum peruvianum differs from A. procumbens by having nerved glumes (1-2 mm long), narrow leaf blades (0.2-1.2 mm wide), shorter culms (14-24 cm a aus eS x 13mm a shorter lemmas (2.2-3.5 mm lone and shorter anthers 2-2.9 mm long). A key and t g g g A. peruvianum and A. p l E RESUMEN Sed RI :] ARES 1 : Caral TS Vega, P.M. Peterson Soreng & Laegaard Sp. nov. Į en sitios rocosos cerca del ue MTM en Cajamarca, Perú. Esta nueva la segunda i ida atribuida o al género Aphanelytrum. Aphanelytrum Vi nb diri difiere de A. procumbens en ee tiene e gmas nervadas (1- 2 mm largo), láminas de la hoja estrechas (0.2-1.2 mm MD rtos (14-24 cm alto), ent 3-13 mm largo), lemas más cortas ee 2-3.5mm TN yant tas Q-2.9 mm largo). Se incluye una clave y una tabla de comparación de | aA peruvianum de A. procumbens. The genus Aphanelytrum (Hack.) Hack., established in 1902, was first named without a description by Sodiro (1889) with a single species, “A. decumbens Hack.” (ex Sodiro, nom. nud.). Hackel (1887) initially described it as a subgenus of Brachyelytrum P. Beauv.; B. subgen. Aphanelytrum Hack., including a single new species B. procumbens Hack. Hackel (1902) subsequently recognized Aphanelytrum as a genus with the single spe- cies A. procumbens (Hack.) Hack., all based on a single collection of Sodiro’s from Ecuador (Chase 1916). The placement and evolutionary relationships of Aphanelytrum have been somewhat controversial since its inception. It was originally placed in the subfamily “Festuceae” [Festucoideae], applied in the broad sense of Bentham (1881), Hackel (1887), and Hitchcock (1935). Hackel mistook the individual florets for glume- less single-flowered spikelets and placed Brachyelytrum within tribe Agrostideae, subtribe Stipinae. Agnes Chase (1916) reinterpreted the spikelet morphology, and placed the genus between the subtribes Melicinae and Centothecinae, which at that time were considered adjacent subtribes of the subfamily Festucoideae, tribe Festuceae. More recently, after major realignments of the classification of the Poaceae family (e.g. Clayton & Renvoize 1986), the genus was placed in the much more narrowly defined subfamily Pooideae (syn. Festucoideae), tribe Poeae (syn. Festuceae) near Poa (Clayton & Renvoize 1986). Aphanelytrum and Poa have very similar leaf anatomical characteristics, as well as multi-flowered, membranous spikelets with 5-nerved, keeled lemmas, glabrous ovaries, and caryopses with oval hilums (Clayton & Renvoize 1986). It was specifically placed in subtribe Poinae (Soreng et al. 2003; Soreng et al. 2006) and seems to align near J. Bot. Res. Inst. Texas 1(2): 841 — 845. 2007 842 Journal of the Botanical R h Institute of Texas 1(2) [or on an unusually long branch within; 29 base pair differences in the internal transcribed spacer regions (ITS) of nuclear ribosomal DNA] Poa in preliminary cpDNA and ITS sequence analyses (Gillespie et al. 2007; Soreng et al., In press; Saarela pers. com.). Aphanelytrum is currently recognized as a monotypic genus found at mid to high elevations 2000-4050 m) in humid to montane forests in the Andes of Bolivia, Columbia, Ecuador, and Perú (Hitchcock 1927; Clayton & Renvoize 1986; Jorgensen & Ulloa Ulloa 1994; La Torre 2002; Soreng et al. 2003; Leegaard 2005). Aphanelytrum procumbens is a peculiar grass that has unusual spikelets with minute and unnerved glumes, two or three florets that are widely-spaced because of the long flexuous rachillas Y to % as long as the florets, rachillas that are prolonged above the upper florets, and 5-nerved lemmas (Chase 1916; Hitchcock 1927; Nicora & Rúgolo de Agrasar 1987; Watson & Dallwitz 1992; Leegaard 2005). While collecting specimens for the floristic inventory of “jalca formations” in northern Perú, Isidoro Sánchez Vega and others collected a specimen that was not easily assignable to Aphanelytrum procumbens. Since this specimen morphologically resembles A. procumbens, we are describing it as a new species of Aphanelytrum and are therefor ling the generic description to include additional morphological varia- tion. Aphanelytrum peruvianum Sánchez Vega, P.M. Peterson, Soreng & Leegaard, sp. nov. (Fig. 1A & B, D-L). Tre: PERÚ. Cajamarca: Prov. Cajamarca, Dist. Cajamarca, Cerro Akumullca, al SO de Cajamarca, sobre la cima de la ladera occidental del Valle de Cajamarca (7°14'15"S Lat, 78°29'24"W Long), 3300 msnm, 20 Mar 2003, I. Sánchez Vega 11718, M. Sánchez Montoya, R. Cueva R. & J. Montoya (HoLotyee: CPUN!, isotypes: AAU!, Fl, HAO!, HUT!, LOJA!, MICH!, MOI, SI!, US-3472470!, USM). Se ae procumbens Hack. culmis 14-24 cm longis, internodiis 3-13 mm longis, ligulis 2-3 mm longis, laminis 0.2-1.2 mm latis, glumis 1-2 mm longis, inferioribus glumis 1-innervatus, superioribus glumis 3(4-)-innervatus, lemmatibus 2.2-3.5 mm longis 3- vel 5-innervatis, antheris 2-2.9 mm longis, recedit. Caespitose perennials. Culms 14-24 cm tall, with many culms near base, primary and secondary culms appressed, somewhat decumbent near base with extravaginal branching; internodes 3-13 mm long, nu- merous. Leaf sheaths longer than the internodes, membranous to hyaline, open, slightly keeled; ligules 2-3 mm long, membranous to hyaline, decurrent, apex erose often lacerate; blades 3-7 cm long (flag leaf ca 1.6 mm long), 0.2-1.2 mm wide, flat to loosely involute, linear, apex naviculate. Panicles 1.7-2.5 cm long, few- flowered with 5-10 spikelets; branches flexuous, the lower branches with two spikelets, the upper branches with single spikelet. Spikelets 5-7 mm long, usually 3-flowered, purplish, disarticulating above the glumes and between the florets; lower and middle florets usually staminate; upper florets usually pistillate; rachilla joints 1.2-2 mm long, prolonged above the upper floret; glumes 1-2 mm long, subequal, apex acute, often mucronate; lower glume linear, 1-nerved; upper glume oblanceolate, 3(4-)-nerved, often toothed or irregu- larly lobed minutely bifid; lemmas 2.2-3.5 mm long, ovoid, 3- or 5-nerved, apex mucronate with two acute lobes on each side of the mucro, the mucro 0.1-0.3 mm long; paleas 2-3.2 mm long, 2-keeled, apex bifid; lodicules 7-8 mm long, lanceolate, membranous, glabrous; stamens 3; anthers 2-2.9 mm long, yellowish to purplish; ovaries glabrous with two styles and two stigmas. Caryopses glabrous. Phenology.—Flowering in March. Distribution and habitat —Aphanelytrum peruvianum is known only from the type locality near the west- ern highlands of the Cajamarca Valley and is found on rocky sites associated with jalca vegetation (humid alpine grass ecosystems) at 3300 m DISCUSSION We are here emending the generic description of Aphanelytrum (Watson € Dallwitz 1992) to include: leaf blades sometimes involute; spikelets with glumes 1-2 mm long, nerved; lower glumes 1-nerved; upper glumes 3(4-)-nerved; lemmas 3- or 5-nerved. Festuca reclinata Swallen superficially resembles Aphanelytrum since the former species from the Páramo del Almorzadero in the Cordillera Oriental of Colombia exhibits similar morphologies (Stančík & Peterson 2007). The growth habit, panicles, and spikelet characteristics of F. reclinata are strikingly similar to those <= INS NS NA > i EU SS W AAVA ^ x N SSA Z — <=: Se ART SA S Fic. 1. Aphanelytrum peruvianum [I. Sánchez Vega 11718, M. Sánchez Montoya, R. Cueva R. & J. Montoya (US)]. A. Habit. B. Sheath, ligule, and blade. D. Panicle. E. Spikelet. F. Glumes, showing upper glume 4-nerved with ay hed ifid. H. Lemma. I. Paleas. J. Palea, anthers, and rachilla. K. Lodicul ] y l Ovary Ar h ly p IPM. Rodríguez (US)] C. Sheath, ligule and blade. 844 Journal of the Botanical R h Institute of Texas 1(2) Taste 1. Salient fi paring Aphanelytrum peruvi ith A. procumbens. Characters A. peruvianum A. procumbens Culm height 14-24 cm 30-70 cm Internode length 3-13 mm (15-)24-90 mm Ligule length 2-3 mm 1-22 mm Blade width 0.2-1.2 mm 1.5-4.2 mm Glume length 1-2 mm 0.1-0.3 mm Glume nerves present absent Lemma length 2.2-3.5 mm 3.2-8.4 mm Number of lemma nerves IONS 5 Anther length 2-29 mm 3.5-3.8 mm found in A. peruvianum. Both species have weak, decumbent culms with extravaginal branching, narrow few-spikeleted panicles, spikelets with flexuous rachillas, and small, nerved glumes (lower glumes 1-nerved; upper glumes 3-nerved). In comparison with F. reclinata, A. peruvianum has smaller lemmas (2.2-3.5 mm vs. 7-8.5 mm in F. reclinata), smaller anthers (2-2.9 mm vs. 3.5-3.8 mm), and shorter spikelets (5-7 mm vs. 10-13 mm). Moreover, the strongly keeled lemmas of F. reclinata do not agree with its placement in Festuca. Further study is needed to confirm the connection of F. reclinata with Aphanelytrum. Aphanelytrum peruvi differs from A. procumbens (see Table 1) by several characteristics, most notably: culm height (14-24 cm long in the former versus 30-70 cm long in the latter), internode length [3-13 vs. (15-)24-90 mm], ligule length (2-3 vs. 1-2 mm), leaf blade width (0.2-1.2 vs. 1.5-4.2 mm), glume length (1-2 vs. 0.1-0.3 mm or absent), presence of glume nerves (lower glumes 1-nerved, upper glumes 3- or 4- nerved vs. both glumes unnerved), lemma length (2.2-3.5 vs. 3.2-8.4 mm), number of lemma nerves (3 or 5 vs. 5), and anther length (2-2.9 vs. 3.5-3.8 mm). In addition to these salient characteristics, A. peruvianum has leaf sheaths that are usually longer than the internodes (vs. shorter in A. procumbens), blades that are flat to loosely involute (vs. flat in A. procumbens, see Fig. 1C), and shorter rachilla extensions (1.2-2 mm vs. 1.6-4.2 mm in A. procumbens). A key is presented below to help differentiate between the two species of Aphanelytrum. KEY TO THE SPECIES OF APHANELYTRUM — . Glumes 1-2 mm long, the lower 1-nerved, the upper 3- or 4-nerved; leaf blades 0.2—1.2 mm wide; culms 14—24 cm long; internodes 3-13 mm long; lemmas 2.2-3.5 mm long; anthers 2-2.9 mm long . ____ A. peruvianum 1. Glumes 0.1—0.3 mm long, minute or absent, unnerved; leaf blades 1.5—4.2 mm wide; culms 30-70 cm long; internodes (15—)24—90 mm long; lemmas 3.2—8.4 mm long; anthers 3.5-3.8 mm long ___ ____ A. procumbens ACKNOWLEDGMENTS We thank the National Science Foundation (NSF, Grant 0071506) to Michael O. Dillon and the The Field Museum (Chicago) for financial support; Alice R. Tangerini at the Smithsonian Institution for preparing the illustration; Patricia Gómez Bustamonte for help preparing the Spanish abstract; Alain Touwaide for help correcting the Latin diagnosis; and Diego A. Giraldo Cañas and Jeffery M. Saarela for suggesting improve- ments to the manuscript. REFERENCES BENTHAM, G. 1881. Supplemental papers to Bentham and Hooker's Genera Plantarum [Gramineae]. J. Linn. SOc., Bot. 19:14-134. Chase, A. 1916. The structure of the spikelet of Aphanelytrum. Bot. Gaz. 61:340—343. CLAYTON, W.D. and S.A. Renvoize. 1986. Genera Graminum. Grasses of the world. Kew Bull. Add. Ser. 13:1—389. GILLESPIE, L.J., A. ARCHAMBAULT, and R.J. Sorens. 2007. Phylogeny of Poa (Poaceae) based on trnT-trnF sequence data: major clades and basal relationships. Aliso 23: In press. Ciakas V. + I,A | : f Apl ht r4 D 845 HackeL, E. 1887. Gramineae. In: A. Engler & K. Prantl, eds., Die Natürlichen Pflanzenfamilien 2.2:1-97. Wilhelm Engelmann, Leipzig. HackeL, E. 1902. Neue Graeser. Oesterr. Bot. Z. 52:8-15. HircHcock, A.S. 1927. The grasses of Ecuador, Perú, and Bolivia. Contr. U.S. Natl. Herb. 24(8):201—519. HitcHcock, A.S. 1935. Manual of the grasses of the United States. USDA Misc. Publ. 200. U.S. Govt. Printing Office, Washington. Jorcensen PM. and C. ULLOA ULLOA. 1994. Seed plants of the high Andes of Ecuador—A checklist. AAU Rep. L&GAARD, S. 2005. Poaceae (Gramineae). In: P. Sklenář, J.L. Luteyn, C. Ulloa Ulloa, PM. Jorgensen, and M.O. Dillon, eds. Flora genérica de los páramos. Mem. N.Y. Bot. Gard. 92. Pp. 358—391. La Torre, M.I. 2002. Primer registro de Aphanelytrum (Poaceae: Poideae) para el Perú. Arnaldea 8(2):53—56. Nicora, E.G. and Z.E. RUGOLO DE AGRASAR. 1987. Los Géneros de Gramíneas de América Austral Argentina, Chile, Uru- guay y areas limítrofes de Bolivia, Paraguay y Brasil. Editoria Hemispherio Sur S.A., Buenos Aires, Argentina. Sopiro, PL. 1889. Gramíneas ecuatorianas de la Provincia de Quito. Anales Univ. Centr. Ecuador 3(25):474-484. SORENG, R.J., J.| Davis, and M.A. Voionmaa. In press. A phylogenetic analysis of Poaceae tribe Poeae s.l. based on morphological characters and sequence data from three plastid-encoded genes: evidence for reticulation, and a new classification for the tribe. Kew Bull. SORENG, R.J., P.M. PETERSON, G. Davipse, E.J. Jubziewicz, F.O. ZULOAGA, T.S. FILGUEIRAS, and O. Morrone. 2003. Catalogue of New World grasses (Poaceae): IV subfamily Pooideae. Contr. U.S. Natl. Herb. 48:1—730. SORENG, R.J., PM. PETERSON, G. DAVIDSE, E.J. Jubziewicz, F.O. ZULOAGA, T.S. FILGUEIRAS, and O. Morrone. 2006. Suprageneric Classification. http://mobot.mobot.org/W3T/Search/nwgclass.htm| Stančík, D. and P.M. Peterson. 2007. A revision of Festuca (Poaceae: Loliinae) in South American paramos. Smith- sonian Contr. U.S. Natl. Herb. 57: In press. Watson, L. and M.J. Dallwitz. 1992. The grass genera of the world. CAB International University Press, Cambridge. 846 Journal of the Botanical R h Institute of Texas 1(2) DOOR REVIEWS SYLVAN RAMSEY KAUFMAN and WALLACE KAUFMAN. 2007. Invasive Plants: Guide to Identification and the Impacts and Control of Common North American Species. (ISBN 978-0-8117-3365-6, pbk). Stackpole Books, 5067 Ritter Road, Mechanicsburg, PA 17055, U.S.A. (Orders: www.stackpolebooks. com, orders@stackpolebooks.com, 717-796-0411, 717-796-0412 fax). $39.95, 458 pp., 503 color photos, 8 1/45. 1/21. Touted as “the first-ever field guide to invasive plants in North America”, this book is a thoughtful compendium of over 150 invasive plant species of the U.S. and Canada and the larger issue of invasive species impact and control. The first 36 pages provide an adequate line between "exotic" and “pest”, and the colored history of overview of the e Re on what it means to be invasive, the fine nd those D od sought me their ecological benefits. Suggestions are provided for various manage- l ies, i ical, biological, and chemical methods, and are later revisited relative to each individual des ment and species. All instructions are clear idi concise, as is the overview of common herbicides, their use, and their potential dangers. Natural or eco-friendly herbicides are not mentioned. TT 1 kseguesfromi | ial in Part I to species descriptions in Part II by way of a“ eo (s i Lus in the familiar : Pet E ; : : E MEC biology key format", which, though technically accurate, is hard 11 ] ld! g., bullets numbers, letters). The individual ies d ipti ] l however lerful ff. Each plant is depicted i ] color photograph, along with the specific epithet, various common names, family name, and any hybridizing species or species for which n p may ES EUM In A DUE the id prove no for LE des MD M d features, habitat, and range, as i erica. Species- M management tips and a short list of a are Msc provided for each eine with special mention for sources that bibliography of 25 articles, books, and pa a x credits section, and an index of categories from d Ag i 1 layperson mco recommend and with an annotated key, Latin names this book.—M. Brooke Byerley, Botanical Research Institute 2 Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U. Oscar E CLARKE, DANIELLE SVEHLA, GREG BALLMER, and ArLEE MONTALVO. 2007. Flora of the Santa Ana River and Environs: With References to World Botany. (ISBN 978-1-59714-050-8, pbk.). Heyday Books, PO. Box 9145, Berkeley, CA 94709, U.S.A. (Orders: www.heydaybooks.com). $29.95, 496 pp., 3200 images and illustrations, 6" x 9", Casual observers of the botanical world and experts alike, located both a ix in: z ue Santa Ana River watershed, will enjoy and bs uds this en new 2s The authors SB n: oud ] I I by not un. dd over 1,000 a A } } 1 Spe [E 8 I ecology iated pl ity. As noted by the authors, the Santa Ana River i is one of the largest rivers lying wholly within Southern California, i dynamic plant communities found in three major segment alluvial scrub segment inland valley segment, and dosi M Wand 1 E CNN S t I UNITE Į I species. The nro to the flora of the Santa Ana River is organized his basis and, therefore, [ ial for developing a greater understanding of the pl p identification. Further, it is this very focus that makes the book so broadly applicable to all levels of experti d to al a in UE pe do not padri reside ae the Santa Ana River auras For tl hat do li ithi a River watershed, tl g k for identifying f ] und n f species along the ri li ling envi While the 1 Es £T ape | PA Wa n pa E J Soe | J T J T 1 1-271 cies accounts are b cencralla Bue to visually highlight these key features. To assist E dE in c cue E inu he s have also included a pli d "Botany Basics” section and glossary. E Also of note are the numerous Appendix 2—“Family Character Tables.” These summary tables, as well as the authors’ look at world phylogeny in Appendix 5 ar useful and oe pe poe aa of ba book. This su is sure to not "o increase one's cun iur of e d of the felinda Melinda ummary tables throughout e Book spotlighting family and genera B mq me € VCI Santa Ana River McCoy, Herbarium Volunteer, Botanical Research Institute of Texas, Fort Worth, Texas 76102-4060, USA. J. Bot. Res. Inst. Texas 1(2): 846. 2007 A NEW SPECIES OF CHUSQUEA SECT. VERTICIEEATAE (POACEAE: BAMBUSOIDEAE) FROM ECUADOR Lynn G. Clark, Christopher D. Tyrrell, Jimmy K. Triplett Amanda E. Fisher Department of Ecology, Evolution and Organismal Biology Department of Biological Sciences lowa State University Idaho State University Ames, lowa 50011-1020, U.S.A. Pocatello, Idaho 83209-8007, U.S.A. ABSTRACT Chusquea annagardneriae, a species of Chusquea sect. Verticillatae from Ecuador, is Md as new. It is ieee and compared and contrasted ee Chusqued pd = o to Mo itis most pu CI ished from all other L o O species of Ch 1 1 a 5 : ads ther ÉSTE UBER: 1 T cy 1 n Į pen mpressed spikelets with glumes I and II both greatly reduced and scalelike, and glumes III and IV extending about half of the spikelet length. Chusquea annagardneriae is endemic to the province of Loja, Ecuador, and occurs in dry forest scrub and forest remnants at 2000 to RESUMEN Se describe Chusquea annagardneriae, una especie nueva de Ecuador que pertenece a Chusquea sección Verticillatae. Chusquea an- MA se ilustra y se cds con C. dba la nM más A um Se goers Toen Me de todas las otras J lámina de la hoja P más o menos pee is NUN de ramitas palate en el RM de ramas, una sinflorescencia abierta y iramidal, gl I It I pd reducidas y escamosas, y glumas III y IV extendiéndose por la mitad de if longitud de la espiguilla. CI a provincia de Loja, Ecuador, y g secundaria de bosque seco y en restos de ne a2000a 12250 m.s.n.m. INTRODUCTION The native bamboo diversity of Ecuador is considerable and includes seven genera of woody bamboos (Bam- buseae) and six genera of herbaceous bamboos (Olyreae). Among the woody bamboos of Ecuador, Chusquea Kunth is the most diverse genus, with an estimated 30 pes of which several are undescribed. In this paper we describe and illustrate Chusquea Ineriae, one of the Ecuadorian species new to science. The 30 or so species of Chusquea in MN fall into either Chusquea subg. Swallenochloa (McClure) L.G. Clark or Chusquea subg. Chusquea. The latter subgenus is subdivided into five sections and a few informal groups, with representatives of all five of the formally recognized sections known from Ecuador. This new species clearly belongs to sect. Verticillatae, based on its relatively thin foliage leaf blades, more or less dor- sally compressed spikelets, and extremely reduced glumes I and I (Clark 1989). Chusquea annagardneriae joins C. albilanata L.G. Clark, C. perligulata (Pilger) McClure, C. simpliciflora Munro, and C. uniflora Steud., the other members of this section in Ecuador. The new species most closely resembles C. albilanata, with which it is compared and contrasted. Interestingly enough, the first collections of this species were made only in 1989, and the remaining collections were made in 1996. Fortunately, both vegetative and flowering specimens are available so that we have a relatively complete picture of the species, but it does appear to be uncommon. This area of Ecuador, however, seems to be less well explored botanically so Chusquea annagardneriae may be more widespread than currently available collections suggest. TAXONOMIC TREATMENT Chusquea annagardneriae L.G. Clark, C.D. Tyrrell, Triplett & A.E. Fisher, sp. nov. (Fig. 1). Tyee: ECUADOR. Loja: road Velacruz-Catacocha, ca. km 6, 2040 m, ca. 4°S, 79°35'W, 11 Mar 1989 (fl), B. Ollgaard, L. Ellemann & B. Eriksen 90959 (HOLOTYPE: QCA!; ISOTYPE: AAUD). J. Bot. Res. Inst. Texas 1(2): 847 — 851. 2007 848 Journal of the Botanical R h Institute of Texas 1(2) SE GEL S NA IN uS : J A Cul | f J 7 hes. D. Synflorescence. E. Detail of branch compl t base, showing fibrillar branchl | izes of subsidiary branches. F. fl G. Detail of culm leaf pex showing pseudopetiolate blade, abaxial view. (A, C, E-G based on dark & Asimbaya 1437; B, D based o on \Ollgaard et al. 90959). Clark et al., A pecies of Cl from Ecuad 849 Culmi 2.5-4 m alti, 0.5-1.1 cm diametro, scandentes; Pacis 20-32 cm longa, MN ue Folia dex 24.5-28 cm lam ntes longa, ut videtur persistentia; vaginae 22-25cm ongae, 6-10- pl i ngi r quam la pubes in quarta parte superiore, abaxialiter glabrae, tessellatae pro p | ingulum 0.2-1 mm latum, pubescens; ze. 2.2-4 cm longae erectae, lanceolatae pl Į 1 laxiali brid abaxialitet glabrae g ; : : A IED ics) EEC Pees 1 D EE EN E : 1 m ES : dal | HERI 3-5; v 1ae glabra l pilosa I 3 b L E A TS > maculatae; laminae 2.9-8.5 cm longae, 0.4-1.3 cm latae, ratio long.:lat. 5-8.5, lanceolatae, adaxialiter glabrae, abaxialiter glabrae, non tessellatae vel interdum infirmissimus tessellatae, basibus auriculatis interdum cordatis. Synflorescentia 3-9 cm longa, aperta, pyramidalis; rami primarii infimi 1.5-4.5 cm longi. Spiculae 5-5.7 mm longae, dorsaliter compressae; glumae I et II squamiformes, minimae; glumae III et IV plerumque ad dimidium longitudem spiculae, acutae, plerumque breviter mucronatae; lemma 4.6-5 longum, subacutum vel breviter mucronatum. Woody bamboo. Culms 2.5—4 m tall, 0.5-1.1 cm diam., scandent; internodes 20-32 cm long, tuberculate, more or less terete, glabrous, a narrow band of pubescence often present just below the nodes in new shoots, but absent from the mature internodes. Culm leaves 24.5-28 cm long (only two complete examples seen), apparently persistent, juncture of the sheath and blade more or less horizontal; sheaths 22-25 cm long, 6-10 times as long as the blade, adaxially antrorsely scabrid-pubescent for at least the upper one-fourth, abaxially glabrous, tessellate on the upper two-thirds, margins free, at least the overlapping one ciliate toward the summit; girdle 0.2-1 mm wide, pubescent; inner ligule ca. 2 mm long, dark brown, abaxially pubescent, ciliate; outer ligule sometimes visible as a ciliolate rim to 0.5 mm long; blades 2.2-4 cm long, erect, lanceolate, adaxially antrorsely scabrid, abaxially glabrous, base rounded, narrower than the sheath summit and thus more or less pseudopetiolate, apex subulate. Nodes at mid-culm with one triangular central bud subtended by 40—50 subsidiary buds in several rows in a U-shaped crescent, nearly surrounding the central bud; nodal line dipping slightly below the bud/branch complement; supranodal ridge more or less pronounced. Branching um subsidiary prn: numerous, of two sizes, usually two larger and J 1 rebranching, the several to many smaller subsidiary | ginto short, curly fibrillar branchlets or longer leaf branches. Foliage leaves 3—5 per complement; sheath glabrous or pilose, mottled, only To weakly keeled toward the apex, the overlapping margin and summit ciliate; outer ligule a minute ciliolate or ciliate rim 0.1-0.2 mm long; inner ligule 0.5-0.8 mm long, truncate to rounded or slanted, margin ciliate; pseudopetiole 0.5-1 mm long, distinct, glabrous; blades 2.9-8.5 cm long, 0.4-1.3 cm wide, L:W = 5-8.5, lanceolate, adaxially glabrous, abaxially glabrous, not tessellate or occasionally very weakly tes- sellate in patches, midrib centric, margins glabrous to finely serrulate, base usually auriculate, sometimes cordate, apex acuminate-subulate. Synflorescences 3-9 cm long, open, pyramidal, well exserted from the subtending sheath; rachis flattened toward the base, becoming angular toward the apex, scabrous-pubescent; branches flattened to angular, pubescent, the primary ones strongly spreading to slightly reflexed, pulvinate at the base, the pulvinus area tomentose to villous, the lowermost primary branches 1.5-4.5 cm long, the secondary branches and pedicels also spreading; pedicels 1-3 mm long, angular, scabrid-pubescent, often sinuous. Spikelets 5-5.7 mm long, dorsally compressed; glumes I and II scalelike, no more than 1/25 the spikelet length, obtuse, glabrous, nerveless, glume I ca. 0.1 mm long, glume II ca. 0.2 mm long; glumes III and IV usually about Y the spikelet length, triangular, acute, usually shortly mucronate, margins cili- olate apically, glume III 2-3 mm long, 3-nerved, glume IV 2.2-3.3 mm long, 3- or 5-nerved; lemma 4.6-5 mm long, navicular, subacute to shortly mucronate, pubescent toward the apex; palea 4.6-5.1 mm long, bimucronate, 2-keeled, sulcate only toward the apex, 4- or 6-nerved, sulcus pubescent toward the apex. Stamens 3; anthers ca. 2.7 mm long. Lodicules 3, the anterior pair 1-1.2 mm long, asymmetrical, ciliate, the posterior one ca. 0.9 mm long, symmetrical, ciliate. Fruit unknown. Etymology.—Chusquea annagardneriae is named for our friend and colleague Anna B. Gardner (1958- 2006) in honor of her contributions to agrostology, particularly to the Grasses of lowa project (Grasses of lowa 2004) and to bambooillustration, in the form of both line drawings and website development (Bamboo Denon o listinguished from any other Chusquea by its auriculate foliage leaf blades (Fig. 1F). It is ie NE by its more or less pseudopetiolate culm leaf blades, presence of fibrillar branchlets, subsidiary branches of two sizes, and its delicate open synflorescences that bear relatively small, dorsally 850 Journal of the Botanical R h Institute of Texas 1(2) Taste 1. A gi | f Ineriae and C. albilanata Character C. annagardneriae C. albilanata Culm diameter (cm) 0.5-1.1 1-1.5 Culm height (m) 2.5-4 - Internode pubescence glabrous band of white, woolly pubescence 0.2-1 cm wide just below the nodal lines Culm leaf blade length (cm) 2.2-4 3-74 Culm leaf sheath: blade ratio 6-10 1.7-5.5 Foliage leaf blade length (cm) 2.9-8.5 8-17 (+22) Foliage leaf blade width (cm) 0.4-1.3 1.8-3.7 (4 Foliage leaf blade base auriculate to rarely cordate rounded to od rounded Foliage leaf sheath pubescence overlapping margin and summit ciliate glabrous o length (cm) 3-9 11-15 ost branch length (cm) 1.5-4.5 7-9 Vd secondary dry forest scrub and montane forests, secondary remnant forest patch vegetation Distribution s. Ecuador c. and n. Ecuador, Colombia compressed spikelets. Another unusual feature is the presence of an outer ligule in at least some of the culm leaves, but since so few complete examples were seen, it is not clear how consistent this character might be. This species is known from only five collections from the province of Loja in southern Ecuador, where it persists in secondary scrub from bosque seco (dry forest), around pastures and on rocky slopes, and at the edges of forest remnants at elevations of 2000 to 2250 m. Based on current knowledge, C. annagardneriae is endemic to Loja, of restricted distribution, and uncommon if not rare. It may, however, be more widely distributed in Loja and neighboring El Oro province, and could perhaps be expected in northern Peru. Among the Ecuadorian Chusqueas, C. annagardneriae is one of three species with fibrillar branchlets present in the branch complement (Fig. 1E); Chusquea lehmannii Pilger and C. albilanata are the other two. A fourth species, Chusquea scabra Soderstr. & C. Calderón from Costa Rica, also has fibrillar branchlets that are caused by an ascomycetous fungus (Soderstrom & Calderón 1978) and we presume that this is true for the three other species as well. Despite the presence of fibrillar branchlets, these species do not appear to be closely related except for the presumed sister relationship between C. annagardneriae and C. albilanata. Among the species of Chusquea sect. Verticillatae in Ecuador, C. annagardneriae and C. albilanata share tuberculate internodes, the presence of fibrillar branchlets in the branch complements, open pyramidal synflorescences bearing small delicate spikelets, and spikelets with glumes III and IV about half the spikelet length. Besides the distinctive auriculate foliage leaf bases of C. annagardneriae, the two species differ in several features summarized in Table 1. Most notably, mature internodes in C. albilanata retain a band of hair just below the nodal line and the foliage leaf blades are larger. The synflorescences of C. albilanata are also larger overall and the lowermost branches are longer than those of C. annagardneriae. The two 1989 collections were made from flowering plants and one of the 1996 collections (Clark & Asimbaya 143) was also made from a flowering plant, although the synflorescences were not mature. The other two 1996 collections, which were from the same area, were completely vegetative. This suggests that either C. annagardneriae has a short flowering cycle of approximately seven years, with one population simply not flowering yet (or perhaps out of synchrony with the others) or this species is a sporadic bloomer. Anu Additional specimens examined. ECUADOR. Loja: 7.7 km SE of Velacruz and 11.3 km NE of Catacocha, 2150 m, 4%01'S, 79°32'W, 12 Feb 1996 (fD, L. Clark & P. Asimbaya 1436 (AAU, ISC, LOJA, MO, Q, QAP, QCA, QCNE, US); 2 km SW of Velacruz on the road to Catacocha, 2170 m, 3°59'S, 79°35'W, 12 Feb 1996, L. Clark & P. Asimbaya 1437 (AAU, ISC, LOJA, MO, Q, QAP, QCA, QCNE, US); ca. 5 E of Velacruz, road Velacruz-Catamayo, 2250 m, 3?58'S, 79°32'W, 12 Feb 1996, L. Clark & P. Asimbaya 1438 (AAU, ISC, LOJA, QCA, QCNE, US); carretero Velacruz-Olmedo, Km 18, 2050 m, 3°55-59'S, 79°34-38'W, 11 Mar 1989 (fl), A. Freire Fierro 1270 (QCA). Clark etal., A pecies of CI from Ecuad 851 ACKNOWLEDGMENTS Field work in Ecuador by Clark was supported by National Science Foundation (NSF) grant DEB-9218657 and was conducted under the auspices of the Pontifícia Universidad Católica del Ecuador (Herbario QCA). Clark thanks the staff of QCA for their generous assistance and for access to the herbarium. Final preparation of the manuscript was supported by NSF grants DEB-0515712 to Clark and DEB-0515828 to S. Kelchner (Idaho State University). We thank Anna B. Gardner for the fine illustration of this species. REFERENCES BAMBOO Biopiversity. 2005. http;//www.eeob.iastate.edu/research/bamboo/index.html CLARK, L.G. 1989. Systematics of Chusquea section Swallenochloa, section Verticillatae, section Serpentes, and section Longifoliae (Poaceae: Bambusoideae). Syst. Bot. Monogr. 27:1—127 Grasses or lowa. 2004. http://www.eeob.iastate. edid SODERSTROM, T.R. and C.E. CALDERÓN. 1978. Chusquea and Swall (Poac Bambusoid g icrelation- ships and new species. Brittonia 30:297—312. 852 Journal of the Botanical R h Institute of Texas 1(2) BOOK REVIEWS PAUL MARTIN BROWN (TEXT) and STAN ForsoM (Artwork). 2008. Field Guide to the Wild Orchids of Texas. (ISBN 978-0-8130-3159-0, pbk). University Press of Florida, 15 Northwest 15th Street, Gainesville, FL 32611-2079, U.S.A. (Orders: www.upf.com, 352-392-1351, 352-392-7302 fax, 1-800-226-3822). $29.95, 316 pp., 286 color plates, 7 water color plates, 91 b/w illustrations, 67 maps, 6" x 9". Once again Paul Martin Brown and partner artist Stan Folsum have produced a gem of a regional flora. Readers wh familiar with their work are already acquainted with the well established f it of the volume: initial visual beauty accompanied by straightforward introductions and explanatory materials, logical progression in presentation of materials, concise and germane data, detailed regional maps, botanical glossary for non-botanists, appendices, and much more—all in a convenient-size volume to tuck into your backpack as you go searching for those plants in situ both experienced and novice seekers of the wild orchid in its natural habitat, spotting and identifying the plant becomes For much simpler with the aid of this field guide. Maps, photos, and pM. in addition to the carefully included usual flowering times 1 1 T Ery pe es E: T 1 Es 1 TNS LI S | 1 : A 1 1 1 1 I guag identify “finds.” Simplifyi l he line drawings and pl c act 1 lcl 1 log necessary to separate and eee the plants. Even first time venturers into the field on peri gl d excit t to become avid enthusiasts With the upcoming holidays, this exquisite volume is an excellent gift choice.—Helen Jeude, Botanical Research Institute of Texas, Fort Worth, TX 76102-4060, U.S.A. ARTHUR M. SHAPIRO. 2007. Field Guide to the Butterflies of the San Francisco Bay and Sacramento Valley Regions. (ISBN 978-0-520-24469-6 hbk.; 978-0-520-24469-8, pbk.). University of California Press, Berkeley, CA. (Orders: CPFS c/o University of California Press, 1095 Essex Street, Richmond, CA 94801, U.S.A.; 609-883-1759; 609-883-7413 fax) $30.00 hbk., $18.95, pbk., 346 pp., 31 color plates 11 x18 em. One would expect excellence in a work by this well-known and accomplished lepidopterist, and, indeed, this field guide is among the best. It is also important for its coverage of regions where loss of habitat and therefore host pu continue to io a number of species and subspecies. The introduction alone i us Bs price of the book wit geog of the areas E a. anda ire readable and detaile p NN flying seasons, and larval host a ants—and gan more. Notes iin: dE. ET : fa 4 P 1 E SOS g mation that could otherwise be gleaned ES gh i hi ] curr aaa literature. Still, itis a practical guide for the amateur butterfly watcher as well as pe with intense serious interests. cda ed o: good ES DEN butterflies as the viewer sees them—in flight or pausing to nectar, with a depicti g tion. Added features are checklists of butterflies of the Bay Area and regional lists of those of the Sacramento Valley and Sutter Buttes. is field guide can be recommended for any one interested in butterflies whether or not one frequents the regions covered. It is "xs and informative reading.—Joann Karges, (TCU Library, o m Research Institute of Texas, Fort Worth, TX, 76102-4060, J. Bot. Res. Inst. Texas 1(2): 852. 2007 ESTUDIOS EN LAS ACOCTNACEAE NEOTROPICALES XXX: TRESNUEVAS ESPECIES ANDINAS:DE MANDEVILLA (APOCYNOIDEAE: MESECHITEAE) J. Francisco Morales Instituto Nacional de ud (INBio) Apartado Posta Santo Domingo, idm ee RICA RESUMEN Se describen Mandevilla mequatonaliss M. pieds y M. lejana: tres nuevas especies de los Andes de Ecuador y Perú. Se incluyen descripciones, ilustracion las especies más afines. PALABRAS CLAVE: Ápocynaceae, Apocynoideae, Mesechiteae, Mandevilla ABSTRACT Mandevilla aequatorialis, M. niam 2 M. lojan, ME new Vp from the Andes of Ecuador and Peru, are described and il- lustrated and their relationshi L La concepción y circunscripción del género Mandevilla Lindl., el más grande de la subfamilia Apocynoi- deae, ha variado notablemente desde la última monografía del género (Woodson 1933). En aquel entonces, Mandevilla se trató como una unidad genérica distinta de Macrosiphonia Müll. Arg., con alrededor de 120 especies. Pichon (1948) no aceptó dicha propuesta taxonómica y propuso unir de nuevo ambos géneros, lo cual prácticamente no fue seguido y paso desapercibo en afios subsecuentes. Zarucchi (1991) propuso el género monotípico Quiotania, un grupo similar en varios caracteres morfológicos a Mandevilla, del que difería principalmente por el tamaño de las corolas. En años posteriores, Henrickson (1996), elevó a nivel genérico una de las dos secciones de Macrosiphonia, proponiendo el género Telosiphonia, grupo restringido principalmente a México y el S de Estados Unidos. De esta manera, Mandevilla se volvió un género con alre- dedor de 3 géneros satélites cercanamente relacionados (Macrosiphonia, Quiotania, Telosiphonia) y otros cuya relación aparentemente no era tan estrecha (Allomarkgrafia Woodson, Mesechites Múll. Arg., Tintinnabularia Woodson). Como resultado de estudios cladísticos derivados del análisis de varios indicadores molecula- res y morfológicos, Simóes et al. (2004, 2006, 2007) redefinieron los límites genéricos de Mandevilla, de manera que Macrosiphonia, Quiotania y Telosiphonia fueron reducidos a la sinonimia. Esta propuesta, elevó el nümero de especies de Mandevilla a más de 170, convirtiéndolo en el género más grande de las Apocynaceae neotropicales. La evidencia filogenética provista en los trabajos de Simões et al. (2004, 2006) demostró que la clasificación infragenérica de Woodson (1933) es altamente parafilética, mientras que la propuesta por Pichon (1948) es monofilética. Sin embargo, la mayoría de los taxones infragenéricos propuestas por ambos taxonómos son polifiléticos, por lo que se necesita mayor evidencia y un estudio más detallado para poder establecer una nueva clasificación intragenérica. Continuando con el desarrollo de una nueva monografía de Mandevilla, se describen tres nuevas espe- cies, restringidas a los Andes de Ecuador y Perú. Mandevilla aequatorialis J.E Morales, sp. nov. (Fig. 1). Tiro. ECUADOR. Imsasura: Cotacachi, parroquia Plaza Gutiérrez, Tabla Chupa, 12 May 1992 (fl), G. Tipaz et al. 976 (notoriro: MOI, fotografía, INB!; isoripo: QCNE!). A Mandevilla jasminiflora Woodson cui affinis, foliis basi cordatis, sepala longiribus, et antheris auriculatis acuminatis differt. Liana con tallos subleñosos, ramitas aplanadas cuando jóvenes, subcilíndricas en la madurez, moderada y diminutamente puberulentas cuando jóvenes, el indumento más esparcido en tallos viejos, los coléteres J. Bot. Res. Inst. Texas 1(2): 853 — 857. 2007 854 Journal of the Botanical R h Institute of Texas 1(2) interpeciolares cortos e inconspicuos, cónicos, ca. 1 mm de largo, formando una línea interpeciolar inconspicua. Hojas: pecíolos 10-23 mm de largo; láminas 2,5-6,3 x 1,1-2,8 cm, ovadas a ovado- elípticas, firmemente membranáceas, el ápice gradualmente acuminado o agudo, la base leve o conspicuamente cordada, glabras en la superficie adaxial, glabrescentes a densa y diminutamente puberulentas en la superficie abaxial, sin doma- cios a lo largo del nervio central en la superficie abaxial, coléteres del nervio central en grupos 3 6 4, ca. 1 mm de largo, agrupados en la base del nervio, venas secundarias más o menos ar- cuadas. Inflorescencias más larga o igualando en longitud las hojas adyacentes, axilar, densa a moderadamente puberulentas, con muchas flores, pedúnculo 14-28 mm de largo, pedicelos 8-10 mm de largo, brácteas 1-3 x 0,5-1 mm, escariosas; sépalos 3-5 x 1-1,5 mm, angostamente ovados, acuminados, glabrescentes, a veces con unos cu- antos pelos esparcidos cerca de la región apical y a lo largo del nervio medio, con 2 6 3 coléteres en la base de la cara adaxial, ubicados cerca de los márgenes, ca. 0,5 mm de largo, irregularmente lacerados apicalmente; corola hipocrateriforme, verde amarillenta, glabra exteriormente (excepto por la cara adaxial de los lóbulos), los cinco lóbu- los con la aestivación dextrorsa, tubo 3-4 mm de largo, 2 mm en diámetro en la boca, recto y no inflado, con una corona anular rudimentaria y con prolongaciones basales sobre cada lóbulo de la Fic. ialis. A. Ramit fl las. B. Detallede corola, ca. 0,5 mm de largo o menos, lóbulos 2,5-4 las uina del nervio vio cent Vito. C C. oe dab dide x 2-2,5 mm, angostamente obovados, extendidos, a adaxia con dos lobDulos yd e pero algo reflexos distal y marginalmente, densa Gineceo, most- ando las anteras, disco y ovario. e Detalle de la cabeza estigmática (G. — Y diminutamente papilado-tomentulosos en la mam 976, MO). superficie adaxial; estambres insertos en la mi- Y E tad distal, anteras ca. 3 mm de largo, glabras, los ápices exsertos, la base sagitada, ovario ca. 1 mm de largo, glabro, estilo recto, cabeza estigmática 1,5-2 mm de largo, el apículo cerca de 1-1,4 mm de largo; nectario variada e irregularmente pentalobulado, más corto que el ovario. Folículos desconocidos. Distribución, habitat y ecología.—Esta especie es endémica a Ecuador, donde se conoce de los departa- mentos de Pichincha e Imbabura, donde crece en bosques húmedos montanos, en elevaciones de 2300-3000 m. Floración ocurre en mayo y septiembre. Mandevilla aequatorialis es morfológicamente algo similar a M. jasminiflora Woodson, pero se puede separar fácilmente por sus hojas levemente cordadas basalmente (vs. redondeadas a obtusas), láminas foliares con las venas secundarias arcuadas (vs. dispuestas ca. en ángulo recto en relación al nervio central, flores con los sépalos más largos (3-5 x 1-1,5 mm vs. 1-1,5 x 1-1,5 mm) y anteras con las aurículas estériles sagitadas, largas y cortamente acuminadas. El epiteto de este nuevo taxón hace referencia al hecho de que la especie es conocida por el momento solo en Ecuador. Morales, NM H A M a HII A E Al y Perú 855 Especímenes adicionales examinados. ECUADOR. Pichincha: Quito, carretera Nono-Guarumos-Tandayapa, ent y Cerro Guarumos, 1 Sep 2001 (fD, A. Freire-Fierro et al. 3143 (QCNE) Mende frigida J.E Morales, sp. nov. (Fig. 2). Tiro. PERÚ. Piura: Cerro Aypate, E de Ayabaca, 22 Sep 1991 (fl), A. Gentry al. 74988 (HoLoTIro: MO}, fotografía, INB!; isoriros: INB!, USM!). A Mandevilla assimilis (K. Schum.) J.F. Morales, cui affinis, foliis glabris, subtus in axillis nervi medii foveolatis et antheris 3,5-4 mm longis differt. Liana, ramitas inconspicusmente aplanadas cuando jóvenes, cilíndricas o subcilíndricas cuando viejas, glabras; coléteres interpeciolares i | ca. 1 mm de largo. Hojas: pecíolos 5-8 mm de largo; láminas 2,2—5,4 x 1,2-2,4 cm, elípticas, e membranáceas, abruptamente corto-acuminadas apicalmente, obtusas a anchamente cuneadas apicalmente, glabras, usualmente con domacios a lo largo del nercio central en la superficie abaxial, coléteres del nervio central en grupos de 1 ó 2, ca. 1 mm de largo, agrupados en la base del nervio, venas secundarias más o menos arcuadas. Inflorescencia igualando las hojas adyacentes, axilar, algunas veces pareciendo terminal o subterminal tallos jóvenes, glabras, con pocas flores, pedún- culo 35-47 mm de largo, pedicelos 10-14 mm de largo, brácteas 3,5—4,5 x 0,5 mm, escariosas; sépalos 2,5-3,5(-4) x 1-1,5 mm, angostamente ovados, agudos, muy diminuta e inconspicuamente puberulentos, coléteres enteros, subenteros o apicalmente lacerados; corola hipocrateriforme, blanca, muy diminuta e inconspicuamente puberulenta exteriormente, tubo 10-11 mm de largo, 2-2,5 mm en diámetro en la boca, abultado sobre la posición de los estambres, lóbulos 8-9 x 2 mm, angostamente elípticos, erectos a suber- ectos, diminutamente puberulentos por ambas caras; estambres insertos en ca. la mitad de la longitud del tubo de la corola, anteras 3,5—4 mm de largo, glabras, incluidas, la base subauriculada; ovario ca. 1,5 mm de largo, glabro, estilo levemente curvado, cabeza estigmática 2,5-3 mm de largo, el apículo ca. 2 mm de largo, nectarios 2, ca. 1 mm de largo. Folículos desconocidos. Distribución, habitat y ecología.—Endémica a Perú, donde es conocida únicamente de la localidad tipo, en bosques montanos nubosos, en elevaciones de 2700-2800 m. Especímenes con flores han sido recolectados en septiembre. Mandevilla frigida se encuentra relacionada con M. assimilis (K. Schum.) J.F. Morales, conocida de Ec- uador y Perú, de la que difiere por sus láminas foliares con la superficie abaxial glabra y con domacios a lo largo del nervio central (vs. tomentulosa a densamente puberulenta y sin domacios) y anteras más cortas (3,5-4 mm vs. 4,9—5,2 mm). El epiteto de este nuevo taxón hace referencia al hecho de que la especie crece en sitios fríos de elevaciones altas de la Cordillera de los Andes. Mandevilla lojana J.E Morales, sp. nov. (Fig. 3). Tro. ECUADOR. Loja: Cerro Villanaco, carretera Loja-La Toma, 17 Dic 1988 (fl, fr), J. Madsen 75938 (HoLotipo: QCA!, fotografía, INB!; isoriro: QCNE!, fotografía, INB!). A Mandevilla ligustriflora Woodson, cui affinis, tubo longiribus et corollae lobis 5-7 mm longis differt. Liana, ramitas tiernas aplanadas y anguladas, cilíndricas o subcilindricas cuando viejas, glabras; coléteres interpeciolares inconspicuos, ca. 0,5 mm de largo. Hojas: pecíolos 4-8 mm de largo, láminas 3,5-5,6 x 1-1,8 cm, elípticas a obovado-elípticas, firmemente membranáceas, agudo-mucronuladas apicalmente, cuneadas a obtusas basalmente, glabras en ambas superficies, usualmente con domacios a lo largo del nervio central en la superficie abaxial, coléteres del nervio central en grupos de 2, ca. 0,5 mm de largo, agrupados en la base del nervio, venas secundarias más o menos arcuadas. Inflorescencias más corta que las hojas adyacentes, axilar, glabras, con muchas flores, pedúnculo 27-29 mm de largo, pedicelos 14-16 mm de largo, brácteas 1 x 0,5-1 mm, escariosas; sépalos 2 x x 1 mm, ovados, agudos, glabros a glabrescentes, coléteres irregular- Jem Top. mente lacerados; corola me, blanca, diminuta e inconspicuamente puberulenta exteriormente, parte EM M del tubo de la corola 3-4 x 2 mm, parte superior angostamente cónica, 6-6,5 mm de largo, 5-6 mm en diámetro en la boca, corona reducida a vestigios coronales, leve y tenue, lóbulos 5-7 x 2,5-3,5 mm, ovados, erectos a suberectos, diminutamente puberulentos en ambas superficies; estam- bres insertos en la base de la parte superior del tubo, anteras ca. 4,5 mm de largo, glabras, incluidas, la base subauriculada, ovario ca. 1 mm de largo, glabro, estilo recto o levemente doblado, cabeza estigmática 33,2 856 Journal of the Botanical R h Institute of Texas 1(2) mm de largo, el apículo 2-2,2 mm de largo, nectarios dos, casi tan largos como el ovario. Folículos 12,5-25 x 0,2-0,4 cm, levemente moniliformes, glabros; semillas 10-11 mm de largo, glabras, coma 7-14 mm de largo, crema. Distribución, habitat y ecología.— Esta especie es endémica al S de Ecuador, conocida por el momento únicamente en la localidad tipo, en bosques montanos nubosos y veg- etación paramosa, en elevaciones de 2500 m. Especímenes con flores y frutos han sido recolectados en diciembre. Mandevilla lojana está cerca- namente relacionada con M. ligus- triflora, pero se puede separar por sus flores con sus partes (sépalos, tubo, lóbulos, anteras) más largas y grandes, así como por sus hojas con los domacios distribuidos a lo largo del nervio central en la superficie abaxial (vs. restringidos a la base del nervio). El epiteto de este nuevo taxón honra a la provincia de Loja, sitio peculiar e interesante en Ecua- dor de la que numerosos endemismos son conocidos. AGRADECIMIENTOS Quiero agradecer a los curadores de los herbarios COL CUVC- HUA, Fic 2. Mandevilla frigida AR it infl ias. B. D | b i à HUQ, JAUM, K, MEDEL, MO, Q, leni | |. C. Detalle delos d D. Cáliz, pedicelo, bráctea y E H i E m uu uu OG OCNE TOPES SUS e corola. E Sé| | vista adaxial. F. Tul lla " E 7 | Icañalad flecha). G. Antera, por permitirme facilidades para el estilo, ovario y nectario. H. Detalle de la cal tigmática (Gentry et al. 74988, MO). uso de colecciones. Asimismo, se J agradece al personal del herbario de la Universidad de Antioquia (HUA), por el apoyo logistico continuo brindado durante mis multiples visitas a Medellin, Colombia, asi como por la asistencia en el trabajo de campo desarrollado. Se agradece también grandemente a Steffen Matezki, de la Universidad de Bayreuth, Alemania, por suministrar y autorizar el uso de las fotografias de M. ligustriflora. La asistencia de las siguientes personas en facilitar la visita de sus respectivos herbarios es grandemente agradecida: Alberto Agudelo (HUQ) en Armenia, Quindio, Colombia, Homero Vargas (QCNE) en Quito, Pichincha, Ecuador, Philip Silverstone-Sopkin (CUVC) en Cali, Valle del Cauca, Colombia y Ricardo Callejas (HUA), en Medellin, Antioquia, Colombia. M H A M PI HII A E Al Morales Fic. 3 Mandevillalojana A. Ramit infl ias. B. D del i tral enl fi laxial. C. Detall | ios. D. Sépalo y coléter, vista adaxial. E. Caliz, pedicelo, bractea y corola. F. Tubo dela corola abierto, mostrando las Hu * H rj A Y J £ Ļ Y G n Hu Jal L : fet H. Anteras, estilo, ovario y nectarios. I. Folículos. J. Semillas (Madsen 75938, QCA). 857 REFERENCIAS HENRICKSON, J. 1996. Studies in Macro- siphonia (Apocynaceae): generic recognition of Telosiphonia. Aliso 14:179-195. PicHoN, M. 1948. Classification des Apocy- nacées. X. Genre Mandevilla. Bull. Mus. Hist. Nat. (Paris), sér. 2, 20:211-216. Simoes, A.O., M. ENDRESS, T. VAN DER NIET, L. KINOSHITA & E. Conti. 2004. Tribal and intergeneric relationships of Mes- echiteae (Apocynoideae, Apocyna- ceae): evidence from three not icoding pastid DNA regions and morphology. Amer. J. Bot. 91:1409-1418. Simoes, A.O., M. ENDRESS, T. VAN DER NIET, L.S. KINOSHITA & E. Conti. 2006. Is Mandevilla (Apocynaceae, Mesechiteae) mono- phyletic? Evidence from five plastid DNA loci and morphology. Ann. Mis- souri Bot. Gard. 93:565-591. Simoes, A.O., L.S. KINOSHITA & M. ENDRESS. 2007. New combinations in Mandev- illa Lindley (Apocynaceae). Novon 17:89-90 Woobson, R.E. 1933. Studies in the Apocy- naceae. IV. The American genera of Echitoideae XXVI. Ann. Missouri Bot. Gard. 20:605-790. ZARUCCHI, J.L. 1991. Quiotania: a new genus of Apocynaceae-Apocynoi- deae from Northern Colombia. Novon 1:33-36. 858 Journal of the Botanical R h Institut f Texas 1(2) BOOK REVIEW GERHARD GOTTSBERGER and ILSE SILBERBAUER-GOTTSBERGER. 2006. Life in the Cerrado: a South American Tropical Seasonal Vegetation Vol 1. Origin and Structure, Dynamics and Plant Use. (ISBN 3- 00-017928-3, hbk.). Reta Verlag, Ulm, Germany. (Orders: ilse.silberbauer@uni-ulm.de; Botanischer Garten und Herbarium, Universitaet Ulm, Hans-Kreb-Weg o.n., D-89081 Ulm, Germany). $64.60, 280 pp., numerous color figures, maps, ques and tables, 29.5 x 22.5 em. The book Life in the Cerrado: a South American Ti ER 1E o eu 1 P sd O. the following: definition of a cerrado, location soil types, origin of ph g diversity of flora life history, and natural and human influences. This book is full of very c olorful ima ges, maps and charts that cl l expla in the flora found in the cerrado. The authors begin by defining B term as a Pd x um type that is drastically different from the vegetation found in a typical savanna; these differ | ition, and fl . This vegetation is mostly found in central Brazil and has changed contac ened nS. The soil composition varies and has very poorly developed mineral composition. Cerrado soils are well drained, and soil moisture is found to mace influence the ees of a hera are five basi do f , distinguishable by vegetation height and density. In order from t these are: cerradáo, cerrado sensu stricto, campo cerrado, canes sujo, and campo limpo. ee all five forms exhibit some of i e domi m nant species, floristic diversity of th varies o locality. Differences can occur over relatively short distances, often abruptly changing from open to more closed canopy mt ed: of non- pod cad nd bs cerrado also occurs, sometimes forming ecotones or transitional areas in the process. T alc needed 11 order to fully understand t I iti and geographical r gi ns of th rrad The noe of the cerrado consists of trees, lianas, palms, VA vines, ps au few vascular epiphytes. Trees often appear stunted k, fissured, and corky bark and are adapted for rapid g lopment. piel ay appear quite old, but longevity of fth I 30-40 years. A g | f other pl l hil d life, many with thicl ] i Į water ODD and me high aluminum content of the soil Ecological and cli f hat infl 1fl il ilability, fire, frost, and seasonality. Moisture is of found until a soil depth of 2.0— de meters o iain, longer this water dur- ing the dry season. To combat the nutrient-deficient soil, other pl I f f nitrog ] phospl Fire pow a Pp 2 influence on the NA 2 deterring non-native or p tablish t ] fl aiding in g g f dlings. Frost is very similar to fire in that i damage plant but l l i getation with tl tion of new leaves. Fire is found throughout th do f ti idered more of a local eS The region ane just two seasons: dry and wet. Flowering and fruiting of ee dd die is determined by he parti and nati ue Sad on these seasonal patterns. lly i I g There are many ful I l ] aluable vood, fruits ani seeds, oils, cork, fiber, and pigments. However, th ] ion tl hi ded i s to peed understand the medicinal uses of various plants and the potential economic resources available. Adding to its overall value and importance, this area ee also been es asa is che ola dis au motel i eps has drastically decreased in size over the last decade. The 1 F population. This was a very i ing ee o ighly ] i li ical 1 or pi 1 1 € 1 j 3l I 1 1 fuli charts tables, and maps to help better explain the cerrado. The "d will take away a better understanding of the specific vegetation found in this region and the natural and anthropogenic influences that have effected its growth.—Keri McNew, MS Biol Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A ogy, Project Manager, J. Bot. Res. Inst. Texas 1(2): 858. 2007 ESTUDIOS EN LAS APOCYNACEAE NEO TROPICALES XXXI: EL COMPLEJO DE MANDEVIELA HIRSUTA Y CUATRO NUEVAS ESPECIES J. Francisco Morales Instituto Nacional de o (INBio) Apartado Postal 22 Santo Domingo, Heredia, a RICA RESUMEN a] Landevilla hi d ib orrida (Perú), M. inexperata (Ecuador), M. megabracteata (Colombia y Guyana) y M. similaris (Venezuela). Se A ilustraciones, descripciones, datos de distribución, especímenes examinados y se comentan las relaciones con los taxones más relacionados. PALABRAS CLAVE: Apocynaceae, Apocynoideae, Mesechiteae, Mandevilla ABSTRACT i ENEA y P Pes 1 f, Following a study of the M w species are described: M. horrida (Perú), M. Mic Adm (pruden. M. megabracteata (Colombia and Guyana) and M. Tu (Venezuela). Illustrations, descriptions, distributi ined, as well as discussions about closely related taxa are included Mandevilla Lindl. (Apocynoideae, Mesechiteae), es el género más grande de las Apocynaceae con más de 170 especies distribuidas desde México y las Antillas hasta el N de Argentina. La percepción del mismo ha cambiado notablemente desde la última monografía (Woodson 1933), ya que varios de los géneros cercana- mente relacionados y considerados como grupos distintos desde hace décadas (e.g., Macrosiphonia), no han sido apoyados por estudios moleculares recientes (Simoes et al. 2004, 2006, 2007). En términos general ndevilla es un grupo complejo por varias razones, principalmente por la gran cantidad de taxones que lo conforman, la similaridad vegetativa de algunas especies, así como por la alta variación intraespecífica presentes en algunos grupos, en cuanto a la densidad y distribución del indumento en estructuras florales y vegetativas se refiere. Mandevilla hirsuta (Rich.) K. Schum. es la especie con el mayor rango de distribución geográfica en el género, el cual se extiende desde el S de México hasta Brasil, Bolivia y Paraguay (Morales 2005). Esta especie presenta una alta variación intraespecifica en cuanto a la densidad del indumento de sus partes vegetativas, de manera que existen especimenes con hojas densamente hirsutas en su superficie abaxial (Meneses et al. 693, INB, LPB), hasta otros en los cuales el indumento esta casi ausente y las hojas son prácticamente glabras (Melo et al. 827, INB, UB). Sin embargo, existen una serie de variaciones o estados intermedios que no permiten reconocer variedades basado en la densidad del indumento. A pesar de esta variabilidad, los caracteres florales son relativamente constantes, de manera que es posible reconocer fácilmente una sola unidad taxonómica en todo el rango de distribución de la especie. 1 * a A través del tiempo y basado en la variabilidad del indumento, otras esp con ese taxón, han sido confundidas como M. hirsuta, grupo en el cual podemos distinguir a M. bracteata (Kunth) Kuntze, M. krukovii Woodson, M. lancibracteata Woodson, M. moritziana (Múll. Arg.) Donn.Sm., M. sagittarii Woodson y M. steyermarhii Woodson. Este grupo, el cual conoceremos como el complejo de M. hirsuta, ende en ies Je] 186 (A beg al común inflorescencias con brácteas foliáceas o subfoliáceas, corolas i mes y moniliformes. En general, este complejo no ha sido estudiado con detalle y estudios preliminares recientes han revelado varias novedades taxónomicas (e.g., Morales 2005, 2006) Como resultado de una revisión detallada del complejo de M. hirsuta, se proponen cuatro nuevas espe- a] cies. Adicionalmente, se brinda la sinonimia completa de M. hirsuta, incluyend ipciones, ilustraciones y datos de distribución para cada taxón. J. Bot. Res. Inst. Texas 1(2): 859 — 869. 2007 860 Journal of the Botanical R h Institute of Texas 1(2) Una clave para las especies del subgénero Exothostemon con brácteas florales foliáceas se brinda a con- tinuación. Algunos datos de esta clave han sido tomados de una nueva monografía de Mandevilla (Morales, datos sin publ.) por lo que pueden diferir de otros publicados anteriormente (e.g., Woodson 1933). 1. e de la corola hipocrateriforme Lóbulos de | la bl la gargant illa; láminas foli 7-16 mm de ancho, las venas ebd usualmente no evidente aridana J.F. Morales Lóbulos de la a amarillos, la garganta usualmente ele a anaranjada; laminas foliares 15- - mm anchor las venas terciarias evidentes al menos abaxialm 3. Corola muy diminuta y es id pul | lal glal te ext te, la pubescen- E | | Al P | nen 15) a os glabros M. rugellosa (Rich.) L. Allorge Ps pilosa a pilosulosa externamente; inflorescencia densa a moderadamente pilosulosa; folículos rcidamente hirsutos a da M. villosa (Miers) Woodson 1. Tubo de la po infundibulifo 4. Láminas foliares prem. con E base obtusa, cuneada a redondeada _M. javitensis (Kunth) K.Schum. 4. Láminas foliares usualmente con la base cordada a subcordada. 5. Hojas sésiles a subsésiles, con el peciolo 1-2 mm de largo M. tristis J.F. Morales 5 Hojas eee [eco neus eon el pue a " d mm de largo. GA d a lo largo del nervio central y de las venas secundarias en la superficie adaxial sépalos 1-2 mm de largo M. moritziana (Müll.Arg.) Donn.Sm. 6. Láminas foliares no bulladas; coléteres distribuidos solo a lo e ac) nervio SO o si ocasional- mente presentes en venas secundarias (M. lancibracteata) | del tub de 9 mm de largo; sépalos 2,5-18 mm de largo 7. Parte inferior del tubo de la corola 7-14 mm. 8. Sépalos 7,5-9 mm de largo, parte inferior del tubo de la corola 12-14 mm de largo M. ar EM rales 8. Sépalos 3-3,8 mm de largo, parte inferior del tubo de la corola 7-9 mm de largo ____M. E 7. Parte inferior del tubo de la corola 18-35 mm. 9, 7- lar mm de algo: in tal s5 Q l | pall sl puberulentos cuando jóvenes, glabrescentes con la aes M. similaris J.F. Morales 10. Venas terciarias evidentes; jeudi | hirsutos, híspidos a hispidulosos i Brácteas florales i 681 mm de Tu M. megabracteata J.F. Morales . Brácteas florales 11-25 mm de largo. 12. Parte superior a tubo de la corola 15-21(-25) mm de largo, con un diámetro de (1 6-21 mm en el orificio; folículos libres, solo unidos en el ápice — — M. hirsuta (Rich) K. Schum. 12. Parte superior del tubo de la corola 27-31 mm de largo, con un diámetro de 7-9 mm en el orificio; folículos fusionados a lo largo de su entera longitud _ M. sagitarii Woodson 9. Sépalos 2,5-6 mm de largo. 13. Tallos con una conspicua línea interpeciolar en cada nudo; ramitas jóvenes pelea glabrescentes, el indumento no evidente steyermarkiiWoodson 13. Tallos careciendo de una conspicua línea interpeciolar; ramitas hirsutulas, psa pu- berulentas a híspidas 14. Láminas foliares con el ápice obtuso-mucronado a redondeado-mucronado; tubo de la corola con un diámetro de 14-22 mm en la fauce — — — M. bracteata (Kunth) Kuntze 14. Láminas foliares con el ápice caudado-acuminado, corto-acuminado o cortamente cuspidado; tubo de la corola con un dpi. de En 3 mm en la fauce 15. Tallos esp | li lentos; brá ae 17-28 mm de largo; Mit inferior del bo 30- 357 mm de largo ___ la Woodson 15. Tallos hispidos; bracteas florales 32-46 mm de largo; parte inferior del tubo 2 5-23 mm de largo M. ais J.F. Morales M H dal | PA pee | J sila hi + 861 Morales Mandevilla hirsuta (Rich.) K. Schum., Nat. Pflanzenfam. 4(2):171. 1895. (Fig. 1). aa Du Rich., Actes Soc. Hist. Nat. Paris 1:107. 1792. Echites richardi Roem. € Schult., Syst. Veg. 4:391. 1819, nom. illeg. Ambl ) Miers Apocyn. S. Amer. 185. 1878. Tiro: GUYANA FRANCESA. Cayenne, 1792 (fl), Leblond s.n. (Lectotiro, designado por Allorge-Boiteau (1998), P-LA [foto, INB]; isoLecToTIPOS, C, G-DC [foto F neg. 26863], P [2 cartulinas]). Echites tomentosus Vahl, Symb. Bot. 3:44. 1794. Ed tomentosa du Miers, pun S. Amer. 213. 1878. Mandevilla tomentosa (Vahl) Kuntze, Revis. Gen. Pl. 2:416. 1 M K. Schum., Nat. Pflanzenfam 4(2):171. 1895, non. inval. Tipo: GUYANA FRANCESA: Cayenne, fecha ined (fl), von Rohr 63 (HoLotipo: C [2 usum isoriPOs: BM [2 cartulinas, fotos, INB]). us UE Nov. Gen. 2: ee ed. e 3:218. 1818 [1819]. Exothostemon macrophyllum (Kunth) G. Don, Gent. Hist. 4:82. Sc at. Pflanzenfam 4(2):171. 1895. Tiro. VENEZUELA. Amazonas: río Orinoco, cerca Echites ibn e 1837.) n Borja, Carchi Isla Panunamae, May año A fl), Humbol i ER 1214 (HoLotiro: P-HB; isotirO: B, destruido). m campestris Vell., Fl. Flumin. 113. 1829. Amblyanthera campestris (Vell.) Müll.Arg., Fl. Bras. 6(1):149. 1860. Rhabdadenia camp- estris (Vell.) Miers, Apocyn. S. Amer. 121. 1878. Tipo: BRASIL. Rio DE JANEIRO: Vell., Fl. Flumin., Icon 3:tab. 43. 1827 (uecroripo, esignado aquí). Echites hispidus Willd. ex Roem. & Schult., Syst. Veg. 4:795. 1819. Amblyanthera hispida (Willd. ex Roem. € Schult.) Müll.Arg., Fl. 1):147. 1860. Mandevilla hispida (Willd. ex Roem. & Schult.) Hemsl., Biol. Cent.-Amer., Bot. 2(10):316. 1881. Mandevill tomentosa var. hispida (Willd. ex Roem. & Schult.) Kuntze, Revis. Gen. Pl. 2:416. 1891. Tiro: BRASIL. Datos perdidos, Hoffmannsegg qu oriPo: B-W 5166 [foto F neg. 4490]). Echites neg Pohl ex Stadelm., Flora 24(1 Beibl.):25. 1841. Mandevill iculata (Pohl ex Stadelm.) K. Schum., Nat. Pflanzenfam. 40 1. 1895. Tipo: BRASIL. DE JANEIRO: Serra de Macacú, fi dida (fl), Pohl 657 (HoLotipo: M; IsOTIPO: NY). Echites Mu Ruiz & Pav. var. n. ius Stadelm., Flora 24(1 Beibl.): 27. 1841. Tipo: BRASIL. Bania: Datos perdidos (fl), Martius s.n. (HoLoTiPo: M [fotocopia, INB Echites hirsutus Ruiz & Pav. var. angustifolius Stadelm., Flora 24(1 Beibl.):28. 1841. Echites lasiocarpus A. DC. var. angustifolius (Stadelm.) A. DC., Prodr. 8:463. 1844. Tipo: BRASIL. Rio DE Janeiro: Porto de Estrella, fecha perdida (fl), Martius s.n. (Lectotipo, designado aquí, M [foto, INB]). Echites almadensis Stadelm., Flora 24(1 Beibl.):28. 1841. Amblyanthera palustris Müll.Arg. var. almadensis (Stadelm.) Múll.Arg., Fl. Bras. 6(1):146. 1860. Tiro: BRASIL. Banu: Itahypé, Almada, fecha perdida (fl), Martius s.n. (HoLotipo: M, [fotocopia, INB, foto F neg. 20150] hi lel ex Stadelm., Flora 24(1 Beibl.):29. 1841. Amb |i dapib (A. DC.) Müll.Arg. var. stadelmeyeri (Mart. ex TIPO: no localizado). Mart Stadelm.) Müll.Arg., Fl. Bras. 6(1):149. 1860. Tipo: BRASIL. Minas Gerais: datos perdidos, Ackermann s.n. (HOLO Minas Gerais: Lavras, Reserva Poco Bonito, 14 Nov 1987 (fl), Gavilanes & — designado aqui, INB; IsoNEoTIPO, ESAL) Echites ciliatus Stadelm., Flora 24(1 Beibl.):32. 1841. Amblyanthera ciliata (Stadelm.) Müll.Arg., Fl. Bras. 6(1):145. 1860. Tiro: BRASIL. Bania: Caiteté, fecha perdida (fl, fr), Martius s.n rbi ca M [fotocopia, INB, foto F neg. 20143]) Echites hirsutus Ruiz & Pav. var. angustifolius Stadelm., Flora 24(1):Beibl. 28. 1841. Echites lasiocarpus A. DC. var. angustifolius (Stadelm.) DC., Prodr. 8:463. 1844. Tiro: BRASIL. Rio DE JANEIRO: Porto d'Estrella, fecha perdida (fl), Martius 162 (Lectotiro, designado por Morales (2005), pr Echites e Ruiz var. yu ius Stadelm., Flora 24(1):Beibl. 27. 1841. Tiro: BRASIL. Bania: Datos perdidos (fl), Martius s.n. (HOLOTIPO: M HR INB]) Echites fluminensis A. DC., Prodr. 8:452. 1844. Amblyanthera fluminensis (A. DC.) Múll.Arg., Fl. Bras. 6(1):148. 1860. Mandevilla fluminensis Donn.Sm., Enum. Pl. Guatem. 2:47. 1891. Tiro: BRASIL. Datos perdidos, Fl. Flumin. Icon 3:pl. 44. 1831 (1827) [LECTOTIPO, designado aquí]. Echites fluminensis A. DC. var. claussenii A. DC., Prodr. 8:452. 1844. Amblyanthera claussenii (A. DC.) Miers, Apocyn. S. Amer. 187. 1878. Tipo: BRASIL. Minas Gerais: Caxoeira do Campo, 1840 (fl), Claussen s.n. (HoLoTIPO: G-DC; Isotipo: K) Echites tomentosus Vahl var. laticordatus A. DC., Prodr. 8:463. 1844. Amblyanthera palustris Müll.Arg., Fl. Bras. 6(1):145. 1860. Temnadenia palustris (Mull.Arg.) Miers, Apocyn. S. Amer. 213. 1878. pa palustris (Müll.Arg.) Hemsl., Biol. Cent.-Amer., Bot. 2(10):317. mann s.n. (LEcTOTIPO, designado aquí, G-DC [foto F neg. 26864]; 1881. Tiro: BRASIL. Bania: localidad perdida, 1827-1830 (fl, fr), S ISOLECTOTIPOS, CGE, HAL [2 cartulinas], K, MO, P [3 cartulinas]). Echites lasiocarpus A. DC. Prodr. 8:463. 1844. Temnadenia lasiocarpa (A. DC.) Miers, Apoc. S. Amer. 210. 1878. Mandevilla lasiocarpa (A. DC.) Malme, Bih. Kongl. Svenska Vetensk.-Akad. Handl. 24 (3/10):25. 1899. Tiro: BRASIL. Banta: Cuiabá (citado como Cuyabá), fecha perdida (fl, fr), Manso 29 (HoLotipo: G-DC; isoripo: HAL). hites lasiocarpa A. DC. var lobbiana A. DC., Prodr. 8:464. 1844. Temnadenia lobbiana (A. DC.) Miers, Apocyn. S. Amer. 209. 1878. Tipo: BRASIL. Rio DE JANEIRO: montaña de Orgaos, fecha perdida (fl), Lobb s.n. (notorio: G-DC; isoriro: K) ips hispida (Willd. ex Roem. & Schult.) Müll. Arg. var. tomentosa Müll.Arg., Fl. Bras. Ta pl. 44. f. 3. 1860. Tipo: BRASIL. o Grosso: localidad perdida, 1833 (fl, fr), Gaudichaud 105 (LecrotiPO, designado aquí, P). P ovata Miers, Apocyn. S. Amer. 188. 1878. Tiro: BOLIVIA. Datos perdidos E Miers s.n. (HoLoTIPO: BM [foto, INB]). Temnadenia pallidiflora Miers, Apocyn. S. Amer. 211. 1878. Tiro: BRASIL. Rio DE JANEIRO: entre Magé y Freichal, sin fecha (fl), Miers s.n. (HOLOTIPO: BM). Mandevilla rusbyi Britton, Bull. New York Bot. Gard. 4:409. 1907. Tipo: BOLIVIA. Datos perdidos (fl, fr), Bang 2843 (HoLotipo: NY; ISOTIPOS: F [foto F neg. 51185], K, MO, US [2 cartulinas]). 862 Journal of the Botanical R h Institute of Texas 1(2) Mandevilla denticulata S.E Blake, Contr. Gray Herb. 7. Tiro: BELICE. New Haven, 8 Mar 1907 (fl), Peck 696 (HoLoTiPo: GH [foto, INB]; isoripo: K). os a Lianas; ramitas var iadamente RA a hirsutas o glabras; coléteres a in- conspicuos, ca. 0,4 mm de largo. Hojas opuestas; pecíolos 11-55 mm de largo; láminas foliares 4,2-14,4 x 2,5-7,8 cm, elípticas, ovado-elípticas a obovado- elípticas, el ápice caudado-acuminado a cuspidado, la base cordada a sub- cordada, raramente redondeada, los coléteres irregularmente distribuidos a lo largo del nervio central, membraná- ceas, esparcidamente estrigillosas a glabrescentes o más raramente glabras en la superficie adaxial, esparcida a moderadamente hirsutulas a tomen- tosas en la superficie abaxial, el indu- mento más denso sobre los nervios, más raramente glabras o glabrescentes, no revolutas, las venas secundarias im- presas en ambas caras, las venas terci- arias impresas en la superficie abaxial, evidentes o inconspicuas en la superfi- cie adaxial. Inflorescencia más corta o igualando las hojas subyacentes, axilar, hispidulosas, estrigosas, puberulentas, Fic. 1. Mandevilla hirsuta (A-H de Morales & Lápiz 2695, INB; I-J de Morales et al. 2411, NB)... tomentulosas a glabrescentes o glabras, M it ; A fl bn la B in d fi ial de la hoja, mostrando los con muchas flores, pedúnculo 13-63 col A lo largo de a (se las cn feas € C. Cáliz y pedicelo, con la mm de largo, pedicelos 3-8 mm de EE E OR UV PA BIS largo, brácteas 11-21 x 1,5-8 mm de giboso. F. Antera, vista dorsal. G. Cabeza estigmática. H. Nectario y ovario. I.Foliculos.J. largo, ovadas, foliáceas a subfoliáceas; Semilla. sépalos 7-12(-15) x 1-3 mm, angosta- mente ovados, ovado-elípticos a linear- ovados, esencialmente iguales, acuminados a acuminado- d M apicalmente, el ápice usualmente no a} J 1 reflexo, foliáceos a subfoliáceos, densa a , hirsutulos, tomentosos a glabrescentes o glabros externamente, el coléter solitario, entero, peter ai | roso apicalmente; corola infundibuliforme, el tubo crema a verde-crema externamente, algunas veces con tonos rojizos, los lobulos amarillos, anaranjado-amarillentos, blanco-crema a blancos, la garganta usualmente roja internamente, raramente blanca, variadamente hirsutas, el indumento más denso en botones florales, el tubo basalmente giboso, la parte inferior 25—32(—35) x 2-5,8(-7) mm, la parte superior 15-21(225) mm de largo, cónica a estrechamente cónica, con un diámetro de (11-)16-21 mm en el orificio, el ápice del botón floral acuminado a cortamente apiculado; lóbulos (11—)22-29 x (92)18—25 mm, obovados, extendidos, e irregularmente reflexos marginalmente; estambres insertos en la base de la parte superior del tubo, anteras 4,5—5,2 mm de largo, glabras dorsalmente, la base auriculada, con las aurículas redondeadas a subtruncadas, cabeza estigmática 2-2,2 mm de largo; ovario 1,5-1,9 mm de largo, glabro o glabrescente; nectario anular, irregularmente Morales, N iac dal SEPT RAR NEAR a pentalobulado, ca. la mitad de la longitud total del ovario. Foliculos 10,8-19,3(-21,2) cm x 1,5-4,5 mm, usualmente no fusionados longitudinalmente, usualmente solo unidos en su región apical, más raramente fusionados a lo largo de su entera longitud, variadamente hispídulosos o puberulentos, raramente glabres- centes, moniliformes; semillas 6,5-9(-13) mm de largo, usualmente diminuta y densa a moderadamente puberulentas, más raramente glabrescentes, coma 1,3-2,5 cm de largo, amarillo-canela. Distribución, hábitat y ecología.—N de México hasta Brasil, Paraguay, y Bolivia, así como en Trinidad y en las Antillas menores (St. Thomas), creciendo en bosques, sabanas, vegetación asociada a afloramientos rocosos y formaciones de cerrado, en elevaciones de 0-1800 m. Mandevil delind n partes vegetativas y florales, de manera que, aunque el indument á usualmente presente == a hirsuta es una de las especies más variables en el género, principalmente en el tipo y densidad en prácticamente todas las partes de la planta, M con carencia total de indumento también son ESTE encontrados. Estos especimenes con hojas e infl mol j idénticos a la típica variedad pubescente y no difieren en ningún otro carácter, mos lo que el ET aislado de estos rangos de variación puede llevar a la errónea conclusión de que varias especies pueden ser reconocidas (a nivel espe- cífico o varietal). Sin embargo, el análisis de la pubescencia en todo el rango de distribución geográfica, ha demostrado que no existe una discontinuidad (ni siquiera a nivel geográfico) que justifique tal separación. Además, antes de cualquier intento de segregación de este taxón, es importante tomar en cuenta la alta va- riación en la presencia y densidad del indumento en otras especies del género (e.g., M. pohliana (Stadelm.) A. H Gentry), así como la alta intergradación presente entre especímenes con indumento denso a aquellos que carecen totalmente de este. En términos generales, Mandevilla hirsuta ha sido fácilmente confundida con M. sagittarii Woodson otro taxón morfológicamente muy similar, del que difiere principalmente por la forma del tubo de la corola. Amblyanthera hispida var. tomentosa Müll.Arg. es lectotipificada y la colección Gaudichaud 105 selec- cionada como el lectotipo, ya que este especimen tiene flores y frutos y se encuentra en buen estado de preservación. El otro sintipo citado por Müller (1860) es la colección tipo de Echites lasiocarpus A. DC. (Morales 2005). Tres sintipos fueron citados en la descripción de Echites hirsutus Ruiz & Pav. var. angustifolius: dos cor- responden a colecciones de Martius sin número de los estados de Bahia y Rio de Janeiro y la tercera, a una colección de Pohl del estado de Minas Gerais, Brasil. Acá se selecciona la colección de Martius recolectada en el estado de Rio de Janeiro como el lectotipo, pues es la que se encuentra en mejor estado de preservación. En la lectotipificación de Echites tomentosus Vahl var. laticordatus A. DC. se selecciona la colección de Salzmann s.n. como el lectotipo, ya que este especimen está representado en diferentes herbarios, mientras que el otro sintipo, una colección realizada por Perrottet en Guyana, es conocido por un unico especimen i nd: en el herbario de París (P). Un i (Ackermann s.n.) fue citado en la descripción de Echites stadel i Mart. ex Stadelm., el cual no ud ser localizado en los T herbarios europeos. Por lo tanto, se procede a designar un neotipo que concuerde con los caracteres mencionados en la descripción original. Mandevilla horrida JE Morales, sp. nov. (Fig. 2): Tiro. PERÚ. Cajamarca: San Ignacio, San José de Lourdes, 23 nov 1999 ), R. Vásquez & Flores 26349 (HoLoTIPO: INB; IsoTipo: MO). A Mandevilla hirsuta (Rich) K. Schum., cui affinis, bracteis 32-46 x 10-19,5 mm (vs. 11-20 x 1,5-8 mm), calycis laciniis 4-6 mm longis (vs. 7-12(-15) mm), et tubo proprio 20,5-23 mm longis (vs. 25-32(-35) mm) differt. Lianas; ramitas densa a moderadamente híspidas; coléteres interpeciolares inconspicuos, ca. 0,6 mm de largo. Hojas op ; pecíolos 14-48 mm de largo; láminas foliares (7,3-)8,6-15,9(-17,3) x G23,7-7,1C 7,8) cm, elípticas, obovado-elípticas a obovadas, el ápice caudado-acuminado, la base cordada, los coléteres ir- regularmente distribuidos a lo largo del nervio central, membranáceas, esparcidamente estrigillosas en la superficie adaxial, densa a moderadamente hirsutas en la superficie abaxial, el indumento más denso a lo largo de los nervios, no revolutas, las venas secundarias impresas en ambas caras, las venas terciarias impre- sas en la superficie abaxial, apenas evidentes o inconspicuas en la superficie adaxial. Inflorescencia usual- 864 Journal of the Botanical R h Institute of Texas 1(2) mente más corta que las hojas subyacentes, axilar, densa a moderadamente hirsutas, con muchas flores, pedúnculo 1,7-5,4 cm de largo, pedicelos 1,5—3(-5) mm de largo, brácteas 32-46 x 10-19,5 mm, ovadas a ovado-elípticas, foliáceas; sépalos 4-6 x 1,4-2 mm, angostamente ovados a elíptico- ovados, esencialmente iguales, acuminados apicalmente, el ápice no reflexo, foliáceos, esparcidamente hispídulosos, el coléter solitario, subentero a levemente erosos api- calmente; corola infundibuliforme, el tubo rojizo, los lóbulos blancos, blanco-rosados, amarillos o rojos, densa a esparcidamente pilosa externamente, el indumento más denso en botones florales, el tubo basal- mente giboso, la parte inferior 20,5-23 x 1,9-4 mm, la parte superior 19-26 mm, campanulada a urceolado-campanulada, con un diámetro de 1-1,3 cm en el orificio, el ápice del botón floral agudo a cortamente acuminado; lóbulos 13-16 x 10,5-13 mm, obovados, relativamente extendidos; estam- ies} bres insertos en la base de la parte superior del tubo, anteras 3,9-4,1 mm de largo, glabras dorsalmente, la base auriculada, con las aurículas truncadas o subtrunca- das, cabeza estigmática 1,1-1,3 mm de largo; ovario 1,4-1,7 mm de largo, glabro; nectario anular levemente pentalobulado, Fic. 2. Mandevilla horrida (A-H de Vásquez & Flores 26349, INB; I-J de Gentry etal. ca. la mitad de la longitud total del ovario. 61230, INB). A. Ramita con infl B. Detalle de la superficie adaxial de | Folículos 16,8-23 cm x 1,5-5,2 mm, no hoia J I Aa I I P p t ime Sal J £ L X 3 f : ) 3 ^ A fusionados longitudinalmente, usualmente C. Cáliz y pedicelo, con la bráctea floral ida. D. Vista adaxial de dos sépalos, m : m alee: la base. E. Botón floral parcial berto mostrando solo unidos en su región apical, híspidos o eee Sane eee el tubo basalmente aiboso. F. Antera, vista dorsal. G. hirsutos, moniliformes; semillas 14-15,2 Cabeza estigmática. H. Nectario y ovario. I. Folículos. J. Semilla. mm de largo, glabras, coma 1,4-2,3 cm de largo, amarillo-canela. Distribución, habitat y ecología.—En- démica al departamento de Cajamarca, en el N de Perú, donde crece en formaciones de selva andina, en elevaciones de 1500-2050 m. Especímenes con flores y frutos han sido recolectados entre enero y marzo, junio, julio y noviembre. Mandevilla horrida ha sido confundida con la común M. hirsuta, taxón con una morfológia bastante similar. En forma general, las especies relacionados con M. hirsuta (e.g., M. sagittari, M. steyermarkii) son mor- fológicamente muy similares entre sí y es fácil confundirlos cuando sus caracteres florales no son estudiados en detalle y su identificación se basa principalmente en los caracteres de las hojas y tallos. Sin embargo, M. horrida se puede diferenciar con facilidad de M. hirsuta por sus inflorescencias con las brácteas florales mucho más grandes (32-46 x 10-19,5 mm vs. 11-20 x 1,5-8 mm), sépalos ligeramente más pequeños (4-6 mm vs. 7-12(-15) mm), así como corolas con la parte inferior del tubo de 20,5-23 mm de largo (vs. 25-325) Morales, N iac dal BERT AAA NN TER - mm) y la parte superior campanulada a urceolado-campanulada (vs. cónica a estrechamente cónica). En forma general, estos caracteres han sido analizados cuidadosamente en el material de M. hirsuta disponible ix en día y no se a localizar especimenes que representen existen estados intermedios. E inados: PERÚ. Amazonas: Imaza, Yamayakat, río Marañón 28 jul 1994 (fD, Jaramillo et al. 183 (INB, MO). PUE San Ignacio, Huarango, El Triunfo, 14 jul 1996 (fD, Campos & Díaz 2965 (INB, MO, USM); San José de Lourdes, Camana, 4 mar 1997 (fl), Campos & Corrales 3396 (INB, MO, USM); San Ignacio, La Copa, Vista Florida, 18 jun 1997 (fl, fr), Campos & García 4036 (INB, MO); San Ignacio, Huarango, 11 mar 2000 (fl, fr), Campos et al. 6565 (INB, MO, USM); San Ignacio, San José de Lourdes, campamento Zural, 28 ene 1999 (fD, Díaz et al. 10550 (INB, MO); La Palma, NO de Chirinos, 5 feb 1988 (fD), Gentry et al. 61230 (INB, MO, USM); San Ignacio, San José de Lourdes, 21 ene 1999 (fl, fr), Díaz et al. 10411 (INB, MO, USM). Mandevilla inexperata J.E Morales, sp. nov. (Fig. 3). Tio: ECUADOR. Esmeraldas: San José, jun 1865 (fl), J. Isern 1110 (HoLoTIPO: MA [fotos, INB]). A Mandevilla hirsuta (Rich) K. Schum., cui affinis, tubo proprio 12-14 mm longis (vs. 19-3 t i 9 mm diametro (vs. 12-25 mm) differt. Liana; ramitas diminuta y moderadamente estrigosas, el indumento algo uncinado apicalmente; coléteres interpeciolares inconspicuos, hasta 0,8 mm de largo. Hojas opuestas; pecíolos 12-14 mm de largo; láminas foliares 7-11 x 3,6-6,1 cm, elípticas, el ápice cuspidado-acuminado, la base subcordada, los coléteres irregu- larmente distribuidos a lo largo del nervio central en la superficie adaxial, membranáceas, inconspicuamente puberulentas a lo largo de las venas en ambas caras a glabrescentes, membranáceas, no revolutas, la venación secundaria impresa en ambas caras, las venas terciarias usualmente solo impresas en la superficie abaxial, a veces apenas visibles en la superficie adaxial. Inflorescencia más corta que las hojas subyacentes, axilar, diminuta y moderadamente estrigosa, el indumento algo uncinado apicalmente, con muchas flores, pedún- culo 8-32 mm de largo, pedicelos 2,5-4 mm de largo, brácteas 19-35 x 3-6,5 mm, dne elípticas a ovado-elípticas, foliáceas; sépalos 7,5-9 x 1,7-2 mm, ang d l , el ápice no reflexo, esparcida y diminutamente puberulentos, el coléter solitario, D a ula y levemente lacerado apicalmente; corola infundibuliforme, el color desconocido, glabra o glabrescente externamente, el tubo giboso basalmente, la parte inferior 12-14 x 1,9-3,1 mm, la parte superior 12,5-13,5 mm, angos- tamente cónica, 8-9 mm de diámetro en la boca, el ápice del botón floral cortamente acuminado; lóbulos 10-12 x 9-10 mm, obovados; estambres insertos en la base de la parte superior de la corola, anteras 4-4,3 mm de largo, glabras dorsalmente, la base auriculada, con las aurículas obtusas, cabeza estigmática 1,9-2,3 mm de largo; ovario 1,8-2,2 mm de largo, glabro, nectario anular, levemente a moderada e irregularmente pentalobulado, ca. la mitad de la longitud total del ovario. Folículos desconocidos. Distribución, hábitat y ecología. —Endémica a la provincia de Esmeraldas, en el N de Ecuador, donde se conoce ünicamente de la localidad tipo, en bosques muy hümedos bajo los 300 m de elevación. Flores han sido recolectadas en junio y noviembre. Mandevilla inexperata difiere de M. bracteata (Kunth) Kuntze M. hirsuta y M. steyermarkii Woodson por sus corolas con un diámetro en la boca de 8-9 mm (vs. 12-25 mm) y tubo más pequeño, con la parte inferior de 12-14 mm de largo (vs. 19-36 mm) y la superior de 12,5-13,5 mm de largo (vs. 14-24 mm). Mandevilla krukovii Woodson, M. lancibracteata Woodson y M. moritziana (Múll.Arg.) Donn.Sm. difieren de M. inexperata por sus sépalos mucho más cortos (2-5 mm vs. 7,5-9 mm). Finalmente, en M. sagittarii, otra especie con corolas con una boca angosta, las hojas tienen un indumento persistentemente tomentoso y glauco abaxialmente y corolas más largas en relación a M. inexperata. Mandevil uncinado de sus ramitas e inflorescencias (vs. hirsuto o hirsutulo y erecto o suberecto), láminas foliares incon- c a inexperata difiere de M. megabracteata (descrita acá) por el indumento estrigoso y apicalmente spicuamente puberulentas a glabrescentes abaxialmente (vs. esparcidamente hirsutas), brácteas florales más pequeñas (19-25 mm vs. 44-68 mm) y corolas con la parte inferior más corta (12-14 mm vs. 23-25 mm). Es muy llamativo el hecho que esta especie fue recolectada hace más de 140 años y el espécimen tipo paso inadvertido en las colecciones del Real Jardín Botánico en Madrid, Espafia, las cuales han sido estu- diadas por varios especialistas en la familia. 866 Journal of the Botanical R h Institute of Texas 1(2) ECIITANOD Loja: entre Yangana y Valladolid, 5 nov 2002 (f1), Grant et al. 02-4293 (MO). Mandevilla els piod J.E Mo- rales, sp. nov. (Fig. 4). Tiro: GUY- ANA. Región cud montañas Pakaraima, Mt. Wokomung, cabeceras de Wusupubaru creek, 17 feb 1993 (fl, fr), Henkel et al. 1447 (noLotipo: INB; IsoTIPOS: A Mandevilla steyermarkii Woodson, cui affinis, ramulis hirsutis (vs. glabris vel glabrescentiis), petiolis 8-10 mm longis (vs. (1421-62 mm), calyces laciniis 9-10 mm longis (vs. 2,5-3 mm), et corollae faucibus 10-12 mm diametro (vs. 17-25 mm) differt. Liana; ramitas moderada a esparcidam- ente hirsutas; coléteres interpeciolares inconspicuos, hasta 0,5 mm de largo. Hojas opuestas; pecíolos 8-10 mm de largo; láminas foliares 8,5-12 x 3,2-4,1 cm, elípticas, obovado-elípti- cas a obovadas, el ápice abruptamente corto-acuminado, la base cordada, los coléteres irregularmente distribuidos a lo largo del nervio central en la superfi- cie adaxial, membranáceas, glabrescen- tes en la superficie adaxial, con pelos largos y espaciados en las nervaduras únicamente, esparcidamente hirsutas en la superficie abaxial, el indumento Fic. 3. Mandevilla inexperata (Isern 110, MA). A. Ramitacon flores. B. Detalle de la pul denso a lo largo de los nervios, no rev- = J 1 tall C Niatall J I E J E | olutas, la venación secundaria impresa lol lel i I (senalad fled ).D Cáliz y pedicelo, la brá fl | removida, E Sépalo y coléter, vista adaxial F. Tubo de un botón parcialmente abierto, M fici giboso. G. Antera, vista dorsal. H. Cabeza usualmente solo visibles en la superficie en ambas superficies, las venas terciarias estigmática. L Nectario y ovario. | abaxial. Inflorescencia conspicuamente más larga que las hojas subyacentes, axilar, el pedünculo hirsuto, el resto moderadamente hirsutula, con muchas flores, pedúnculo 18-23 mm de largo, pedicelos 3-4 mm de largo, brácteas 44-68 x 7-11 mm, angostamente elípticas, conspicuamente foliáceas, verdes; sépalos 9-10 x 0,9—1,1 mm, angostamente ovados, largamente acuminados apicalmente, el ápice a veces ligeramente reflexo, subfo- liáceos, esparcidamente hirsutulos cerca de la base externamente, en el resto el indumento muy esparcido, el coléter solitario, irregularmente lacerado apical te; corola infundibuliforme, el tubo verde externamente, los lóbulos blancos a crema, densa o moderadamente hirsuta o hirsutula en botón, tornándose inconspicua y muy esparcidamente hirsútula a glabrescente externamente en la antésis, el tubo giboso basalmente, la parte inferior 23-25 x 2-3,5 mm, la parte superior 14-16 mm de largo, angostamente campanulada, con un diámetro de 10-12 mm en la boca, el ápice del botón floral cortamente acuminado; lóbulos 11-15 x 9-13 mm, obovados, apenas extendidos; estambres insertos en la base de la parte superior del tubo, anteras 3,9-4,1 mm de largo, glabras dorsalmente, la base subtruncada y subauriculada, con las aurículas apenas visibles, truncadas a subtruncadas, cabeza estigmática 2-2,2 mm de largo; ovario 1,8-2,4 mm de largo, Morales, N iac dal PREC AAN NN TER - glabro; nectario anular, leve e ir- regularmente pentalobulado, 1-1,2 mm de largo. Folículos 11,6-16,1 cm x 1,2-4,1 mm, esparcidamente hirsutulos o puberulentos, monili- VW formes; semillas inmaduras 10-11 W mm de largo, diminuta y densamente EB wo 1 puberulentas, la coma 1,6-2,1 cm de largo, canela. Distribución, hábitat y ecología.— Restringida al N de Colombia (Depar- tamento de Cesar y el E de Guyana, donde se encuentra en bosques muy hümedos y vegetación secundaria relacionada, en elevaciones de 800— 1150 m. Material con flores ha sido recolectado entre febrero y marzo. El único espécimen con frutos fue recolectado en febrero. Mandevilla megabracteata es morfológicamente similar a M. stey- ermarkii, ya que ambas comparten inflorescencias con brácteas florales foliáceas y conspicuas, que ocultan los botones florales. Sin embargo, M. megabracteata puede ser fácilmente distinguida por sus ramitas moderada aesparcidamente hirsutas (vs. glabras o glabrescentes), pecíolos 8-10 mm de largo (vs. (14)21—62 mm de largo), láminas glabrescentes en la superficie adaxial y esparcidamente hirsutas ab- Fic. 4. Mandevilla megabracteata (Henkel et al. 1447, INB). A. Ramita con flores y frutos. B. ~ OD lladal Erin ab D. Cáliz y pedicelo, con la bráctea floral removida. E. Sépalo y coléter, vista adaxial. F. Tubo d botón floral trando el tubo gib G. Antera, vista dorsal. H. Cab tiomáti axialmente (vs. glabras), sépalos 9-10 mm de largo, angostamente ovados y I. Nectario y ovario. J. Semilla. largamente acuminados apicalmente (vs 2,5—3 mm, ovados y agudos apicalmente), corola con los lóbulos blancos a crema y con un diámetro de 10-12 mm en la boca (vs. lóbulos amarillos, anaranjado-amarillos o anaranjados y con un diámetro de 17-25 mm), así como anteras de 3,9-4,1 mm de largo (vs. 7,5-8,5 mm) y folículos esparcidamente hirsutulos o puberulentos (vs. glabros). Especímenes adicionales examinados: COLOMBIA. Cesar: La Jagua de Ibirico, vereda Alto de las Flores, 9 mar 1996 (fl), Fernández- Alonso et al. 13186 (COL). Mandevilla similaris J.E Morales, sp. nov. (Fig. 5). Tiro: VENEZUELA. BoLivar: Cumbre del Cerro Guaiquinima, 25 may 1978 (£D, Steyermarh et al. 117347 (HoLotipo: INB; isoripo: VEN). A Mandevilla lancibracteata Woodson, cui affinis, calycis laciniis 8-10 mm longis (3-3,8 mm) et corollae faucibus 18-23 mm longis (vs. 7-9 mm) differt. Liana; ramitas esparcidamente puberulentas cuando jóvenes, glabrescentes con la edad; coléteres interpe- ciolares inconspicuos, hasta 0,5 mm de largo. Hojas opuestas; pecíolos 5-8 mm de largo; láminas foliares 4,5-10 x 2-4,8 cm, elípticas, obovado-elípticas a obovadas, el ápice abruptamente corto-acuminado, la 868 Journal of the Botanical R h Institute of Texas 1(2) base cordada o subcordada, los coléteres irregulamente distribuidos a lo largo del nervio central en la superficie adaxial, mem- branáceas, esparcidamente puberulenta a glabrescentes en la superficie adaxial, densa a moderadamente pilosulosas a glabres- centes abaxialmente, no revolutas, venas secundarias impresas en ambas caras, las venas terciarias usualmente no evidentes. 3 Rer más laroa aue lac hoias eanbva- eo b J centes, axilar, esparcidamente puberulenta a glabrescente, con muchas flores, pedún- culo 40—45 mm de largo, pedicelos 2,5 mm de largo, brácteas 25-48 x 4-9 mm D de largo, angostamente elípticas, elípticas a ovado-elípticas, foliáceas; sépalos 8-10 x 0,9-1,4 mm, angostamente ovados, acu- minados apicalmente, el ápice no reflexo, escariosos, i r r a glabrescentes externamente, el coléter solitario, entero a subentero apicalmente; corola infundibuliforme, el tubo verde-pá- lido, los lóbulos amarillos o crema, esparcida a moderadamente hirsutulas, sobretodo en botones florales, el tubo basalmente giboso, la parte inferior del tubo de la corola 18-23 x 2-3 mm, la parte superior 14-17 mm de largo, angostamente campanulada, con un diámetro de 10-13 mm en el orificio, el Fic. 5. Mandevilla similaris (Steyermark et al. 104449, VEN). A. Ramita con flores. B. Spe del botón floral agudo; lóbulos 14-16 x 10-12 mm, obovados, extendidos; estam- b J ). C. Cáliz y pedicelo, con la bráctea floral ida.D. bres insertos en la base de la parte superior Lleawalad flanel Sépalo y coléter, vista adaxial. E. Tubo de un botón trand giboso. F. Antera, vista dorsal. G. Cabeza estigmática. H. Nectario y ovario. del tubo, anteras 3,5-4 mm de largo, dor- salmente glabras, la base inconspicuamente J 1 J auriculada, con las aurículas a subtruncadas, cabeza estigmática 1,5-1,7 mm de largo; ovario 1,5—2 mm de largo, glabro a inconspicuamente hirsutulo en su región basal sobre el nectario; nectario anular, irregularmente pentalobulado, 2/3 a la mitad de la longitud total del ovario. Folículos desconocidos. Distribución, hábitat y ecologta.—Endémica al Estado de Bolivar en Venezuela, donde crece formaciones de Aq sabana y vegetación arbustiva asociada a rocosos en elevaciones de 900-1600 m. Especimenes con flores han sido recolectados en enero, mayo, agosto y diciembre Material con frutos fue recolectado en septiembre. Mandevilla similaris conforma un complejo junto con M. lancibracteata, ya que ambos taxones son morfológicamente muy similares y fácilmente pueden ser confundidos entre sí, sobretodo por compartir inflorescencias con las brácteas florales foliáceas y corolas con la parte superior del tubo angosta, con una diámetro alrededor de 1 cm. Sin embargo, M. similaris se puede distinguir de M. lancibracteata por sus láminas foliares con los coléteres restringidos al nervio central (vs. distribuidos también en las venas secundarias), Morales, N iac dal BERE ARAN TN TER - peer 1 d Y con las venas terciarias no evidentes (vs. usua Imente sépalos de 8-10 mm de largo (vs. 3-3,8 mm) y corolas con la parte inferior de 18-23 mm de longitud (vs. 7-9 mm de largo). Especímenes adicionales pacas VENEZUELA. Bolivar: faldas del Auyan-tepuí, ene 1949 (fD, Cardona 2754 (VEN); Final de La Escalera, 8 ago 1979 (fl), Morillo € 38 (VEN); La Escalera, al S de El Dorado, 13 feb 1980 (st), Morillo 8142 (VEN); parque nacional Canaima, La Escalera, km 88 entre odo 8 sep 1987 (fr), Ramírez et al. 167 (VEN); drenaje del río Cuyuní, al S de El Dorado, 22-28 dic 1970 (fD, Steyermark et al. 104489 (\ ; drenaje del río Cuyuní, a lo largo del río Anawaray-parú, al S de El Dorado, 25 dic 1970 (fD, Steyermark et al. 84937 (VEN); Gran Sabana, entre El Dorado y p Elena, 30 Dic 1974 (fD, Steyermark 111289 (VEN). AGRADECIMIENTOS Quiero agradecer a los herbarios BM, B-W, C, CGE, COL, ESAL, F, G, G-DC, GH, HAL, K, M, MO, P, P-HB, P-LA, UB, US, USM, VEN, W, Z por el envío de material en préstamo, así como por facilitar el acceso a sus colecciones. Asimismo, agradezco la colaboración de Eliana Noguera, Shingo Nozawa y Sirli Leython que permitió revisar material en el Herbario Nacional de Venezuela (VEN). Finalmente, Barry Hammel (MO) realizó la traducción del resumen al inglés. REFERENCIAS ALLORGE-BorTEAU, L. 1998. New combinations in Odontadenia and Mandevilla (Apocynaceae). Phytologia 84: Morales, J.F. 2005. Estudios en las Apocynaceae Neotropicales Xl: una nueva especie de Mandevilla (Apocynoi- deae, Mesechiteae) para Sur América, con un nuevo reporte para las Apocynaceae de Paraguay. Sida 21:1549-1556 Morales, J.F. 2006. Estudios en las Apocynaceae Neotropicales XXV: novedades y nuevos reportes en las Apocy- naceae (Apocynoideae, Rauvolfioideae) de Venezuela. Sida 22:355-365 MULLER ARGOVIENSIS, J. 1860. Apocynaceae. En: C.EP. von Martius, ed. Flora Brasiliensis 6(1):1-180. Simoes, A.O., M. ENDRESS, T. VAN DER Niet, L.S. KinosHita, & E. Conti. 2004. Tribal and intergeneric relationships of Mes- echiteae (Apocynoideae, Apocynaceae): evidence from three noncoding pastid DNA regions and morphol- ogy. Amer. J. Bot. 1:1409-1418 Simoes, A.O., M. ENDRESS, T. VAN DER Niet, L.S. Kinoshita, & E. Conti. 2006. Is Mandevilla (Apocynaceae, Mesechiteae) monophyletic? Evidence from five plastid DNA loci and morphology. Ann. Missouri Bot. Gard. 93:565-591. Simoes, A.O., L.S. KinosHITA, & M. ENDRESS. 2007. New combinations in Mandevilla Lindley (Apocynaceae). Novon 17:89-90. Woobson, R.E. 1933. Studies in the Apocynaceae. IV. The American genera of Echitoideae XXVI. Ann. Missouri Bot. Gard. 20:605-790 — WS Journal of the Botanical R h Institute of Texas 1(2) 870 BOOK REVIEW WiLLOw ZucHowskt. 2007. Tropical Plants of Costa Rica: A Guide to Native and Exotic Flora. (ISBN 978- 0-8014-7374-6, pbk.). Cornell University Press, Sage House, 512 East State Street, Ithaca, NY, 14850, U.S.A. (Orders: www.cornellpress.cornell.edu). $35.00, 532 pp., 540 color photographs, 2 maps, 56 line la IE: PHAR 535-49 ts of Costa Rica i 11 f ling osta Rica ph lOLOBI aph Th; or with a general interest in Wd) ans os descriptive species accounts, wonderful color and conspicuous plants M in one Rica, the author certainly fulfills d. i :1 11 Turid Forsyth, and illustrations for 4 s both useful and dd small e to carry and utilize in the fiel Perfect for the lay e or first-time visitor to Costa Rica, Spe- her goal to create a guide th O p E la 1 Tals gnig 8 cies accounts in the ene are organized mus the g ] ings within which a traveler the plants. Each chapter of the l setting id d by the Mn titles: Painted Treetops, Other Common Trees, Roadside mi Garden OUS Fruits Er d ing F d Reforestation, Special Habitats, Typical pee Groups, and Conspicuous Grasses. Individ le the followi bo b pan ish and English), other a name to which the species bs physical description, fl ing/fruiting distribution, identification of related species, and comments. Th I f the speci lay f. ginf he natural history ial, medicinal, and ae d unique cl teristics of each sp Sidel i a f1 1 A 1 1 pA such titles as "Sex Life of Bs 2 ie Wasps, 5 “The ae xn in Costa Rica,” o does Chan come from?” and “Drift Seeds” to a few A 1 a Ge A S e aut t g both scientifi d common nam Melinda McCoy, Herbarium name ju a list m. by family, a complete bla and an index with Research Institute of Texas, Fort Worth, Texas 76102-4060, U.S.A. Volunteer, Botanical BOOK NOTICES Two Books on Florida Plants RicHARD P. WUNDERLIN and Bruce E Hansen. 2003. Guide to the Vascular Plants of Florida (ed. 2). (ISBN 0-81302632-6; hbk.). Univ. Press of Florida, Gainesville. University Press of Florida, 15 Northwest 15th Street, Gainesville, FL 32611-2079, U.S.A. (Orders: www.upf.com, 352-392-1351, 352-392-7302 fax, 1-800-226-3822). $39.95, 787 pp., 6" x 9". We've all been using the book for several years, but this review gee perhap rl the more than 4100 taxa of ferns, fern allies companion to the multivolume Flora of Florida ceda & Hansen 2000 and in progress).” ls botanists that it’s still for sale. It is “a means to lized in Florida. It serves as the field guide C. Rrrcuie Beit. and Bryan J. Tavior. 1982. Florida Wild Flowers and Roadside Plants (3" printing, 1998). (ISBN 0-9608688-3-6, pbk.). Laurel Hill Press, 5000 Walnut Cove Rd., Chapel Hill, NC 27516-8168, U.S.A. (Orders: www.laurelhillpress.com, 919-942-9533, 919-942-9533 fax). $19.95, numerous color photos, map, D x B 500 species are illustrated by a color photo. For each, there is a brief di ic d iption, comments on habitat and distribution, and a dot map (with the state divided into quarters). Useful book and a great stam J. Bot. Res. Inst. Texas 1(2): 870. 2007 BOERHAVIA TRIQUETRA VAR. INTERMEDIA (NYCTAGINACEAE); A NEW COMBINATION AND VARIETAL STATUS FOR THE WIDESPREAD SOUTHWESTERN NORTH AMERICAN B. INTERMEDIA Richard Spellenberg rd of Biology, MSC 3AF co State University Las To wM 88003-8001, U.S.A. AB OT edu ABSTRACT The name for the poorly known, narrowly distributed, Boerhavia triquetra S. Wats. predates the name B. intermedia M.E. Jones, a well- T i 1 = £41 Sod A 1 E 1 A £1 1 i $ xj 1 11 J á E $ type locality of B. triquetra at Bahía de los Angeles in Baja California, shows the two taxa to thoroughly intergrade. The nomenclatural combination Boerhavia triquetra S. Wats. var. intermedia (M.E. Jones) Spellenb. is proposed. A key distinguishing this variety and the var. triquetra is presented. RESUMEN El nombre de la poca conocida Boerhava triquetra S. Wats., de distribución limitada, precede el nombre B. intermedia M.E. Jones, una especie ios conocida y de o extensa en el suroeste árido de Norte América. Una revisión de muestras de Boerhavia en her- barios y | g n localidad del ue en cae b los ao Baja California, demuestran qu los do S. Wats. var. ee (M. E. Jones) Pi Se p t ] disti variedad y la var. triquetra. L o In 1902 M.E. Jones described an annual Boerhavia from El Paso, Texas, U.S.A., as Boerhavia intermedia M.E. Jones. Prior to that time this taxon was included in a broadly conceived B. erecta L., the name under which Jones first distributed his collections of B. intermedia. For example, Gray (1853) concluded his review of Wright’s collections of Boerhavia by noting that other species in his collection, apparently with annual roots, had been distributed as B. erecta L. The outward similarity of this desert annual, B. intermedia, to the more mesic and tropical B. erecta, is often noted (e.g., Kearney and Peebles 1960) and has resulted in a combination of the former as a variety in the latter (B. erecta var. intermedia (M.E. Jones) Kearney & Peebles). Molecular " evidence suggests, however, that B. erecta and B. inter not particularly closely related (Norm Douglas, pers. comm. 2007). Boerhavia intermedia as understood by Standley (1918) and Spellenberg (1993, 2003), among other authors of regional floras, is widespread and variable, occurring throughout the warm, arid regions of northwestern Mexico and southwestern United States. The only attempt to taxonomically address the variation through taxonomic division of this taxon was B. universitatis Standl., from Tucson, Arizona (Standley 1909), since shortly after its inception a name consistently considered a synonym of B. intermedia. Though not explicitly stated by Jones (1902) or Standley (1909), B. intermedia has been understood to have truncate fruits with five angles separated by rugose sulci. In 1889 S. Watson described B. triquetra S. Wats. from a collection of plants made by E. Palmer, the type originating from near Bahía de los Angeles, Baja California, Mexico. It was distinguished from other Boerhavia by its 3-4-angled truncate fruits with rugose sulci. Since that time the distinction between B. intermedia and B. triquetra in regional floras has not been clear. Both are distinguished from other Boerhavia, and allied to one another, by their annual habit and narrowly obdeltate (in profile) ribbed (not winged) fruits with pedicels that join in a terminal umbel. Those with umbels reduced to solitary fruits still differ from other Boerhavia by the habit and fruit shape, though the distinction between B. intermedia and B. erecta is then subtle. Wiggins (1964) separated the two species in his key primarily by flowers solitary or in umbellate or J. Bot. Res. Inst. Texas 1(2): 871 — 874. 2007 872 Journal of the Botanical R h Institute of Texas 1(2) close cymules. He noted the fruit to be 3-5 angled in B. triquetra, 5 angled in B. intermedia, the fruit of the former slightly broader. Annual Boerhavia of the very dry Mojave and western Sonoran deserts of south- eastern California and adjacent Sonora often have one fruit per peduncle and have been called B. triquetra on specimens. Spellenberg (1993) attempted to distinguish the two taxa by numbers of fruit per cluster, and a subjective aspect of inflorescence structure, making no reference to rib number. In 2003 Spellenberg maintained B. intermedia and B. triquetra, noting in discussion that the distinction between the two seemed moot. The hesitancy to combine the two names in 2003 into one taxon was reinforced by Spellenberg not having seen B. triquetra (which has priority over B. intermedia) in the field. In addition, the application of B. triquetra, a name for a poorly known taxon from a very limited region, was not descriptive for the widespread well-known species with usually 5-sided fruits. Nevertheless, in preparing the treatment of the Nyctaginaceae for the Jepson Manual, 2*d edition, Andrew Murdock (at UC Berkeley) came to the conclusion that the two names apply to one taxon in California (pers. comm. 2006). In October, 2006, I spent several days in the vicinity of Bahía de los Angeles. Late summer rains had been good, and small collections of Boerhavia were made, as were observations. Those collections have been deposited at HCIB and NMC. Including these specimens and those deposited in herbaria, 71 collections from throughout the range of both taxa were examined for numbers of fruit per terminal cluster, fruit size, rib number per fruit, and pedicel length. The following observations were made Pedicels vary from 0.2 to 3.2 mm throughout the range, the shortest occurring within the confines of the Sonoran Desert, but longer (+ 1.7 mm) common within the region also. Watson (1889) noted the pedicels of B. triquetra to be very short (ca. Y line [= 1/24” = ca. 1 mm]). My own collections from around the Bahía de los Angeles had pedicels 0.271 mm long; to the north and to the south along the eastern side of the peninsula in this region pedicels ranged to 1.3-1.9 mm long. Elsewhere they were as short as 0.3 mm. On plants with sufficient fruiting peduncles the number of fruits in 10 umbels was counted. Numbers of fruits per umbel ranged from 1-22. Those with single fruits per umbel occurred throughout the range, but were particularly common in the deserts of southern California. Those with the highest number were from the coast of Sonora. Number of fruits per umbel commonly was 1—5 in the vicinity of Bahía de los Angeles; one plant which had consistently 1 fruit/umbel had only 5-sided fruits (Spellenberg 13790, below). No particular difference between fruit number in this region versus other areas was obvious, except that in Chihuahua, Coahuila and Durango fruit number commonly was 3-7 per umbel, with 1-fruited umbels less frequent. Plants with primarily, or entirely, 1-fruited umbels from throughout the Southwestern warm deserts are represented by the following collections (here and throughout all specimens at NMC unless otherwise cited (herbarium citations from Holmgren and Holmgren, http://sciweb.nybg.org/science2/IndexHerbari- orum.asp): AZ, Pinal Co., San Tan Mts. Regional Park, Damrel and Damrel 24716; BC, Mcpio. Ensenada, on road to Bahia de los Angeles 37 km E of Hwy 1, Spellenberg 13790; Roos s.n., CA, San Bernardino Co., Big Morongo Reserve, in Big Morongo Canyon, Helmkamp B-19, UTEP; Riverside Co., Chuckawalla Mts. 2.5 mi SE of Desert Center, 23 Oct 1963, UTEP; NM, Dofia Ana Co., ca 10 mi E of Las Cruces, W base of Organ Mts., Spellenberg 2921. All these had only five-sided fruits. Three- and four-sided fruits were found on Sanders 051 and Van Devender & Reina G. 2000-939 (citations end of next paragraph), both of which had few fruits per umber (1-2 fruits, av. 1.6; 1-4 fruits, av. 2.5, respectively). Of note is that the very earliest inflorescences on many plants have 1 or few fruits, as often do very late inflorescences. The preponderance of few-fruited umbellate plants on the Mojave and western Sonoran deserts may simply be selection for this early-development feature in an area of unpredictable and reduced summer moisture. Fruit size did not seem to differ much from one part of the range to the other. Within the Bahia de los Angeles region fruits were 2.0-2.8 mm long; slightly to the south fruits reached 3 mm long. Elsewhere fruits were 2.03.1 mm long. The range of fruit width is less in the vicinity of Bahía de los Angeles (the length/ width ratio equalling 1.5-2.0) than for plants from throughout the range of B. intermedia (l/w = 1.3-2.7). The proportionately broadest fruits are from a plant that has only 3- and 4-sided fruits from Sonora (SON, Mcpio. Hermosillo, Playa Esthela just N of Bahia de Kino, Van Devender & Reina G. 2000-939). A similar D intermedia 873 Spellenberg, ] plant with only 4-sided fruits (and 1 fruit with a poorly developed 4" side, i.e., almost 3-sided) had fruits with l/w = 1.9 (CA, Imperial Co., Julian Wash, E of Peter Kane Mt., Sanders 051). Numbers of sides on the fruits varies throughout the range of Boerhavia intermedia. An attempt was made to count all scorable fruits on a plant unless there were hundreds available, and then counting was haphazard, i.e. without intentional bias. Outside the Sonoran Desert plants bear mostly 5-sided fruits, but there are oc- casional fruits with 4 sides. In the following the numbers of fruits counted on any one plant are given in the sequence 5-, 4-, and 3-sided as, for example, CHIH, 1 mi S of Camargo, (53, 10, 0) Henrickson 7735; CHIH, 7 mi S of Ojinaga, (74, 1, 0), Spellenberg & Spellenberg 3699; COAH, 4 km S of Las Margaritas, (38, 2, 0), Chiang et al. 9519c; 14 mi NW of San Pedro, (56, 3, 0), Henrickson 5975; NM, Doña Ana Co., 15 mi N of Las Cruces, (92, 7, 0), Spellenberg 2649; TX, Presidio Co., 24.9 mi E of Redford, (31, 1, 0), Spellenberg 3710. In the Sonoran Desert, excluding central Baja California (reviewed below), plants have 5-sided fruits, or mostly 5-sided fruits with some 3- or 4-sided fruits. Examples of those with some 3- and 4-sided fruits are: AZ, La Paz Co., (49, 1, 1), Spellenberg 7864; Pinal Co., (152, 3, 0), Spellenberg 13271; SON, Mcpio. Em- palme, 1.9 km NW of MEX 15 on Empalme-Guaymas bypass, (79, 8, 0), Reina G. et al. 2002-1055; 25 km S of Empalme (98, 14, 1 [1 fruit with 9 ribs]), Spellenberg & Douglas 13317. Mcpio. Guaymas, 1 mi E of San Carlos Bay, (45, 2, 0) Spellenberg et al. 2688; 4 mi. W of San Carlos, (35, 2, 0) Spellenberg et al 2693; 4 mi W of San Carlos, (30, 2, 1), Spellenberg et al. 2694; Guaymas, Colonia La Peninsula, (71, 1, 0), Spellenberg & Willson 3632; 4.6 km W of San Carlos, (45, 4, 0), Spellenberg 13811. Mcpio. Hermosillo, ca.10 mi S of Hermosillo, (90, 6, 0), Spellenberg et al. 2683. Mcpio. Moctezuma, 18.9 km SSE of jct with Moctezuma-Huásabas Hwy, (43, 1, 0), Van Devender & Reina G. 2006-801. Plants with 3- and 4-sided fruits, usually among 5-sided fruits on the same plant, are most common at low elevations on the Sonoran Desert around the northern portion of the Gulf of California, and plants with the greatest proportion of 3- and 4-sided fruits are found here also. To the north and south of Bahía de los Angeles, on the eastern side of Baja California, numbers of sides per fruit also varies widely: Mcpio. Ensenada, ca 16 km S of Puertocitos, (22, 0, 0) Spellenberg 13783; ca. 30.5 km S of Puertocitos, (38, 0, 0) Spellenberg 13784; 13.5 km N of Punto Bufeo, (92, 4, 1) Spellenberg 13785. Mcpio. Mulegé, 8 km W of Santa Rosalia, (29, 0, 0) Spellenberg 13803; ca. 1.5 km S of Santa Rosalia, (7, 14, 6), Spellenberg 13805; ca. 1.5 km S of Santa Rosalia, (41, 0, 0) Spellenberg 13806; 4.6 km N of Mulegé, (plant A = 158, 0, 0; plant B = 56, 2, 0) Spellenberg 13807. It is noteworthy that 13805 and 13806 were within 100 m of each other. Other collections from the general region: Mcpio. La Paz, Isla San José, León de la Luz 9409a, (25, 0, 0); Sierra la Giganta, Moran 18860 (60, 2, 0). Mcpio. Mulegé, Sierra la Giganta, Carter 4859, (47, 2, 0). These results are compared with sampling from in the immediate Bahía de los Angeles (BLA) region on the eastern side of Baja California: BC, Mcpio. Ensenada, road to BLA 37 km E of Hwy 1, (57, 0, 0) Spellenberg 13790; 4.3 km W of BLA, (82, 51, 6) Spellenberg 13791; BLA, (30, 4, 0) Spellenberg 13792; 4.5 km S of BLA (33, 95, 39) Spellenberg 13793; 1.6 km N of BLA, (70, 16, 0) Spellenberg 13795; 6.4 km W of BLA (plant A = 4, 15, 21; plant B = 17, 8, 3) Spellenberg 13799. This last collection was taken from a crowded population of plants in an arroyo bottom that did not differ noticeably among one another with regard to habit or fruit number per umbel, but these were selected from among those that bore mostly 5-sided fruits. Counts from the type specimens (Palmer 521, Lower California, Los Angeles Bay in 1887) are: (1, 18, 21) US 933319; (8, 60, 89 — from a count of fruits in packet made in 2001) US 933313; (1, 24, 23) US 22954; (0, 4, 6) UC 101261. Though I was able to find many plants that had a habit very similar to the types of B. triquetra around Bahía de los Angeles, none that I found had proportions of 3- and 4-sided fruits as high as in the types. A broad survey of plants known as B. intermedia and B. triquetra revealed no differences that unequivo- cally distinguish the two. Fruit size, rib number and rugosity are similar, and differences are variable and overlapping. Stems may be more widely spreading in some populations from Baja California, but erect plants occur there also. Plants may be more strongly ascending to the east, but widely spreading plants also occur. It is proposed that the two names be considered to apply to a single species with two highly intergradient varieties, requiring a new combination: £L Dat PRA D hi PEA 874 Journal of f Texas 1(2) Boerhavia triquetra S. Wats. var. intermedia (M.F. Jones) Spellenb., comb. nov. Boerhavia intermedia M.E. Jones, Contr. W. Bot. 10:41. 1902. Boerhavia erecta L. var. intermedia (M.E. Jones) Kearney & Peebles, J. Wash. Acad. 29:475. 1939. Tyre: TEXAS: El Paso, 10 Sep 1883, M.E. Jones 4173 (Lectotype, designated by Reed, Flora of Texas 211. 1969; US 223742! IsoLEcToTYPES: POM!). Reed od this name in his treatment of Nyctaginaceae for Texas when he specifically cited the US specimen as the type. Boerhavia universitatis Standl. Contr. U.S. Natl. Herb. 12:380. 1909. Type: ARIZONA: Tuscon, Campus Univ. of Arizona, alt. 740 m, 2 Sep 903, Thornber s.n. (HOLOTYPE: ARIZ! isotypes: MO! USD. The two varieties may be distinguished as follows. The var. triquetra is restricted to low elevations (0-300 m) in the Sonoran Desert around the Gulf of California. The var. intermedia is widespread in Mojave, Sonoran, and Chihuahuan deserts, including low elevations around the Gulf of California. Though the key below seems to allow for a potentially large number of intergradient plants, no plants in the sample that I observed fall within the range of bearing 50-80% 5-sided fruits. 1. 50% or more of the fruits on a plant 4- or 3-sided B. triquetra var. triquetra 2. 80% or more of the fruits on a plant 5-sided B. triquetra var. intermedia Four other named Boerhavia are either part of, or very closely related to, this complex. Two are narrow endemics with the ribs expanded into broad wings, B. alata S. Wats., from the islands around Guaymas, Sonora, and B. megaptera Standley, from south-central Arizona. Two others are poorly distinguished from B. triquetra var. intermedia. One, B. maculata Standley, from the coast of southern Sonora and adjacent Sinaloa, has larger and proportionately broader 5-ribbed fruits and comparatively large flowers in 1-2-flowered umbels. Except for its larger flowers, a feature common among these annual Boerhavia that grow along the Sonoran coast, it resembles other Boerhavia specimens with few-flowered umbels taken at low-elevation from around the Gulf of California. The other, B. lateriflora Standley from the vicinity of Guaymas, Sonora, has large inflorescences of dense head-like cymes or compound umbels. A suitable disposition of these names may be revealed by critical study of molecular and population variation. ACKNOWLEDGMENTS I am grateful to Rob Soreng for counting ribs on fruits on the three types at US. Rob noted (pers. comm.), “Could be a few more 5s but I did not want to pry the 4rs up to check, but most 4s were clearly 4s.” Andy Murdock (UC) provided helpful comments during final preparation of the manuscript. Richard Felger and an anonymous reviewer graciously reviewed the manuscript. Discussions with Norm Douglas (DUKE) regarding annual species of Boerhavia in the Sonoran Desert have been very helpful. REFERENCES Gray, A. 1853. New genera and species of Nyctaginaceae, principally collected in Texas and New Mexico. Amer. J. Sci. Arts, ser. 2, 15:259-263, 319-324. Jones, M.E. 1902. Nyctaginaceae. Contr. W. Bot. 10:34-54. Kearney, T.H. and RH. Peestes. 1964. Arizona flora, 2"4 ed., with supplement by J.T. Howell, E. McClintock and col- laborators. Univ. Calif. Press, Berkeley. Reep, C.F. 1969. Nyctaginaceae. Flora of Texas 2:151—220. Texas Research Foundation, Renner. SPELLENBERG, R. 1993. Nyctaginaceae. In: The Jepson manual: higher plants of California, J.C. Hickman, ed. Univ. Calif. Press, Berkeley. Pp. 768-774. SPELLENBERG, R. 2003. Boerhavia. In: Flora of North America north of Mexico, vol. 4, Flora of North America Editorial Committee, eds., Oxford Univ. Press, New York. Pp. 17-28. STANDLEY, P.C. 1909. The Allioniaceae of the United States with notes on Mexican species. Contr. U.S. Natl. Herb. 12(8):303-389. STANDLEY, PC. 1918. Allionaceae. N. Amer. Fl. 21(3):171-254. WATSON, S. 1889. VI. Contributions to American botany. 1. Upon a collection of plants made by Dr. E. Palmer, in 1889, about Guaymas, Mexico, Muleja and Los Angeles Bay in Lower California, and on the island of San Pedro Martin in the Gulf of California. Proc. Amer. Acad. Arts Sci. 24:36-87. INERASPECIPIC TAXONOMY AND NOMENCLATURE OF ELEOCHARIS ACUTANGULA (CYPERACEAE) David J. en and Stephan L. Hatch Richard Carter Tracy Herbarium Herbarium, D f Biolo Department E ET Science & Management Valdosta State University Texas A&M Universit) Valdosta, Georgia 31698, U.S.A. 4^ College Station, Texas 77843-2126, U.S.A. ABSTRACT A taxonomic study of Eleocharis acutangula (Roxb.) Schult. was conducted in order to better define this poorly understood and variable po eee Multivariate statistical NE and ecological and ord data of worldwide collections of E. acutangula E acutangula, E tang bsp. breviseta D.J. Rosen, subsp. nov., and E ] bsg eacop] Rosen seep nov. N ] I i ] tl I f E. acutangula and a Bei synonym, E. fistulosa Schult. var. robusta Boeck. A i fE gula is provided that includes a key to the subspecies, detailed P illustrations, and notes on habitat and distribution RESUMEN Se realizó un estudio taxonómico de Eleocharis acutangula n Schult. para definir mejor esta especie pantropical variable y pobre- mente conocida. Un análisi adísti Mp yd lógicos y de distribución a nivel mundial de E. acutangula fueron la iónen E bsp. acutangula, E. aene dnd pins ae Rosen, subsp. nov., y E. deutingum saben necito D] Rosen SIS nov. Lai 5 totipifi E. ¢ ng de un sinónimo heterotípico, E. fistulosa Schult. var. robusta Boeck. Se aporta un tratamiento taxonómico ide jn M que ed una clave de subespecies, descripciones detalladas, ilustraciones, y notas sobre el hábitat y distribución. Eleocharis R. Br. is a cosmopolitan genus of about 200 species and over 600 published names with a center of diversity in the Neotropics (González-Elizondo & Tena-Flores 2000). Eleocharis subg. Limnochloa (P. Beauv. ex Lestib.) Torr. (= Eleocharis ser. Mutatae Svenson) comprises over 35 species occurring in season- ally wet to permanently flooded habitats from principally tropical regions, and is distinguished from other Eleocharis by a combination of the following morphological characteristics: (1) cartilaginous, un-keeled (rarely obscurely-keeled), many-veined floral scales; (2) generally large culms that are often as thick as the cylindrical spikelet; and (3) biconvex (rarely trigonous) achenes usually with epidermis of large, conspicuous polygonal cells (Svenson 1929; González-Elizondo & Peterson 1997). Five new species in subg. Limnochloa have recently been described from the New World: E. eglerioides S. González & Reznicek and E. liesneri S. González & Reznicek from Venezuela (S. González-Elizondo & Reznicek 1996), E. yecorensis Roalson from Mexico (Roalson 1999), E. laeviglumis R. Trevis. & Boldrini from Brazil (Trevisan & Boldrini 2006), and E. steinbachii D.J. Rosen from Bolivia (Rosen & Hatch in press). However, no comprehensive study of subg. Limnochloa has been published since the seminal work of Svenson (1929, 1939). Eleocharis acutangula (Roxb.) Schult. is the most widely distributed species of Eleocharis subg. Limno- hloa (Svenson 1939 [as E. fistulosa Schult.]). In the New World it is reported from near sea level to eleva- tions over 2200 m from various habitats including cloud forests, forest depressions, savannahs, grasslands, palm swamps, lake margins, borrow pits, and roadside ditches. Old World habitats include swamps, forest depressions, streams, savannahs, grasslands, borrow pits, lake margins, and rice paddies. Several authors have reported considerable variation in E. acutangula (Svenson 1929 [as E. fistulosa], 1939; Haines & Lye 1983; Browning et al. 1997). Svenson (1929, 1939) indicated E. planiculmis Steud. and E. fistulosa Schult. var. robusta Boeck. were potential segregates of E. fistulosa, which is treated herein as a synonym of E. acutangula. Hess (1953) described Heleocharis pseudofistulosa H. Hess based on plants he collected in Angola, and stated that they differed from E. fistulosa in surface characteristics of the achene. Hess (1957) later provisionally J. Bot. Res. Inst. Texas 1(2): 875 — 888. 2007 876 Journal of the Botanical R h Institute of Texas 1(2) reported H. cf. pseudofistulosa from South America (Brazil), which differed in having terete rather than sharply three-angled culms; this is presumably E. obtusetrigona (Lindl. & Nees) Steud. Svenson (1939) included E ing et al. (1997) described variability among specimens of E. acutangula from different geographical areas in = eocharis fistulosa among five poorly defined tropical African taxa, and Brown- southern Africa. Our research reported here, including observations of live plants in the field and a study of herbarium specimens from a broad geographical area, shows considerable variability within E. acutangula, thus confirming the work of Svenson (1939) and Browning et al. (1997). A critical examination of over 600 specimens of E. acutangula suggested sufficient variation existed to warrant recognition of three infraspecific taxa: E. acutangula subsp. acutangula, E. acutangula subsp. breviseta, and E. acutangula subsp. neotropica. The objectives of this research were: (1) to investigate the morphological variation within E. acutangula and (2) to review all the apposite nomenclature in order to typify E. acutangula and its synonyms. METHODS Specimens were borrowed from herbaria that could provide loans yielding broad geographical representa- —= tion of Eleocharis acutangula including types and authentic specimens. Over 600 specimens were examined from the following herbaria (acronyms follow Holmgren et al. 1990): BM, BRI, BRIT, C, CIIDIR, CM, E, F, FTG, GA, GH, IBE, ICN, K, LL, M, MEXU, MICH, MO, NH, NU, NY, P, PH, PRE, RSA, TAES, TEX, US, USF, VSC, WIS, Z, and ZT. Selected for multivariate analysis were 198 mature herbarium specimens (including types) complete for all morphological characters measured. Specimens studied originated from Africa, Aus- tralia, Bolivia, Brazil, China, Colombia, Cuba, Dominican Republic, Ecuador, El Salvador, Guyana, India, Japan, Madagascar, Malaysia, Mexico, Panama, Peru, United States, Venezuela, and Vietnam. Because of the limited number of specimens complete for all morphological characters, duplicate specimens collected by the same collector were measured. A complete citation of all specimens examined during this research can be found in Rosen (2006). Quantitative and qualitative vegetative characters (e.g., culm height, width, texture, and cross-sec- tional shape; leaf sheath texture and structure; rhizome length and diameter) are highly plastic. Although these features are of some use in Eleocharis at the infrageneric level, they are of no value in distinguishing infraspecific taxa. In Eleocharis subg. Limnochloa, significant variation in culm anatomy in response to en- vironmental conditions has been reported (Edwards et al. 2003; Baksh and Richards 2006). Svenson (1929) emphasized achene characters and perianth bristle texture in differentiating species of Eleocharis. A review of the literature reveals a tendency of workers investigating closely related species and infraspecific varia- tion in Eleocharis to rely primarily on characters associated with the achene (Hines 1975; Larson & Catling 1996; Gregor 2003). Indeed, achene-related characters are important in taxonomic limits in Eleocharis at all levels (Menapace 1991). oy ed E were selected for initial evaluation (Table 1). For each specimen a ma- d from near the base of a spikelet. Each specimen measured was complete for all rae so that the data matrix contained no missing values. One measurement per character was taken from each specimen, and 198 specimens (114 of Eleocharis acutangula subsp. acutan- gula, 67 of E. acutangula subsp. breviseta, and 17 of E. acutangula subsp. neotropica) were analyzed utilizing principal component analysis (PCA). The raw morphometric data were standardized and analyzed using NTSYSpc 2.11Q, and the principal components were generated using a correlation matrix (Rohlf 2000). A final analysis comprising six characters (Table 2) was run, and a scatter plot of the first two principal components was generated in an effort to depict morphological relationships. RESULTS The first three principal components represented 87.3% of the total variance (50.3%, 24.3%, and 12.7% for PCI, PC2, and PC3 respectively; Table 2) of 198 specimens scored for six morphological characters. Prin- cipal component 1 is most influenced by high positive loadings of LONBRSTL, TBRCL, BRSTLNACHNL Rosen et al., T f Eleoci g | 877 TABLE 1. Initial 20 fi Fleact gul Symbol Character ACHNL achene length (from base to constriction at neck) ACHNLW ratio of achene length to width (achene shape) ACHNMAX ratio of achene length to distance from achene base to widest point (determines if achene is widest above, at, or below middle) ACHNSCAL ratio of achene length to floral scale length ACHNW achene width (at widest point) BRSTACH number of perianth bristles longer than summit of achene BRSTLACHNL ratio of length of longest perianth bristle to achene length BRSTLNACHNL ratio of number of perianth bristles longer than summit of achene to total number of perianth bristles BRSTN number of perianth bristles LONBRSTL length of longest perianth bristle LONROW number of longitudinal rows of cells on achene face NECKWACHNW ratio ofachene neck width to achene width NECKW achene neck width SCALEL floral scale length SCALELW ratio of floral scale length to width SCALEW floral scale width TBRACHW ratio of tubercle width to achene width TBRCL tubercle length TBRCLW ratio of tubercle length to width (tubercle outline shape) TBRCW tubercle width (Table 2). Principal component 2 is most influenced by a high positive loading of NECKWACHNW and a high negative loading of TBRCIW (Table 2). Although there are vary- ing degrees of overlap among the three taxa, specimens from each 1 subspecies tinct groups (Fig. 1). Specimens of Eleocharis acutangula subsp. breviseta and E. acutangula subsp. neotropica are almost completely separated along principal component axes l and 2 (Fig. 1). Specimens of E. acutangula subsp. acutangula overlap slightly with E. acutangula subsp. breviseta along principal component axis l and E. acutangula subsp. neo- tropica along principal component axis 2. The relatively small area of the graph (Fig. 1) occupied by E. acutangula subsp. breviseta and E. subsp. neotropica compared to that occupied by E. acutangula subsp. acutangula is presumably the result 1.5 | t t t t yy yo v vv vy 1.0 + n e vy L : Y" o v % 0 0.5 + j No) (e) | IE o Ke together into dis- e O o O i O O Q 0.0 2 “5° j AX O UT e Oo o) e oO [e] a rabo 008 98% 7 e $$ "o d o dr. e... $ » i mos Ry -0.5 + id e e? o? - BE ... ? EE o e O O o O -1.0 + o + -1.5 t | t t | -1.5 -1.0 -0.5 0.0 0.5 1.0 1.5 PC 1 Fi 1.8 pl ££ Ce a | f, Df ^ €, iables from 198 specimens ircles), E. acutangula subsp breviseta (close b £ El n ; 4. nj } 4. lat 9 r circles), and E. acutanaula subsp an wA F 878 Journal of the Botanical R h Institute of Texas 1(2) TABLE 2. El | ] | i |l h principl | loadi hefi i used in PCA of 198 speci f Eleocharis gula subsp. acutangula, E. acutangula subsp. breviseta, and E. acutangula subsp. neotropica Character PC1 PC2 PC3 Eigenvalue 3.018 1.459 0.760 Percent variance 50.302 24,317 12.675 LONBRSTL 0.862 0.365 0.030 TBRCL 0.832 -0.348 -0.290 BRSTLNACHNL 0.828 0.324 0.040 TB 0.695 -0.643 -0.259 ACHNL 0.644 0.220 0.576 NECKWACHNW 0.015 0.799 -0.524 El as I, acutanaula. E acutangula subsp. breviseta and E acuranaula subsp. neotronica. M Y Y | | | TABLE 3. Select ct isons fi + 1 standard deviation) are provided for quantitative characters. Character subsp. acutangula subsp. breviseta subsp. neotropica achene length (mm) 1.8(1.6-2) 1.6(1.4-1.7) 1.7(1.6-1.8) achene width (mm) 1.4(1.2-1.6) 1.3(1.2-1.4) 1.4(1.3-1.5) achene color at maturity shiny dark amber shiny dark brown shiny yellow-green (tinged (dark brown) with amber) tubercle width (mm) 0.7(0.6-0.9) 0.7(0.6-0.8) 0.9(0.8-1) chene neck width (mm) 0.6(0.4—0.7) 0.5(0.4—0.6) 0.8(0.7—0.8) tubercle length (mm) 0.6(0.5—0.8) 0.4(0.3-0.5) 0.5(0.4—0.5) ratio of achene neck width 0.4(0.3—0.5) 0.4(0.3-0.4) 0.5(0.6-0.7) to achene width description of perianth usually all overtopping usually few-none all overtopping tubercle; bristles summit of achene and overtopping summit of coarsely retrorse nearly to sometimes the tubercle; achene; only afew short, | the base coarsely retrorse nearly salient retrorse spinules to the base or less often near the tips completely smooth length of longest perianth 2.6(2-3.3) 1.2(1-1.5) 3.6(3.2-4) bristle (mm) ratio of length of longest 1.4(1.1-1.7) 0.8(0.6-0.9) 2(1.9-2.3) perianth bristle to achene length ratio of tubercle length 0.8(0.6-1.1) 0.6(0.5-0.8) 0.5(0.4-0.6) width of less morphological variability in E. acutangula subsp. breviseta and E. subsp. neotropica, which perhaps stems from their relatively limited geographical distributions when compared with the more morphologically variable and widespread E. acutangula subsp. acutangula. This could also indicate active speciation (local adaptation) in E. acutangula subsp. acutangula. DISCUSSION AND CONCLUSION Multivariate analysis and thorough examination of ca. 600 specimens, including types, warrants the recognition of three infraspecific taxa within Eleocharis acutangula. The presence of several conspicuous morphological differences between the three taxa along with a relatively cohesive geographic distribution of E. acutangula subsp. breviseta and E. subsp. neotropica (Fig. 2) suggests subspecies is an appropriate rank for classification (Stuessy 1990). A summary of the characters accounting for most of the variability in the multivariate analysis and our observations of achene color and the length and texture of perianth bristle Rosen et al., T f Eleochari t | 7 J 879 spinules indicate that features of the mature achene and perianth are essential for identification of the sub- species of E. acutangula (Table 3). Eleocharis acutangula subsp. breviseta is distinguished from E. acutangula subsp. acutangula by its shorter achenes and tubercles and short perianth bristles with only a few short retrorse spinules near the tips. Eleocharis acutangula subsp. neotropica differs from E. acutangula subsp. acu- tangula by its long, soft, flexuous perianth bristles, weakly constricted achene apex, and the tubercle being usually as wide to wider than long. Differences between E. acutangula subsp. breviseta and E. acutangula subsp. neotropica are summarized in Table 3. Greater variability was observed in specimens referable to Eleocharis acutangula subsp. acutangu pany a for several of the parameters used in the multivariate analysis (Fig. 1; Table 3). Variation was also observed in achene epidermal cell shape, often in achenes from the same herbarium specimen (Rosen 2006). Svenson (1929, 1939) suggested that E. fistulosa var. robusta and E. planiculmis may represent taxa distinct from E. acutangula. However, a critical examination of the types indicates they are only minor expressions of highly variable E. acutangula subsp. acutangula. Specimens from Madagascar [DuPuy 2429 (MO, K, P); Bathie 17929 (P, US); and Bathie 2722 (P)], including type material of E. fistulosa, exhibited obtusely trigonous culms rather than sharply wing-angled triquetrous culms observed in all other specimens of E. acutangula examined. Indeed, the protologue of S. fistulosus describes the culms as “subtriquetro”. These specimens are otherwise referable to E. acutangula subsp. acutangula for the characters used in the multivariate analysis. We do not propose segregation on the basis of a single, highly plastic vegetative character and with such a limited number of specimens examined. TAXONOMIC TREATMENT KEY TO SUBSPECIES OF ELEOCHARIS ACUTANGULA . Longest perianth bristle 3.2-4 mm long, soft, ee re spinulose to below the middle (nearly to the base); achene neck weakly constricted, 0.6-0.7 times achene width; tubercle 0.4-0.6 times long as wide; mature achenes yellow-green (tinged with les distribution limited to northwest South America subsp. neotropica . Longest Suid bristle 3.2 mm long or shorter, a Des leia only at the tips to near m base Bern or sometime Spoon eas neck aS constricted, 0.6 achene width or less; tubercle 0.5-1.1 times long a e; ! ch s dark amber to us brown distribution more aa 2. Perianth nice: ee achene or rarely few to all reac spinules restricted to the distal half or more commonly only near the tip; achene 1 ie 7 E mm long, dark brown; 3 bercle 0.3-0.5 mm lon subsp. breviseta erianth bristles | than ach | ,Spin ems Adel to base or rarely HT omens, Piu MEAT 6-2 mm s qd abet or rarely dark brown; tubercle 0.5-0.8 mm long N subsp. acutangula jua . Eleocharis acutangula (Roxb.) Schult. subsp. acutangula (Fig. 3 a—b). Scirpus acutangulus Roxb. Fl. Ind. 1:216. 1820. Eleocharis acutangula (Roxb.) Schult. Mant. 2:91. 1824. Limnochloa acutangula (Roxb.) Nees. Contr. Bot. India 114. 1834. Tver: INDIA, Roxburgh s.n. (Lectotype here designated: BM [BM000847992]!). Scirpus medius Roxb. Fl. Ind. 1:216. 1820. Limnochloa media (Roxb.) Nees. Contr. Bot. India 114. 1834. Tyre: INDIA, Roxburgh s.n. (not found) Scirpus fistulosus Poir. Encyclopédie Méthodique, Botanique 6:749. 1804. nom. illeg., non Scirpus fistulosus Forssk. 1775. Eleocharis fistulosa Schult. Mant. 2:89. 1824. Type: MADAGASCAR, Poiret s.n. (HOLOTYPE: P [Herbier du Petit-Thouars., P00376392]!; ISOTYPE: Herb. Poiret in Herb. Moquin-Tandon [P00370140]9. Eleocharis pu osa Schult. var. robusta Boeck. Flora 62:563. 1876. Heleocharis jua o Hess. Ber. Schweiz. Bot. Ges. 63:331. cum descr. ampl. Type: AFRICA, Africa centralis, Seriba Ghass g ep 1869, Schweinfurth 2326 (LECTOTYPE here designated: GH!; ISOLECTOTYPES: Z [000006263, OI Eleocharis planiculmis Steud. Syn. Pl. Glumac. 2:80. 1855. Tyre: Java, Zollinger 281 (notorvee: P [P00368895]!; isotypes: P [P00368896, P00368897]!, K [K000290949, onto Heleocharis pseudofistulosa H. Hess. Ber. Schweiz. Bot. Ges. 63:329. 1953. Type: ANGOLA: Provinz Huila, Guanhama, Tümpel, 15 km südlich Cubango an der Strasse pu Cassinga, 14 Jan 1952, Hess 52/220 (HoLotyre: ZT photo! [based on Hesss designation of “Typus”]; isorvre: BOL, K, TAES!, Z 880 Journal of the Botanical R h Institute of Texas 1(2) Fic. 2. Distributi f Eleochari ti I b ti la ( irdes), E. acutanaula subsp breviseta (closed circles), and E. acutangula subsp | \ g \ n g Plants perennial. Roots coarse, fibrous, drab-brown to reddish, small storage structures present in care- fully collected plants, cylindrical-reniform, brown; primary rhizomes caudex-like, thick, hard, ascending, concealed by roots and persistent culm bases (occurring only in carefully collected specimens); secondary rhizomes elongated, to 4 mm thick, scales to 17 mm long (few seen). Culms triquetrous (a few specimens from Madagascar trigonous) distally, (25—)38-81(-135) cm long x (1.2-)2.1-4.4(-6.5) mm wide, soft, internally spongy, with incomplete transverse septa, smooth, green, finely longitudinally striate when dry. Leaves 2, reduced to sheaths, apically oblique, membranous, loose, friable, proximally pinkish to dark maroon (dark purplish), distally drab, apex acute. Spikelets cylindric, (11-)21-39(-56) mm long x (2.5-) 3.2-4.8(-6) mm wide, acute; proximal scale with flower, obtuse, amplexicaul-clasping, appearing as a continuation of culm, remaining floral scales conspicuously spirally arranged, appressed to somewhat spreading at maturity, ovate-oblong, 2.5—)3.8-5.3(-6) mm long x (1.7-)2.3-3.4(-4.8) mm wide, cartilaginous, abaxially greenish to stramineous lly, j ginally, sparsely red-maculate and sometimes the veins or other areas reddish or pinkish (purplish) tinged, usually with a fine dark band near apex, adaxially sparsely to copiously red-maculate, apex acute (rounded), distal 0.1-0.5 mm translucent hyaline-erose, central area nearly flat, coarsely many veined, only mid-vein conspicuous in adaxial view. Flowers with 6—7(-8) perianth bristles; bristles sub-equal, usually 1.1-1.8 times the length of the achene (rarely one or few just reaching its summit or slightly shorter), retrorsely spinulose nearly to base or rarely completely smooth (both conditions can occur in same population), stramineous or pinkish to dark maroon; stamens 3; anthers (1.1-)1.3—2.2(-3.2) mm long, stramineous; style 3-fid. Achenes biconvex, very broadly obovoid to obovoid, the shoulders and sides near the apex usually straight and forming an obtuse angle, or sometimes rounded, (1.4-)1.6-2.0(-2.2) mm long x (1.02)1.3-1.6(-1.8) mm wide, with (11212-15(-19) longitudinal rows of deeply concave trans- versely oblong to linear polygonal cells visible through transparent periclinal layer on each achene face, dull yellow-buff maturing to shiny dark amber (dark brown), apex constricted to a distinct neck about 0.3-0.5 times the width of achene. Tubercle dorsoventrally compressed, shallowly triangular-deltate (triangular), 0.5-0.8(-1.1) mm long x (0.5-)0.6-0.9-1.2) mm wide, stramineous, maturing to dark brown. Rosen et al., T f Eleochari t | 881 Lectotypification of Eleocharis acutangula Eleocharis acutangula was described by Roxburgh (1820) as Scirpus acutangulus based on plants from India. As is the case with apparently all Roxburgh names, no type specimen was designated (Forman 1997). Schultes transferred S. acutangulus to Eleocharis without indicating a type. A literature search revealed no reference to a particular type specimen although a number of authors indicate the “type” is from India (e.g., Haines & Lye 1983; Gordon-Gray 1995; Browning et al. 1997). Typifying Roxburgh names can be difficult since his specimens were widely distributed, making locating specimens annotated by him or known to be associ- ated with him challenging (Forman 1997). Almost all of Roxburgh’s nearly 2600 species were illustrated by color drawings prepared by local Indian artists; the original set is at CAL, and a duplicate set at K (Sanjappa et al. 1991). Forman (1997) indicated that the Flora Indica drawings were often superior to the correspond- ing Roxburgh specimen (if one can be found), and in some instances make a better choice for a type. From the set of drawings at Kew a high resolution digital photograph was obtained of the front and back of the drawing of S. acutangulus. The drawing, a stylized depiction of an immature plant, was annotated in what the first author interprets as Roxburgh’s hand. We made queries to curators at key herbaria indicated by Forman (1997) in an effort to locate an au- thentic Roxburgh specimen. Mark Spencer (BM) presented a specimen (BM-000847992) that he considers to have been associated with Roxburgh, the most compelling evidence being annotations on the verso and front of the specimen. The verso is annotated “Ind. Orient Roxburgh” in an unknown hand, indicating that the specimen was received from Roxburgh, and the front was annotated “72” in what the first author inter- prets to be Roxburgh’s hand. We selected this specimen as the lectotype of Scirpus acutangulus since it fits the description in the protologue and is thought with reasonable certainty to have been used by Roxburgh. Problems with Typification of Scirpus medius Roxburgh described Scirpus medius as being similar to S. acutangulus but having shorter culms with smooth, rounded angles. Nees (1842) transferred the name to the genus Limnochloa. Roxburgh’s description of the culms as having rounded angles is troublesome, as all Asian specimens of Eleocharis acutangula examined during this research had triquetrous culms. No specimens annotated as S. medius were seen by us, and no specimens were located in herbaria where Roxburgh’s specimens were distributed. Thus, we follow Svenson (1929) and others (Blake 1939; Koyama 1985; Gordon-Gray 1995) in placing S. medius in synonymy under E. acutangula. Clarification of the Authorship of Eleocharis fistulosa Scirpus fistulosus Poir. is illegitimate because of an earlier homonym, S. fistulosus Forsskal. Eleocharis fistulosa Link is also invalid because Link failed to associate the specific epithet with the name of the genus or species, or with its abbreviation, as mandated by the Art. 33.1 of the ICBN (McNeill et al. 2006). Thus, E. fistulosa Schult. is the correct author citation (See ICBN Articles 58.1; 7.5; and 33, Note 2.). Since the priority of E. fistulosa does not date back to the publication of Poiret's illegitimate use, E. acutangula (Roxb.) Schult. has priority as the oldest legitimate name for the species. Lectotypification of Eleocharis fistulosa var. robusta Eleocharis fistulosa var. robusta was described by Boeckeler based on Schweinfurth 2326 from Central Africa. Boeckeler's types were at B, and, if the holotype of E. fistulosa var. robusta was ever extant at B, it was destroyed by the fire of 1943 (Robert Vogt, B, pers. comm .). In this case, Schweinfurth 2326 from GH is designated as lectotype, and two duplicates from Z become isolectotypes. Excluded Name Eleocharis fistulosa var. micrantha Chermezon was described from specimens from Senegal (Chermezon 1936). Attempts to locate type specimens cited in the protologue have been unsuccessful thus far. Svenson (1939) relegated this name to synonymy under E. nupeensis Hutchinson & Dalziel based on the description, a temporary solution adopted here. 882 Journal of the Botanical R h Institute of Texas 1(2) Sa SEED ESOS L F + 1 dicts] E E " f£. b LAM HI E 4 I I Fic. 3. A-B Eleocharis acutangula subsp. acutangula: A abaxial view of act B apical view of achene. C-H E. acutangula subsp. breviseta: C habit; | p ; ; | l-K E. acutangula subsp pi [ iew of ad 1 K apical view of act A-B from Hooper € Gandhi 2373, C-F from Rosen & Carter 3206, G-H from Howard & Howard 9862, and I-K from McDaniel & Rimachi 18552. Drawn by Neva Mikulicz. P lecnibAl I Aer nf | A A Representative Specimens Examined: NORTH AMERICA. MEXICO: Chiapas: E side of Pueblo Solistahuacan, Municipio of Pueblo Nuevo Solistahuacan, elev. 1700 m, 26 Oct 1971, Breedlove 21527 (MO, NY). Hidalgo: Lake Atexca below Molango, 09 Nov 1946, Moore 1938 (GH). Jalisco: I Guadalajara, 1888, Pringle 2061 (NY). Nayarit: near Lake Labor, ca 15 mi SE of Tepic, 25 Sep 1960, McVaugh 19426 (MICH). Tabasco: km 64 rumbo de Huimanguillo a Fransisco Rueda, 35 msnm, 06 Nov 1979, Orozco & Zamudio 2187 (MO). Veracruz-Llave: Mpio. Las Choapas, ca 5.4 km S of the town of Las Choapas, along rural road to El Chichon, 13 Jul 2006, Rosen et al. 3870 (CIIDIR, GH, K, MICH, MO, TAES, TEX, US, VSC, WIS). CENTRAL AMERICA. GUATEMALA: Alta Verapaz: E of Tactic, alt 1,300 m, 20 Feb 1942, Steyermark 43970 (F). Chiquimula: between Chiquimula and La Laguna, alt. 500-1000 m, 27 Oct 1939, Steyermark 30713 (F). Huehuetenango: vicinity of Maxbal, ca. 17 mi N of Barillas, Sierra de los Cuchumatanes, alt. 1500 m, 15-16 Jul 1942, Steyermark 48770 (E). Izabal: near Puerto Barrios, sea level, 25 Apr 06 May 1939, Standley 72862 (F). Jutiapa: SE end of Potrero Rosen et al., T y of Eleocharis acutangul 883 Carrillo, 13 mi NE of Jalapa, alt. 1500-1700 m, 12 Dec 1939, Steyermark 33099 (F). Santa Rosa: 4 mi N of Barberena, 18 Feb 1951, Fassett 28844 (F). BELIZE: Toledo District: near junction of Southern Highway and Pine Hill, 22 Nov 1998, Holst et al. 7064 (MO). HONDURAS: Comayagua: beu of Siguatepeque, ca. 1050 m, 25 Mar 05-Apr 1947, Standley & Chacon 6595 (F). Copan: 14 Jul 1971, Harmon & Fuentes 6445 (MO, NY-2 sheets). Francisco Morazan: near Las Mesas, 900 m, 10 Sep 1950, Standley 26634 (GH). Olancho: Santa Maria del BN 23 mi NE of San Esteban along road to Bonito Oriental, 03 Jul 1994, Davidse et al. 35564 (CIIDIR). EL SALVADOR: Ahuachapan: Lagunita las Ninfas, Apaneca, 28 Jan 1951, Fassett 28721 (GH). NICARAGUA: Comarca del Cabo: Bihmona, 7 Jul 1972, Seymor 5707 (CIIDIR, GH, MO-mixed with E. interstincta). Esteli: Reserva Natural Miraflor, Municipio de Esteli, Comunidad los Volcancitos, 10 Jul 1999, Rueda et al. 11643 (MO). Zelaya: Cano Manso Awalka Tingni, reached by Geodesia turn on road between Torre 7 and Bismuna Tara, ca. 11.9 km SW of Bismuna Tarra, 19 Apr 1978, Stevens 7704 (CIIDIR-2 sheets, MO). COSTA RICA: Isla De Cocos: Macollas en suelo humedo, a la orilla del par tano, Bahia de Wafer, ca. nivel del mar, 31 Jul 1981, Gomez-Laurito 6915 (F, MO). Alajuela: 6 km W of Venicia, elev. 450 m, 15 Oct 1968, Davidse & Pohl 1307 (F, MO). Cartago: Laguna Dona Anacleta, Canton Paraiso, Lago Crater, 22 Aug 1983, Novelo 1209 (MO). Guanacaste: upper N fork of Rio Sabalito, just N of San Joaquin de Coto Brus, 13 Sep 1985, Grayum et al. 6011 (MO). Puntarenas: San Joachim de Sabosa, just N of San Vito, 22 Feb 1982, Barringer & Gomez 1688 (F). PANAMA: Chiriqui: S of El Boquete, 01 Mar 1918, Killip 4569 (NY). Cocle: El Valle de Anton and vicinity, 500—700 m, 23-27 Jul 1935, Seibert 476 (MO, NY, US). Veraguas: vicinity of La Mesa in sunny muddy bottom in pasture, 28 Dec 1968, Tyson 6054 (MO). CARIBBEAN BASIN. JAMAICA: Clarendon Parish: Mason River Field Station, 4 mi W of Kellitts, 2300 m, 27-29 Jul 1979, Thomas 2146 (MICH). Saint Catherine Parish: Charlton, near Ewarton, 03 Apr 1903, Harris 8513 (NY). DOMINICAN REPUBLIC: San Cristóbal Province: between Duarte Hwy. Kl. 28 and Haina, 11 Oct 1947, Allard 15976 Cee Santa Domingo Province: 8 km from La Batata on road to Mata de Piedra and La Catalina, 09 Dec 1980, Mejia & Zar . DOMINICA: Lesser Antilles, St. George Parish, vicinity of freshwater lake, NE of Laudat, E side of Morne Macaque, loccaly common, 2500’, 20 Mar 1991, Hill et al. 22119 (GH, R regimiento de Puerto Valdivia, km 11 de Pto a mina uc Oro "Canarias", colecciones en escombreras de 2 anos, 14 May 1987, Callejas et al. 3486 (MO-—2 sheets, NY). Risaral La Virginia-Cerrito, extremo norte de parte ancha del Valle del Rio Cauca, lomas bajas, 22 Aug 1989, Silverstone- ae 5504 (MO). Cauca: Chisquio, Finca Los Derrumbos, alt. c. 1700 m, 11 May 1940, Asplund 10577 (LL). Huila: 3 km W of Garzon, upper basin of Rio Magdalena, 17 Feb 1959, Mason 13888 (GH, US). Meta: ca. 17 km SW of Puerto Lopez, along road between La Balsa and Bocas del Guayuriba, 17 Jan 1970, Schuyler 4165 (PH). Valle del Cauca: Calima, on Rio Calima, 14-15 Sep 1922, Killip 11247 (GH, NY, PH). VENEZUELA: Apure: Guanare, Esteros y pantanos cerca de los AA cano Mau 25 Oct 1980, Stergios 2387 (MO). Aragua: El Limon, near Maracay, in Morass, 29 Jan 1922, Pittier 10116 (GH). Bolivar: ato Sta. Teresa, Mar 1946, Tamayo 3211 (F, US). Guarico: Orituco, 25 km SW de la Estacion Biologice de la Clanus (sic) Edo Gu 19 o 1982, Montes 1343 (MO). Portuguesa: terrenos de la Unellez, 06 Sep 1984, LU 7051 (MO). GUYANA: Upper Takutu-Upper Essequibo Region: Rupununi Distr., Shea Village, 09 Feb 1994, Jansen-Jacobs et al. 3634 (NY, US). SURINAM: in Maurisie swamp, W of 4-Gebroeders Mts., 27 Sep 1968, Oldenberger et al. 194 (NY). ECUADOR: ao Amazonica, Archidona, Coca km 9.3 roadside, 15 Apr 1988, Laegaard & Renvoize 70909 (MO, NY). Pastaza: Amazonica, Hacienda San Antonio de Baron von Humboldt, 2 km al Nede Mera, 20 Feb-20 Mar 1985, Palacios et al. 144 (MO). BRAZIL: Distrito Federal: Brasilia, 27 Jun 1979, Heringer 1652 (NY). Bahia: 37 km N from Correntina, on the Inhaumas road, 29 Apr 1980, Harley 21957 (NY). Mato Grosso: 17 Oct 1968, Harley et al. 10711 (NY). Minas Gerais: without location, 1816-1821, Catal 616 (K, P). Sao Paulo: Butantan, S. Paulo, with- out date, Gehrt 5403 (GH, NY). Amazonas: lagoa permanente, 500 m ao Sul da BR 230 km 4, 15 Aug 1980, Janssen & Gemtchujnicov 514 (M). Maranhao: Brejo, Ets. Ecologica UFMG, 02 Apr 1991, Neto 461 (CIIDIR). Parana: Rolandia, Fazenda Conquista, area alagada, dentro da lagoa, no. 14, 11 Mar 2003, Vanzela 35.42 (CIIDIR). Rio de Janeiro: Goias, Formosa, Bisual, 20 Oct 1965, Pereira & Duarte 9414 (NY). Rio Grande do Norte: near Bento Fernandes, 70 km W from Natal, shallow at pond’s edge, 28 Aug 1987, Tsugaru & Sano B-1273 (GH). Rio Grande do Sul: M. Rio Pardo, Riniao Reserva, Feb 1923, Jurgens s.n. (US). Without location, 1844, Weddele 1195 (P). BOLIVIA: La Paz: Iturralde, Luisita, sabana humeda, W del rio Beni, Palmar, 12 Sep 1984, Haase 540 (NY). Santa Cruz: Andres Ibanez, NE side of Viru-Viru Pampa and property of Aeropuerto Internacional, along road to Chuchio, 4.5 km E of turnoff from highway from Santa Cruz to Warnes on road to Chuchio, 15 May 1998, Nee 49365 (CIIDIR, TEX). PARAGUAY: Departamento Central: Estero del Ypoa, Villeta, Puerto Guyrati, 4.5 km S of Villeta, 02 Dec 1992, Zardini & Aquino 34134 (CIIDIR, US). Caazapa: Tavai, Enrramadita, 05 Dec 1988, Mereles 2067, 2069, 2070, 2081 (MO). la Cordillera: 1 km E of Nueva Colombia on road to Atyra, 09 Jun 1990, Zardini & Velazquez 20917 (MO). Misiones: Estancia La Soledad, Santiago, 30 Apr 1961, Pedersen 6029 (US, GH, MO, NY, TEX). Paraguari: Estero Ypoa, between Nueva Italia and Yuquyty ona hill, 18 Mar 1992, Zardini & Aquino 31333 (CIIDIR). Amambay: Ao. Estrella, Prop. De Heisecke, 08 May 1989, Soria 3749 (MO). ARGEN oncepcion, Carambola, Estancis “Buena Vista”, 19 Feb 1985, Pedersen 14072 (MO, NY). Misi Aira, 1913, Rodrig 3 (GH). Chaco: Dep lo de Mayo, Colonia Benite balsado burger, 16 Dec 1943, Schulz 4118 (F, GH). AFRICA. ANGOLA: Benguela: Gebirge sudlich Ganda, Tumpel bei Calusipa, 30 km sudlich Chicu- ma, 1580 m, 24 Dec 1951, Hess 51/419 (Z). Bie Bie: Baixo Cubango, 28 km nordlich Caiundo in der Umgebung der Missao cat. Capico, flacher sumpf, 31 Jan 1952, Hess 52/525 (Z). BOTSWANA: Ngamiland District: Moremi Wildlife Reserve, N Okavango swamp, Kwani River floodplain, Jul 1964, Tinley 1057 (NU). BURKINA FASO: I’ ene Mare de Bidi, 20 Sep 1996, Madsen 5749 (NY). Boulgou: some km SE of Tenkodogo, 31 Aug 1996, Madsen 5455 (NY). B JNDI: B Plaine Rusizi km 14, 800 m, 16 Mar 1975, Reekmans 4390 (MO). Bujumbura: Bujumbura, plaine Rusizi km 14, 780 m, 13 Feb 1972, Reekmans 1539 (MO). Provense ya Bururi: Giho Gihara, 20 May 1980, Reel 205 (MO). CAMEROON: Nord: ca. 15 km NE of Maroua, along road to Waza, 12 Sep 1964, Wilde 3215 (K). CENTRAL AFRICAN REPUBLIC: Prefecture de la Sangha-Mbaere: Sangha Economique, Dzanga-Sangha Reserve, 40 km S of 884 Journal of the Botanical R h Institute of Texas 1(2) Lidjombo on tributary of Keine, 26 Oct 1988, Harris & Fay 1488 (MO, PRE). COMORO ISLANDS: Mayotte, Grande Terre, Ouangani, Coconi, Valarana, 26 Fev 2002, Barthelat 6 Sifari 708 (P). CONGO: Vallie Uruanda, 26 Oct 1953, Liben 852 (K, PRE). ETHIOPIA: Kaffa Province: d ca. 5 km E of Jimma, along bi b to Addis Ababa, 02 Nov 1970, Friis et al. 38 (C, K). GABON: Nyanga: à plus ou moins 7km sur la route de Doussala vers B direction Nord-Ouest, Petit nds 25 Mar 2000, Sosef 1016 (MO). Haut- yon Bateke Plateau, Mpassa River watershed, 4. 2 km N of station of the Project de Protection Gorilles, 27 Nov 2001, Walters et 1. 982 (MO). GHANA: Brong-Ahafo Region: 1 m S of Atebubu, 16 Nov 1970, Hall & Duodu 42128 (MO). Guinea-Bissau: 08 Dec 1944, o 1594 (MO). Ivory Coast: d’ Abidj f Abidjan, 21 Oct 1963, Wilde 1109 (Z). LIBERIA: Grand Bassa County: Sanokwele Dist., Ganta, 02 Dec 1935, Harley 781 (NY, US. Nimba County: Mt. Nimba, Crete, Marc a’, 04 Jul 1974, Adam 28878 (MO). MADAGASCAR: Antananarivo Province: Ankazobe, Jun 1927, Bathie 17929 (P, US). Fianarantsoa Province: Ambatofinandrahana, Itremo, petite vallee a l'ouest du Massif de l'itremo, 1680 m, 26 Nov 1993, Du Puy & Andriantiana 2429 (MO, K, anga Province: 10 km E Antsalova, 20 Mar 1993, Villiers et al. 4855 (K). Toamasina province, Marais de Didy, voir Joncacee ef aa2m, 21 Feb LE Cours 1758 (P). Malawi: C l Region: Kasungu National Park, Angombe Hill, 03 Sep 1970, Hall-Martin 1712 (PRE). Nigeria: Kano: 12 Sep 1973, Jackson & Apcjoye 10-12973 (MO). Plateau: in vicinity of Bukuru, near Jos, 29 Jun 1970, Blum 2488 (WIS). Rhodesia: Hartley: Avondale farm dam, 25 Feb 1969, Mavi 983 (NU). S.W.A. (Southwest Africa?): 8 km S of Makuri vlei on road to Gimsa, 03 Mar 1985, Hines 361 (PRE). Senegal: Kaolak Region: Kaolak, Nov 1824, Berhaut s.n. (Z). Sierra Leone: Elliot 4453 (GH). SOUTH AFRIC Province: Transvaal, Witklip S bos, Nelspruit Dist., in water in Witklipdam, Kruid, 27 Jan 1976, Kluge 862 (PRE). KwazalacNacal: North coast, Lake Nhlabane area, W corner of North Lake, 19 Sep 1991, Ward 11378 (PRE, NU, NH red SUDAN: nan. Nyany, nr. Maar, 80 km N of Bor, 01 Feb 1981, Lock 81/10 (K). Swaziland: Malolotja Nature Reserve, Yers dam, stream, 17 Dec 1985, Heath 406 (PRE). Tanzania: Dar es Salaam Region: Mbezi, 2km WNW of Dar es Salaam SM (by cattle E to Tanzania packers), 12 Jun 1974, Wingfield 2752 (MO). Iringa District: T7, km 13 Ufinda-Sao Hill Rd., E side of road, 10 Jun 1996, Faden et al. 96/130 (K, US). Ruvuma Region: Ruanda, Urundi, Vallie Uruanda, Oct 1953, Liben 852 (M). Zaire: P Equateur: Bikoro, 01 Oct 1957, Thonet 7 (M). Kasai-Occidental: Kabinda, 26 Jul 1934, Becquaert 62 (GH). Katanga: River Kalule, pres de la ferme Rostenne, Elisabethville, 27 Mar 1963, Symoens 10155 (K). Province Orientale: 1940, Germain 171 (M). Zambia: Central Province: Mkushi Dist., David Moffat's farm, Munchiwemba dambo, 20 Sep 1995, Bingham & Nkhoma 9711 (PRE). Luapula Province: Lake Bangweulu, S part, swamps between Ncheta Island and Chibambo Lagoon, 11 Feb 1996, Renvoize 5585 (K). Northern Province: 8 km N of Kasama, 22 Jan 1961, Robinson 4296 (K, MO, NU). Western Province: ca. 10 mi NE of Mongu, 18 Nov 1959, Drummond & Coohson 6597 (MO). Zimbabwe: Manicaland: Mare Dam, Rhodes Inyanga National Park, 06 Jan 1972, Gibbs Russell 1210 (M, MO-2 sheets, K). Matabeleland North: Wankie National Park, Ngamo Pans 54 mi SE of main camp, 17 Apr 1972, Russell 1645 (NU). Salisbury District: 6 mi spruit, 4800’, 10 Jan 1932. ASIA. CHINA: Huebi: Central China, 1885-1888, Henry 4102 (GH, US-2 sheets). Yunnan: 1530 m, May 1936, Wang 73552 (GH). INDIA: Bangladesh: East Bengal, 1863-64, Griffith 6235 (NY). Karnataka: near Station, 10 Nov 1971, Hooper & Gandhi 2373 (MO, NY). Kerala: Malappuram Dist., between Tirurangadi and Parappanangadi, almost sea level, 12 Nov 1993, Cook & Camenisch 5169 (Z). Maharashtra: Pashan, near Poona, lake margin, 30 Dec 1971, Hooper 112 (K). Nepaul: without date, Hook & Thomson s.n. (NY). Tamil Nadu: Dharmapuri, Denl luk to Jowalagiri, to Karareddy pond, 18 Dec 1978, Matthew & Venugopal 2041 . INDONESIA: Alor: 1938, Jaag? s.n. (ZT). Java: Meester Cornelis, 1991, unknown 23139 (K). Jawa: Barat, Banten, 1936, Hachenberg 1 (GH). JAPAN: Hondo: Shinjo in Kii, 11 Oct 1953, } i Nagaitani valley, 15 Oct 1953, Koyama 5885 (MO, NY). Kyushu-chiho: Hondo, E in Kii, 11 Oct 1953, F 838 (BRIT, NY, US, WIS). Yoron-jima: Liukiuensis, 30 Aug 1921, Uyehara s.n. (US). MALAY PENINSULA: Langkowi, ricefields near Kueh, 14 Nov 1941, Comes? 37973 (K). MALAYSIA: Malacca: Kampong Bukit Piatu, 02 Apr 1955, es 40551 (K). PHILIPPINE ISLANDS: Lanao Mindanao: in 6” of water pocket in grassland, 04 E 1938, Zwickey 50 (GH, US). SIAM: growing in open fields, 13 Jul 1968, Kerr 15798 (K). SRI LANKA: North Eastern Province: Amparai Dist., Helawe Piya ca. 7 miS m Panama, E of Helawe Lagoon, sea level, 08 Feb 1971, Koyama et al. 14026 (GH, NY). VIETNAM: Quang Nam-Dà Nang: T ( ): Annam, Mount Bani, in tl ca. 25 km from Tourane (Da Nang), May—Jul 1927, Clemens & Clemens 4050 (E, X, MO, NY, PH, US, US, Z). OCEANIA. AUSTRALIA: Queensland: Cook Dist., Abattoir swamp, 4.5 km N of Mount Molloy, 20 May 1995, Clarkson 10317 (BRD. PAPUA-NEW GUINEA: Morobe District: vicinity of Kajabit Mission, n» 800-2000 ft, Aug-Dec 1939, Clemens 10600 (GH, US). National Capitol District: Hohola Port Moresby, 14 Nov 1973, White 37815 (BRI, GH, K, M, US) Distribution. —Pantropical; in México from the states of Chiapas, Hidalgo, Jalisco, Nayarit, Tabasco, and Veracruz-Llave. In Central America known from Belize, Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua, and Panama. In the Caribbean Basin known from Dominica, Dominican Republic, Grenada, and Jamaica. In South America known from records in Argentina, Brazil, Colombia, Ecuador, Guyana, Paraguay, Surinam, and Venezuela. Some previous reports of Eleocharis acutangula from the Galápagos Archipelago are based on misidentified specimens of E. obtusetrigona (Stewart 1911). Other reports were not verified (e.g., Jergensen & León-Yánez 1999), and no authentic specimens of E. acutangula from the Galá Archipelago have been seen. Perhaps most widespread and occurring in more variety of habitats in Sd Africa, with records from Angola, Botswana, Burkina Faso, Burundi, Cameroon, Central African Republic, Comoro Is- lands, Congo, Ethiopia, Gabon, Ghana, Gunea-Bissau, Ivory Coast, Liberia, Madagascar, Malawi, Nigeria, Rosen et al., T f Eleochari t | 885 Rhodesia, Senegal, Sierra Leone, South Africa, Sudan, Swaziland, Tanzania, Zaire, Zambia, and Zimbabwe. Of sporadic distribution in Asia and Oceania with records from Australia, China, India, Indonesia, Japan, Malaysia, Papua-New Guinea, Philippine Islands, Siam, Sri Lanka, and Vietnam. —Various disturbed and natural freshwater herbaceous and forested wetlands including marshy open grasslands, coastal savannas, and tropical forests from sea level to 2300 m. Reportedly forms expansive stands on a variety of soil types usually associated with other aquatic plants. Weedy in rice and other crop rota- tions and aquatic habitats, and used as a fiber crop in Borneo, Brazil, and Sumatra (Simpson & Inglis 2001). Note.—Eleocharis acutangula subsp. acutangula as treated here remains a variable taxon and includes forms meriting additional systematic study. Of particular interest are plants reviewed from Madagascar (including the type of E. fistulosa) with obtusely trigonous culms. 2. Eleocharis acutangula (Roxb.) Schult. subsp. breviseta D.J. Rosen, subsp. nov. (Fig. 3 c-h). Tyee: DOMINI- CAN REPUBLIC: El Seibo Province, 3-7 Nov 1946, Howard & Howard 9862 (HoLoTYPE: GH!; Isotype: NY-2 sheets!, P!, US). A Eleochari la (Roxb.) Schult. subsp l ianthii setis | ioril inuli i i i acheniis co) E E E E E E parvibus et lona brevioribus recedit. Plants perennial. Roots coarse, fibrous, mostly maroon (a few drab-brown), small storage structures pres- ent in carefully collected plants, cylindrical-reniform, white; primary rhizomes caudex-like, thick, hard, ascending, EU AME by roots and persistent culm bases (occurring only in carefully collected specimens); secondary rl elongated, to 3 mm thick, scales to 9 mm long (few seen). Culms triquetrous, (19-)30— 712134) cm long x (1.12)1.5—3.7(-7) mm wide, soft, internally spongy, with incomplete transverse septa, smooth, green when fresh, finely longitudinally striate when dry. Leaves 2, reduced to sheaths, apically oblique, membranous, loose, friable (upper distil portion disintegrating when submerged), proximally dark maroon, distally drab, apex acute. Spikelets cylindric, (10217-34(-49) mm long x Q.22)2.7-4.1075.5) mm wide, acute; proximal scale with flower, obtuse, amplexicaul-clasping, appearing as a continuation of culm; remaining floral scales conspicuously spirally arranged, appressed to somewhat spreading at maturity, ovate-oblong, (3.1-)3.3—-4.5(-5.9) mm long x (1.4-)1.8-3(-4.0) mm wide, cartilaginous, abaxially greenish centrally, stramineous marginally and sometimes reddish or pinkish tinged, with a fine dark band near apex, adaxially sparsely to copiously red-maculate, apex acute (rounded), distal 0.1-0.4 mm translucent hyaline- erose, central area nearly flat, coarsely many veined, only mid-vein conspicuous in adaxial view. Flowers with (5—)6-7(-8) perianth bristles, bristles sub-equal, (0.4—)0.6-1(-1.2) times the length of achene (rarely few-all bristles overtopping achene summit) with only a few short, salient retrorse spinules near tips (rarely spinules present in distal half), stramineous or pinkish to dark maroon; stamens 3; anthers (0.921.1-1.9(- 2.7) mm long, stramineous; style 3-fid. Achenes biconvex, broadly obovoid, the shoulders and sides near the apex usually straight and forming an obtuse angle, (1.3-)1.4-1.822.1) mm long x (1.121.2-1.4C1.6) mm wide, with (9211-14(-16) longitudinal rows of deeply concave transversely oblong (linear) polygonal cells visible through transparent periclinal layer on each achene face, dull yellow-green maturing through amber to shiny dark brown, apex constricted to a distinct neck about 0.4 times width of achene, in the field achenes sometimes persistent after the floral scales have shed giving spikelet a beaded appearance. Tubercle dorsoventrally compressed, shallowly triangular, (0.22)0.3-0.5(-0.6) mm long x (0.5-J0.6-0.8=1) mm wide, light brown tinged with green, maturing to dark brown. Specimens examined: NORTH AMERICA. U.S.A: Florida: Lee Co.: 4 km SW of the intersection of Hwy. 82 and Green Meadows Rd., SE of Fort Myers, 12 Nov 2004, Rosen 3206 & Carter (CIIDIR, GH, K, MEXU, MICH, MO, NY, P, TAES, TEX, US, VSC, WIS), S side of Griffin Rd., just S of entrance to Pinewood Lakes in Gateway, 28 Oct 1993, Orzell and Bridges 22526 (BRIT, FTG, USF). MEXICO: Campeche: a aprox. 10 km al sureste de la ciudad de Campeche, Mun. Campeche, alt. 80 m, 10 Nov 1980, Novelo & Zetina 721 (TEX), 14 km N of Ocozocoautla on road to Mal Paso, Municipio of Ocozocoautla de Espinosa, 07 Oct 1974, Breedlove 38254 (MEXU, MO). Chiapas: 96 km S of Mexican Hwy. 190 on road to Nuevo Concordia, 10 Oct 1974, Breedlove 38516 (NY). Guerrero: 1.5 km al NW del Rincon de la Via, 28 E 1988, Verduzco 389 (MEXU). CENTRAL AMERICA. PANAMA: vicinity of El Llano, 7-8 Sep 1962, Duke 5526 (MO, USF), Near the big he Rio Tecumen Province, 11 Dec 1923, Standley 26509 (MO). Canal Zone: Laguna de Portala, near Chepo, province a Oct 1911, Pittier 4602 (NY). Panama: Sabanas near Chepo, 30 m, 20 Jan 1935, Hunter & Allen 87 (MO), 1.6 km W of Juan Diaz, 10 Oct 1917, Killip 4090 (PH, RSA, US), Camino del Boticario, near Chepo, altitude 30 to 50 meters, Oct 1911, 886 Journal of the Botanical R h Institute of Texas 1(2) Pittier pr (GH, NY, US), near Matias Hernandez, wet ne 30 Dec 1923, Standley 28909 (US), near Matias Hernandez, wet field, 30 Dec 984 (US). CARIBBEAN BASIN. CUBA La Habana: Vedado-Habana, Sabana de Monasterio, 23 Jun 1920, Leon 9215 (NY). a location, 1860-1864, Wright 3376 (GH, mixed with E. mutata, MO, NY, P). SOUTH AMERICA. COLOMBIA: Magdalena: Rincon Hondo, Magdalena Valley, 10 Aug 1924, Allen 357 (MO). Vaupes: Rio Vaupes, Mitu y alrededores, 08 Sep 1951, Schultes & Cabrera 13977 (GH), Rio Vaupes, Mitu and vicinity, 09 May 1953, Schultes & Cabrera 19257 (GH-2 sheets). VENEZUELA: Bolivar: 27 km SW of Caicara along Hwy. 19 to Ciudad Bolivar, 22 Nov 1973, Davidse 4355 (MO). Tachira: between La Rochela and La Espuma, SW of Santo Domingo, 31 Jul 1979, Steyermark & Liesner 119299 (MO). Zulia: Perija, carretera Calle Larga-San Felipe-Jaguacita, km 25 al SE de San Felipe, 09 Oct 1977, Bunting 5656 (NY), ca 50 km SSW s pu by air, 19 km W of main road, 26 Mar 1982, Liesner & Gonzalez 13183 (NY). GUYANA: Upper Takutu-U Baboon Hill ee Tau) 1.5 km S of Sand Creek Village, 21 Jun 1989, Gillespie et al. 1803 (NY). ECUADOR ane Rios: p e San Juan, Vince, 07 Mar 1988, Laega 70652 (K, NY). BRAZIL: Mato Grosso do Sul: Pantanal do Miranda-Abobral, Passo do Lontra, Rodovia MS 122, e Sao Bento, depois da 2a porteira, 11 Jul 1997, Rodrigues et al. 3 (K). Rondonia: Guapore, Porto Velho, 1952, Cordeiro & Silva 270 (US). Amapa: Rio Macacoari, Municipio de Macapa, 05 Aug 1951, Froes & Black 27231 (US). Parana: Curitiba, Paso do Lontra O Miranda) Mato Grosso, 13 Oct 1972, Hatschbach & Scherer 30441 (NY, Z). Rio de Janeiro: Rio de Janeiro, Jan 1914, Hoel 3 ). Rio Grande do Sul: Jari, estrada do Caracuru, Jari, estrada do Caracuru, campo alagado, 09 Aug 1969, Silva 2636 (NY). Roraima: borrow pit close to road from Furo do Maraca to SEMA research station, 10 Mar 1987, Edwards 2529 (K). BOLIVIA: Beni: Ballivian, la zona de influ del rio Yacuma, 09 Mar 1980, Beck 3248 (NY), Ballivian, espiritu en la zona de influencia del rio Yacuma, 13 Apr 1980, Beck 3354 (NY). AFRICA. SOUTH AFRICA: KwaZulu-Natal: Hlabisa, St. Lucia, E shores, 30 Nov 1959, Feely & Ward 15 (K, M, NU-2 sheets]), Near Howick, 1990, Taylor 131 (NU), Ingivauvima distr., near Salumhlanga, Ndumu Game Reserve, 22 Dec 1972, Pooley 1624 (NU), Greater Durban area, Mlazi Valley, 15 Mar 1992, Ward 11925 (NU, PRE), Greater Durban area, Mlazi Valley, 15 Mar 1992, Ward 11926 (NU, NH photo), Transvaal, Waterberg, 13.2 m NW of Warmbaths, 19 Mar 1965, Acocks 23562 (K, PRE), near Maputa, D near Nyinyani, 29 Oct 1980, Cunningham s.n. (NU), Karkloof floodplain near junction with Kusane River, Jan 1977, Kotze s.n. (NU). TANZANIA: Singida Dist.: T. 5, M. 12.7 from Issuna e Singida-Manyoni Road, 4,800 ft, 13 Mar 1964, Greenway & Polhill 11543 mum T4, Sumbawanga Dist., goli Mbuga, 5 km S ga, 19 Jun 1996, Faden et al. 96/302 (US), T5, Manyoni Dist., Chaya Lake, S of Itigi-Tabora track, 16 km W of Kazikazi, 02 Jul 1996, Faden et al. 96/522 (US). ZIMBABWE: Gokwe, Sengwa Nature Resreve, Jan 1966, Jacobsen 73 (NU), District Gokwe, Sengwa research station, 09 May 1966, Jacobsen 3218 (PRE). Distribution.—In the U.S.A. known only from Lee County, Florida. In México known from the states of Campeche, Chiapas, and Guerrero, and in Central America known only from Panama. In the Caribbean Basin known only from Cuba and the Dominican Republic. Most widespread in South America with re- cords from Bolivia, Brazil, Colombia, Ecuador, Guyana, and Venezuela. In Africa known from South Africa, Tanzania, and Zimbabwe. Habitat.—Disturbed and natural freshwater wetlands including marshy open grasslands and tropical forests; reported from 0-1400 m Etymology.—The subspecific epithet is indicative of the short perianth bristles of this taxon. 3. Eleocharis acutangula (Roxb.) Schult. subsp. neotropica D.J. Rosen, subsp. nov. (Fig. 3 i-k). Tee: PERU. DEPARTAMENTO DE LoRETO: Maynas, Iquitos, prolongacion Yavari, Versailles-Paina, open annually burned grassland, 23 Mar 1974, McDaniel & Rimachi 18552 (HOLOTYPE: MO!; isotypes: IBE-2 sheets [photos!], NY! A Eleocharis acutangula (Roxb.) Schult. subsp. acutangula perianthii setis brevioribus mollibus flexuosis longissimis 1.8—2.4-plo lon- gitudo achenii, achenii apicibus constrictis infirme usque ad 0.6—0.7-plo latitudem achenii et stylopodiis plerumque latioribus quam dit. longioribus rece Plants perennial. Roots coarse, fibrous, drab-brown; primary rhizomes caudex-like, thick, hard, ascending, concealed by roots and persistent culm bases; secondary rhizomes elongated, to 3.2 mm thick, scales to 14 mm (few seen). Culms triquetrous, (44-)56-86-106) cm tall x (2—)2.7-3.9(-4.3) mm wide, soft, internally spongy, with incomplete transverse septa, smooth, green to drab gray-green and finely longitudinally stri- ate when dry. Leaves 2, reduced to sheaths, apically oblique, membranous, loose, friable, proximally dark maroon, distally drab, apex acute. Spikelets cylindric, (1.5—)2—3.1(23.5) cm long x (3823.2-4.2(-4.5) mm wide, acute; proximal scale with flower, obtuse, clasping, appearing as a continuation of culm; remaining floral scales conspicuously spirally arranged, appressed, ovate-widely ovate, (3.9-)4.1-4.8(25) mm long x (2.3-)2.4-3.2(-3.8) mm wide, cartilaginous, stramineous (faintly greenish centrally), adaxially sparsely red-maculate, apex acute (rounded), the distal 0.2-0.3 mm translucent hyaline-erose, central area nearly flat, abaxially coarsely many veined, the veins raised and visible at 20x, only mid-vein distinguishable in adaxial view. Flowers with (5-)6—7 perianth bristles, bristles sub-equal, (1.8—)1.9-2.3(-2.4) times achene Rosen et al., T f Eleochari t | 887 length, coarsely retrorsely spinulose nearly to base, stramineous, sometimes becoming reddish-brown distally; stamens 3; anthers 0.9-1.8(-2.3) mm long, stramineous; style trifid. Achenes biconvex, broadly obovoid, (1.3-)1.6-1.8 mm long x (1.2-)1.3-1.5(-1.6) mm wide, with 12-14-16) longitudinal rows of deeply concave transversely oblong polygonal cells visible through transparent periclinal layer on each achene face, dull to shiny yellow-green (sometimes tinged with amber), apex constricted to a distinct neck about 0.6—0.7 times achene width. Tubercle dorsoventrally compressed, wider than tall and appearing very shallowly to shallowly triangular, sometimes apex appearing truncate or retuse, (0.3-)0.4-0.5(-0.6) mm long x (0.620.8-1C-1.1) mm wide, light-dark brown. Specimen mined: SOUTH AMERICA. ECUADOR: Pastaza: Villano, Compamento Base de Arco, Pantano al noreste de la pista, jo 29'S, 77? 27’ W, Feb 1994, Palacios 12171 (CIIDIR, MO). Napo: Archidona, Reserva Ecologica Antisana, Comunidad Shamato, En- trada por km 21, Shamato, 00? 43'S, 077? 49'W, 24 Apr 1998, Clark et al. 5122 (MO). PERU: Amazonas: Bagua, along roadside from Chiriaco to Puente Venezuela (3.9 Km NE Chiriaco), elev 600—800 ft, 31 Oct1978, Barbour 4355 (F, IBE-photo, MO). Cusco: Paucar Tawbo, Montaueza Choutachaca, 780-1000 m, 28 Nov 1965, Vargas 16887 (US). Loreto: Puerto Almendras on the Rio Nanay, 30 km N of Iquitos, 600 m, 16 Aug 1981, Moore & Ruiz 114 (F); Prov. E ae de 80 cm, 03 Aug 1967, Torres 340 (GH-2 sheets); Prov. Maynas, Iquitos, Carretera de Zungaro Cocha, cerca a la q 12 Aug 1983, Ri I 08 (IBE-photo, VSC); Maynas, Dtto. Punchana, Rio 22 varadera de; caserio de Padre Cocha, 07 Jul 1994, Rimachi 11004 (IBE—photo, MICH-2 sheets, NY, VSC); Maynas, Inmedia de la Guarnicion militar de Gueppi, sobre la margen izquierda del Rio Putumayo, borde con Ecuador, 26 May 1978, Diaz 368 (F, MO); Maynas, Distrito Iquitos, Caserio Nina Rumy, Rio Nanay, 73° 25’ W, 03° 48’ S, 22 Apr 1988, Ruiz 1262 (MO); Iquitos Region, 26 Jul 1966, Martin & Lau-Cam 1164 (GH). Huanuco: Leoncio Prado, 3 km SE of Pucayacu, on road from Tingo Maria to Tocache Nuevo, ca 75 km NW of Tingo Maria, 10 Dec 1981, Plowman & Rury 11288A (MO, NY). BOLIVIA: Pando: Provincia Nicolas Suarez, Cobija 2 km hacia el Sur, 19 Oct 1988, Beck 17139 (K, US). Distribution.—Known only from northwest South America from Ecuador, Bolivia, and Peru. Habitat —Specimens examined are from various freshwater wetlands including marshy open grasslands, tropical forests, and roadside ditches, reportedly from 0-1000 m. Etymology.—The specific epithet indicates the decidedly neotropical distribution of this taxon. ACKNOWLEDGMENTS We are grateful to the staff of all of the herbaria cited for loan of specimens for study, assistance with questions about the disposition of type specimens and literature, and assistance with nomenclatural questions. The first author is grateful to Socorro González-Elizondo for sharing her expertise and for facilitating field work in Mexico and Eric Roalson for bringing certain literature items to his attention. The helpful comments of J. Wipff and two anonymous reviewers are gratefully acknowledged. We offer special thanks to Neva Mikulicz for preparing the excellent illustration. Financial support for the first author was provided by the Frank W. Gould Award for Graduate Student Research in Plant Systematics and the Dr. Harry Wayne Springfield Graduate Student Endowment. Financial support for field work in Florida by R. Carter was provided by the Valdosta State University Foundation. REFERENCES BAKSH, S.I. and J.H. RicHaros. 2006. An architec | Jel for Eleochari hol | develo tof Eleocharis cellulosa (Cyperaceae). Amer. J. Bot. 93:707-715. Blake, S.T. 1939. A monograph of the genus Eleocharis in Australia and New Zealand. Proc. Roy. Soc. Queensland 50:88-132 BRowNING, J., K.D. GorDON-Gray, and C.J. Warp. 1997. Studies in Cyperaceae in Southern Africa 32: Eleocharis sub- genus Limnochloa section Limnochloa. S. African J. Bot. 63:172-184. CHERMEZON, H. 1936. Contribution à la flore cypérologique du Sénégal: Cypéracées récoltées par M. Trochain. Arch. Bot. Mem. 7:1-32. EDWARDS. A.L., D.W. Lee, and J.H. RicHArps. 2003. Responses to a fluctuating environment: effects of water depth on growth and biomass allocation in Eleocharis cellulosa Torr. (Cyperaceae). Canad. J. Bot. 81:964-975 Forman, L.L. 1997. Notes concerning the typification of names of William Roxburgh's species of phanerogams. Kew Bulletin 52:523-534. 888 Journal of the Botanical R h Institute of Texas 1(2) ELizonDO, M.S. and A.A. ReznicEk. 1996. New Eleocharis (Cyperaceae) from Venezuela. Novon 6:356-365. (SC WIÁLE GONZALEZ-ELIZONDO, M.S. and P.M. Peterson. 1997. A classification of and key to the supraspecific taxa in Eleocharis (Cyperaceae). Taxon 46:433-449. GONZALEZ-ELIZONDO, M.S. and J.A. Tena-FLORES. 2000. Eleocharis (Cyperaceae) in the New World. In: K. L. Wilson & D. A. Morrison (eds.), Monocots: systematics and evolution. CSIRO Publishing, Australia. Pp. 637-643. GORDON-GRAY, K.D. 1995. Cyperaceae in Natal. Strelitia 2. National Botanical Institute, Pretoria, South Africa. GREGOR, T. 2003. Eleocharis mamillata - Distribution and infraspecific differentiation. Folia Geobot. 38:49-64. Haines, RW, and K.A. Lye. 1983. The sedges and rushes of East Africa. East African Natural History Society. Nairobi. Hess, H. 1953. Uber die Gattungen Heleocharis und Carex aus Angola und dem unteren Kongo. Ber. Schweiz. Bot. Ges. 63:317-359. Hess, H. 1957. Beiträge zur Kenntnis der Gattung Heleocharis in Brasilien. Ber. Schweiz. Bot. Ges. 67:91-94. Hines, D.M. 1975. A monograph of the Eleocharis ovata complex (Cyperaceae) in North America. Ph.D. Disserta- tion. University of Michigan, Ann Arbor. HoLmMGREN, P.K., N.H. HoLMGREN, and L.C. Barnetr. 1990. Index herbariorum Part I: The herbaria of the world. New York Botanical Garden. New York. JØRGENSEN, P.M. and S. Leon-YANez. 1999. Catalogue of the vascular plants of Ecuador. Missouri Botanical Garden Press, St. Louis. Koyama, T. 1985. Cyperaceae. In: M.D. Dassanayake & F.R. Fosberg, eds. Flora of Ceylon, Vol. 5., pp. 125-405. Am- erind Publishing Co. Pvt. Ltd., New Delhi. Larson, B.M.H. and P.M. Catuna. 1996. The separation of Eleocharis obtusa and Eleocharis ovata (Cyperaceae) in Eastern Canada. Canad. J. Bot. 74:238-242. McNelt, J., F.R. Barrie, H.M. Bunper, V. DeuouuN, D.L. HAwkswonrH, K. MARHOLD, D.H. NICOLSON, J. PRADO, P.C. Sit va, J.E. SKOG, J.H. Wiersema, and N.J. TunLAND. 2006. International code of botanical nomenclature (Vienna Code). Regnum Veg. 146:1-568. Menapace, F.J. 1991. A revision of the genus Eleocharis (Cyperaceae) section Maculosae. Ph.D. Dissertation. Southern Illinois University, Carbondale. Nees VON Esenseck, C.G.D. 1842. Cyperographia sive descriptio in Brasilia provenientium In: K. F. P. von Martius. Flora Brasiliensis, Vol. Il, Part |. Pp. 1-226. RoALSON, E.H. 1999. Eleocharis yecorensis (Cyperaceae), a new species of spike-sedge from Mexico. Aliso 18:57-60. Rontr, F.J. 2000. NTSYS-pc: numerical taxonomy and multivariate analysis system, version 2.1. Exeter Software: Setauket, NY Rosen, D.J. 2006. A systematic study of select species complexes of Eleocharis subgenus Limnochloa (Cyperaceae). Ph.D. Dissertation. Texas A&M University, College Station. Rosen, DJ. and S.L. HarcH. (in press). A new species of Eleocharis suba. Limnochloa (Cyperaceae) from Bolivia. Brittonia. RoxBURcH, W. 1820. Flora Indica or descriptions of Indian plants. Serampore. SANJAPPA, M., K. THOTHATHRI, and A.R. Das. 1991. Roxburghs flora Indica drawings at Calcuta. Bull. Bot. Surv. India 331-232 Simpson, D.A. and C.A. INaus. 2001. Cyperaceae of economic, ethnobotanical and horticultural importance: a checklist. Kew Bull. 56:257-360. Stewart, A. 1911. A botanical survey of the Galápagos Islands. Proc. Calif. Acad. Sci. IV, 1:7-288. Stuessy, T.F. 1990. Plant taxonomy; the systematic evaluation of comparative data. Columbia University Press, NY. Svenson, H.K. 1929. Monographic studies in the genus Eleocharis |. Rhodora 31:121-135, 167-191. Svenson, H.K. 1939. Monographic studies in the genus Eleocharis V. Rhodora 41:1-19, 90-110. Trevisan, R. and 1.1. BoLorin1. 2006. A new species of Eleocharis R. Brown (Cyperaceae) from Southern Brazil. Novon 16:155-157. FOURTEEN NEW COMBINATIONS IN SEDUM (CRASSULACEAE) Hideaki Ohba Department of Botany Hongo 7-3-1, Tokyo 113-0033, JAPAN ohbagum.u-tokyo.ac.jp ABSTRACT hi : ; lin Sedum L. for tl f, 1 i f+} in Flora of North A th of M. s. Seins S. Watson var. ciliosuni (Howell) H. Ohba, Sradiatum var. denau nessun (R.T. Clausen) H. Ohba, S. wrightii A. Gray var. priscum (R.T. Clausen) H. Ohba, S a var. dencia: rum (R.T. Clausen) H. Ohba, S. obtusatum A. Gray var. boreale RI Clausen) H. Ohba, S. obtusatum var. paradisum (Denton) H. Ohba, S. obtusatum var. retusum (Rose) H. Ohba, S A. Berger var. eastwoodiae (Britton) H. Ohba, S. laxum var. flavidum (Denton) H. Ohba, S. laxum var. heckneri (M. Peck) H. Ohba, S. laxum var. latifolium (R.T. Clausen) H. Ohba, S. stenopetalum Pursh var. monanthum (Suksd.) H. Ohba, S. lanceolatum Torr. var. subalpinum (Fröd.) H. Ohba, and S. oreganum Nutt. var. tenue (R.T. Clausen) H. Ohba RESUMEN Se necesitan catorce combinaciones nuevas en Sedum L. en el tratamiento del género para la Flora of North America north of Mexico: S. radiatum S. Watson var. ciliosum (Howell) H. Ohba, S. radiatum var. depauperatum (R.T. Clausen) H. Ohba, S. wrightii A. Gray var. priscum (R.T. Clausen) H. Ohba, S. wrightii var. densiflorum (R.T. Clausen) H. Ohba, S. obtusatum A. Gray var. boreale RI Clausen) H. Ohba, S. obtusatum var. paradisum (Denton) H. Ohba, S. obtusatum var. retusum (Rose) H. Ohba, S A. Berger var. eastwoodiae (Britton) H. Ohba, S. laxum var. flavidum (Denton) H. Ohba, S. laxum var. heckneri (M. Peck) H. Ohba, S. laxum var. latifolium (R.T. Clausen) H. Ohba, S. stenopetalum Pursh var. monanthum (Suksd.) H. Ohba, S. lanceolatum Torr. var. subalpinum (Fröd.) H. Ohba, y S. oreganum Nutt. var. tenue (R.T. Clausen) H. Ohba. The following new combinations are required for the forthcoming treatment of Sedum (Crassulaceae) in volume 8 of Flora of North America north of Mexico. Sedum radiatum S. Watson var. ciliosum (Howell) H. Ohba, comb. & stat. nov. Basionyu: Sedum ciliosum Howell, Fl. N.W. Amer. 214. 1898 Sedum radiatum S. Watson var. depauperatum (R.T. Clausen) H. vue comb. & stat. nov. Basionym: Sedum radiatum subsp. depauperatum R.T. Clausen, Sedum N. Amer. 258, figs. 59-60. Sedum wrightii A. Gray var. priscum (R.T. Clausen) H. Ohba, comb. & stat. nov. Basioxvw: Sedum wrightii subsp. priscum R.T. Clausen, Bull. Torrey Bot. Club 106:215. 1979 Sedum wrightii A. Gray var. densiflorum (R.T. Clausen) H. Ohba, comb. & stat. nov. Basionym: Sedum wrightii subsp. densiflorum R.T. Clausen, Variat. Sp. Sedum Mex. Cordill. Plateau 13. 1981. Sedum obtusatum A. Gray var. boreale (R.T. Clausen) H. Ohba, comb. & stat. nov. Basowvw: Sedum obtusatum subsp. boreale R.T. Clausen, Bull. Torrey Bot. Club 69:32. 1942 Sedum obtusatum A. Gray var. paradisum (Denton) H. Ohba, comb. & stat. nov. Basionyu: Sedum obtusatum subsp. paradisum Denton, Brittonia 30:256, fig. 2. 1978. Sedum obtusatum A. Gray var. retusum (Rose) H. Ohba, comb. & stat. nov. Basionvw: Gormania retusa Rose in Britton & Rose, New N. Amer. Crassul. 31. 1903 Sedum laxum (Britton) A. Berger var. eastwoodiae (Britton) H. Ohba, comb. & stat. nov. Basioxvm: Gormania eastwoodiae Britton in Britton & Rose, New N. Amer. Crassul. 903. Sedum laxum (Britton) A. Berger var. flavidum (Denton) H. Ohba, comb. & stat. nov. Basionym: Sedum laxum subsp. flavidum Denton, Brittonia 30:233, fig. 1. 1978 Sedum laxum (Britton) A. Berger var. heckneri (M. Peck) H. Ohba, comb. & stat. nov. Basionvm: Sedum heckneri M. Peck, Proc. Biol. Soc. Wash. 50:121. 1937 J. Bot. Res. Inst. Texas 1(2): 889 — 890. 2007 890 Journal of the Botanical R h Institute of Texas 1(2) Sedum laxum (Britton) A. Berger var. latifolium (R.T. Clausen) H. Ohba, comb. é stat. nov. Basionvm: Sedum laxum subsp. latifolium R.T. Clausen, Bull. Torrey Bot. Club 69:38. 1942. Sedum stenopetalum Pursh var. monanthum (Suksd.) H. Ohba, comb. & stat. nov. Basionym: Sedum monanthum Suksd., Werdenda 1:19. 1927, based on Sedum uniflorum Howell, Fl. N.W. Amer. 213. 1898 not Sedum uniflorum Hook. & Arn Bot. Beechey Voy. 263. 1838. Sedum lanceolatum Torr. var. subalpinum (Fród.) H. Ohba, comb. & stat. nov. Basionvm: Sedum stenopetalum Pursh var. subalpinum Fród., Acta Horti Gothob. 10(App.):70, figs. 521-527, pl. 43, fig. 2. 1935 [1936] Sedum oreganum Nutt. var. tenue (R.T. Clausen) H. Ohba, comb. & stat. nov. Basionvu: Sedum oreganum subsp. tenue R.T. Clausen, Sedum N. Amer. 355, fig. 91. 1975. ACKNOWLEDGMENTS Ithank James Zarucchi, Craig Freeman, and Nancy Morin (Flora of North America) for assistance with get- ting these new combinations published. This study was supported by a Grants-in-aid for Scientific Research from the Japan Society for Promotion of Science (Nos. 17570073 and 18255004). A NEW GYPSOPHILOUS SPECIES OF ERIGERON (ASTERACEAE: ASTEREAE) FROM NORTHEASTERN MEXICO Guy L. Nesom Botanical Research Institute of Texas 509 Pecan Street Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT Erigeron heleniae Nesom, sp. nov., is described from a gypsum outcrop northwest of Monterrey in Nuevo Leon, Mexico. It is char- acterized by its diminutive, suffruticose habit, thickened taproot, hirsute-glandular vestiture, linear and subclasping leaves, and tiny, solitary heads with white, filiform rays RESUMEN Exigeron ADR. Nesom, T nov., se describe de un o das en el noroeste » MO en Nuevo León, México. Se I raiz engrosada, i glandular, hoj subapretadas, y cabezuelas ST aoa Js ; x 121 PL Gypsum outcrops in northeastern Mexico have fostered the evolution of many endemic species. Yet another is described here, known only from a single collection by George S. Hinton, whose botanical acuity and persistence (along with that of his father, Jaime Hinton) have greatly expanded knowledge of the region's plants. Erigeron heleniae Nesom, sp. nov. (Figs. 1-2). Tree: MEXICO. Nuevo Leon: Mpio. Mina: N of Los Molina, 2690409.23" N, 100?4427.57" W, gypsum hillside, [open gypsum exposures, flats and gentle slopes, with scattered ocotillo and other low perenni- als], 737 m elev, 4 Sep 2005, George S. Hinton 28412 (HoLotypE: TEX). Erigeronti gypsovero N ilis sed diff li lari li itulis minoril llis radii brevioribus filiformibus, foliis sub- amplectentibus, vestimento hirsuto-glanduloso, et pappo externo squamarum prominentium. mE MT Suffruticose[ athickened, woody taproot, individual plants up to 25 cm wide; stems intricately branched proximally, 5-8 cm high, hirsute to hirsute-villous, hairs strongly deflexed on proximal third, spreading above; leaves, and phyllaries hirsute with white, erect-arching hairs 0.4-1 mm long, hairs arising from thickened, orange-resinous basal cells, minutely stipitate-glandular, the phyllaries densely so. Leaves linear to linear-oblanceolate, flat, 7-10(-15) mm long, 0.8-1 mm wide, even-sized to immediately below the heads, slightly but distinctly auriculate and subclasping. Heads solitary on bracteate peduncles 3-7 mm long; involucres turbinate-campanulate, 3.5-5 mm wide; phyllaries in 3(-4) subequal series, elliptic-lanceolate, 2.53 mm long, with a dark orange-resinous midvein, densely minutely glandular. Ray florets 22-30, white or slightly pink-tinged, 3-3.5 mm long, ligules 0.2-0.4 mm wide, apparently not reflexing or coiling. Disc florets 1-1.6 mm long. Achenes obovate-oblong, 0.7 mm long, 2-nerved, sparsely strigose; pappus of 10-12 persistent bristles 0.8-1 mm long, outer pappus of prominent scales 0.1 mm long. Etymology.—Erigeron heleniae is named for Helen Hinton, mother of George S. Hinton and wife of Jaime Hinton. She has been close witness to the excitement of discovery by her family of so many of the region's beautiful and unusual species. The new species occurs on gypsum outcrops about 55 kilometers northwest of the city of Monterrey. There are two large gypsum outcrops in the immediate area, about 5.5 kilometers apart and covering a total of about 3.5 hectares. Essentially the same set of gypsophilic species occurs on both outcrops; Erigeron heleniae is common on both and prefers slopes to flat areas. As with many other gypsophilic endemics from this region of Mexico, this area of outcrops probably is the total extent of the range of the new species. Erigeron heleniae is similar in its diminutive habit and narrow leaves to E. gypsoverus Nesom, which is J Bot. Res Inst Texas 1(2): 201 — 894. 2007 892 A A Journal of p MOST A \ SAN \ N TN N) | ESA Xx (Sd A \ (t N NAAA j N 4 1 l \ M A Y Nesom, À new gypsophilous species of Erigeron 893 894 Journal of the Botanical R h Institut f Texas 1(2) relatively common in the open pine woods in the gypseous region around Galeana, Nuevo León, and known from numerous collections. Erigeron heleniae occurs about 140 kilometers north of the closest population of E. gypsoverus and at a lower elevation. Both species apparently are obligate gypsophiles, but the two probably are not even closely related within the genus. Contrasts below distinguish them. 1. Stems, leaves, and phyllaries closely strigillose, a taproot slender; proximal branches elongate, stolon-like; leaves not at all basally auriculate or claspir ES involucres 5-7 mm wide; ray florets 5-6 mm outer ofin nconspici Ious setae; 1050- n m ele = = Erigeron gypsoverus . Stems, pum and phyllaries hirsute, minutely m CERTES taproot thickened; proximal branches intricately branched; leaves slightly but distinctly auriculate and subclasping; involucres 3.5-5 mm wide; ra florets 3.5-5 mm long; outer pappus of prominent scales; ca. 750 m elev. ray rigeron heleniae Morphological features shared by Erigeron heleniae and E. dryophyllus A. Gray (of central Nuevo León and adjacent Tamaulipas) suggest that these two species may be closely related: subclasping leaves and hispid- villous vestiture, the hairs sharply deflexed on proximal portions of the stem. Erigeron dryophyllus differs most conspicuously in its rhizomatous habit, broad and lobed leaves, much larger heads on long peduncles, and habitats primarily over limestone. e ACKNOWLEDGMENTS George Hinton provided details regarding the locality and habitat of the new species as well as a review of the manuscript. Tim Lowrey also reviewed the manuscript. The illustration is by Linny Heagy. FOLIA TAXONOMICA 3. PASSIFLORA DAVIDII (PASSIFLORACEAE), ANEW SPECIES IN SUBGENUS PASSIFLORAANDA KEY TO THE SECTIONS OF SUPERSECTION SHIPULATA Christian Feuillet Department of Botany, MRC-166 Smiths sonian Institution PO. Box 3712 Washington, D.C. 20013-7012, U.S.A. feuillec@si.edu ABSTRACT E The first key t Stipulata Feuillet =e M. dida is Phone Dt davidii Pn nov. is described from French Guiana. Its foliaceous sup and b I I g I Stipulata section Granadillastrum Triana & Planch. RÉSUMÉ La premiere clé d de la supersection Stipulata Feuillet Bu M. cud UE est fournie. rod aii iu nov. est décrite de Guyane francaise. En raison d tipul 1 | p le sous-genre Passifl I ion Stipulat ti dillastrum Triana & Planch. In the new classification of Passiflora (Feuillet & MacDougal 2003; MacDougal & Feuillet 2004) replacing the system proposed by Killip (1938), Passiflora subg. Passiflora supersect. Stipulata Feuillet & J.M. MacDou- gal is a large coherent group which includes more than 95 species sharing foliaceous, usually asymmetric stipules. KEY TO THE SECTIONS OF SUPERSECTION STIPULATA . Flowers in pseudoracemose inflorescences, 25-75 cm long. (P racemosa Brot). sect. Calopathanthus Harms : Flowers solitary or in pair in leaf axils. 2. Bracts setaceous or narrowly ovate, less than 2.5 mm wide, scattered along the pedicel. (4 species) ermesinae (Cervi) Feuillet & J.M. MacDougal 2. Bracts foliaceous, usually more than 2.5 mm wide (but often deeply dissected in sect. Dysosmia), usually verticillate, sometimes scattered or loosely grouped 3. Stipules and bracts lacerate-dentate or laciniate to 1-4-pinnatisect. (20 sp.) sect. Dysosmia DC. 3. Stipules and bracts entire to serrate. 4. Hypanthium cylindric, as long as the sepals. (4 sp.) sect. Tacsonioides DC. 4. Hypanthium campanulate or rarely short-tubular and then shorter than the sepals. (> 66 sp.) sect. Granadillastrum Triana & Planch. The members of Passiflora sect. Granadillastrum are herbaceous or woody climbers, with leaves petiolate and with two or more glands on the petiole; the laminas are unlobed or lobed; the venation palmate or sometimes pinnate with 1—2 pairs of minor submarginal veins at base. Their flowers are solitary or rarely two in the leaf axils. The hypanthium is campanulate or rarely short-tubular. Triana and Planchon (1873) described Passiflora section Granadillastrum for one species, P. semiciliosa Planch. & Linden published in the same paper. In Killip (1938), Granadillastrum was raised to the rank of subgenus and some species were added that all belong now in supersect. Tacsonia. In addition to P. semiciliosa, we place in section Granadil- lastrum species formerly placed in subgenus Passiflora (subg. Granadilla of Killip) ser. Kermesinae Killip ex Cervi (in part), Imbricatae Killip ex Cervi, Simplicifoliae (Harms) Killip ex Cervi, Lobatae Killip ex Cervi, and Menispermifoliae Killip ex Cervi. Those series were not described by Killip. His key to the series suggested a strong emphasis placed on leaf shape, separating species with lobed leaves and those with unlobed leaves J Bot. Res Inst Texas 1(2): 205 — 898. 2007 896 Journal of the Botanical R h Institute of Texas 1(2) into informal series. Since then the series were validated by Cervi (1997), but several species proved to be variable in leaf shape. Although the species in this group should be separated in subcategories, at this point itis premature to propose formal series. It is in 1987 that I saw the new species, P. davidii, for the first time. The young plant had two stems climbing on a rope in the camp at Montagne des Nouragues in French Guiana (Feuillet 4395). Its stipules with an egg-mimic yellow swelling at the apex and the unlobed leaves with several yellow swellings on the margin set it apart from the other species of subgenus Passiflora. One stem had young flower buds, the other was sterile. I saw this species again twice, in French Guiana, with David Rignon on Montagnes Tortue and near Cacao, but without flower or fruit. No specimens were collected. After nearly 20 years, David collected it in bloom in February 2006, confirming it was a new species and providing enough information for a formal description. Passiflora davidii Feuillet, sp. nov. (Fig. 1). Tv: FRENCH GUIANA: cultivated 1 km from the village of Cacao, along the road to Cayenne, 0-50 m, 52°25'W, 4°33'N, 23 Feb 2006, D. Rignon 16 (HoLoTYPE: US; isotypes: CAY, P). 4n Passiflora davidii i be. Passifl Stipulata sect. Granadillastro per H TESTIS EDS 1 ex tinens. Passiflorae actiniae Hook. affinis; stipulis cum mu- LE Al 1 1 PE E 1 $ 5 E I ZI ec pcc [e] jT o Į E dictincta. A specieb i il ipulis foliacei j d pice inflatum differt. S F de Vine, size unknown, climbing with tendrils, short-pubescent throughout except where noted; stems terete, in some places striate. Stipules foliaceous, 1.8-2 x 1 cm, half-cordate at base, rounded and short mucronate at apex, the awn or mucro 1.5-2 mm long, blunt, with the apical half to 2/3 swollen and yellow, margin entire, paler abaxially when fresh, midrib slightly to clearly excentric, but straight, other veins reticulate. Leaves: petiole 3.8-4.2 cm long, 6-9-glandular; glands sessile, swollen, orange-yellow, spread along the petiole with 2 in subapical position; blade simple, unlobed, ovate, 10 x 6.5-7 cm, widely rounded, slightly peltate at base, acute to right angled at apex, margin entire, briefly recurved, narrowly glandular with scat- tered yellow small swellings, glabrous adaxially, paler abaxially when fresh, venation palmate at the very base with 5-7 veins, forming a submarginal wavy vein, apical half pinnate, with 6-7 main lateral veins ascending and toward the margin also included in the formation of a submarginal vein, minor lateral and most secondary veins more or less perpendicular to the midrib, tertiary veins reticulate. Flowers axillary, one per leaf axil, pedicel 272.8 cm long below the bracts, stipe 5-6 mm between the bracts and the flower; bracts 3, one larger, verticillate, green, lanceolate, 2-3 x 1-1.3 cm, rounded, slightly convex even around the bud, briefly cuneate at base, acute to narrowly-rounded at apex with a short swollen mucro less than 0.01 x 0.01 cm, paler abaxially when fresh; hypanthium campanulate, about 0.3 x 0.4 cm deep, sepals white and glabrous adaxially, yellow-green abaxially in parts exposed in bud, narrowly oblong, 3 x 1.5 cm, with apical awn 1-2 mm long, petals white, narrowly oblong, 3.5 x 1-1.3 cm, glabrous, corona with micro-trichomes on the outer filaments, outer filaments spreading to cover the perianth, 2-3.5 cm long in the same flower, white in their proximal half, blue violet in their apical half, thin, numerous, in 2-3 not clearly separated rows, inner filaments 0.3-0.4 mm, white, about as thick as the previous ones, in several rows, operculum membranous, limen membranous, androgynophore 1.2 cm long, glabrous, stamens filaments 0.8-1.0 cm long, flat, green speckled with red, glabrous, anthers 0.7-0.8 x 0.25-0.30 cm, yellow, glabrous, gynophore ca. 0.1 cm long, ovary obovoid, 0.4 x 0.25-0.27 cm, styles green speckled with red, glabrous, wider toward apex, stigmas hemispherical, pale yellow, glabrous. Fruit unknown. The description is based on the holotype that has two nodes, each with a leaf and a flower. Therefore the variation in measurement is the variation between 2 consecutive nodes. The sterile specimen (Feuillet 4395) consists in the apex of a stem actively growing, the stipules 1.5-2.5 x 0.5-1 cm are already fully de- veloped, some are even longer than on the holotype, but the petioles 0.4—1.5 cm, and the leaf blades 4.5-7 x 2.5—4 cm, are not fully grown. The early development of the stipules, displaying the egg-mimic mucros, apparently acts as a deterrent for the females of the solitary egg-laying species of heliconiid butterflies, and protects the apical meristem from grazing by their caterpillars. In the key by Killip (1938), Passiflora davidii would be close to P. actinia, a species from southeastern Feuillet, Passiflora davidii, ies in sul Passifl 897 Fic. 1. Passiflora davidii Feuillet. Top: Rignon 16 (photo M.-F. Prévost), Lower left: Feuillet 4395 (photo C. Feuillet). Brazil. It can be distinguished from P. actinia by the short pu- bescence on all vegetative parts and the stipules only slightly asymmetric with a mucro that is swollen, yellow, and act as an egg-mimic. The specimen looks like P. loretensis in general ap- pearance, but can be recognized by the 6—9 glandular petiole. Contrary to P. actinia and P. loretensis, P. davidii has a pubescent ovary. The flower is very similar to that of P. retipetala, that, in les with a longer contrast with P. davidii, is glabrous and has stipu awn. In the Guianas, P. davidii can be separated from the other members of supersection Stipulata with the characters used in the key below. The epithet of the new species honors David Rignon who collected the type specimen. He lives in French Guiana near the village of Cacao and is interested in passionflowers and butter- flies. David and his family were great field companions in 2003, when we saw P. davidii without flowers near the type locality. lairiére du C lesN 52°42'W, 4°03'N, 100-120 a qu SES o m, 28 Aug 1987, st., Feuillet 4395 (CAY). 898 Journal of the Botanical R IRI f Texas 1(2) KEY TO THE SPECIES OF SUPERSECTION STIPULATA IN THE GUIANAS & VENEZUELAN GUAYANA 1. Stipules laciniate, cleft to base, not deciduous. (sect. Dysosmia). 2. Ovary and fruit glabrous. 3. Stem glabrous; fruit red etida var. orinocensis (Killip) Feuillet 3. Stem hispid; fruit yellow orange P. foetida var. hispida (DC. ex Triana & Planch.) Killip 2. Ovary and fruit indumentum scarce to dense. 4. Stem stiffly hirsute; ovary and fruit hirsute 4. Stem softly pilosulous 5. Bracts 2-3-pinnatisect, 2-3 cm long 5. Bracts 1(-2)-pinnatisect, less than 2 cm long 1. Stipules entire or serrulate. (sect. Granadillastrum). 6. Stipules acute at apex 6. Stipules obtuse to rounded at a y: T with an apical awn or arista more than 5 mm long. 8. Stipule awn swollen at tip 8. Stipule arista not swollen at ti 7. Stipules with an apical arista or mucro less than 5 mm long. 9. Leaf blade reddish beneath; bracts petiolate 9. Leaf blade pale green or glaucous beneath; y bracts sessile 10. Leaves 3-lobed, red- glandular at ma P. garckei Mast. 10. Leaves unlobed, margin glandular yellow swellings P. davidii Feuillet P. foetida L. var. foetida P. foetida var. dip in (Ham.) Mast. P. moritziana Planch. P. exura Feuillet P. retipetala Mast. P. stipulata Aubl. P. picturata Ker ACKNOWLEDGMENTS I am indebted to my friend and colleague Fanchon Prévost (CAY) for processing the plant collected by David Rignon into herbarium specimens and for the detailed labels and great photographs. I would like to thank the curators and directors of the herbaria CAY, NY, P, U, US whose collections have been made available for my work when writing this paper. This is number 128 in the Smithsonian's Biological Diversity of the Guiana Shield Program publication series. John MacDougal and Peter M. Jorgensen provided critical reviews for which Iam most appreciative. REFERENCES Cervi, A.C. 1997. Passifloraceae do Brasil. Estudo do género Passiflora L., subgénero Passiflora. Fontqueria FeuiLLET, C. and J.M. MacDoucaL. 2003 [May 2004]. A new infrageneric classification of Passiflora L. (Passifloraceae). Passiflora 13(2):34-38. Kup, E.P. 1938. The American species of Passifloraceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 19:1—613. MacDouca,, J.M. and C. FeuiLLET. 2004. 2. Systematics. In: T. Ulmer and J.M. MacDougal, eds. Passiflora, Passionflow- ers of the world. Timber Press, Portland. Pp. 27-31. TRIANA, J.J. and J.E. PLANCHON. 1873. Prodromus florae Novo-Granatensis. XIX Passifloreae. Ann. Sci. Nat., Bot., sér 5, 17:121-186. — FOUR NEW COMBINATIONS IN MOSIERA (MYRTACEAE) FROM THE CARIBBEAN Andrew Salywon Desert Botanical Garden 1201 N. Galvin Parkway Phoenix, Arizona 85008-3437, U.S.A. asalywon@dbg.org ABSTRACT FM L New combinations for four C ly assigned to Eugenia L. or Psidium L. are proposed: Mosiera andro- i d for Mocier d i siana, M. cuspidata, M. gracilipes, and M xerophytica. A | de RESUMEN ee : : de TENA i i d Eugenia | o Psidium L.: Mosiera a M. cuspidata, M. ales yM. xerophytica. Se PE un lectotipo para Moe androsiana. Recent taxonomic and molecular studies of Mosiera (Landrum 1992; Salywon 2003; Salywon et al. 2004; Salywon & Landrum in press) have clarified the boundaries of the genus. However there are no morpho- logical synapomorphic characters that differentiate Mosiera from related genera. It is therefore separated from other genera by a combination of characters including: tetramerous flowers, lustrous or glandular seed coats one to six cells thick, and C-shaped embryos with cotyledons that are ca. 4 the length and about the same width as the hypocotyls. Landrum and Kawasaki (1997) and Salywon and Landrum (in press) provide keys to distinguish Mosiera from similar genera. The genus contains approximately 18 species distributed mainly in the Caribbean, with two species in Mexico and one of these in Guatemala. New combinations are required for four Caribbean species previously placed in Eugenia or Psidium that can be assigned to Mosiera, in order to make the names available for a manuscript in preparation on the molecular systematics of the family (Salywon et al. in prep.). Mosiera androsiana (Urban) Salywon, comb. nov. Basiosvw: Eugeni iana Urban, Fedde, Rep. Sp. Nov. 13:467. 1915. Psidium androsianum (Urban) Correll, J. Arnold Arbor. 58:41. 1977. Tree: BAHAMAS: Andros, Mangrove Cay, coppice, near Lisbon Creek, 16-19 Jan 1910, Small & Carter 8496 (LeEcTOTYPE, selected here: NY [vascular plant type image library # 84,503]; IsoLECTOTYPES: E US [2 sheets]). The holotype in Berlin is lost, and the NY specimen is here chosen as the lectotype. Mosiera androsiana is morphologically most similar to M. longipes (O. Berg) Small but differs in having leaves 0.5-2.1 x 0.15-1 cm (vs. 1.1-5.2 x 0.2-3.8 cm) with the secondary venation not visible (vs. secondary venation visible), and seeds ca. 3 mm long (vs. ca. 2 mm long). The leaf morphology of this species is among the most variable in the genus. Mosiera cuspidata (Alain) Salywon, comb. nov. B Psidi pidatum Alain, Brittonia 20:159. 1968. Tyee: HISPANOLA [DOMINICAN REPUBLIC]: Boca del Infierno, Los Haitises. Samaná Prov., on limestone cliffs, near the edge of water, 24 Jun 1930, Ekman H. 15427 (HOLOTYPE: US; isotypes: MICH, S) Mosiera cuspidata is a shrub or small tree to 4 m tall with distinctively large leaves for the genus. It is ecologically interesting because it grows on vertical limestone cliffs and on small emergent limestone outcrops in the ocean with little or no soil. Mosiera depu id (Alain) Salywon, comb. nov. B Psidi ilipes Alain, Phytologia 25:269. 1973. Tye: DOMINICAN BLIC. [Cordillera Central]: [Prov. La Vega, Municipio de eee), T Redonda, Ciénaga de la Culata, Constanza, in cloud AA 1700-2000 m, 30 Nov 1969, Alain 17138 (HoLotYPE: NY; isotypes: JBSD, US). J. Bot. Res. Inst. Texas 1(2): 899 — 900. 2007 900 Journal of the Botanical R h Institute of Texas 1(2) This species is known only from three collections, all from the same locale, and may be of conservation concern as clearcutting for farmland and wood has deeply encroached into the cloud forest where this species grows. Additional fieldwork is needed to determine if it is distributed on other mountain tops nearby. Mosiera xerophytica (Britton) Salywon, comb. nov. Basionw: Ms ee ee Britton, Bull. Torrey Bot. Club 51:11. 1924. b, bar ERTO RICO: Cayo Muertos, limestone rocks, shrub, 2 m, g , 9-12 Mar 1915, Britton, Cowell & Brown 4982 (HOLOTYPE: NY; IsoTYPE: US). In the early 1990s, Myrtaceae specialist Bruce Holst (presently at Selby Botanical Gardens), while work- ing at Missouri Botanical Garden, first recognized that this species was out of place in Eugenia and sug- gested it might belonged in Psidium. He therefore brought it to the attention of Les Landrum, at Arizona State University, who is conducting revisionary work on the genus. Subsequently, Landrum and Bonilla (1996) in a study of anther glandularity in the American Myrtinae determined that this species should be transferred to Mosiera, but they did not make the new combination. Instead they deferred doing so until the genus could be worked on as a whole. ACKNOWLEDGMENTS I express my appreciation to Leslie Landrum for his valuable insights and support in my studies in Myrta- ceae. I am very indebted to Frank Axelrod (UPRRP), and the staff at JBSD, especially Daisy Castillo de Vásquez and Martín de la Cruz, who helped make my herbarium work and fieldwork possible. I thank the curators of the following herbaria for kindly providing access to specimens for this study: ASU, BM, F, FLAS, JBSD, MICH, MO, NY, S, UPRRP and US. The comments of Bruce Holst and Donald Pinkava helped improve this paper. I gratefully acknowledge funding support from the Friends of the Center for Latin American Studies at Arizona State University, American Society of Plant Taxonomists Graduate Student Research Award, National Council of State Garden Clubs, Arizona Federation of Garden Clubs, Sigma-Xi, and the Explorers Club. REFERENCES LANDRUM, L.R. 1992. Mosiera (Myrtaceae) in Mexico and Mesoamerica. Novon. 2:26-29. LANDRUM, L.R. and J. Bonita. 1996. Anther glandularity in the American Myrtinae (Myrtaceae). Madroño 43: 508-68 LANDRUM, L.R. and M.L. Kawasaki. 1997. The genera of Myrtaceae in Brazil: and illustrated synoptic treatment and identification keys. Brittonia 49:508-536. SALYWON, A. 2003. A monograph of Mosiera (Myrtaceae). PhD. Dissertation, Arizona State University, Tempe, U.S.A. SALYWON, A., N. Snow, M.F. WoJciecHowski, J. CsizwADI, and L. Lanbrum. 2004. Phylogenetic relationships of Myrtaceae as inferred from nrDNA ITS sequences. Botany 2004, abstract 38-5. SALYWON, A. and L.R. Lanbrum. Curitiba (Myrtaceae): a new genus from the Planalto of southern Brazil. Brittonia (in press). NEW COMBINATIONS IN PHILADELPHUS (PHILADELPHACEAE) James Henrickson Plant Resources Center University of Texas Austin, Texas 78712-0471, U.S.A. ABSTRACT T] T ; d T pq ESTE Philadolnt H hell A. Gray var. madrensis (Hemsley) Henr eos comb. & stat. nov.; " Philadelphus ropas A. Gray var. TET cto) Henrickson, comb. & stat. nov.; and Philadelphus microphyllus A. Gray var. stramineus (Rydb.) Henrickson, comb. & sta RESUMEN e ; Dita ida] TT DPhiladolnl : hell A. Gray var. madrensis (Hems- les) o comb. & stat. nov. “Philadelphus microphyllus A. Gray var. argyroealya (V Weston) Henrickson, comb. & stat. nov., y Philadelphus microphyllus A. Gray var. stramineus (Rydb.) Henrickson, comb. & stat. n Preparation of a treatment of Philadelphus for Flora North America Vol. 8 necessitates the following new combinations. Philadelphus microphyllus A. Gray var. madrensis (Hemsley) Henrickson, comb. & stat. nov. Basowvw: Philadelphus madrensis Hemsley, Kew Misc. Inform. Bull. 1908:251. 1908. Type: MEXICO. Duranco: Sierra Madre, B.C. Seemann 2167 (HOLOTYPE: K!). Philadelphus microphyllus A. Gray var. argyrocalyx (Wooton) Henrickson, comb. & stat. nov. Basionyw: Philadelphus argyrocalyx Wooton, Bull. Torrey Bot. Club. 25:452. 1898. Tyre: U.S.A. New Mexico. Lincoln Co.: Eagle Creek, 14 Aug 1897, E.O. Wooton 524. (HOLOTYPE: US!). Philadelphus microphyllus A. Gray var. stramineus (Rydb.) Henrickson, comb. & stat. nov. Basionym: Phila- delphus stramineus Rydb., N. Amer. Fl. 22:172. 1905. Type: U.S.A. CALIFORNIA. Mono Co.: White Mts., Aug 1888, WH. Schockley s.n. (HOLOTYPE: NY!; isotypes: A! F! US). & Justification of the new combinations will be published at a later time in this journal. J. Bot. Res. Inst. Texas 1(2): 901. 2007 902 Journal of the Botanical R h Institute of Texas 1(2) BOOK REVIEWS ANN Fow er RHOADS and Timotny A. Brock (illustrations by A. Auisko). 2007. The Plants of Pennsylvania: An Illustrated Manual (ed. 2). (ISBN 978-0-8122-4003-0, hbk.). Univ. of Pennsylvania Press, 3905 Spruce Street, Philadelphia, PA 19104-4011, U.S.A. (Orders: Univ. Penn. Press Warehouse, PO Box 5030, Baltimore, MD 21211, U.S.A.; custser@pobox.upenn.edu; 800-537-5487; 410-516-6998 fax). Bae 1088 pp., 2,645 line d maps qe TOt Co E [2] 1 s A 1 1 1 «1 Tahal xl Ttf iJi In and genera E oyed in the first edition with family Ads that best SUE our current desque of phylogenetic relationships within the vascular plants. ... We drew Hes on m Flora of North America (published volumes and website) and the work of the Qu d e . and many of the references cited therein in arriving at the [order] for the second edition. The text Plant S Phylogenetic Approach, Second Edition, by Judd et al. (2002) was also an important source." An exceptional fo) value for 1088 pages, m nearly 3400 species, and including numerous additions to the flora beyond the first edition of 2000. Wu ZHENGYI, PETER H. Raven, and Howc Deyuan (eds.). 2007. Flora of China, Vol. 13. Clusiaceae through Araliaceae. (ISBN-Vol. 13: 978-1-930723-59-7, hbk.). Science Press (Beijing) and Missouri Botanical Garden Press (St. Louis), PO. Box 299, Saint Louis, MO 63166-0299, U.S.A. (Orders: www.mbgpress. org, orders@mbgpress.org, 314-577-9547, 314-577-9594 fax). $125.00, 548 pp., 8 3/4" x 11 1/4". This is “the 13'^ of a 25-volume work. It includes 33 families, 151 genera, and 1288 species, among which 11 genera and 649 species (5096) are endemic to China, and three families, 14 genera, and 76 species are introduced to China." Alangiaceae, Ancistrocladaceae, E Ir Araliaceae, Begoniaceae, Bixaceae, Cactaceae, Caricaceae, Cistaceae, Clusiaceae, Combretaceae, Crypteroniaceae, Cynomoriaceae, Elaeagnaceae, Elatinaceae, Flacourtiaceae, Frankeniaceae, Haloragaceae, Hippuridaceae, Lecythidaceae, Lythraceae Mere corneas Myrtaceae, Nyssaceae, Onagraceae, Passifloraceae, Rhizophoraceae, Stachyuraceae, Tamaricaceae, Tetramelaceae, Thymelaeaceae, Trapaceae, Violaceae. Chinese and American authors collaborate. WALTER S. JUDD, CHRISTOPHER S. CAMPBELL, ELIZABETH A. KELLOGG, PETER E STEVENS, and MICHAEL J. DONOGHUE. 2007. Plant Systematics: A Phylogenetic Approach (ed. 3). (ISBN 978-0-87893-407-2, hbk.). Sinauer Associates, Inc. 23 Plumtree Road Sunderland, MA 01375. (Orders: orders@sinauer.com, publish@ sinauer.com, www.sinauer.com). $94.95, 620 pp., ca. 325 illustrations, 25 color plates, 8 1/2" x 11". This justifiably much-used book is noted to be "appropriate for any course e ] f angi vascular plants and, secondarily, for local flora courses. The text assumes no prerequisite r than E botany or biology” New to the 3"4 edition (beyond 2002 2"4 ed): “color throughout the book, including m more ae two dozen pl l variation in the vee pa new ibaa f maximum likelihood and B ly revised treatments eo 17 families that in ci ll cl d to the many cladograms, taking into account recent taxonomic UR and M Án accompanying CD contains over 3,100 color photographs depicting a wide variety of plants from over 185 families and over 1900 species. Many species are oda by pend Dn um showing different views of the plant, its flowers, its fruits, and its habit. Also on s CD Gwith links to photos) and three the text, according to HM APG, nas and mE Cone: in Dil by chapters: The Science of Plant Systematics; Methods and Principles of Biological Systematics; Classification and Systems in Flowering Plants-Historical Background; Taxonomic Evidence-Structural and Biochemical Characters; Molecular Sys- tematics; The Evolution of Plant Diversity; An Overview of Green Plant Phylogeny; Lycophytes, Ferns, and Phylogenetic Relationships of Angiosperms. Two appendices: Botanical Nomenclature; Specimen Preparation and deb aon. J. Bot. Res. Inst. Texas 1(2): 902. 2007 FIVE NEW VARIETIES OF AGAPETES (ERICACEAE) FROM INDIA D. Banik M. Sanjappa Botanical Survey of India Botanical Survey of India Central National Herbarium CGO Complex, 34 MSO Building PO: Botanic Garden, Howrah Block E, 5% Floor, DF Block, Sector-1 71 a , INDIA Salt Lake City, Kolkata—700 064, INDIA banikdipanwita@yahoo.com m_sanjappa@yahoo.co.in ABSTRACT Five new varieties of the genus Agapetes D. Don ex G. Don from Arunachal Pradesh, Mizoram and Nagaland, India are described and illustrated Key Wonps: Agapetes, new varieties, Arunachal Pradesh, Mizoram, Nagaland, India RESUMEN Se describen e ilustran cinco nuevas variedades del género Agapetes D. Don ex G. Don de Arunachal Pradesh, Mizoram y Nagaland, India The genus Agapetes D. Don ex G. Don (Ericaceae) comprises ca. 95 species (Mabberley 1997) or ca. 80 species (Ruizheng & Stevens et al. 2005) and is distributed from the E Himalayas through SW China and Indochina to SE Asia. In India, the genus comprised ca. 34 species (Santapau & Henry 1973). Presently in India there are 58 species and 15 varieties under the genus Agapetes. A revision of the genus Agapetes for the Flora of India project has involved extensive field surveys in different states of North East India and studying herbarium specimens at various Indian herbaria. During this critical study, five new varieties of Agapetes came to light; they are described and illustrated here. 1. Agapetes acuminata D. Don ex G. Don var. tipiensis Banik € Sanjappa, var. nov. v. (Fig. D. Tyee: INDIA. ARUNACHAL PRADESH: West Kameng district, on the way from Tipi to Sessa, 1050 m, 23 Apr 2002 ISOTYPES: ASSAM, CAL). A varietate typica inflorescentia subumbellata, pedunculo breviore 2-3 mm longo, pedicello breviore 5-5.5 mm longo differt. Epiphytic straggling shrubs, sometimes terrestrial, 0.5-1 m long. Lignotubers basal, amorphous in epiphytic ones. Stems robust, terete, obscurely striate, rugose, glabrous, densely lenticellate even when young; perulae 5-9, closely to loosely alternate, 4-6 mm apart, deltoid to linear, 0.5-2 x 0.2-1 mm, serrate at margins, acuminate at apex, scaly, caducous. Leaves loosely 2-3-stichous, 1.5-3 cm apart; petioles 0.7-1.5 cm long, obscurely rugose, slightly pulvinus at base, glabrous; lamina 10.5-25.5 x 2.5-7 cm, lanceolate to ovate-lanceolate, broadly cuneate to rounded at base, bluntly serrate (serratura 10-18 pairs) at margins, acutely acuminate (acumen 1-1.5 cm long) at apex, chartaceous, glabrous; veins depressed above, brochidodromus, secondary veins 10-18 pairs at 45°-60°, marginal vein 0.5-4 mm away from margins. Inflorescences axillary or cauline, subumbellate, 7-9-flowered, sparsely puberu- lous; peduncles 2-3 mm long; peduncular bracts 5-11, deltoid or boat-shaped, 0.5-1.5 x ca. 0.4 mm, acute at apex, scaly, caducous; pedicels crimson red, 5-5.5 mm long, obconical, ca. 3 mm broad at apex, fleshy, puberulous, articulate at the base of calyx; pedicellar bract 1, linear, 1-1.5 x 0.5-0.6 mm, serrate at margins, acuminate at apex, midvein raised outside, scaly, caducous; bracteoles 2, opposite, 0.5-1 x 0.2-0.3 mm, similar to bracts. Calyx crimson red, 2.53 x ca. 3.1 mm, puberulous; lobes deltoid ca. 1.3 x 1.5 mm, acute at apex. Corolla crimson red, urceolate, 3.5-4 x 4-4.5 mm; lobes 1-1.3 x 1-1.5 mm. Stamens whorled, 3-3.5 mm long; filaments linear, 1-1.5 mm long, slightly incurved, sparsely pilose; anthers 2-2.5 mm long including 1-1.1 mm long thecae, sparsely granular, tail obscure and tubules J Bot Res. Inst Texas 1(2): 003 — 912. 2007 904 Journal of the Botanical R h Institute of Texas 1(2) Hw di TE i p "d A K e s Del. D. Banik Fic. 1. A t ji D. Don ex. G. Don var. IR F 3 FF b = 2 mm; c = 3 mm; d-e = 1 mm (a-e, D. Banik 27953). habit; b. flower; c. calyx and style; d-e. stamens. Scale bars: a = 1 cm; Banik and Sanjappa, New varieties of Agapetes from India 905 1.3-1.4 mm long. Style cylindric, 3.5-4 mm long, glabrous, slightly impressed inside ovary, caducous; stigma glandular-truncate. Fruit not seen. Notes.—This variety differs from the typical variety in having subumbellate inflorescences, shorter peduncles (23 mm long) and pedicels (5-5.5 mm long) whereas the typical variety has corymbose in- J 1 florescences, 3.5-7 mm long p and 1-1.6 cm long pedicels. Variety tipiensis is found in Arunachal Pradesh whereas the type variety occurs in Meghalaya (India), Bangladesh and Myanmar. Distribution and Habitat.—India: Arunachal Pradesh. Common in the subtropical forests at altitudes ranging from 1050-1200 m. Flowering.—April-June. Etymology.—This variety is named after the type locality. PARATYPE: INDIA. Arunachal Pradesh: West Kameng district, Aka hills, Jun 1934 (fl), Bor 15741 (ASSAM). 2. Agapetes flava (Hook. f.) Sleumer var. nagensis Banik & Sanjappa, var. nov. d 2). Tyee: INDIA. NAGALAND: Naga (Barail) hills, above Konoma, 2100 m, Jul 1886 (fl), D. Prain s.n. (HoLoTYPE: CAL, 264391) A varietate t lycis lobis multo breviore 4-6 mm longo, ovato; foliis serrato, basi cuneato differt. E Epiphytic shrubs. Lignotubers basal, amorphous, 7-10 x 5-6 cm. Stems slender to robust, terete, stri- ate, lenticellate, glabrous but sparsely puberulous when young. Leaves: pseudowhorls 2.2—4.5 cm apart, 2-3-leaved; petioles 3-6 x 1.5-2.5 mm, puberulous; lamina lanceolate to broadly elliptic, 7-8.5 x 1.7-3.8 cm, obtuse to cuneate at base, serrate and recurved at margins, acutely acuminate (acumen 0.8-1.7 cm long) at apex; veins depressed above, midvein puberulous at base, brochidodromus, 13-14 pairs of secondary veins at 60-85". Inflorescences axillary or cauline, corymbose or 5 — 6 flowers in fascicles, puberulous; peduncles 1-2.5 mm long; peduncular bracts 2-3, closely alternate, ovate to linear, 0.5-1 x ca. 0.5 mm; pedicels slender, 1.2-1.8 cm long, puberulous; pedicellar bract 1, triangular 1-1.5 x ca. 0.5 mm, puberulous; bracteoles 2, linear, 2-2.5 x ca. 0.5 mm, acuminate at apex, puberulous. Calyx winged, 8-10 x 4—4.5 mm, puberulous; lobes ovate, 4—6 x 2.5—3.5 mm, acute at apex. Corolla yellow, tubular- urceolate, (1.4—1.5)20—22 x 0.5-0.6 cm, puberulous; lobes broadly triangular, obscure or ca.1 mm long. Stamens (1.4)1.8-2 cm long; filaments 1-2 mm long, ellipsoid, glabrous to sparsely puberulous outside; anthers (1.3)1.7-1.8 cm long including 4-4.5 mm long thecae, granular, tail ca. 0.5 mm long, slightly recurved, granular and tubules (0.9)1.3-1.4 cm long, obscurely calcarate, spurs obscure, basal to tubule. r Ovary 3-4.5 mm in diameter; style slender, 1.4-1.6 cm long; stigma truncate. Fruit not seen. Distribution and Habitat.—India: Nagaland. Grows in subtropical forests at ca. 2100 m. Flowering.—June—August. Etymology.—This variety is named after the type locality. Notes.—This variety differs from the typical variety in having much shorter calyx lobes 4-6 mm long, ovate and leaves serrate, cuneate at base whereas the typical variety has calyx lobes 8-11 mm long, elliptic-lanceolate and leaves obscurely crenate, rounded at base. In typical variety the corolla tube al- most is covered by calyx. Variety nagensis is found in Nagaland whereas the type variety is known from Arunachal Pradesh. PARATYPES: INDIA. Nagaland: Naga (Barail) hills, above Konoma, Aug 1886 (fl), D. Prain s.n. (BSIS, 17874); D. Prain s.n. (£D, (CAL, 264390); D. Prain s.n. (f1) (CAL, 264392). 3. aD megacarpa WW. Sm. var. lohitensis Banik & Sanjappa, var. nov. Duis 3). Tyre: INDIA. ARUNACHAL 10 ) ESH: Lohit district, Mailiang to Chipra, ca. 1400 m, 16 May 2003 (fl), D. Banik 27 G AL; isotypes: ASSAM A varietate typica calycis lobis multo longioribus lineari-ellipticis 1.6-2.1 cm longis 3-5 mm latis differt. Epiphytic or lithophytic shrubs, erect, 2-3 m high. Lignotubers basal, amorphous. Stems robust, terete, striate, lenticellate, glabrous; perulae 6-12, close to loosely alternate, ovate to linear-lanceolate, 5-10 x 2.5-6 mm, serrate at margins, acuminate at apex, caducous to persistent. Leaves: pseudowhorls 2.5-5 cm apart, 3-6-leaved; lamina sessile, elliptic or broadly elliptic, 12-18 x 2.7-6 cm, auriculate to 906 Journal of the Botanical R h Institute of Texas 1(2) | - 1 2" A. { — AA " Del. D. Banik Fic. 2. Agapetes flava (Hook. f.) Sleumer var. nag jap 5 mm; d-e = 3 mm [a-e, D. Prain s.n. (Herb. Acc. No. 264392)]. 907 Banik and Sanjappa, New varieties of Agapetes from India Del. D. Banik e. stamens; f. basal portion of stamens; ry and stula: d— yle; 7 habit; b flower; c caly 5 mm (a-g, D. Banik 27534). =1 cm; d, e, 9 = N. Sm. var. lof F 2 Jr g. apical portion of tubules. Scale bars: a-c, f Fic. 3. A 908 Journal of the Botanical R h Institute of Texas 1(2) cordate at base, entire to serrate at margins and obscurely recurved towards apex, acuminate to acutely acuminate (acumen 1-1.5 cm long) at apex, coriaceous, glaucous above, dull below; veins raised above, brochidodromus, 10-12 pairs of laterals at 45°-80°. Inflorescences axillary or pseudoterminal, corym- bose, 6-15-flowered, glabrous, maroon, dealbatus; peduncles maroon, 1-1.5 cm long, 4-5 mm broad, stout, glabrous, dealbatus; peduncular bracts maroon, 5-6, linear, 1-1.5 x 0.5-0.6 cm, prominently veined, glabrous, dealbatus, caducous; rachis 3-3.5 cm long; pedicels 2-3-stichous, maroon, obconical, 23.2 cm long, 0.7-1 cm broad at apex, stout, glabrous, dealbatus; pedicellar bract 1, triangular, 3.5-4 x 1.5-2 mm, serrate, acute; bracteoles 2, opposite, linear, 2-3 x ca. 1 mm, serrate, acuminate, caducous. Calyx maroon, 2.3-2.5 x 1.5-1.6 cm, glabrous, rugose at base, dealbatus; lobes linear-elliptic, 16-21 x 3-5 mm, slightly constricted at base, entire at margins, acuminate to obtusely acuminate at apex, thickly coriaceous, 3-4 pairs of secondary veins at 15°-30°, raised above. Corolla maroon-red, tubular-urceolate, 5-6 x 1.5-2 cm, glabrous, dealbatus, 6-7 pairs of secondary veins at 30°-60°, dark maroon, directed upwards forming closed reticulations; lobes green, ovate-caudate or triangular-caudate, 15-16 x 6—6.5 mm, glabrous, dealbatus. Stamens 10, 2-whorled; filaments spathulate, 4.5—7 mm long, incurved, sparsely hirtellous outside; anthers 4.6—4.8 cm long including 1.1-1.5 cm long thecae, verrucate, tail 0.5-0.7 mm long, verrucate, reflexed and tubules 3.2-3.7 cm long, sparsely verrucate to smooth, calcarate, spurs apical on tubule, 0.5-0.7 mm long, verrucate, ascending and reflexed in adjacent anthers. Style slender, ca. 5.7 cm long; stigma 5-lobed, 23 x 2-2.5 mm. Fruit a berry, maroon, obovoid, 2.5-2.8 x 1.5-1.8 cm, glabrous, dealbatus. Seeds obovoid, 3-5 x 2-2.5 mm, pointed at base; seed coat reticulate. Distribution and Habitat.—India: Arunachal Pradesh. Common in the subtropical forests at altitudes ranging from 1300-1500 m. Flowering and Fruiting.—March-May. Etymology.—This variety is named after the type locality. Uses.— Corolla used as vegetable. Note.—This variety differs from the typical variety in having much longer (1.6-2.1 cm) linear-el- liptic calyx lobes whereas the typical variety has shorter (6.5-11 mm) ovate to lanceolate calyx lobes. Variety lohitensis is found in Arunachal Pradesh whereas the typical variety is distributed in Nagaland (India), China, and Thailand. PARATYPES: INDIA Arunachal Pradesh: Lohit district, Kuibang, 25 Mar 1986 (fD, K. Harid 2909 (APFH); Lailiang to Mailiang, near Lailiang, 1400-1500 m, 16 May 2003 (£D, D. Banik 27533 (CAL). 4. Agapetes odontocera (Wight) Hook. f. var. mizoramensis Banik AD var. nov. (Fig. 4). Tee: INDIA: Mizoram, Thaltlang phui, 16 May 1990 (fl), B.M. Wadhwa & K.P Singh 96834 A i Į fl ifi 1 la differt E Small trees. Stems robust, terete, obscurely striate, lenticellate, glabrous; perulae many, compactly al- ternate to loosely alternate (5-8 mm apart), triangular to linear-triangular. Leaves: pseudowhorls 4-5.5 cm apart, 3-5-leaved; petioles 1-2 mm long, glabrous; lamina narrowly elliptic-oblanceolate, 14-15 x 2.3-3.8 cm, cuneate to obtuse at base, serrate at margins, acutely acuminate at apex, coriaceous; veins raised above, brochidodromus, 15-16 pairs of secondaries at 45°—60°. Inflorescences axillary or cauline, flowers 3-6 in fascicles, puberulous; inflorescence bracts closely alternate; pedicels slightly obconical, 1.8-2 cm long, puberulous; pedicellar bract 1, deltoid, ca.1 x 1 mm, scaly, caducous; bracteoles 2, op- posite, linear, ca.1 x 0.5 mm, similar to bract. Calyx ca. 5 x 2.5 mm, puberulous; lobes ovate-triangular, ca. 3.5 x 2 mm, acuminate at apex. Corolla tubular, 2.5-2.8 x 0.8-1 cm, puberulous; lobes triangular, ca. 1 x 0.5 mm, acuminate at apex. Stamens 10, in 2 whorls, ca. 2.5 cm long; filaments spathulate, 2-2.5 mm long, slightly incurved, puberulous to pilose outside; anthers 1.8-2.2 cm long including 4.5-5.5 mm long thecae, granular, tail obscure and tubules 1.4-1.6 cm long, sparsely granular, calcarate, spurs ca. 1 mm long, linear, reflexed. Style slender, 2.5-2.7 cm long; stigma truncate. Fruit not seen. Distribution.—India: Mizoram. Banik and Sanjappa, New varieties of Agapetes from India 909 Del. D. Banik J Minha U I . D TOC Fic. 4 Ag ji (Wight) F. var. j habit; b. flower; c calyx and style Scale bars: a = 1 cm; b-c = 5mm (a-c, B.M. Wadhwa & D.K. Singh 96834). 910 Journal of the Botanical R h Institute of Texas 1(2) Flowering.—May. Etymology.—This variety is named after the type locality. Notes.—This variety differs from the typical variety in having uniformly puberulous inflorescences whereas the typical variety has totally glabrous inflorescences. Variety mizoramensis is recorded from Mizoram whereas the typical variety is widely distributed in Arunachal Pradesh, Assam, Meghalaya, Mizoram, Manipur and Nagaland. 5: sa ite pe C.B. Clarke var. glanduliflora Banik € Sanjappa, var. nov. (Fig. 5). Tyee: INDIA. ARU- ESH: Lower Dibang valley district, Mehao lake to Roing, 27 Nov 2000 (fl), D.K. Singh & Party 9319 (HoLotTyPE: CAL; ISOTYPE: Rum A varietate typica inflorescentia glandulosopubescenti vel glandis subsessilibus in pedunculo, pedicellis, calyce et corollae costis dif- fert Epiphytic shrubs, erect, 0.5-1 m tall. Stems robust, terete, 5-8 mm in diameter, striate, glabrous, lenticellate when mature; perulae 3-6, closely to loosely alternate, 1—4.5 x 1-1.2 mm, deltoid-ovate to linear-elliptic, entire to glandular at margins, acute to acuminate at apex; midnerve raised, glabrous, persistent. Leaves: pseudowhorls 5.5-8.5 cm apart, 8—9-leaved; petioles 4-8 mm long, glabrous, winged, slightly rugose outside, slightly pulvinus at base; lamina 6-16 x 1-2.5 cm, linear-elliptic, attenuate at base, entire at margins, acuminate at apex, coriaceous, glaucous above, pale below; veins slightly raised above, brochidodromus, 12-16 pairs of secondary veins, alternate at 30-80”. Inflorescences axillary or cauline, corymbose, erect, 9-16-flowered, glandular hairy; peduncles dark red, 3.5-5.5 mm long, glan- dular hairy; peduncular bracts 3-6, closely alternate, broadly deltoid, obscure, glabrous, membranous, caducous; pedicels dark red, 4.5-7 mm long, striate, glandular hairy; pedicellar bract 1, 1-1.5 mm long, linear, persistent; bracteoles 2, linear, ca.1 mm long, similar to bracts. Calyx dark red, 5-6.5 x 3.5-4.2 mm, glandular hairy or with subsessile glands; lobes ovate, 2.83.5 x 1.3-2.5 mm, acuminate at apex, membranous, veins raised above, glandular hairy. Corolla dark red to orange red, tubular-urceolate, 1.5-2 x 0.4-0.5 cm, angular, membranous, reticulate in dark red, secondary veins 5-6 pairs, at ca. 60°, midveins glandular hairy; lobes green or greenish- yellow, 1.5-2 x ca. 1.5 mm, ovate, acute to obtuse at apex, glandular hairy. Stamens 2-whorled, 1.5-1.8 cm long; filaments 2.5-3 mm long, linear, incurved, puberulous outside, hyaline; anthers 1.2-1.5 cm long including 5-5.5 mm long thecae, verrucate, tail bilobed, recurved and tubules ca. 10 mm long, glabrous. Ovary 33.5 mm in diameter, striate, glabrous; style slender, 1.7-1.9 cm long, ca. 1 mm deeply impressed inside ovary; stigma glandular capitate, 5- lobed, ca. 0.5 x 0.6 mm. Fruit a berry, globose, 4—6.5 x 4.5-5 mm, sparsely glandular hairy or with sessile glands. Seeds obovoid, 1-1.2 x 0.5-0.7 mm; seed coat reticulate. Distribution and Habitat.—India: Arunachal Pradesh. Common in the subtropical forests between 1300 and 1400 m altitude. Flowering and Fruiting.— October-November. Etymology.—This variety is named for the glandular pubescence of inflorescence. Notes.—This variety differs from the typical variety in having glandular hairy inflorescence or with subsessile glands on peduncles, pedicels, calyx and midveins of corolla whereas the typical variety has glabrous and eglandular inflorescence. Both varieties are distributed in Arunachal Pradesh. PARATYPES: INDIA. Arunachal Pradesh: Lower Dibang valley district, Tiwari gaon to Roing, 1300 m, 3 Nov 2002 (fl), D. Banik 27413 (CAL); Mehao lake to Baldi camp, 1400 m, 16 Nov 2002 (fD, D. Banik 27432 (CAL); Lohit district, Hawa camp to Udayak pass, 1400 m, 24 Apr 2003 (fr), D. Banik 27508 (CAL). ACKNOWLEDGMENTS The authors are thankful to J.F. Veldkamp, Nationaal Herbarium Nederland, Leiden, the Netherlands for the Latin diagnoses of the new varieties. Pakshirajan Lakshminarasimhan and one anonymous re- viewer provided helpful review comments. One of us (DB) is grateful to the Director, Botanical Survey of India, for granting fellowship, to D.K. Singh, the Additional Director; Botanical Survey of India for Banik and Sanjappa, New varieties of Agapetes from India 911 > LESS > = (= 3 ea 2 = o a d IG. 5 C.B. Clarke var. g Banik & Sanjappa: a. habit; b. flower; c. caly yle; d. stamen; e-h. peduncular bracts; i. t.s. of ov. 2an;b 5 d = 3 mm; e-h = 2 mm; I = 1 mm (a-i, D.K. Singh & Party 9319). 912 Journal of the Botanical R h Institute of Texas 1(2) specimens; and to the Joint Director, Central National Herbarium, Botanical Survey of India, Howrah, for facilities. The acronym APFH is used in the text for The Herbarium, State Forest Research Institute, Arunachal Pradesh, Van Vihar, Chimpu, Itanagar 791 111, Arunachal Pradesh, India. REFERENCES Massertey, D.G. 1997. The plant book.2™ ed. (Reprinted with corrections 1998, 2000). Cambridge University Press. RUIZHENG, F. and PF. Stevens. 2005. Agapetes. In: Editorial Committee, eds. Flora of China, vol. 14. Science Press, Beijing and Missouri Botanical Garden Press, St. Louis. Pp. 504-516. SANTAPAU, H. and A.N. Henry. 1973. A dictionary of the flowering plants in India. (Reprinted in 1998). National Institute of Science Communication, New Delhi. PHYLOGENETIC ANALYSES OF THE GENUS BAPTISTONIA (ORCHIDACEAE: ONCIDIINAE) SENSU LATO BASED ON MORPHOLOGICAL CHARACTERS Guy R. Chiron biers, Université Claude Bernard - Lyon 1 9 rue roe Dubois, 69622 Villeurbanne Cedex, FRANCE ofr chiron@wanado ABSTRACT Pelar; 1 ip shin R l ; ; ; 11 1 pl ; Uem 1 £47 pl ] gi Tel The analyses support monophyly of Baptistonia, but without strong | p support. S praspecifi pported by th l di d RÉSUMÉ On présente, sur la base d'une analyse Pune que et ona de 47 caracteres morphologiques, les relations de parenté entre les espèces de Baptistonia. L'étude soutient 1 genre, avec cependant de faibles valeurs de bootstrap. Quelques taxons supraspécifiques, suggérés par l'analyse, sont den Key Worbs: Baptistonia, Brazil, Gomesa, Orchidaceae, Oncidiinae, phylogeny INTRODUCTION The genus Baptistonia was founded in 1877 by Barbosa Rodrigues based on a single species: B. echinata Barb. Rodr. Soon after the concept was published, it was generally neglected. Kránzlin (1922), in his revision of the entire subtribe Oncidiinae, placed the species in the genus Oncidium Swartz. However, Pabst and Dungs (1977), in their work on the Orchidaceae of Brazil, conserved Barbosa Rodrigues' monospecific genus. Chiron and Castro Neto (2004) noted that the characteristic vegetative and floral structures of the plants of the section Waluewa (Regel) Schltr. of the genus Oncidium are close to those of Baptistonia echinata, and that all species involved are endemic to the southeastern part of Brazil. Consequently they transfered the species of section Waluewa to the genus Baptistonia. The thus enlarged genus now comprises twenty one species and two natural hybrids. DNA sequence data has shown that the species involved are distinct from those belonging to the true genus Oncidium (Williams et al. 2001; Chase et al. 2005). This data also seems to indicate that these species form a monophyletic group which is distinct from its neighboring groups (Faria 2004). It should however be noted that the work of Chase et al. and Faria is based on a limited number of species and that the variations observed in the analyzed sequences are extremely small. At this time, a study based on the trnS-G region of a large number of members of the genus is in progress. Parallel to this study, however, I also wanted to explore the utility of morphology and anatomy in respect to phylogenetic relationships. Faria (2004) published morphological phylogenetic of seven Baptistonia species (less than one-third of the group) but included representatives of various other groups of Brazilian oncidi- ums. The objective of my study is to complement the work of that author and to estimate the interspecific relationships of all naturally occurring Baptistonia taxa at the species level, the hybrids being excluded. MATERIALS AND METHODS The anatomical and morphological characters that are presented and discussed below have mainly been obtained through the observation of several live plants of each species. Herbarium material was only utilized whenever live material was not available in any sufficient quantity. The observations were supplemented by data deduced from the original descriptions of the taxa. The genus Baptistonia is closely related to the Gomesa alliance, a Brazilian Oncidiinae group compris- ing the genera Gomesa R. Br., Rodrigueziella Kuntze and Rodriguesiopsis Schltr. In this study, I included four J. Bot. Res. Inst. Texas 1(2): 913 — 931. 2007 914 Journal of the Botanical R h Institute of Texas 1(2) Gomesa species as outgroup: G. alpina Porsch, G. crispa Klotzsch ex Rchb.f., G. recurva R. Br. and G. sessilis Barb. Rodr. The trees were rooted with three species of the genus Oncidium: O. altissimum Swartz (the type species of the genus, originating from the Antilles), O. nebulosum Lindl. (a mesoamerican species belonging to Oncidium section Oblongata Kraenzlin) and O. baueri Lindl. (another species of the section Oblongata, native of the Amazonian forests.) Furthermore, I have added two species that I suspect to be close to the species of the genus Baptistonia: — Carriella colorata (Kóniger & J.G. Weinm.) V.P. Castro & K.G. Lacerda, which was placed in Oncidium section Waluewa by Senghas (1997) but elevated to a monospecific genus by Castro Neto and Lacerda (2006), and —Oncidium trulliferum Lindl., close to the plants of the genus Baptistonia in respect to its vegetative characters as well as in respect to some of its floral structures, but generally placed in Oncidium section Rostrata Rolfe. In view of the work of Faria (2004), no specimens were included belonging to any of the other groups of the Gomesa clade (a monophyletic group consisting of the Gomesa alliance, Baptistonia and many of the Brazilian oncidiums). All specimens that were studied, with the exception of those belonging to the external Oncidium group, are epiphytic plants originating from the moist forests of the Mata Atlantica. A list of the 217 specimens studied is given in Appendix 1. The second important issue to discuss relates to the pertinence of the characters that have been se- lected for the analysis. Chase (1986) stated that several morphological characters, such as the angle between the labellum and the column, the fusion of the lateral sepals, the general shape of the floral segments, the column arms, and the callus of the labellum, cannot be used for an objective phylogeny. Faria (2004), on the other hand, disagrees with Chase's opinion on this matter. Burns-Balogh and Funk (1986) as well as Freudenstein and Rasmussen (1999) discussed a great number of morphological characters in their cladistic analysis of Orchidaceae. In these articles, however, the utility of the characters was evaluated only in respect to generic relationships, not interspecific. Each of the observable characters may either be invariant within al (e) the groups studied (and therefore are of no use for pl ic analysis), may vary among species, or—as a third possibility—may even vary within a given species, thus being inconsistent and therefore unsuitable for species-level phylogenetics. Thus, this pertinence of the 69 selected vegetative and floral characters, either microscopic or macroscopic, needs to be discussed, in order to (a) eliminate the unsuitable ones, and (b) decide how many different states of each character retained can be used in the present study. In accordance with the recommendations given by Garcia-Cruz and Sosa (2006) for quantitative characters, I employ the Gap-Weighting (GW) coding method as proposed by Thiele (1993), with a limited number of statistical conditions. In this way, the danger of overweighting certain characters is eliminated. Polarity of character coding is based upon pleisiomorphy of Oncidium. According to van den Berg (unpublished), the Oncidiinae evolved, at the beginning of the Miocene, about 23 million years ago, from an ancestor belonging to the tribe Maxillarieae, itself having diversified from a Mesoamerican ancestor. At that time, South and North America were still separated from each other, the Panamean isthmus having been formed during the Pleiocene. It is generally assumed (Por 1995) that there was an inland sea from the east of what is now Colombia to the Pantanal, isolating the entire eastern part of South America, the sea not being connected with the Atlantic Ocean until the Miocene. Therefore, the Brazilian Oncidiinae, especially the Gomesa clade, would have diversified much later, on the basis of an ancestor in Central America. The species of Oncidium sensu stricto have their center of distribution in Central America, and they are probably much closer to the common ancestor than the species belonging to the genus Baptistonia. Morphological and anatomical characters considered According to the above discussion, the following twenty characters were considered but excluded: spacing, Chiron, Phylogenetic study of Baptistonia 915 diameter and surface structure of the pseudobulbs; length, shape, color of the leaves, consistency of the lamina, distribution of the stomata, shape of the epidermal cells; length and diameter of the peduncle of the inflorescence, length of sterile bracts, flower density; fusion of the lateral sepals of the flowers, outline and apical shape of the lateral sepals, shape and position of the lateral lobes of the lip, curvature of the column, shape of the stigmatic cavity. Some of them correspond to what Chase (1986) calls “unreliable characters.” A. Pseudobulbs (mature but young) * Height (character 01) Within any given species, the height of the pseudobulbs may vary. Consequently, I used the maximum height observed. The GW Method, applied with the number of states limited to 2, allows setting a limit of 7 cm to differentiate between species with small or large pseudobulbs. 01-maximum height of pseudobulbs: 1 up to 7 cm—2 more than 7 cm Assumed evolution: 1 — 2 * Shape in vertical section (character 02) The distinctly elongated shape of the pseudobulbs is characteristic for the genus, but certain species of Baptistonia do have pseudobulbs that are elongated-ovoid as in Gomesa. For the evaluation of this character I have chosen to use the ratio of height over diameter (H/D), the diameter being measured at the thickest part of the pseudobulb. The scoring, according to the GW method, has three states: 02-Vertical section of the pseudobulbs: 1 ovate (H/D«2)—2 elongated ovate—3 cigar shaped (H/D»4) Assumed evolution: 1 > 2 > 3 * Shape in horizontal section (character 03) Generally, the horizontal section of the pseudobulbs is rounded, but, in Baptistonia, it is sometimes somewhat flattened although never strongly flattened (elliptical). This character is evaluated by the ratio (D/d) of the large diameter over the small diameter of the section, calculated according to the GW method accepting two states: 03—horizontal section of the pseudobulbs: 1 strongly flattened (D/d > 1.5)—2 slightly flattened (D/d « Assumed evolution: 1 > 2 * Margin of pseudobulb in horizontal section (character 04) The pseudobulb in an horizontal section is generally ribbed to entire and this character seems to be a per- tinent one at the generic level, although not at the specific level. 04—Margins of the horizontal section: 1 angular—2 intermediate—3 rounded Assumed evolution: 1 > 2 3 * Color (character 05) When observed in their natural state, the plants belonging to the genus Baptistonia are easily differentiated from the plants of the genera Oncidium or Gomesa by the color of their pseudobulbs and leaves. The shade of the green of the pseudobulbs and leaves varies distinctly among these genera. Under cultivation at the same spot, this difference remains, showing that the various shades of green are not adaptations to chang- ing environmental conditions. 05—color of the pseudobulbs and leaves: 1 bright green—2 dark green Assumed evolution: 1 — 2 B. Leaves Williams (1974) studied the morphology and the anatomy of the leaves of 80 species belonging to a total of 22 genera of the Oncidiinae. Of those 80 species only 4 belonged to the Gomesa clade. He concluded that the leaves offer very few interesting morphological characteristics, but that some anatomic features can be very useful. For the evaluation of those characters in the genus Baptistonia, see Chiron and Guiard (in preparation). 916 Journal of the Botanical R h Institute of Texas 1(2) * Basal sheaths (character 06) At the generic level this character is useful as, in Oncidium, the sheaths are long and leafy, hardly shorter than the real leaves. On the other hand, in Gomesa, they are leafy but short and, in Baptistonia, they are non-leafy. 06—Basal sheaths: 1 long and leafy—2 short and leafy—3 non-leafy Assumed evolution: 1 > 2 > 3 * Number of leaves (character 07) The majority of the species of Baptistonia show a varying number of leaves per pseudobulb. Therefore, for the present study, I considered the maximum number of leaves for any given species. 07-Maximum number of leaves: 1 more than one—2 a single leaf. Assumed evolution: 1 > 2 * Stomata density (character 08) The density of the stomata on the abaxial surface of the leaves in the genera Baptistonia, Oncidium and Gomesa varies from one species to the others (Chiron & Guiard, in preparation). Although this density also varies within any given species, the intraspecific variation is much lower than the variation among the species. The density of the stomata in most of the species is between 20 and 45 stomata per mm?; in the other spe- cies studied, the density was higher than 55 stomata per mm". This character therefore is pertinent at the generic level. 08-stomata density: 1 2 55 per mm?—2 « 45 per mn? Assumed evolution: 1 — 2 * Shape of the stomata (character 09) The shape of the stomata as observed in Gomesa and Baptistonia is sub-circular to elliptic, whereby the ratio major axis/minor axis can attain 1.4. Notwithstanding the fact that this character can vary considerably within any given species, the differences among the different species allow scoring for three different states. 09—Ellipticity of the stomata: 1 € 1.05—2 > 1.05 and < 1.23 > 1.2 Assumed evolution: 1 + 2 > 3 C. Inflorescence * Total length (character 10) The total length of the inflorescence varies from one species to another. This character must be measured on adult plants during a normal flowering. 10-length of the inflorescence: 1 distinctly longer than the growths—2 as long as or shorter than the growths Assumed evolution: 1 2 * Direction (character 11) The inflorescences are generally erect, later becoming arched under the weight of the flowers. However, in certain species of the genus Baptistonia, they are abruptly folded downward directly above the base and long before the buds begin to form. 11—direction of the inflorescence: 1 upward, more or less arched—2 directly folded downward Assumed evolution: 1 > 2 * Form (character 12) Generally, the inflorescences are branched. They are, however, unbranched in Gomesa and in some Baptis- tonia species. 12- inflorescence: 1 branched -2 unbranched Assumed evolution: 1 — 2 * Formation of the buds (character 13) In general, the buds start to develop after the inflorescence has more or less reached its final length. This is not so in the genus Gomesa and in some other species of the Gomesa clade. In those cases, the buds start to Chiron, Phylogenetic study of Baptistonia 917 develop as soon as the inflorescence starts to develop within the protection of the leafy sheaths. 13—Formation of the buds: 1 when the inflorescence has attained its length—2 as soon as the inflorescence starts to develop Assumed evolution: 1 > 2 D. Flowers * Form (character 14) Depending on the species that is under observation, the flowers of Baptistonia can be more or less spread or more or less closed and as such globular. I consider this character potentially pertinent. 14—flower: 1 fully spread (more or less in one plane)—2 quasi spherical (petals and dorsal sepal directed forward). Assumed evolution: 1 > 2 e Outline (character 15) The outline of the flowers can be distinctly circular or vertically ovate. This was determined by using the ratio (H/W) between height and width of the flowers. 15—Flower outline: 1 circular (H/W < 1.1)—2 vertical (H/W > 1.1) Assumed evolution: 1 — 2 * Color (character 16) Originally, I had based my color evaluation of the flowers on the visual absence of yellow, red, and green pigmentation. However, even when we attribute but two states to each color, meaning “visible” and “non- visible", we obtain a total of 8 possible states. This is too many states for this type of character. Finally, I have decided to retain three states. 16-Flower color: 1 mainly colored bright yellow and/or dark red—2 pale yellow to whitish with reddish- violet spots—3 yellowish or greenish without any red spotting Assumed evolution: 1 2 2 > 3 * Floral bracts (character 17) Within the genus Baptistonia, this characteristic does not vary much, but long flower bracts are typical for the species of the genus Gomesa. The GW coding allowing for 3 states gives: 17—Floral bracts: 1 very short («1/5 pedicel)—2 short—3 long (> 1/2 pedicel) Assumed evolution: 1 — 2 3 D1. Tepals * Outline of the dorsal sepal (character 18) This character is partially pertinent within the genus Baptistonia. 18-Outline of the dorsal sepal: 1 obovate—2 spathulate Assumed evolution: 1 > 2 * Degree of concaveness of the dorsal sepal (character 19) Several species of Baptistonia have a spoon-shaped dorsal sepal. Therefore, this character seems to be per- tinent within the genus Baptistonia as well as at the generic level. 19—dorsal sepal: 1 plane—2 cucullate Assumed evolution: 1 — 2 * Outline of the petals (character 20) In order not to overrate this character, I have coded for only two states. 20—Petals: 1 obovate or claviform—2 notched spathulate to panduriform. Assumed evolution: 1 > 2 * Apex of the petals and of the dorsal sepal (character 21) The form of the apices of the petals and of the dorsal sepal seems to be characteristic within Baptistonia. 918 Journal of the Botanical R h Institute of Texas 1(2) 21- Apex of the petals and of the dorsal sepal: 1 angular—2 rounded or retuse Assumed evolution: 1 > 2 D2. Labellum * Fused (character 22) The labellum may be fused to the base of the column or not. 22-labellum: 1 fused—2 articulated Assumed evolution: 1 > 2 e Claw (character 23) The presence of a long claw is characteristic for the genus Baptistonia. This character, evaluated by the factor R— being the ratio of the length of the claw over the length of the labellum—was GW coded for 3 states. 23-Labellum claw: 1 absent or nearly so (R< 1/20)—2 short (1/20 1/5) Assumed evolution: 1 => 2 3 * Shape of the lateral lobes (character 24) 24—Lateral lobes of the labellum: 1 somewhat elongated (rounded, triangular or square)—2 wider than long-—3 non-existent—4 very elongated (linear, elongated triangular, or linguiform) Assumed evolution: 1 >2 5 3 and 1 > 4 * Direction of the lateral lobes within the labellum plane (character 25) 25—Direction of the lateral labellum lobes: 1 to 90° away from the lip axis—2 toward the base of the lip—3 toward the apex Assumed evolution: 1 2 and 1 3 * Direction of the lateral lobes in respect to the position of the labellum plane (character 26) 26-Lateral lobes: 1 more or less within the plane of the labellum or slightly curved forward—2 at least their basal half distinctly directed backward Assumed evolution: 1 — 2 * Width of the midlobe (character 27) Evaluated in respect to the width of the labellum at the level of the lateral lobes, this character is variable within the genus Baptistonia. 27-Width of the labellum at the level of the midlobe: 1 greater—2 similar—3 smaller in comparison to the width measured at the level of the lateral lobes Assumed evolution: 1 > 2 > 3 * Shape of the midlobe (character 28) or of the labellum if it is entire. Besides being variable within the genus Baptistonia, this character seems to have a certain degree of importance in respect to generic differentiation. 28-Shape: 1 wider than long—2 just about circular or square—3 longer than wide Assumed evolution: 1 > 2 > 3 * Apex of the labellum (character 29) 29-Apex: 1 entire—2 split Assumed evolution: 1 > 2 e Isthmus of the labellum (character 30) I have accorded a great importance to this character as it is a potential differentiating character at the spe- cies level in Baptistonia. In fact, the character is stable and specific for groups of species. It overlaps slightly with character 24 (state 3 of character 24 is the same as state 5 of character 30). 30-Isthmus of the labellum: 1 elongate rectangular—2 trapezoidal—3 short and wide with rounded sinus—4 short with triangular sinus—5 absent (labellum undivided) Assumed evolution: 1 > 2 > 3 > 45 Chiron, Phylogenetic study of Baptistonia 919 * Margins at the sinus (character 31) A small number of species show a sinus with indented margins and this character appears to be stable and characteristic. 31-Margins at the sinus: 1 entire—2 irregular Assumed evolution: 1 2 * Callus (characters 32 to 34) Within the genus Baptistonia, the callus of the labellum is generally made of three parts: a basal part (on the claw), a center part (between the lateral lobes) and an apical part (on the isthmus and on the midlobe). According to Chase (1986), this is a derived condition in the Oncidiinae. In the species that do not have a claw, the callus has no basal part. In Oncidium, the callus does not extend onto the median labellum lobe. In Gomesa, the callus is made up of two parallel, longitudinal lamellae, followed by two crests in the apical part of the labellum. The diversity ofthe structures and their remarkable stability within each species have caused me to retain three distinct characters and have made me accord special importance to the median part. 32—Basal part: 1 absent or not differentiated, not ending in two teeth—2 ending in two well differentiated teeth Assumed evolution: 1 > 2 33-Median part made up of: 1 a compact construction of several well-developed teeth—2 some kind of verrucose and/or sheeted plate—3 a thick, non-verrucose plate, depressed in the center—4 a thin, smooth, inverted V-shaped plate—5 an inverted Y-shaped crest—6 a construction of small elongated wart-like struc- tures, more or less spaced in two rows—7 two large, low crests—8 two thin, high crests Assumed evolution: 1 > 2 => 3 > 4 —> 5and: 1 > 65758 Other character state polarities have been evaluated. This one has been retained because it is most congru- ent with other characters. 34—Apical part of the callus: 1 absent—2 short (< 1/3 of the midlobe)—3 long (> 1/3 of the midlobe) Assumed evolution: 1 > 2 > 3 D3. Column e Slenderness (character 35) This character is variable within the genera studied and seems to be pertinent for the genus Baptistonia. It is quantified by the ratio H/W between the height of the column and its width measured below the stigmatic cavity and coded according to the GW method for three states. 35-Column: 1 medium (2.5 € H/W «4.5)—2 stocky (H/W « 2.5)—3 slender (H/W > 4.5) Assumed evolution: 1 > 2 and 1 => 3 * Positioning in relation to the labellum (character 36) This character does not seem to be pertinent within the genus Baptistonia where it is remarkably stable, but it is useful to differentiate among the genera. 36-column: 1 in the same line as the labellum—2 at an angle of 90? to the labellum—3 tight against the basal part of the labellum Assumed evolution: 1 — 2 3 * Pubescence (character 37) Pubescence of the column is a character of all species within the genus Baptistonia, whereby the degree of pu- bescence varies. Furthermore, this character is useful for the study of the relationships among the genera. 37-column: 1 glabrous—2 slightly pubescent—3 strongly pubescent Assumed evolution: 1 > 2 3 * Tabula infrastigmatica (character 38) The absence of the tabula beneath the stigmatic cavity is common to all species of Baptistonia (although with a regression in B. uhlii). It marks an evolutionary step in relation to other genera. 920 Journal of the Botanical R h Institute of Texas 1(2) 38-tabula infrastigmatica: 1 well-developed—2 slightly developed—3 absent Assumed evolution: 1 > 2 > 3 * Shape of the wings (character 39) 39-wings: 1 independently of their shape, wider than long—2 absent or reduced to two minuscule points—3 longer than wide (sometimes becoming wider at the apex) Assumed evolution: 1 + 2 and 1 > 3 * Spreading of the wings (character 40) 40—wings: 1 opened to 90?—2 absent or opened to 180% —3 parallel or as an arch directed to the front Assumed evolution: 1 > 2 and 1 > 3 * Hairiness of the stigmatic cavity (character 41) The hairiness of the margins of the stigmatic cavity varies between nearly absent and very dense. 41—hairiness: 1 hairs nearly or fully absent—2 hairs rare and/or short—3 hairs long and dense Assumed evolution: 1 2 2 > 3 * Presence and shape of an appendix to the anther (characters 42 to 44) The apex of the anther can be rounded or elongated, more or less recurved upward and more or less split into two parts. I have retained three characters to describe this appendix where its absence or presence is incorporated into those three characters. 42—appendix: 1 absent to short—2 long Assumed evolution: 1 > 2 43-appendix: 1 simple—2 bifid—3 double Assumed evolution: 1 > 2 > 3 44—appendix: 1 straight —2 recurved Assumed evolution: 1 > 2 * Viscidium (character 45) The form of the viscidium, measured as the ratio D/d of the large and small diameter of the ellipse, seems to be pertinent at the generic level. 45-viscidium: 1 circular or ovate (D/d«3)—2 elliptic (D/d23) Assumed evolution: 1 — 2 * Tegula (character 46) 46-tegula of the pollinarium: 1 larger on the side of the viscidium or equally large on both sides—2 spoon- shaped (larger on the side of the pollinia) Assumed evolution: 1 > 2 * Margins of the clinandrium (character 47) Well-developed, irregular margins are characteristic for Baptistonia but little intrageneric variation can be observed. 47-margins of the clinandrium: 1 only slightly developed—2 extending beyond the anther Assumed evolution: 1 > 2 The evaluation of these 47 characters has made it possible to develop the data matrix shown in Table 1. This matrix was subsequently exploited in two different methods: parsimony and distances. Maximum parsimony analysis.—For the first method the phylogeny for the group was obtained by us- ing the software package PHYLIP (see Felsenstein 1989, 1993), which is composed of SEQBOOT version 3.570, MIX version 3.572c and CONSENSE version 3.56c. The coding of the characters corresponding to the evolutionary hypothesis delineated above resulted in a data matrix (not shown here) of 0 and 1. The software parameters were set as follows: SEQBOOT: number of replications set to 1000— morphological data Chiron, Phylogenetic study of Baptistonia 921 MIX: Successive parsimony method according to Wagner and to Camin-Sokal—set of 1000 replica- tions CONSENSE: Oncidium altissimum set as tree root. Neither the introduction of a threshold within the algorithm nor the random selection of the species had any significant impact on the results. Distance analysis. —For the second approach, I have used the software ADE-4 (see Thioulouse et al. 1997) to calculate a distance matrix on the basis of the data matrix given in Table 1 by means of the Manhattan method as performed by the DMAUtil-Quantitative Variables utility, after which the minimal length tree was computed by means of the UPGMA method as provided by the NGStat-Minimal Spanning Tree utility. RESULTS In respect to the two non-strict consensus phylogenetic trees obtained with the parsimony method (one for the Wagner parsimony and one for the Camin-Sokal parsimony), it can be noted that: (a) as a general rule, the bootstrap values are weak, some branches being supported by values below 5096; (b) there is a distinct separation of the genera Gomesa and Baptistonia within the Gomesa clade; (c) there are, among the species of Baptistonia, 6 small groups of species supported by relatively good bootstrap values (25096), groups that are referred to as complexes; (d) these diagrams suggest a low level of reliability for the other relationships (bootstrap «4096). I have attempted to define these 1 $ L: :u of taxa was reduced by replacing! the species by the corresponding complexes. The characters attributed to even further in a second step. For this purpose, the number any given complex were reconstructed on the basis of the characters of the constituent species as well as on the basis of the evolutionary analysis resulting out of these first diagrams. The topology of the initial trees is respected and the relationships among the different parts are consolidated (higher bootstrap values). Figure 1 represents the semi-strict consensus tree obtained by the combined operations described above. The bootstrap values for the complexes are those obtained in the first step using the Camin-Sokal method, and the bootstrap values of the higher groupings are those obtained in the second step, using the Wagner method. They are labeled above the branches and relate to the node situated to the immediate right. The nodes supported by bootstrap values below 4096 are not reproduced. The numbers related to a branch by a dotted line represent the synamorphies of the node ending that branch. Even though, as is usual when using a large data matrix, the consistency index remains low (0.29), the retention index is relatively high (0.78). This suggests that, in spite of the fact that there is a great deal of homoplasy in respect to the chosen characters, the synapomorphies are rather numerous (cf. Freudenstein & Rasmussen 1999). Figure 2 represents the distance tree obtained as described above. Its overall topology is close to that of the tree shown in Figure 1: (a) there is a large distance between the Oncidium group and the Gomesa group; (b) there is a great distance between these two groups and the rest of the species included in the study; (c) mnlos LS among the latter, we find the group “GMT” as well as th brieniana, silvana, pubes, leinigii, pulchella and truncata shown on Figure 1 (for all the specific names of Baptistonia, the taxonomic authorities are given in Appendix 1); (d) Oncidium trulliferum and Carriella colorata are placed at the base of the genus Baptistonia. The following discussion is based on the examination of Figure 1, but is overall compatible with Figure 2. DISCUSSION The genera Baptistonia and Gomesa share a number of characters which are not found in the members of the genus Oncidium. However, the present study also stresses the numerous synapomorphies shown by the members of each of the two genera. 1A | H £ J " n E J 3 | D £f The consolidation of y the nodes is similar. 1(2) £T IeXads5 £L JUUITIdI OF 922 Tague 1. Evaluated states for the different characters of species studied. sisuapueifou 'g ejeupind g seqnd *g Insqed 'g epniu ‘g lozjal|'g "brul “g Vorsyney *g eueipunojgnb *g eyeuitpe 'g louarewep 'g eyennp ‘g e1ofiuJ05 *g eueiuauq 'g Iouiqje *g ENEC CON MS O UST CON OE SONISCN ONE CG O Sere cn C NI SS ONS USE NTE ON M e e CM ONG e > Ecce SS Rr CN ea NIS iA E AECA RON er E A e A ES BE a cie O Ue NON SS SS E SS r CN EON er a ee a (N CONS LEE C EM ee ER EM rM d CN sce AR CN, cp oe yos €) OQ O ve (N (m st in o 8s GQOrrrrrrerre SEU IET UN AR LE ee neri red 923 Chiron, Phylogenetic study of Baptistonia uniejina 'Q uinuiissnje ‘0 winsojnqau ‘9 uaneq ‘0 SIJISSOS *9 PAINDAI “5 eds “5 euldje *5 eJe10]0) *) !USI6pim *g euelua)JaA 'g Iun *g eje»ung) 'g eUeAJIs “g sapooJes 'g D LE M LU LL LLL LL LM IE LE M LL d E LL a E a E == 47 924 Journal of the Botanical R h Institute of Texas 1(2) B. brieniana B. albinoi B. riograndensis B. pubes B. lietzei 14,18,24,26 | B. silvana 27,30,33 B. gutfreundiana B. cruciata B. pabstii B. velteniana i I I I I I I I I I B. damacenoi l= l l | l l | | | I l I | B. leinigii Baptistonia B. cornigera ! 01,23,24 60 30,32,33 echinata pulchella uhlii 05,23,24,37 widgrenii nitida kautskyi B. B. B. B. MEM B. sarcodes B. B. B. truncata : 07,11,16,25,28, > O. trulliferum —— 02,03,04,21,39 C. colorata G. recurva l fo G. alpina 09,10,12,13,15,16,17 i —Lr G. crispa 22,24,33,36,39,45 GL sessili Gomesa IC. 02,06,33,36,38 O. baueri O. nebulosum groupe externe O. altissimum Cami l . un synapomorphies. Chiron, Phylogenetic study of Baptistonia 55 B. brieniana 13 B. riograndensis B. albinoi B. pubes B. lietzei B. damacenoi B. silvana B. gutfreundiana B. cruciata 41 88 B. velteniana 61 B. leinigii 20 B. pabstii B. cornigera B. nitida B. widgrenii B. sarcodes 31 B. echinata 21 B. uhlii B. pulchella = B. truncata 100 | | B. kautskyi O. trulliferum C. colorata G. sessilis 100 | » crispa G. recurva G. alpina O. baueri O. nebulosum La altissimum 100 GMI r: Fic. 2. Distance tree J £L Dat PRA D hi PEA 926 Journal of f Texas 1(2) Synapomorphies of the genus Gomesa: * QO—elliptic stomata; * 17—floral bracts very long; * 10-short inflorescence; * 22-articulated labellum; * 12—inflorescence not branched; * 24—entire labellum; * 13—formation of buds as soon as the inflorescence — * 33-callus consisting of two longitudinal crests; has been generated; * 36-column pressed against the labellum; * 15—vertical shape of the flowers ovate; * 30—wings almost non-existent * ]6-flowers without reddish violet spots; * 45—viscidium elliptic. Synapomorphies of the genus Baptistonia Gwith the e of Carriella colorata and Oncidium trulliferum): * 03—pseudobulbs faintly flattened; 21-apex of the petals and of the dorsal sepal not * 04—horizontal section of the pseudobulbsrounded; angular; * 05—pseudobulbs and leaves dark green; * 37-column pubescent; * 06—basal sheaths non-leafy; * 39—column wings much longer than wide; * 08—density of stomata low; * 46—tegula spoon-shaped * 47-margin of the clinandrium well-developed. Other synapomorphies may also be relevant, but with reversals in certain taxa: * 02—pseudobulbs cigar-shaped (except in B. uhlii, B. | * 42-appendix of the anther long (except in B. sar- truncata, B. nitida and B. widgrenii); codes, B. nitida and B. widgrenii); * |9—dorsal sepal cucullate (except in B. uhlii, B. * 44—appendix recurved (except in B. nitida and B. widgrenii, B. albinoi and B. riograndensis); sarcodes). * 23-clawed lip (except in the echinata complex); Finally, we may note the evolutionary steps shown by the two genera in relation to the genus Oncidium. However, it is not possible to consider them as synapomorphies of the Gomesa clade. To do so it would be necessary to ascertain whether the other groups that are part of this clade would share these synapomor- phies, a step that would go beyond the scope of this study: * 02—elongation of the pseudobulbs; * 36-diminuation of the angle between the column * 06—shortening of the basal sheaths; and the labellum; * 33—modification of the median part of the callus; — * 38—disappearance of the tabula infrastigmatica. Oncidium trulliferum shares 10 of the 16 synapomorphies of the genus Baptistonia and seems to be integrated into this genus. This grouping, which could possibly be designated as Baptistonia sensu lato, is supported by a bootstrap of 5896. This value is fairly low and is caused by the fact that Carriella colorata is found on some of the most parsimonious trees together with one or another group of species belonging to Baptistonia sensu lato. Carriella colorata is placed within the phylogenetic reconstruction as the sister species of this group, the whole being supported by a bootstrap value of 9696. I will now review the six complexes mentioned above: The “truncata” complex [B. truncata and B. kautskyil: bootstrap value 99%. This complex is defined by five synapomorphies that are found only in this group: * 16-ground color of the flower is pale; * 244+25—the lateral lobes of the labellum are short and directed toward the apex of the labellum; * 30-the isthmus of the labellum is short with triangular sinuses; * 40—the wings of the column are spread at 180? Other characters are shared with some of the other groups: * Ol-pseudobulbs not very tall (shared with the echinata complex, with B. widgrenii and with Carriella colorata); * qe 07—pseudobulbs unifoliate (shared with the pulchella complex and with Carriella colorata); * 11-inflorescence growing downward (shared with B. pulchella and B. uhlii); Chiron, Phylogenetic study of Baptistonia 927 * 28—width of labellum midlobe equal to its length (shared with the brieniana complex); * 33—median part of the callus made of a thin, smooth, inverted V-shaped plate (shared with the brieniana complex, with the pubes complex and with B. cruciata); * 34—callus not extending onto the midlobe (shared with B. cornigera and with B. sarcodes). According to my analysis, this complex is at the basis of the genus Baptistonia, and is separated from the other members by a branch that is relatively well supported (6096) and defined by two synapomorphies: * 41— stigmatic cavity with more or less long hairs; * 43—appendix of the anther bifid or double. The “silvana” complex [B. silvana and B. gutfreundianal: 8696. This group is defined by a combination of shared characters each of which can also be found in other species: * 18-dorsal sepal spathulate (shared with the pubes complex and with B. nitida); * 24—lateral lobes of the labellum much longer than wide (shared with the pubes and brieniana complexes as well as with three other species); * 26—base of the lateral lobes distinctly folded backward (shared with the pubes and pulchella complexes, and with B. cruciata); * 27-labellum midlobe wider than the width of the labellum as measured over the lateral lobes (shared with B. nitida); * 30-isthmus of the labellum short and wide with rounded sinuses (shared with B. sarcodes, B. widgrenii and with the echinata complex); * 33—median part of the callus consisting of a thick, non-verrucose plate that is indented at the center (shared with B. cornigera). The “pulchella” complex [B. pulchella and B. uhlii]: 81%. One synapomorphy: * 43—appendix of the anther is bifid; and shared characters, also found in other species: * O7—pseudobulbs unifoliate (shared with the truncata complex and with Carriella colorata); * |O-inflorescence short (shared with B. truncata and with Carriella colorata); * ]l-inflorescence growing downward (shared with the truncata complex); * 26—base of the lateral lobes of the lip distinctly folded backward (shared with the silvana and pubes com- plexes, and with B. cruciata); B. echinata is positioned as a sister species to this complex (bootstrap of 4796) with one synapomorphy: * 33—median part of the callus made up of two low but wide crests; and several shared characters, that are, however, also shared with other species: * Ol-pseudobulbs not very tall (shared with the truncata complex, with B. widgrenii and with Carriella colorata); * 23-claw very short (shared with B. sarcodes, Carriella colorata and Oncidium trulliferum), * 24—width of the lateral lobes of the lip equal to their length (also found in B. sarcodes and Oncidium trul- liferum); * 30-isthmus short and wide with rounded sinuses (shared with B. sarcodes, B. widgrenii and the silvana complex); * 32—basal part of the callus without teeth. It should be noted, however, that, in respect to the distances, B. echinata is the closest neighbor of B. sarcodes, and this pair of species is found to be a sister group of the pulchella complex. The “brieniana” complex [B. brieniana, B. riograndensis and B. albinoil: 79%. This group also ha synapomorphies; it is defined by the following combination of characters: * 09-circular stomata; * 20—petals spathulate to panduriform (shared with B. widgrenii and the pubescomplex); * 28—width of the labellum midlobe equal to its length (shared by the truncata complex); 928 Journal of the Botanical R h Institute of Texas 1(2) * 30-isthmus of the labellum long and rectangular (shared with the pubes complex as well as with B. nitida and B. cornigera); * 33—median part of the callus made up of a thin, smooth, inverted V-shaped plate (shared with the pubes complex, with the truncata complex and with B. cruciata); * 54—apical part of the callus long (shared with the pubes complex and with B. widgrenii). The “leinigii” complex [B. leinigii, B. pabstii and B. veltenianal: 78%. This group is characterized by two synapomorphies: * 30-isthmus trapezoidal * 33—median part of the callus made up of a verrucose, sheeted plate. The “pubes” complex [B. pubes, B. lietzei and B. damacenoil: 7496. Here again, the group is characterized only by shared characters also found in other species: * 20—petals spathulate to panduriform (shared with B. widgrenii, Oncidium trulliferum and with the brieniana complex); * 30-isthmus of the labellum long and rectangular (shared with the brieniana complex as well as with B. nitida and B. cornigera); * 33-median part of the callus consisting of a thin, smooth, inverted V-shaped plate (shared with the brieniana and truncata complexes and with B. cruciata); * 34—apical part of the callus long (shared with the brieniana complex and with B. widgrenii). In my reconstruction, the pubes and silvana complexes are sister groups, although with a weak bootstrap value (4496). The two groups are also close to each other in the distance tree. The four synapomorphies that characterize this clade are all characters of low importance: * 14—flower form; * 18—dorsal sepal spathulate (shared with the silvana complex, as well as with B. nitida, Oncidium trulliferum and Carriella colorata); * 24—form of the lateral lobes of the labellum; * 26—base of the lateral lobes distinctly folded backward (shared with the silvana and pulchella complexes, and with B. cruciata). Together, the pubes, brieniana, silvana, and leinigii complexes form a monophyletic group, here referred to as « GMI ». This group, although weakly supported (bootstrap value: 4296), is characterized by the following synapomorphies: * 23-nail of the labellum long; * 32—basal part of the callus ending in a pair of diverging teeth (shared with B. echinata). Baptistonia cruciata is included in « GM1 » but the analyses did not allow a precise clarification of its position. Baptistonia cornigera appears as a sister species of this GM1 group (bootstrap of 46%). Finally, it should be noted that, on the initial consensus tree, B. widgrenii appears as a sister species to the == entity formed by B. echinata and the « pulchella » complex, with a very weak bootstrap value (26%) and the shared characters 01 and 30. However, this grouping, besides not supported, would probably be artificial because (a) the structure of the pseudobulbs is very different although the pseudobulbs of B. widgrenii are short, and, like those of the other species in this group, much shorter than the pseudobulbs of the other Baptistonia species; and (b) the lateral lobes of the labellum are very different although the isthmus is the same. Furthermore, B. widgrenii does not share any real synapomorphies with this group. Therefore, the parental relationships of B. widgrenii remain to be clarified. CONCLUSIONS The present study supports monophyly of Baptistonia as defined by Chiron and Castro Neto (2004) as well as its distinctiveness from Gomesa based on numerous morphological and anatomical characters. Chiron, Phylogenetic study of Baptistonia 929 The proper place of the two species added to the bulk of the specimens to be studied, Carriella colorata and Oncidium trulliferum, is difficult to define on the basis of this study alone. Oncidium trulliferum is posi- tioned at the base of the genus Baptistonia and should probably be integrated into that genus (bootstrap of 58%). Carriella colorata should rather be regarded as a sister species to the genus. It is, however, better to wait for the results of molecular studies in progress that may clarify this issue. The phylogenetic relationships within the genus Baptistonia are poorly resolved due to the low number of synapomorphies defining the complexes and to homoplasy. However, some monophyletic groups can be distinguished. Baptistonia kautskyi and B. truncata are strongly supported as sister taxa, and are sister to all other spe- cies of Baptistonia. The majority of the complexes defined above have been discussed in detail by Chiron and Castro Neto (2005a, 2005b, 2006a, 2006b). One could argue that each of these complexes represents nothing but the expression of the existence of different populations of the same, rather variable species. As many of the combinations of morphological characters that distinguish the members of a complex as well as their dis- tribution zones do not form a continuum but rather disjointed conglomerates, I come, however, to another conclusion. APPENDIX 1: SPECIMENS EXAMINED Living material (Vouchers have been deposited in LY (and/or SP when noted) for all plants). Baptistonia albinoi (Schltr) Chiron & V.P. Castro: Chiron 2578, Chiron 0084, Chiron 0085, Chiron 0086. B. brieniana (Rchb.f) V.P. Castro & Chiron: Chiron 04604, Castro Neto s.n., Paraguay s.n. B. cornigera (Lindl.) Chiron & V.P. Castro: Chiron 03017, Chiron 03035, Chiron 02549, Chiron 03069, Chiron 03008, Chiron 03066, Chiron 05206, Chiron 06576, Chiron 07075, Chiron 3051, Chiron 3052, Chiron 3062. B. cruciata (Rchb.f) V.P. Castro & Chiron: Chiron 05441, Chiron 05446, Chiron 05452, Zézé s.n., Uhl s.n., Jardin botanique de Lyon 020539. B. damacenoi Chiron & V.P. Castro: Chiron 03208, Chiron 2273, Chiron 2589 (type, SP), Frey746. B. echinata Barb. Rodr.: AOSP s.n. ex Chiron 0042, AOSP s.n. ex Chiron 0043, Jardin botanique de Sáo Paulo s.n. ex Chiron 0065 and Chiron 0066, Chiron 2582, Chiron 3044, Chiron 3045, Chiron 3055, Chiron 3063. B. gutfreundiana (Chiron & V.P. Castro) Chiron & V.P. Castro: Chiron 2240, Chiron 03414, Chiron 05844, Chiron 05848, Castro Neto s.n. ex Chiron 0058, Chiron 3060, Castro Neto s.n. (type, SP). B. kautskyi (Pabst) V.P. Castro & Chiron: Chiron 03194, Chiron 04813, Chiron 04814, Chiron 04816, Chiron 05769, Chiron 06503, Frey 741, Frey 1079, Caliman s.n. B. leinigii (Pabst) V.P. Castro & Chiron: Chiron 0101, Chiron 07009, Chiron 07011 Chiron 07012, Chiron 07015, Chiron 07017, Chiron 07018, Chiron 07019. B. lietzei (Regel) Chiron & V.P. Castro: AOSP s.n. ex Chiron 0041, AOSP s.n. ex Chiron 0044, AOSP s.n. ex Chiron 0045, Castro Neto s.n. ex Chiron 0046, Chiron 06504, Chiron 06505, Chiron 2273, Chiron 2283, Chiron 07026, Campacci s.n. B. nitida (Barb. Rodr.) V.P. Castro & Chiron: Chiron 2591, Castro Neto s.n. ex Chiron 0039, Castro Neto s.n. ex Chiron 0040, pou 2592, Chiron e Chiron 2596, Chiron 05532, Chiron 06540. B. pabstii (Campacci & C. Espejo) V.P. Castro & ore Chiron 3042, Chiron 3059. B. pubes (Lindl.) Chiron & V.P. Castro: Chiron 3034, Chiron 3036, Chiron 3040, Chiron 3046, Chiron 3047, Chiron 3048, Chiron 3053, Chiron 3054, Chiron 06506, Chiron 2241, Castro Neto s.n. B. pulchella (Regel) Chiron & V.P. Castro: Chiron 05466, Chiron 05487, Chiron 05490, Chiron 05498. B. riograndensis (Cogn.) V.P. Castro & Chiron: Chiron 07060, Chiron 07061, Chiron 07062, Chiron 07063, Chiron 07069, i oe B. oe. leg) Chiron & V.P. Castro: Chiron 04850, Chiron 04852, Chiron 05218, Chiron 06570, Chiron 2242, Chiron li silvana (V.P. Castro & Campacci) V.P. Castro & Chiron: Chiron 2890, Chiron 3049, das os Chiron 05600 Chiron 05807, Chiron 06509, Castro Neto s.n. ex Chiron 0057. B. truncata (Pabst) Chiron & V.P. Castro: Chiron 05427, Chiron 2785, Chiron 2262, Chiron 2264, Chiron 2769, Chiron 2899A, Chiron 2899B, Chiron 2899C. B. uhlii Chiron ave dit Chiron Bur fees SP), Chiron 06511, Chiron 06512, Chiron 06534. B. velteniana V.P. Castro & Chiron: Kautskyi s.n. (type, SP), Uhl s.n. ex Chiron 0028 and Chiron 0029. B. widgrenii (Lindl.) V.P. Castro & Chiron: Chiron 0112, Chiron 2243, Chiron Fn Chiron ne Chiron o oli colorata (Kóniger & J.G. Weinm.) V.P. Castro €: K.G. Lacerda: Caliman s.n., Castro Neto s.n., Chiron 2796, Chiron 06508. Gomesa alpina Porsch: Jardin botanique de Sáo Paulo s.n. (SP), Chiron 05346, Alambari s.n. G. crispa Klot s ex ee Chiron ne Chiron 05481, Jardin botanique de São Paulo s.n. (SP). G. recurva R. Br.: Chiron um Chiron 04683, Chiron 04702, Chiron 04810, Chiron 04811, Jardin botanique de Sáo Paulo s.n. (SP). G. sessilis Barb. Rodr.: Chiron 04695, Chiron 05388, Chiron 05398, Chiron 05702, Chiron 05715. O. altissimum Swartz: Chiron 0834, Chiron 1919, Jardin botanique de Lyon s.n. Oncidium baueri Lindl.: Chiron 02041. O. nebulosum Lindl.: Chiron 99041. O. trulliferum Lindl.: Jardin botanique de São Paulo 7028 (SP), Jardin botanique de Sáo Paulo 11008, Chiron 05249, Chiron 0107, Chiron 0108, Chiron 0109. D, IN A40? .B. UUIO AT a Herbarium material Baptistonia albinoi: MBM 209548, MBM 99608, MBM 6225, MBM 48898, MBM 48896, MBM 29392, MBM 22869, SP 25185, SP 31577, W-R 35126. B. brieniana: W-R 16453 (type de Oncidium brienianum), W-R 25704, W-R 7031, P 00430186 (type de Oncidium verrucosissimum). B. echinata: W-R 10660, W-R 20585, W-R 20586 (type de Oncidium brunleesianum). B. lietzei: LE £L Dag PRA D hi PEA 930 Journal of f Texas 1(2) s.n. (type de Oncidium lietzei), W-R 25721 (type de Oncidium hrubyanum). B. leinigii: SP 334938, SP 341393. B. pubes: W-R 13085, W-R 28099, W-R 25713. B. pulchella: W-R 28923. B. riograndensis: ICN-Dutra 925, SP 363181 (type de Oncidium cassolanum). B. sarcodes: K-Lindley no.84 iud de Oncidium sarcodes), MBM 10148, MBM 218812, W-R 44709, WU s.n. B. widgrenii: K- inue n?50 (type de Oncidium widgrenii), SP 46502, W-R 35291, W-R 10423. Gomesa crispa: W-R 45401, xm R 45361. G. re a: W-R 34429, W-R 14023, W-R 11818. Oncidium trulliferum: MBM 7758, MBM 10717, MBM 48903, MBM 226953, W-R pisce R 6817, W-R 3319. = ACKNOWLEDGMENTS The author is indebted to Philippe Choler, Université Joseph Fourier—Grenoble I, for his cooperation in the computation of the phylogenetic tree in figure 2; to Guido Braem, University of Maryland, University College Europe, for his helpful discussions; and to Mark Whitten and Norris Williams for providing critical comments on the manuscript. The study of the herbarium materials was made possible by the hospitality demonstrated by the curators of ICN, K, LE, MBM, P, SP, W, and WU to whom I extend my sincere gratitude. My stay at Vienna was supported by the SYNTHESYS Grant AT-TAF-2736. Access to live specimens was made possible through the collaboration of my Brazilian friend Vitorino P. Castro Neto and through some of his local contacts, especially: Carlos Régent, Sergio Gutfreund, Sydney Marcal, and Thomas Adamski. I am deeply indebted to each and every one of them. 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Molecular systematics of the Oncidiinae based on evidence from four DNA sequence regions: expanded circumscriptions of Cyrtochilum, Erycina, Otoglossum, and Trichocentrum and a new genus (Orchidaceae). Lindleyana 16:113-139. 932 Journal of the Botanical R h Institute of Texas 1(2) BOOK NOTICES Top E Stuessy, VERONIKA Mayer, and Elvira HORANDL (eds). 2003. Deep Morphology: Toward a Renaissance of Morphology in Plant Systematics. (ISBN 3-906166-07-4, hbk.). Regnum Vegetabile: Vol. 141. Gantner Verlag, Liechtenstein. (Orders: Koeltz Scientific Books, D-61453 Koenigstein, PO. Box 1360, Germany; koeltz@t-online.de; www.koeltz.com). $156.00, 326 pages, illus. ca. illus., 6 1/4" x 9 1/4". After an introduction (Chapter 1), the chapters are divided into three sections: (2-5) Genetics and Development, (6-8) Phylogenetic Analysis, (9-13) Ecology and Adaptation, with a final overview (Chapter 14). Table of Contents 1 MA 7] E T I l g and why is it time for i i i dd ics? (Weber). 2.Th SE : TE RE T6 fd ES " nd z Gailing). 3. A m effects on floral M and function: a revie (Digg le). 4. Floral iR AM features ane med na in US M ag and p 5C l hyl 6. o and ee evolution. (Williams and di 7. What should a *complete" pasion Ee ds a dd 8. Beyond morphoclines and trends: f (Hufford and McMahon). and Neinhui 10. Toward a deeper und li d df ion i llen grains: the sporoderm (Hesse). 11. Ecological adaptions and deep phylogenetic splits—evidence and ens n the Nn xylem (Baas, Jansen and Wheeler). 12. The potential of plant biomechanics in functional biology and systematics (Speck, Rowe, Civeyrel, ClaBen- Bockhoff, Neinhuis, and Spatz). 9. Epicuticular waxes and vascular huh systematics: integrating micr NOTOS ood and TE data (Bart Theisen, Borsch s). 13. How a better understanding of adaptions can yield better use of morphology in plant systematics: toward Eco-Evo-Devo (Givnish). 14. Morphological data in pl ics (S STEVEN R. RADOSEVICH, JODIE S. Hot, and CLaubio M. Guersa. 2007. Ecology of Weeds and Invasive Plants: Relationship to Agriculture and Natural Resource Management (ed. 3). (ISBN 978-0-471-76779-4, hbk.). John Wiley & Sons Inc., One Wiley Drive, Somerset, NJ 08875, U.S.A. (Orders: www.wiley.com, 877-762-2974, 1-800-597-3299 fax). $74.95, 454 pp., b/w photos, line drawings, 6 1/4" x 9 1/4". Table of Contents Weeds and Invasive Plants (definitions and classificati i ducti d wildl F W ildlands). . Principles of Weed and Invasive Plant Ecology. Invasibility of Agricultural and Natural Ecosystems. Evolution of Weeds and Invasive Plants. Weed Demography and Population Dynamics. Plant-Plant Association Weed and Invasive Plant Management Approaches, Methods and Tools. Herbicides Systems Approaches for Weed and Invasive Plant Management. e UU UR ev J. Bot. Res. Inst. Texas 1(2): 932. 2007 MUHLENBERGIINAE (POACEAE: CHLORIDOIDEAE: E YNODONTEAE): FROM NORTHEASTERN MÉXICO Paul M. Peterson Jesus Valdés-Reyna Department of Botany e de Botánica National Museum of Natural History Univerisid ma Agraria “Antonio Narro” mithsonian Institution Buenavista, Saltillo diis DC 20013-7012, U.S.A. Coahuila 25315, MÉXICO eterson@si.edu jvaldeseuaaan.mx Yolanda Herrera Arrieta Instituto Politécnico Nacional CIIDIR Unidad Durango-COFAA Durango 34220, MÉXICO yherreragipn.mx ABSTRACT A taxonomic treatment of the subtribe ardid for northeastern México (Coahuila, Nuevo León, and Tamaulipas) is given. A total o dy area in four genera: Aegopogon (1), Blepharoneuron (1), Lycurus Q), and Muhlenbergia (35). The names, aa medía var. ramosa ees and Muhlenbergia spiciformis Trin., are lectotypified. A key for piacere the ee x distributions illustrations, synonymies, and a E n M és 3 o RESUMEN Se presenta el estudio taxonómico la subtribu Muhlenbergiinae para el noreste de México (Coahuila, Nuevo León y Tamaulipas). En total se reconocen 39 ti n el área de estudio pertenecientes a cuatro géneros: Aegopogon (1), Blepharoneuron (1), Lycurus (2), y Muhlenbergia (35). n nombres, Muhlenbergia glomerata var. ramosa Vasey y Muhlenbergia spiciformis Trin., son lectotipificados. Se incluyen una clave para determinar las especies así como descripciones, distribuciones, especimenes examinados, ilustraciones y sinonimia para cada especie. Se incluye una breve discusión indicando las relaciones entre todas las especies de Muhlenbergia para el noreste de México. Coahuila, Nuevo León, and Tamaulipas or northeastern México covers an area of 291,955 km? or 15 96 of the total land of México. This area includes portions of two natural regions known as the Chihuahuan and Tamaulipan Deserts. These regions are considered a center of origin and diversification of arid and semi-arid plant species (Davila-Aranda et. al. 2004; Peterson et al. 2007). As part of the current revision of the grass flora of northeastern México, an examination of the taxonomy and distribution of the species of subtribe Muhlenbergiinae, was begun to aid the agriculture and livestock industries. This study treats 39 species currently placed in four genera: Aegopogon (1 sp.), Blepharoneuron (1 sp.); Lycurus (2 sppJ, and Muhlenbergia (OD spp.). ae Muhlenbergiinae is characterized by having spikelets perfect, staminate or sterile; occasionally with cleistogenes in the leaf sheaths; inflorescence paniculate of spicate main branches or a single raceme; spikelet bearing axis disarticulates falling entire or is persistent; spikelets solitary, paired, or in triplets, occasionally second, 1—3(-6)-flowered; glumes awned or unawned, 1-4-nerved; lemmas 3-nerved (rarely ]-nerved), awned or unawned; and a base chromosome number of x = 8-10. An ongoing phylogenetic study of the Muhlenbergiinae based on nuclear and chloroplast DNA sequences is currently in progress (Columbus & Peterson, in prep). Ina preliminary analysis, Muhlenbergia appears to be paraphyletic since the remaining nine genera in this subtribe are nested within its clade (Peterson et al. 2001a; 2004; 2007). Subtribe Muhlenbergiinae (Duvall et al. 1994; Peterson 2000; Peterson et al. 1995, 1997, 2001b, 2007) currently consists of 10 genera: Aegopogon with four species; Bealia mexicana Scribn., a monotypic genus J. Bot. Res. Inst. Texas 1(2): 933 — 1000. 2007 934 Journal of the Botanical R h Institute of Texas 1(2) from northern Mexico (Peterson 1989; Peterson et al. 1993); Blepharoneuron with two species (Peterson & Annable 1990, 2003); Chaboissaea with four species, three from central México and C. atacamensis (Parodi) P.M. Peterson € Annable from Argentina and Bolivia (Peterson & Annable 1992; Peterson & Herrera Ar- rieta 1995; Sykes et al. 1997); Lycurus with three species, including the amphitropical disjuncts, L. setosus (Peterson & Morrone 1997); Muhlenbergia with 147 species centered in northern Mexico and the southwest- ern United States (Herrera Arrieta & Peterson 2007; Peterson 2000, 2003), containing the important range grass M. montana and the amphitropical disjuncts M. arenicola and M. torreyi (Peterson & Ortíz-Diaz 1997); Pereilema with four species, two from western, central, and southern México; Redfieldia flexuosa (Thurb. ex A. Gray) Vasey, a monotypic genus and southwestern United States endemic of probable hybrid origin; Schaffnerella gracilis (Benth.) Nash, a monotypic genus known only from San Luis Potosí, México (Columbus et al. 2002); and Schedonnardus paniculatus (Nutt.) Trel., a monotypic genus and amphitropical disjunct with spicate primary branches. As currently understood for the New World Chloridoideae, tribes Cynodonteae and Zoysieae are sister, and sister to this clade are members of the Eragrostideae (Columbus et al. 2007; Peterson et al. 2007). All three tribes are supported by DNA sequence data from the chloroplast (matK, trnL—F region) and nuclear (ITS) genomes (Hilu & Alice 2000, 2001; Columbus et al. 2007). Relationships among the ten subtribes within Cynodonteae, as well as the 25 unplaced genera, are uncertain pending additional study. At this point we have no clear idea as to the relationships among these ten Cynodonteae subtribes. However, we do have good molecular support for maintaining the tribe Cynodonteae and morphological support for all of the ten subtribes. The following | contains a key for determining the species, descriptions, distribution, specimens d, illustrations, synonymies, and a brief discussion indicating hypothesized relationships among all native and adventive species of Muhlenbergiinae in northeastern México. This study is based on the examination of herbarium specimens from ANSM, ARIZ, B, BA, BAA, BM, CIDIIR, COCA, ENCB, GH, HINT (George S. Hinton's personal herbarium), IBUG, INEGI, K, LE, MEXU, MICH, MO, MSC, NMC, NMSU, NY, P, RSA, SL SEPM, TAES, TEX, UAT, US UT, UTC VT, and W; including the type specimens of most of the species studied. KEY TO THE SPECIES OF MUHLENBERGIINAE IN NORTHEASTERN MÉXICO 1. Plants annual. 2. Lemmas unawned 3. Ligules with lateral lobes extended into auricles; leaf blades with margins and midveins whitish- es basally, ia mu panicle branches diverging 80-1009 from the rachises; pedicels m long, gl throughout or obscurely puberulent, the hairs ipd m lon Muhlenbergia fragilis Be ees ae Yout | Ho! tended int da NI. ith | | lol longer than the body; fl i i le branches diverging 25- 80° from the rachises; pedicels 2-7 mm long; ee a near Ks apex, the hairs 0.1-0.3 mm long 19. Muhlenbergia minutissima DE s awned, the awns 3-40 mm lon 4. Spikelets in clusters of three, the 20 spikelet perfect and the two lateral spikelets ano and staminate or sterile; lemmas usually with lateral nerves extending into aw . Aegopogon tenellus 4. DET. not in clusters of three, or if in clusters then in pairs; lemmas usum lateral n tendi into a 5. ne glumes 2-nerved, minutely to deeply bifid, the teeth aristate or with awns to 1.3 mm long; e often in sessile-pedicellate pairs; disarticulation at the base of the pedicels; MS with s 6-15 mm long, sti 8. Muhlenbergia depauperata 5 p n 1 Da or eos [IDE eu MED. ot with a single mucro or awn; spikelets | inal -40 mm long, delicate straight Ung to flexuous. 6. Glumes 1.2-2.8 mm long, acute to acuminate; upper glumes 1.5-2.8 mm long __ 34. Muhlenbergia tenuifolia Peterson et al., Muhlenl ii f tl t M 935 6. Glumes 0.3-1.3 mm long, obtuse to acute; upper glumes 0.5- 7. Cleistogamous panicles with 1 open and not densely flowered, 1 the rachises; ligules truncate to o 1.3 mm long. -3 pus present in the axils of the lower sheaths; puc m wide; primary branches ir en to hle naa microsperma 7. Cleistogamous panicles absent in i axils of the lower sheaths; panicles narrow, contrac 0.6-2.8 cm wide; primary branches spreading up to 30? from the rachises; ligul acu 32 DUREE spiciformis . Plants perennial. 8. Plants rhizomatous, rhizomes oft 9. Upper g stout, scaly, and creepin g. lumes always 3-nerved, the apex usually with 2-4 small teeth; old sheaths flattened, ribbon-like or papery, sometimes spirally coiled lumes greenish-plumbeous, the surface scabrous, usually with a few short hairs below; pan- icles 0.5-2 cm wide, the primary branches appressed to spreading up to un from the ra Muhlenbergia quadridentata 10. Glumes whitish, stramineous or SUE green sometimes with EE M the surface shiny, usually glabrous to rarel belo OW; panicles 0.5-6 cm wide, the branches appressed to spreading up to 45? from the rachises Muhlenbergia virescens per glumes edil ]-nerved or sometimes 2- or 3-nerved, but the apex never ms old sheaths MAN flattened 11. T the awns 1-20(-25) mm 12. Lemma awns 0.1-3(-5) mm long, straight; eaei ein the internodes — 13. Muhlenbergia glauca 12. Lemma awns 10-20(-25) mm long, flexuous; leaf sheaths mostly shorter than internodes 21. Muhlenbergia polycaulis 11. Lemmas unawned or mucronate, the mucros to 1 mm long; anthers yellow, purple, greenish or reddish. 13. Panicles narrow Ic Open 4- lo cm wide. Panic le bran to 40° fi red, | not capillary, ightly appressed or loosely spreading up | UVES VLG and along the margins on proximal 2/3 to % 15. Muhlenbergia jaime-hintonii 14. Panicle branches capillary, diverging 30-90(-100)* from the rachises at maturity; paleas glabrous. 1 5. Ligules 0.5-2 mm long, hyaline, with well-developed lateral lobes or P bla margins and midveins prominent, whitish, thicken aa arenacea 15. Ligules 0.2-1 mm long, ciliate, without lateral lobes or ‘or auricles; sera margins an idveins not conspicuously thickened, greenish 13. Panicles contracted, narrow, 0.1-3.9 cm 16. Spikelets 1.4-2.4 mm lon 17. Culms 44-82 cm Ms leaf blades 5-22 cm long, 1 long 15. Muhlenbergia jaime-hintonii 17. Culms 4-30 cm tall; leaf blades 0.2-1.8 mm wide; panicles 1-5 cm long 18. Lemmas and paleas glabrous or with minute appressed pubescence, the hairs about 0.1 mm long; leaf sheaths 0.3-2.4 cm long; panicles partially included i the upper sheaths; ligule apices truncate Muhlenbergia utilis 18. Lemmas and paleas densely villous, the hairs less 0.4-1 mm long; leaf sheaths 15 cm long; panicles usually on an exserted peduncle; ligules apices acute 37. Muhlenbergia villiflora var. villiflora 7. Muhlenbergia asperifolia wide 8-5 mm wide; panicles 13-34 cm 16. Spikelets 2.4-8 mm long. 19. Glumes 3-8 mm long anang the awns), 1.3-2 times longer than us lemma; lea blades 2- UN vide, fl 0.4-0.8 mm lon 19. Glumes 1 Muhlenbergia racemosa -36mm Io Y to as long as the lemma; leaf blades 0.5- 6 mm wide, flat to mou anthers 0.7- ib mm eng mas with ta 0 m long on the lower 1⁄2 along the midveins Pa margins; leaf Ps m 42 cm AAA not arcuate; anthers 1.8- ri mm lo orange 20. Lemmas glabrous or 3. Muhlenbergia glauca the hairs less than 0.1 mm long; leaf blades 0.4-6 cm long, somewhat arcuate spreading; anthers 0.7-1.4 mm long, yellow to purplish _ 27. Muhlenbergia repens 936 Journal of the Botanical R h Institute of Texas 1(2) 8. Rhizomes absent 2] T glumes 3-nerved; old sheaths flattened and sometimes spirally twisted near base No 2. Upper glumes 3-toothed and 3-awned, the teeth (including the awns) 1/3 to Y p en of the glume, and the awns up to 1.7 mm long pets montana 22. Upper glumes 2-4-toothed, the teeth small and unawned, less than 1/6 T length oft glume. 23. Glumes a plumbeous, the surface scabrous, usually with a few short hairs below; panicles 0.5-2 cm wide, the primary branches appressed to spreading up to 30° from the n 25. Mahlenbergia quadridentata 23. Glumes whitish, stramineous or grayish ith pluml the surface shiny, usually glabrous below; panicles 0.5-6 cm wide, the branches a: " ps ° from the rachises hlenbergia virescens . Upper glumes usually 1-nerved (rarely 2- or 3-nerved); old sheaths usually not T or spirally twisted near base. 24, Panicles wide, open, oy contracted to diffuse, (2-)3-20 cm wide; panicle branches usually widely spreadir «M LU loosely aov ci dit Y. 25. Lemmas unawned, mucronate (mucros < 1 mm long), or awned, with awns 1-4 mm long. 26. Culms (65-)80-230(-300) cm tall; leaf blades 20-100 cm long, 2-7 mm wide, flat or ded. Sos O . Leaf sheath auricles absent; ligules 10-25 mm long; lemmas eL along the midvein and margins on lower 5-34 hlenbergia emersleyi . Leaf p auricles present (-1)2-4(-10) mm long, linear EE to phe tri- Susp du above, straight or twisted, firm below; ligules 2-10(-12) mm long; | pubescent with scattered hairs on lower Y __ 30. Muhlenbergia busta N ^ 26. Culms 10-70 cm tall; leaf blades 1-16 cm long, 0.3-2.5 mm wide, filiform, involute, flat or olded. 28 Boma Mende SERE inn no vous meee to ee lemmas ie a densely villous nes en nerves 2. Blepharoneuron tricholepis 28. Lemmas with awns 0.5-4 mm long; spikelet pedicels erect, not flexuous nodding to reflexed; lemmas without a thick covering of tawny to shiny silky hairs on the midvein and margins but with appressed pubescence, the hairs scattered; paleas sparsely pubescent between the nerves. 29. Leaf blades not arcuate, 1-2.2 mm wide, 4-10(-16) cm long; 1 or more culm nodes exposed; leaf blades reaching 14—12 of the plant height______ 6. Muhlenbergia arenicola 29. Leaf blades arcuate, 0.3-0.9 mm wide, 1-3(-5) cm long; usually no culm no exposed; most leaf blades reaching no more than 1/5 of the Ta height 5. Muhlenbergia torreyi 25. Lemmas awned, with awns 5-30(-40) mm long. 30. Plants conspicuously branched, especially from the middle and lower nodes, loosely caespitose; culms O geniculate, I 31. Plants distinctly bushy with ase; panicles 6-15 cm wide; anthers 1.5- 25 mm | ng; caryop ceeds compresse ed, Xo sh brown 22. Muhlenbergia porteri . Plants not distinctly bushy in appearance; plants short-lived perennials to annuals with delicate bases; panicles 1-6.5 cm wide; anthers 0.9-1.5 mm ds Car narrowly fusiform, terete, brownish 34. Müller gs tenuifolia 30. Plants not Sour branched, caespitose; culms erect to rarely slightly decumbent ar bas 2 n eae heaths compressed-keeled or flattened; glumes longer, as long or a little dud the lemm eaf sheath eae present 0.4-2.6 cm long on lower portions and up to 6.4 cm d above; lemmas glabrous or margins pubescent on the lower 1/3, LE lower 1/3 with scattered hairs; glumes usually with faint, pios, scattered hairs, the hairs less en 0. l mm long aa rd 33. Leafsheath aur pul along the midvein and m on the mon 34; glumes without widely scattered hairs — 11. MUERE UE emersleyi Peterson eta | MA ll L H4 £, 41 + M 32. Basal leaf sheaths ; glumes usually much shorter than the lemma. 34. Spikelets 3.5-5.1 mm long; lemmas 3.5-5.1 mm long; paleas ki mm a 35. | | | | not +l Pehia 1-3 mm wide, flat or involute, not falcate; culms usually with la Aes 937 node; anthers purplis Muhlenbergia rigida 35. Lemmas glabrous, smooth and shining, stramineous; leaf Tn 0.2-1.2 mm wide, tightly involute, falcate; culms usually with 2-4 bon, anther ers reenish 31. n E setifolia 34. Uu -)2.4-3.6 mm long; lemmas 2.3-3.6 mm long; paleas (2-)2.4-3.6 m lon 36. Cums25- n call el MAU ju guid and m long; caryopse s1- | 2mm long, fusiform to ovoid __ 36. Culms 74-150 cm tall; leaf blades 20-65 cm long, r not aen lemmas green and glabrous; an glabrous between the nerves; ur mm longi ONORE I -1.7 mm long, fusiform___ 24. Panicles narrow, contract ted, and or spikelike 0.2-3 cm wide; m branches uoud lly CIU CU UU loosely spreadit | 34. cen Pr 0.2-1.2 mm wide, densely cios sometimes interrupted near base; AM branches tightly appressed, the branches 0.1 m lon . Culms (3550-200 cm tall; panicles 15-60 cm E e rounded near base. 39. Lemmas awned, the awns 10-20 mm long; leaf blades 0.5-1.5 mm wide, tightly involute, falcate, and densely pubescent above; glumes puberulent on upper 2/3 14. 39. Lemmas mucronate, the mucro less than 1 mm long; leaf blades 1.5-6 mm wide, flat or involute, never falcate, and without hairs above; glumes without hairs, scabrous. 40. Glumes 3.4-5.6 mm long, usually longer than the lemma, apex unawned and not mucronate; ligules (5-)8-40(-50) mm long; anthers 1.5-2. a pls greenish 40. Glumes 1.8-3.2 mm long, shorter than the lemma, vae ind mucronate or short-awned, the awn up to 1.7 mm long; ligules 0.5-2(-3) mm long; anthe ER purpusii dark MS e stricta Muhlenbergia gypsophila Muhlenbergia macroura -1.8 yellow to purple 28. aos rigens 09) CO . Culms 15-60 cm tall; panicles (2-)4-16 cm long; leaf sheaths compressed-keeled near 41. Lower glumes 2- or 3-nerved and awned, the awns 1-5 mm long; spikelets paired on a branch, the lower spikelet staminate or zs and De Men Rond b 42. Upper leaf blades a SU V. 1.5- 3m usu ly with triangular | lol . | to vee 3. os phleoides 42. Upper leaf blades acuminate with slender awn-like seta 3-8(-10) mm long; ligules (2-)3-12 mm long, sometimes with lateral lobes the same length as the ligule; culms erect 4. Lycurus setosus 41. Lower glumes all 1-nerved, unawned or mucronate, the mucros 0.5-1 mm long; spike- lets borne singly on a branch, or if in pairs, the upper and lower spikelets both bisexual. 43. Lemmas awned, the awns 10-20 mm long; spikelets 3.5-4.3 mm long, stra- mineous to purplish; glumes 1.2-2.5 mm, the apex a ks erose, not K Qo . Lemmas mucronate, the mucros 0.5-1 mm long; spikelets ‘i p mm lon green or plumbeous; glumes Du mm long, apex acute to obtuse 2 reos the mucros 0.5-1 mm long; lemmas without distinct, raised lateral nerv 37. Panicles narrow, os- )1-3 cm wide, loosely contracted to narrowly open; primary branches loosely appressed to spreading up to 70° from the rachises, the branches 0.4-8.5 cm lon 44. Lemmas unawned, mucronate (mucros < 1 mm long), or awned, the awns 1-6 mm long. 45. Spikelet pedicels flexuous nodding to reflexed, capillary; lemmas with a thick covering of tawny to shiny silky hairs located on the midvein and margins; ligules 2. Ble mucronate; lemmas with distinct, lateral nerves — ossis gypsophila g, dar Muhlenbergia wrightii (0.3-)0.7-2(2-2.7) mm long, hyaline to opaque throughout == pharoneuron tricholepis 938 Journal of the Botanical R h Institute of Texas 1(2) 45. Spikelet pedicels erect, not fl a thick covering of tawny to sl tl lvei | ins but without hairs, puberulent throughout or X dense villous on the margins b lo ;ligules 4-35 mm long, E gua ag or SIME 46. Spikelets «anti 1.5-2.2 mm lona, gree- ish; T not decurrent and brownish below alien dubia 46. Spikelets 2.4-3.5 mm long; basal leaf sheaths compressed ea or flattene s 1.1-1.6 mm long, yellowish or purplish; ligules deccurent, bou below. 47. Glumes longer than the florets, glabrous and without hairs, scabrous or smooth; lemmas glabrous, scabrous or smooth, rarely uM ligules 10-35 mm long hlenbergia lindheimeri 47. Glumes shorter than the florets, pubescent manm near m base; lem- mas villous on the lower Y, and margins below; ligules 5-13 mm long 23. Muhlenbergia pubigluma 44. Lemmas d the awns 10-40 mm long. . Glumes 0.3-1.1 mm long, < % as long as the lemma, apices obtuse to acute. 49. Cums udis ub a 1 a none ligules 4-10 mm long, decurrent, firm i on the upper 2/3; lemmas short pilose oed: 2.3-3.1 mm long, apices acute; lemma awns 10-20 mm d anther mm long; caryopses 1-1.2 mm long 4. Mublenbergia purpusii 49. ree 8 nodes; ligules 1-3 mm long, not decurrent ish below; glumes glabrous; lemmas sparsely appressed-pubescent on the lower 1⁄4, 2.8-4 mm long, apices acuminate; lemma awns (10-)20-40 mm lon anthers 0.9-1.6 mm long; caryopses 2-26 mm long _____ 32 MURIS Bere spiciformis 48. Glumes (1-)1.2-2.8 mm long, > Y as long as the lemma (at least the upper glume), apices acute to acuminate. 50. Culms stigulose below the nodes; anthers 1.5-2 mm a E lemmas elliptic, widest near the middle Meier a polycaulis 50. Culms scaberulous below the nodes; anthers 0.9-1.5 mm > yellowish; le- mas lanceolate, widest near base 34. Muhlenbergia tenuifolia Aegopogon Humb. & Bonpl. ex Willd., Sp. Pl. 4(2):899. 1806. Tr: Aegopog hroides Humb. & Bonpl. ex Willd. Slender annuals or perennials, caespitose to sprawling, occasionally with stolons; hermaphrodites or func- tionally andromonoecious. Culms 2-55 cm tall, erect, geniculate or decumbent, herbaceous; internodes hollow. Leaf sheaths open; ligules membranous; blades flat. Inflorescences terminal, open to somewhat contracted, racemelike, l-sided panicles; each branch with 3 spikelets; disarticulation at the base of the branches. Spikelets 1-flowered; lateral spikelets pedicellate, staminate or sterile, sometimes rudimentary; central spikelets nearly sessile or short-pedicellate, laterally compressed, perfect; glumes shorter than the florets, equal, 1-nerved; lemmas membranous, 3-nerved, the central and lateral nerves usually extending into awns; paleas 2-nerved, the nerves usually extending into awns; lodicules 2; ovary glabrous, styles free to their bases, stigmas 2; stamens 3. Caryopses fusiform to ellipsoid, laterally compressed. Chromosome number x = 10. Four species, three in México and one endemic to South America. Etymology.—Name from the Greek aix, ‘goat’, and pogon, ‘beard’, alluding to the many awns of the spikelets. Comments.—In a preliminary molecular analyses all species a placed in Aegopogon fall into a “Muhlenbergia subg. Muhlenbergia” clade that has anatomical characteristics consistent with PCK (phospho- enolpyruvate carboxykinase) subtype of C, photosynthesis (Peterson 2000; Peterson et al. 2001b, 2004; Peterson & Herrera Arrieta 2001; Columbus & Peterson, in prep.). Anatomical features of this clade include: loosely arranged chlorenchyma with tabular cells that are indistinctly radiate and continuous between the vascular bundles [PCK subtype, defined as centrifugal/evenly distributed photosynthetic carbon reduction (PCR) cell chloroplasts (with grana), XyMS+ and presence of PCR cell wall suberized lamella, in Hattersley and Watson's (1992) sense], shield-shaped (narrower than deep) central bulliform cells, and primary vascular bundles with non-sclerosed phloem (Peterson & Herrera Arrieta 2001). Peterson et al., Muhlenl ii f T í Mavi 35 1. Aegopogon tenellus (DC.) Trin., Gram. Unifl. Sesquifl. 164. 1824. (Fig. 1, A & B). Lamarchia tenella DC., Cat. Pl. Horti Monsp. 120. 1813. Tyee: Cultivated at Montpellier, De Candolle s.n. (HoLotyPe: MPU?; Isotype: US-75926 fragm!.). Hymenothecium unisetum Lag., Gen. Sp. Pl. 4. 1816. Aegopogon unisetus (Lag.) Roem. & Schult., Syst. Veg. 2:805. 1817. ibo gemini- florus var. unisetus (Lag.) E. Fourn., Mexic. Pl. 2:71. 1886. Tree: MÉXICO: cultivated from seed, D. Sessé s.n.(HOLOTYPE: MA). Aegopogon gracilis Vasey, Bull. Torrey Bot. Club 13(12):230. 1886. Tyee: MÉXICO. Chinuanua: Aug 1885-Nov 1885, E. Palmer 28 (uo- 53050). Aegopogon geminiflorus var. abortivus E. Fourn., Mexic. Pl. 2:71. 1886. Tree: MÉXICO. Veracruz: Orizaba, 1864, Weber & Thomas s.n. (SYNTYPE: P; IsosyNTYPE: US-75961 fragm. ex P!). Escamela, M. Bourgeau 750 (sYNTYPE: P; IsosyNtyPE: US-75303 fragm!). VERACRUZ: Orizaba, Botteri 41 (syNtypE: P). Orizaba, M. Botteri & A.L. Sumicrast 84 (syntypE: P). San Luis Porosi: Sep 1877, Schaffner 7 (SYNTYPE: P; ISOSYNTYPE: GH) > Aegopogon geminiflorus subvar. purpureus Griseb. ex E. Fourn., Mexic. Pl. 2:71. 1886. Type: MEXICO. Distrito Feperat: Tacubaya, Schaffner 168a (SYNTYPE: P, IsOSYNTYPE: US-75962!). PANAMA. Seemann s.n. (SYNTYPE: K). Aegopogon cenchroides var. abortivus E. Fourn., Mexic. Pl. 2:72. 1886. Aegopogon tenellus var. abortivus (E. Fourn.) Beetle, Univ. Wyoming 19. 1948. Type: MÉXICO. Veracruz: M. Botteri & A.L. Sumicrast 1187 (HOLOTYPE: P!; isotype: US-75949 fragm. ex P!). Aegopogon imperfectus Nash, N. Amer. Fl. 17(2):138. 1912. Tyee: MÉXICO. Cutnuanua: Arroyo Aucho, Sierra Madre, cool mossy ledges, 15 Oct 1887, C.G. Pringle 1408 (HOLOTYPE: NY; isotype: US-740860!). Slender often sprawling, caespitose annuals. Culms (2—) 6-30 cm tall, glabrous below the nodes; internodes 0.6-6 cm long, glabrous to pilose. Leaf sheaths mostly 0.5-4.8 cm long, shorter than the internodes, gla- brous to sparingly pilose; ligules 0.6-1.5 mm long, apex mostly truncate, lacerate; blades 1.5-6 cm long, 0.5-1.5(C1.7) mm wide, flat, scaberulent and pubescent above, smooth beneath. Panicles 2-6 cm long, 0.5-1.2 cm wide, open, loosely-flowered, racemose primary branches 2-4 mm long, excluding the awns, one per node. Spikelets 1.5-3.2 mm long, often greenish or purplish, the clusters with one short-pedicelled spikelet (bisexual), the pedicels 0.2-0.6 mm long and the other two spikelets (staminate or sterile) short- pedicelled, the pedicels about 0.7-1.5 mm long; glumes (1-)1.3-2 mm long, oblong and wider distally, apex deeply notched, entire or mucronate, the mucro 0.2-1 mm long, lobes obtuse; lemmas 2.5-3.2 mm long, 3-awned, the central awns 3-8(-11) mm long, lateral awns usually 0.2-2 mm long or missing; paleas 2.2-3 mm long, puberulent, apex 2-mucronate, the mucros less than 0.8 mm long; anthers 0.5-0.8 mm long, yellowish. Caryopses 1.1-1.3 mm long, obovoid, light brownish. 2n = 20, 60. Phenology.—Flowering August through November. Distribution and habitat.—Moist slopes, cliffs, barrancas, canyons, roadsides, and along or near springs usually in shaded areas associated with Pinus spp., Quercus spp., Salvia sp., Ceanothus sp., Juniperus spp., Arbutus sp., Holodiscus sp., and Acacia spp.; 1300-2860 m. Aegopogon tenellus ranges from southern Arizona throughout most of México and Central America. Specimens examined. MEXICO. Tamaulipas: Municipio de Gémez Farias, ee San José, M.E. Crespo-Ovalle 523 (ANSM). Nuevo Leon: Municipio de Galeana, km 12 carr. Galeana-Zaragoza, G. E Blepharoneuron Nash, Bull. Torrey Bot. Club 25: 88. 1898. Tee: Blepharoneuron tricholepis (Torr.) Nash. Caespitose perennials and slender annuals, much branched near base, rhizomes absent. Culms 10-70 cm 1 tall, hollow, occasionally pithy, to strigose below the nodes. Leaf sheaths open, glabrous, usually longer than the internodes; ligules membranous or hyaline, truncate to obtuse often lacerate, the margins entire, decurrent; blades flat to involute, short pubescent above, glabrous to scabrous below. Inflorescences open or often somewhat contracted panicles with ascending to spreading branches; the pedicels often capil- lary, wiry and flexuous or nodding to reflexed, minutely granular just below the spikelet. Spikelets 1-flow- ered, slightly compressed, disarticulating above the glumes; glumes subequal, ovate to obtuse, occasionally lanceolate, faintly 1-nerved, grayish-green (olivaceous), glabrous; lemmas 3-nerved, firmer than the glumes and slightly longer, lanceolate, grayish-green, with densely appressed, tawny to shiny silky hairs located on the midvein and margins, the apex acute to obtuse, occasionally mucronate; paleas about same length as lemmas or slightly longer, 2-nerved, densely villous between the nerves; lodicules 2, club-shaped, truncate, the lateral margins thin; stamens 3; styles not at all united at the base; stigmas plumose. Caryopses fusiform to ellipsoid, brownish. Chromosome number x = 8. Two species in México, one of these endemic. 940 Journal of the Botanical R h Institute of Texas 1(2) Fic. 1. Aegopogon tenellus [FW. Gould 10391 (TAES, US)]. A. Habit B. Panicle branch with tl pikelets, central spikelet bi | (perfect) and the two | | spikel i terile. Muhlent j [PM. Peterson 5703 & CR. Annable (US)]. C. Habit. D Ligule E. Glumes. F. Florets. G J Lodicules, stamens, and pistil. Peterson et al., Muhlenl ii f +1 vd Mavi 25 Name from the Greek blepharis, ‘eyelash’, and neuron, ‘nerve’, a reference to the silky hairs located on the margins and midvein. Comments.—In a preliminary molecular analyses Blepharoneuron tricholepis and B. shepherdii are sister, or are within a grade that includes Muhlenbergia filiformis (Thurb. ex S. Watson) Rydb., M. ligularis (Hack.), and M. vaginata Swallen (Peterson et al. 2001b, 2004; Columbus & Peterson, in prep.). 2. Blepharoneuron tricholepis (Torr.) Nash, Bull. Torrey Bot. Club 25:88. 1898 (Fig. 2, A-J). vilfa tricholepis Torr., Pacif. Railr. Rep. 4(5):155. 1857. Sporobolus tricholepis (Torr.) J.M. Coult., Man. Bot. Rocky Mt. 411. 1885. Type: U.S.A. New Mexico. Bernadillo Co.: Sandia Mountains, 10 Oct 1853, Bigelow s.n. (LecrotyPE: NY! designated by Peterson & Annable, Syst. Bot. 15:522. 1990; isoLecTroTYPE: MO!) Densely tufted, caespitose perennials. Culms 10-70 cm tall, 0.3-0.7 mm diameter just below the inflores- cence, glabrous to scabrous just above and below the nodes, slender, erect; internodes 1.2-10 cm long. Leaf sheaths 1.8-9.5 cm long, usually glabrous, crowded near base, the margins glabrous, occasionally scabrous, whitish, shorter or longer than the internodes; ligules (0.3—)0.7-2(-2.7) mm long, hyaline to opaque, truncate to obtuse, the margins entire, decurrent, the apex erose-dentate to finely ciliate and irregularly toothed; blades 1-15 cm long, 0.6-2.5 mm wide, filiform, involute, arcuate, scabrous above and below, the midvein evident, raised on the abaxial surface. Panicles 3-25 cm long, 1-10 cm wide, a narrow or open panicle with ascending primary branches spreading 0—50? from the rachises; primary branches 1-6 cm long, 1-3 per node; pedicels 2-9 mm long, slender, capillary, minutely granular just below the spikelet, flexuous nod- ding to reflexed; nodes per panicle 5-25. Spikelets 2.3-3.4(-3.8) mm long, grayish-green; glumes nearly equal to subequal in length, grayish-green (olivaceous) and occasionally purplish tinged, glabrous, ovate to oblong-elliptical, the apex obtuse to acute often irregularly toothed; lower glumes (1.5-)1.8-2.6(3) mm long; upper glumes (1.7—)2-3.2(-3.7) mm long, broader and often appearing 3-nerved by its characteristic folding, however, only the midvein contains a vascular trace; lemmas (2-)2.33.4(3.8) mm long, lanceolate, grayish-green (olivaceous), often purplish tinged; apex acute to obtuse with a thick covering of appressed to spreading tawny to shining silky hairs on the midvein and margins, the hairs up to 1.2 mm long; pa- leas (22)2.2-3.5(-3.9) mm long, densely villous between the nerves; anthers 1.2-2.1 mm long, brownish. Caryopses 1.2-1.4 mm long, fusiform to elliptic, the embryo with a dorsal ridge extending 2/3 of the grain, light brownish. 2n = 16. Phenology.—Flowering mid August through November. Distribution and habitat.—Dry rocky to sandy slopes, canyon walls, rock outcrops, dry meadows, and open woods in pine-oak-madrone forests with Arctostaphylos spp., pinyon-juniper woodlands, spruce-fir forests with Ceanothus sp., Cupressus sp., and Cornus stolonifera, aspens groves, and pine woodlands with Artemisia and Garrya; central México north throughout the Sierra Madre Occidental, Sierra Madre Oriental, and through the Rockies to Utah and Colorado, U.S.A. (Peterson & Annable 1990); 700-3660 m. Specimens examined. MEXICO. Coahuila: Municipio de Arteaga, 4 km N de Las Vigas, R. Banda-Silva s.n. (ANSM); Las Vigas, Cañón de ee Sierra de Arteaga, J.A. Villarreal Q. 1794, M.A. Carranza P. & J. Valdés-Reyna (ANSM); Municipio de Múzquiz, Madera del Carmen, ooded canyon above Campo El Dos, P.M. Peterson 18908 & J. Valdés-Reyna (US). Nuevo Leon: 11.4 mi W of Dieciocho de Marzo up n towards Cerro Potosi, P.M. Peterson 13335 & M.B. Knowles (US); 17.5 mi W of Dieciocho de ^ f Cerro Potosi, P.M. Peterson 13355 & M.B. Knowles (US); Municipio de Aramberri, Cerro El Viejo, G.S. Hinton 23968 (ANSM, MEXU, TEX); Municipio de Galeana, Sierra Madre Oriental: ascent of Infernillo, 15 mi SW of Galeana, C.H. Mueller & M. T. Mueller 926 (MEXU, TEX); Municipio de General Zaragoza, Peña Nevada, M.H. Cervera-Rosado 336 (COCA). Lycurus Kunth, Nov. Gen. Sp. 1:141. 1816. Wee: Lycurus phleoides Kunth, (Lecrorvee: designated by Hitchc. U.S.D.A. Bull. 722:139. 1920). Caespitose perennials; rhizomes absent. Culms 10—60 cm tall, erect or decumbent, usually branched. Leaf sheaths open above, laterally compressed; ligules membranous to hyaline, decurrent; blades flat or folded, margins prominent. Inflorescences contracted spikelike panicles, terminal and axillary; each branch with a pair of unequally pedicellate spikelets or a pedicellate spikelet and a short secondary branch bearing two spikelets; disarticulation at the base of the pedicels or branch, the paired spikelets falling as a unit. Spikelets Botanical R h Institut f Texas 1(2) 942 Journal of tl Fic. 2. Blepharoneuron tricholepis [PM. Peterson 5567 & C.R. Annable (US)]. A. Habit. B. Ligule. C. Panicle. D. Spikelet. E. Glumes. F. Floret. G. Floret. H. Palea. I. Lemma, ventral view. J. Lodicules, stamens, and pistil. Peterson et al., Muhlenl ii f T í Mavi T 1-flowered; usually the lower spikelet in each pair staminate or sterile and the upper spikelet bisexual; glumes subequal, awned, 1-or 2(3-)-nerved; lower glumes usually 2-nerved, 2-awned; upper glumes 1(2-)-nerved, 1(22-awned; lemmas membranous, 3-nerved, lanceolate, puberulent along the margins, apices awned, the awn usually shorter than the lemma; paleas about as long as the lemmas, apex acute, sometimes the nerves extending as two mucros, margins hyaline; stamens 3. Caryopses fusiform. Chromosome number x = 10. Name from the Greek lykos, ‘wolf’, and oura, ‘tail’, a reference the the spikelike inflorescence. Comments.—Possession of paired spikelets and 2-nerved lower glumes that are 2-awned, although not common within the Muhlenbergiinae, is not unique to Lycurus. Muhlenbergia brevis and M. depauperata exhibit morphological features that suggest a close relationship with Lycurus. Mez (1921) indicated a rela- tionship with Lycurus when he transferred M. shaffneri E. Fourn., considered a synonym of M. depauperata, to Lycurus. Muhlenbergia brevis and M. depauperata share many morphological features with Lycurus, most importantly: spikelets borne in pairs and lower glumes that are 2-nerved and 2-awned (Peterson 2000). Schaffnerella gracilis is apparently sister to the three species of Lycurus for trnL-F and ITS sequence analyses, and this Lycurus-Schaffnerella clade is found in a well-supported trnL-F-derived clade that includes M. wrightii, all four species of Chaboissaea, and Schedonnardus paniculatus (Peterson et al. 2004; Columbus & Peterson, in prep.). Allozyme studies were useful is determining the biogeographical history of the amphitropical Lycurus setosus and it seems likely that this species has recently dispersed to South America because populations there contain less genetic variation (Peterson & Morrone 1997; Peterson 2000). 3. Lycurus palon Kunth, Nov. oo: Sp. No ed. ) 1:142, pl. 45. 1816. (Fig. 3, A & B). Wee: MÉXICO: inter Guanaxuato et Ti a Buffa, alt. 1030 hexap., Sep, EWH.A. Humboldt & A.J.A. Bonpland s.n. (HOLOTYPE: P!; isotypes: BAA-fragm.!, US-610840 fragm. ex P-Bonpl!, US-610841 fragm.!). A Loosely caespitose perennials. Culms 20-60 cm tall, erect t umbent at base, scabrous to puberulent just above or below the nodes. Leaf sheaths usually 0.5—3 cm long, glabrous to pubescent, compressed-keeled near base; ligules 1.5-3 mm long, acuminate and erose, usually with triangular lateral lobes 0.5-2.5 mm long, the lateral lobes decurrent below; blades 2-8 cm long, 1-1.5 mm wide, flat or folded, glabrous to puberulent below and puberulent to hispidulous above, margins and midvein whitish-thickened, apex acute or some- times with a narrow bristle 0.5-3 mm long on the upper blades. Panicles (2—)4-10 cm long, 4-8 mm wide, contracted, narrow, densely flowered, and spikelike; primary branches 0.1-0.3 cm long, tightly appressed. Spikelets 3-4 mm long, 1-flowered; glumes 1-2 mm long, scaberulous near apex, awned, the awns 1-5 mm long; lower glumes usually 2(3-)-nerved; upper glumes 1(2-)-nerved, awns 2-5 mm long; lemmas 3-4 mm long, puberulent along the margins, awned, the awns 1-3 mm long; paleas 3-4 mm long, puberulent between the nerves; anthers 1.5-2 mm long, yellow. Caryopses 1.8-2.2 mm long, brownish. 2n = 40. Phenology.—Flowering July through October. Distribution and habitat. —Rocky slopes, canyons, bajadas, and flats often occurring in calcareous soils and associated with Quercus gregii, Larrea tridentata, Pinus cembroides, Juniperus flaccida, Salvia regla, Cowania plicata, Bouteloua gracilis, B. uniflora, Dasylirion longissimum, Y. carnerosana, Agave lecheguilla, Jatropa dioica, Muhlenbergia spp., Berberis sp., Opuntia spp, Selaginella sp, Hechtia sp., Brahea sp., and Mimosa sp.; northern and central México in Aguascalientes, Chihuahua, Distrito Federal, Durango, Guanajuato, Hidalgo, Jalisco, México, Michoacán, Morelos, Nayarit, Oaxaca, Puebla, Querétaro, San Luis Potosí, Sonora, Tlaxcala, and Zacatecas to southwestern U.S.A. in Texas and New Mexico (Espejo Serna et al. 2000; Peterson et al. 2001b; Powell 1994; Reeder 1985; Sánchez & Rügolo de Agrasar 1986); 1200-2500 m. Specimens examined. MÉXICO. Coahuila: Sierra El Pino, 9.6 km SW of Rancho El Cimarron, P.M. Peterson 10635 & C.R. Annable (US); 33.5 km W of Rancho El Cimarron, P.M. Peterson 10656 & C.R. Annable n 5 mi W of Chapultepec, P.M. Peterson 13278 & M.B. Knowles (US); 30.6 km SE of Saltillo on HWY 57 towards San Luis Potosí, P. M. Pi 240 & C.R. Annable (US); 6 km S of Saltillo, P.M. Peterson 8347 & J. Valdés-Reyna (US); 35.4 km W of Ocampo at Cuesta Zozaya, P.M. Peterson 8367 & J. Valdés-Reyna (US); western base of Picacho del Fuste, I.M. Johnston 8444 (US); du of La Noria, R.M. pL Municipio de Acuña, del Carmen Mountains, E.G. M Jr. 658 (MEXU); L li XU); Municipio de Arteaga, 2 km NW of Ejido Sierra Hermosa, R. ees n. (ANSM); Sierra Zapaliname, at 2 a E of Saltillo up Camino de Quatro (Las Palapas), P.M. Peterson 17858, J. Valdés- J J te of Texas 1(2) I Lat +l ¿VU Mal OF 944 E euo DOS as SSS WSs nee Lis pela, NUS Ñ rDM D Peterson et al., Muhlenl ii f T í Mavi "T Reyna & R.H. Cardenas (US), P.M. Peterson 18805 & J. Valdés-Reyna (US); Cañón San Lorenzo, Sierra de Zapalinamé, 8 km S of Saltillo & 3.2 km E de Buenavista, R. López Aguillón s.n. (ANSM); 3 mi SE of Saltillo, I.M. Johnston 7250 (US); road to Canyon San Lorenzo, P.M. Peterson 10545 & C.R. Annable (US); Las Vigas, Cañón de la Carbonera, Sierra de Arteaga, J. Valdés-Reyna 1797, J.A. Villarreal Q. & M.A. Carranza P. (ANSM); Predio El Cristal, E. Pérez Torres 61 (COCA): Municipio de Castaños, Paso de San Lázaro, Sierra de la Gavia, 37.6 nclova on Hwy 57, P.M. Peterson 9990, C.R. Annable & J. Valdés-Reyna (ANSM, US); Municipio de Cuatrociénegas, Sierra de la Madera, Cañón Charreteras, Rancho Charreteras, J.A. Villarreal Q. 7350, M.A. Carranza P. & R. Rodríguez L. (ANSM); Cañada Los Pozos, 8 mi SW of Tanque El Tropical along road to Carión del Desiderio, T.L. Wendt s.n. & J. Valdés-Reyna (MEXU); Municipio de Müzquiz, near Santo Domingo, F.W. Lyle 454, C.H. Muller (ANSM, US); Sierra Maderas del Carmén, E. García Aguilera s.n. (ANSM); Municipio de Ocampo, Mesa Grande, 40 km NW of Hacienda de La Encantada, R.M. Stewart 1631 (MEXU); W base of Picacho del Fuste, NE of Tanque Vaionetta, I. M. Johnston 8444 (MEXU); Municipio de Parras, Sierra de Parras, 2-4 km S of Ejido Colorado, 8.8 km al W of Parras, C.P. Cowan 3610 (TEX- m LN de Ramos Arizpe, Sierra de la Paila, Ejido El Cedral, J.A. Villarreal Q. 5330, M.A. Carranza P. & A. Rodriguez G. (ANSM de Saltillo, 20 km E of Saltillo, Carretera Saltillo-Torreón, R. Almeida 1642 (ANSM); 4 km W of Saltillo, Carretera Saltillo- General Ea J. Valdés-Reyna 1553, L.E. Rodríguez G. & R. Vásquez A. (ANSM); 6 mi W of Saltillo along highway 40 towards Torreón, S.L. Hatch 5040 & J. Valdés-Reyna (ANSM); Buenavista, Brigada Coahuila 102, 103, 104 (COCA); Buenavista, 7 km S de Saltillo, Carretera 54 Saltillo- Concepción del Oro, A. Rodríguez-Guillén 101, R.E. Rodríguez-Charúa 107, M. Jiménez-F. s.n., N. Ochoa- R. s.n., R. Palomo-Garza & P. García-Solís s.n. (ANSM); Cañón San Lorenzo, 1.5 km E of Pozo, R. López Aguillón s.n. (ANSM); Cerro del Pueblo, W of La Ciudad de Saltillo, J. Valdés-Reyna 2045, M. Martínez M. & M.A. Carranza P. (ANSM); Hwy 54 towards Concepción del Oro, 6 km S of Saltillo, M.E. Barkworth 5108, P.S. Hoge, J. Valdés-Reyna, & M.A. Carranza P. (ANSM); Rancho Experimental Los Angeles, 48 km S of Saltillo, Carretera 54 Saltillo- Concepción del Oro, J.A. Villarreal Q. 6710, S.L. Hatch 4538, C.W. Morden, J. Valdés- Reyna, M.A. Carranza P. & D.E. Lozano (ANSM); Saltillo, G. Fisher s.n. (UAT); Sierra Madre Oriental, Terrenos de Tranquitas, R.M. Francois 5827 & J. Passini (ANSM); Sierras adyacentes al Paso de Carneros, J.S. Marroquín-de la Fuente 3058 (ANSM); 29 km S of Saltillo on MEX 54 just W of Estacion Caranaros, P.M. Peterson 8413 & M.A. Carranza P. (US); RS de San Pedro, entre Cuatrociénegas & San Pedro, A. Rodríguez- Gámez 945, J J. López & J. L. Elizondo E. (ANSM, TEX); Muni de Si Mojada, Cañon de Tinaja Blanca, W of Santa Elena Mines, R.M. Stewart 1950 (US); Municipio de Torreón, Sierra de Jimulco, 1L 7 km E of Flor de Jimulco, P.M. Peterson 8453, J. Valdés-Reyna, P.A. Fryxell & J.A. Villarreal Q. (ANSM, US); Sierra de Jimulco, Mina San José, J.A. Villarreal Q. 7791 & M.A. Carranza P (ANSM, TEX). Nuevo León: 12.3 mi S of Milagro, P.M. Peterson 17810 ae Valdés- oe & G.S. Hinton ide 16 km NE of Sandia on road to La Ascensión, P.M. Peterson 15826 & J. Valdés-Reyna (US); i Ejido La Encantada, P.M. Peterson 16733, J. Valdés-Reyna & M. Sosa Morales (US); 9.4 mi W of San Antonia de Peña Neda P.M. Peterson 16791, J. Valdés-Reyna & M. Sosa Morales (US); 75.8 km N of Dr. Arroyo on Hwy 61, P.M. Peterson 11144 & C.R. Annable (US); 6.5 mi S of Border of Coahuila and Nuevo Leon on Hwy 57 towards Matehuala, P.M. Peterson 13286 & M.B. Knowles (US); 2 mi W of Dieciocho de Marzo up road towards Cerro Potosí, P.M. Peterson 13325 & M.B. Knowles (US); 36 mi NE of Dr. Arroyo on Hwy 61 towards Linares, P.M. Peterson 13366 & M.B. Knowles (US); N 10 mi E of San Roberto along highway 58, S.L. Hatch 4978, J. Valdés-Reyna 6 P. Dávila A. (ANSM); 12 km SE of San Roberto on eee 58, S.L. Hatch 4580, C.W. Morden & J. Valdés-Reyna (ANSM); Alrededor de Galeana, J.A. Ochoa-Guillemar 1258 (COCA); Hacienda Pablillo, M. Taylor-Edwards 134 (MEXU); Rancho Aguililla, G.B. Hinton 19863 (MEXU, TEX); alrededores del poblado Santa Clara de González, N. Bazaldua- poesia 110 ee Tamaulipas: Municipio de Bustamante, Camino a La Joya de San Francisco, J.L. Ramos-Delgado 99 (COCA); M ] 15 km SW of Miquihuana, P. Hiriart 301, V. Juárez & R. Molczadzki (MEXU); 7 km W of Miquihuana, P. Hiriart 899,V. Tuáre R. Molczadzhi, J.L. López, F. Ojeda & J.A. Gutiérrez (MEXU); 9 km N of Miquihuana, P. Hiriart 335, V. Juárez & R. Molczadzhi (MEXU); 3 mi N of Miquihuana, Stanford 2481, Lauber & Taylor (US); Colonia La Peria, J.F. Iribe-Duarte 106 (COCA); Ejido El Aserradero, J.G. Galván-Infante 37 & R.A. Carranco-Rendon 70 (COCA); Ejido Valle Hermoso, Cañón de La Sierra Miquihuana, R.A. Carranco-Rendon (COCA); Municipio de Palmillas, Ejido Palmillas, R.A. Carranco-Rendon 213 (COCA); Municipio de Tula, La Tapona, M.H. Cervera-Rosado 133 (COCA) 4. Lycurus setosus (Nutt.) C. Reeder, Phytologia 57:287. 1985. (Fig. 3, C—E). Pleopogon setosum Nutt., Proc. Acad. Nat. Sci. Philadelphia 4:25. 1848. Tyre: U.S.A. New Mexico: Santa Fe Co.: mountains near Santa Fe, 1841 or 1842, William Gambel s.n. (HOLOTYPE: K!; isotypes: PH, US-610839 fragm. ex K!). Lycurus alopecuroides Griseb., Abh. Kónigl. Ges. Wiss. Gottingen 19: 255-256. 1874. Type: ARGENTINA: Prov. CATAMARCA: ca. Belén, en el altivalle de las Granadillas, Feb 1872, P G. Lorentz 548 (HoLoTYPE: GOET; IsoTYPES: BA!, BAA!, CORD, SI!, US-996080 fragm. ex GOET!). Ly phleoid glaucifolius Beal, Grass. N. Amer. 2:271. 1896. Type: MEXICO. C ky hills near Chihuahua, 28 May 1885, C.G. Pringle 426 (Lectotype: MSC designated by Hitchcock, Contr. U.S. Natl. Herb. 17:305. 1913, without explicitly citing a specific DI et in a specific herbarium, fully lectotypified by C. Reeder isolectotypes: MO-3701765!, NY, P!, SI!, US-996079!). .1985 , who cited the MSC specimen as “holot otype”; > Caespitose perennials. Culms 30-60 cm tall, erect, scabrous to puberulent just above or below the nodes. Leaf sheaths usually 1-5 cm long, glabrous to pubescent, compressed-keeled near base; ligules (22)3-12 mm long, acuminate to erose, sometimes with lateral lobes the same length as the ligule, the lateral lobes decurrent below; blades 4-10(22) cm long, 1-2.5 mm wide, flat or folded, glabrous to puberulent below 946 Journal of the Botanical R h Institute of Texas 1(2) and puberulent to hispidulous above, margins and midvein whitish-thickened, apex acuminate with slender awn-like seta 3-8(-10) mm long on the upper blades. Panicles 2-)4-10 cm long, 4-8 mm wide, contracted, narrow, densely flowered, and spikelike; primary branches 0.1-0.3 cm long, tightly appressed. Spikelets 3-4 mm long, 1-flowered; glumes 1-2 mm long, scaberulous near apex, awned, the awns 1-5 mm long; lower glumes usually 2(3-)-nerved; upper glumes 1-nerved, awns 2-5 mm long; lemmas 3-4 mm long, puberulent along the margins, awned, the awns 1-3 mm long; paleas 3-4 mm long, puberulent between the nerves; anthers 1.5-2 mm long, yellow. Caryopses 1.8-2.2 mm long, brownish. 2n = 40 Phenology.—Flowering July through October. Distribution and habitat. —Wooded canyons, rocky slopes, desert grasslands, and rock outcrops with Pinus spp. Quercus spp., Bouteloua hirsuta, Chrysothamnus spp., Pseudostuga menziesii, Cupressus spp., Abies spp., Cornus stolonifera, Ceanothus spp., and Yucca spp.; northwestern México in Baja California, Chihuahua, 1 Durango, Sonora, and extending into the southwestern U.S.A., then disjunct in Bolivia (Espejo Serna et al. 2000; Peterson et al. 2001b; Reeder 1985; Sánchez & Rúgolo de Agrasar 1986); 1400-2500 m Specimens examined. MÉXICO. Coahuila: Sierra El Pino, 39.5 km W of Rancho El Cimarron, P.M. Peterson 10672 & C.R. Annable (US); 1219 A NIC Municipio de Acuña, Serranías del Burro, Rancho El Bonito, Cañón El Bonito, 7 mi W de la casa principal, D.H. Riskind 2185 (ANSM Argentina and Municipio de Cuatrociénegas, Sierra de la Madera, Cañada Los Posos, 0.8 mi by road SW from Tanque El Tropical along road to Cañón del Desiderio us Valdés- a 1053 & T.L. Wendt (ANSM), Sierra de la Madera, Cañón del Agua, mouth of canyon un Valdés- NAE 1000, 10126 T.L Orientale, Ejido el Cinco, R.M. F 92 & J. Passini (ANSM); M : Sierra Madre Oriental Ejido El Cinco, R.M. F i 83 & J. Passini (ANSM); Municipio de M quiz, Cuesta de la mE J.A a L. s.n. (ANSM); Sierra del Carmén, R.M. F is 5514 & J. Passini (ANSM); Ejido San men P.M. Peterson 18842, J. Valdés-Reyna & C. Sifuentes (US); Madera del Carmen, 7.2 mi NW of Pilares, P.M. Peterson 18876 & J. Valdés-Reyna (US); Campo El Dos, P.M. Peterson 18910 & J. Valdés-Reyna (US); Municipio de Sierra Mojada, San Antonio de los Alamos, I.M. Johnston 8252a (US). Muhlenbergia Schreb., Gen. Pl. (ed.8) 1:44. 1789. Wee: Muhlenbergia schreberi J.E Gmel., (LecroTvrE: designated by Nash in Britton & Brown, Ill. Fl. N. U.S., ed. 2, 1:184. 1913). Plants annual or perennial; hermaphrodites or occasionally monoecious; often caespitose, sometimes mat- forming, usually rhizomatous, rarely stoloniferous. Culms 2-300 cm tall, erect, geniculate, or decumbent, usually herbaceous, sometimes becoming woody; internodes solid or hollow. Leaf sheaths open, auricles absent; ligules membranous or hyaline, sometimes firm or coriaceous, acuminate to truncate, sometimes minutely ciliolate, sometimes with lateral lobes longer than the central portion; blades narrow, flat, folded, or involute, sometimes arcuate. Inflorescences terminal, sometimes also axillary, open to contracted or spikelike panicles; disarticulation usually above the glumes, occasionally below the pedicels. Spikelets 1(2—3)- flowered, laterally compressed or terete; glumes subequal in length, mostly shorter than the lemmas, or as long or exceeding the lemmas, usually (0)1(2, 3)-nerved, apices entire, erose, or toothed, truncate to acuminate, sometimes mucronate or awned; lower glumes sometimes rudimentary or absent, occasionally bifid; upper glumes shorter than to longer than the florets; lemmas membranous or rather firm, glabrous, scabrous, or with short hairs, 3-nerved (occasionally appearing 5-nerved), callus glabrous or hairy, apices awned, mucronate, or unawned; awns, if present, straight, flexuous, sinuous, or curled, sometimes borne between 2 minute teeth; paleas membranous or rather firm, well developed, about as long as the or shorter than the lemma, 2-nerved, rounded on the back; lodicules 2, short, fleshy, truncate; lateral margins thin; ovary glabrous, styles free to their bases, stigmas 2; stamens (1, 2)3, anthers purple, orange, yellow, or oli- vaceous. Caryopses elongate, fusiform or elliptic, slightly dorsally compressed, usually not falling free from the lemma and palea. Cleistogamous panicles sometimes present in the axils of the lower cauline leaves, enclosed by a tightly rolled, somewhat indurate sheath. Embryo large, with an epiblast, scutellar tail, and elongated mesocotyl; embryonic leaf margins meeting, endosperm hard. Chromosome number x = 10. Etymology.—Named for Gotthilf Henry Ernest Muhlenberg (1753-1815), a Lutheran minister and pioneer botanist of Pennsylvania. Comments.—A good historical account, biogeographical summary, and subgeneric classification of Muhlenbergia was given in Peterson (2000) and Peterson and Herrera Arrieta (2001). Based on population Peterson et al., Muhlenl ii f T í Mavi " genetic studies of the amphitropical disjunct, Muhlenbergia torreyii (Peterson & Ortíz-Diaz 1998), it is clear that migration predominantly occurred from a center of diversity (for Muhlenbergia & Muhlenbergiinae) in southwestern North America (centered in northern Mexico) to northern Argentina-southwestern Bolivia (Peterson & Herrera Arrieta 1995; Peterson & Columbus 1997; Sykes et. al. 1997; Peterson & Morrone 1997; Peterson & Ortíz-Diaz 1998; Peterson 2000). Chihuahua apparently has the highest number of species of Muhlenbergia (59 spp.), followed by Durango with 55 species (Herrera Arrieta 2001; Herrera Arrieta & Peterson 2007). In northeastern México, M. jaime-hintonii is endemic to the region and is known only from Nuevo León. Even though preliminary phylogenetic results of trnL-F and ITS DNA sequences indicate that Muhlen- bergia is polyphyletic, there are monophyletic lineages that correspond to previous delineated taxa (Peterson et al. 2001a, 2004; Columbus & Peterson, in prep.). In addition to all species of Aegopogon and Pereilema, the following seven species placed in Muhlenbergia subg. Muhlenbergia are well supported in the molecular analysis and exhibit PCK-like anatomical characteristics: M. glauca, M. microsperma, M. polycaulis, M. porteri, NAGEL 1 M. racemosa, M. spiciformis, and M. tenuifolia ia distichophylla, M. dubia, M. emersleyi, M. gypsophila, M. jaime-hintonii, M. lindheimeri, M. macroura, M. pubigluma, M. purpusii, M. rigens, M. rigida, M. robusta, M. setifolia, and M. stricta (Muhlenbergia subg. Trichochloa) have anatomical characters of deep adaxial furrows, vascular bundles positioned in two or three levels, and inflated cells located below (abaxial to) the primary vascular bundles; and are found in a well supported molecular clade (Peterson et al. 2001a, 2004; Peterson & Herrera Arrieta 2001). There is good molecular support for M. montana, M. quadridentata, and M. vire- scens, all members of a group referred to as the “Muhlenbergia montana complex” that have 3-nerved upper glumes and sclerosed phloem (Peterson et al. 2001a, 2004; Peterson & Herrera Arrieta 2001). The annuals, M. depauperata, M. fragilis, and M. minutissima, with M. arenicola and M. torreyii form a fairly well supported molecular clade; and Muhlenbergia arenacea, M. asperifolia, M. repens, M. utilis, M. villiflora, and M. wrightii are all members of a fairly well supported molecular clade that includes the M. montana complex, Lycurus- Schaffnerella, Chaboissaea spp., and Schedonnardus paniculatus (Columbus & Peterson, in prep). 5. Muhlenbergia arenacea (Buckley) Hitchc., Proc. Biol. Soc. Wash. 41:161. 1928. (Fig. 1, C-G). sporobolus arenaceus Buckley, Proc. Acad. Nat. Sci. Philadelphia 14:89. 1862. Type: U.S.A. New Mexico/Texas: May-Oct 1849, C.H. Wright 737 (LECTOTYPE: GH designated by Hitchcock, Man. Grass. U.S. 886. 1935, but without citing a specific sheet in a specific herbarium; ISOLECTOTYPE: US-997371! Sporobolus auriculatus Vasey, Contr. U.S. Natl. Herb. 3(1):64. 1892. Tyre: U.S.A. Texas: Duval Co.: Pena, 1890, G.C. Nealley 492 (LECTOTYPE: US-997369! designated by Hitchcock, N. Amer. Fl. 17(6):447. 1935, but witl t in a specific herbarium) o Lat Rhizomatous perennials. Culms 10—30(-40) cm tall, decumbent, terete to compressed-keeled near the base; internodes scaberulous below the nodes. Leaf sheaths about 1⁄2 as long as the internodes, margins hyaline; ligules 0.5-2 mm long, hyaline, with lateral lobes or auricles 1-2 mm long; blades 0.7-4(-6) cm long, 0.5-1.7 mm wide, flat, occasionally folded, tapering, scabrous abaxially, strigulose adaxially, margins and midveins thickened, whitish, apices narrow, often sharp, erose. Panicles 5-15 cm long, 4-14 cm wide, broadly ovoid, open; primary branches 2-8 cm long, capillary, straight to slightly flexuous, diverging 45-80(-100)? from the rachises, never appearing fascicled, naked proximally; pedicels 1-11 mm long, usually longer than the spikelets. Spikelets 1.5-2.5 mm long, occasionally with 2 florets; glumes 0.9-2 mm long, equal, 1-nerved, usually acute to acuminate, occasionally erose and mucronate, mucros to 0.2 mm long; lemmas 1.5-2.5 mm long, lanceolate to oblong-elliptic, plumbeous to purplish, sparsely appressed-pubescent on the lower 15 of the margins and midveins, hairs to 0.3 mm long, apices acute to obtuse, sometimes shallowly bilobed, mucronate, mucros to 0.3 mm long; paleas 1.5-2.6 mm long, lanceolate, glabrous, obtuse to acute; anthers 1-1.5 mm long, yellowish to purplish. Caryopses 1-1.3 mm long, elliptic, brownish. Distribution and habitat—Muhlenbergia arenacea grows in sandy flats, plains, alluvial fans, washes, depressions, and alkaline mesas in open grasslands, at elevations of 1000-2200 m. It ranges from the southwestern U.S.A. to northern México in Chihuahua, San Luis Potosi, and Zacatecas (Espejo Serna et al. 2000; Peterson 2003). 948 Journal of the Botanical R h Institute of Texas 1(2) Specimens examined. MÉXICO. Coahuila: 153.2 km NW of Muzquiz on Hwy 53 towards Boquilla del Carmen, P.M. Peterson 10602 & C.R. Annable (US); Sierra El Pino, 9.2 km SW of Rancho El Cimarron, P.M. Peterson 10624 & C.R. Annable (US); Municipio de Acuña, Sierra del Carmen, E.G. Marsh Jr. 907 (MEXU); Municipio de Ocampo, Rancho el Barranquito, 50 Km de S rumbo a Sierra pde M.A. Carranza P. 576 & F.J. Carranza P. (ANSM); Rancho La Rueda, 87 km NW of Ocampo, D. Ibarra s.n. (ANSM 12 km N of the Coahuila-Zacatecas state line along highway 54 between ed and Concepción del Oro, Zacatecas, S. L Hatch 4516, C. W. Morden & J. Valdés-Reyna (ANSM); 29.2 mi S of Saltillo, carretera 54 Saltillo- Concepción del Oro, Zacatecas, P.M. Peterson 10033, C.R. Annable & J. Valdés-Reyna (ANSM, US); 3 mi E of San Jose, I.M. Johnston 8217 (MEXU); Municipio de Sierra Mojada, Reserva de la Biosfera de Mapimí limite del estado de Coahuila, Chihuahua & Durango, R. Ruíz Esparza R. 552 (ANSM) 6. Muhlenbergia arenicola Buckley, Proc. Acad. Nat. Sci. Philadelphia 14:91. 1862. (Fig. 4, A-E). Podose- mum arenicola (Buckley) Bush, Amer. Midl. Naturalist 7(2):40. 1921. Type: U.S.A. Texas: arid places in western Texas, C. Wright 735 (LecroTYPE: PH designated by Hitchcock, Man. Grass. U.S. 886 (1935), but without citing a specific sheet in a specific herbarium, ISOLECTOTYPE: US-87239 fragm). Caespitose perennials. Culms (15—)20—60(-70) cm tall, somewhat decumbent, 1 or more nodes exposed; internodes hispidulous below the nodes. Leaves somewhat basally concentrated, most blades not reaching more than 34-15 of the plant height; leaf sheaths usually a little shorter than the internodes, not keeled, scaberulous, margins hyaline, basal sheaths rounded, not becoming spirally coiled when old; ligules 2-9 mm long, hyaline, acute, lacerate, often with lateral lobes; blades 4—10(—16) cm long, 1-2.2 mm wide, not arcuate, flat, folded, or involute, scabrous, often glaucous, midveins and margins not thickened, green. Panicles 12-30 cm long, 5-20 cm wide, diffuse; primary branches 1-10 cm long, diverging 30-80? from the rachises, naked basally; pedicels 1-4(-6) mm long, erect. Spikelets 2.5-4.2 mm long; glumes 1.4-2.5 mm long, equal, 1-nerved, apices scaberulous, acute to acuminate, minutely erose, sometimes mucronate, the mucro to 1 mm long; lemmas 2.5-4.2 mm long, narrowly elliptic, usually purplish, scabrous distally, appressed-pubescent on the lower !5—34 of the margins and midveins, apices acuminate, awned, awns 0.5-4 mm long; paleas 2.5-3.5 mm, narrowly elliptic, intercostal region sparsely pubescent, apices acuminate, with 2 short mucros 0.1-0.2 mm long; anthers 1.5-2.1 mm, greenish. Caryopses 1.9-2.3 mm long, fusiform, brownish. 2n = 80, 82. Distribution and habitat—Muhlenbergia arenicola grows on sandy mesas, limestone benches, and in val- leys and open desert grasslands, associated with Bouteloua gracilis, B. uniflora, Larrea tridentata, and Yucca carnerosana at elevations of 600-2135 m. The range of M. arenicola extends to southwestern U.S.A. and the species is a disjunct in northwestern Argentina (Peterson & Ortiz-Diaz 1998). Specimens examined. MEXICO. Coahuila: Sierra El Pino, 9.2 km SW of Rancho El Cimarron; P.M. Peterson 10619 & C.R. Annable (US); 153.2 km NW of Muzquiz on Hwy 53 towards Boquilla del Carmen, P.M. Peterson 10603 & C.R. Annable (US); 0.3 km E of Hwy to Zacatecas up road to Canyon San Lorenzo, P.M. Peterson 10550, C.R. Annable & J. Valdés-Reyna (US); Municipio de Acuña, 2 km N of El Conejo on the winding road to La Huerfanita, M.C. Johnston 9192, F. Chiang C. & T.L. Wendt (ANSM); Municipio de Arteaga, Sierra de Arteaga, Bosque de la Montaña, 10 km del entronque Los Lirios & carretera 57 a Matehuala, rumbo a los Lirios, P.S. Hoge. 257, M.E. Barkworth, J.A. Villarreal Q. € J. Valdés-Reyna (ANSMD; Municipio de Francisco I. Madero, Valle de Buenavista, S de los Remedios, T.L. Wendt 1387 & EJ. Lott (ANSM); Municipio de Múzquiz, La Cuesta del Plomo on the Müzquiz-Boquillas highway, F. Chiang-Cabrera 9207 (MEXU), T.L. Wendt & M.C. Johnston 9207 (ANSM); Municipio de Ocampo, Rancho experimental Santa Teresa de La Rueda, 87 km NE of Ocampo, brecha Ocampo-Boquillas del Carmen, R. Vásquez-Aldape s.n. (ANSM); Rancho experimental Santa Teresa de La Rueda, 87 km NE of Ocampo, brecha Ocampo-Boquillas del Carmen, M. Vásquez-Rodríguez s.n. (ANSM); Rancho La Rueda, 87 km NE of Ocampo, D. Ibarra s.n. (ANSM); Municipio de Parras, Rancho el Tunal, 25 "m ESE of Parras de la Fuente, A. Rodríguez- Gámez 1181 & M.A. Carranza P. (ANSM); ^ Saltillo, 28.3 mi S of Saltillo, carretera 54 a Concepción del Oro, Zacatecas, P.M. Peterson 10026 C.R. Annable & J. Valdés-Reyna (ANSM, US); 47 km S of Saltillo on MEX Hwy 54 to Concepcion del Oro near entrance road to El Salitre Randiera, P.M. Peterson 10032, C.R. Annable & J. Valdés-Reyna (ANSM, m 29.2 mi S of Saltillo, carretera 54 Saltillo- Concepción del Oro, Zacatecas, P.M. Peterson.10032, C.R. Annable & J. Valdés-Reyna (ANSM, US); 30.6 km SE of Saltillo on Hwy 57 towards San Luis Potosi, P. M. Peterson 6239 & C.R. Annable (US); 50 km S of Saltillo, carretera 54 a Concepción del Oro, Zacatecas, entronque El Colorado, J.A. Villarreal 2 2061, M.A. Carranza P. & J. Valdés-Reyna (ANSM, MEXU); 3 mi E of San Jose, I.M. Johnston 8219 (MEXU); Entronque Derramadero, a 20 km S de Saltillo por la carretera Saltillo-Zacatecas, carretera 54, J. Valdés-Reyna & M.A. Carranza P. (ANSM); Rancho experimental Los Angeles, 48 km S de Saltillo, carretera 54 Saltillo- Concepción del Oro, Zacatecas D pr -Aburtor 56, S. L. Hatch. 5019, G.A. Puente-Tristán, J. S. Sierra-Tristán & J. Valdés-Reyna (ANSM); Municipio de Sierra Mojada, 15 km E of San Antonio de los Alamos, I.M. Johnston 8288 (MEXU). Nuevo León: 64.4 km S of San Rafael on d 57, P.M. Peterson 6236 & C.R. Annable (US); 12.3 mi S of Milagro, P.M. Peterson 17815, J. Valdés-Reyna 6 G.S. Hinton (US); ¡Municipio de Galeana , 2 km SE of El Canelito, on road to San Peterson et al., Muhlenl ii f T í Mavi T "d j S A Y b yg K Y | Y A A Vv. =s SS 3 ÉS ESA Fic. 4. Muhlenbergia arenicola [PM. Peterson 6239 & C.R. Annable (US)]. A. Habit B. Ligule. C. Glumes. D. Floret. E. Lodicules, stamens, and pistil. Muhlen- bergia torreyi [PM. Peterson 5608 & C.R. Annable (US)]. F. Habit. G. Ligule. H. Glumes. I. Lemma. 950 Journal of the Botanical R h Institute of Texas 1(2) Juan de Dios, M.C. Johnston 7984, T.L. Wendt (ANSM); Ejido El Tokio, M.L. Avalos-Marín s.n. (UAT, ANSM). Tamaulipas: Municipio de Bustamante, Ejido Felipe Ángeles 2 km NO rumbo a Bustamante, R. Díaz-Pérez s.n. (UAT) 7. Muhlenbergia asperifolia (Nees & Meyen ex Trin.) Parodi, Revista Fac. Agron. Veterin. (Buenos Aires) Osee . (F ig. 5, A-D). vi lfa UA ia Nees & Mon ex Trin., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 6,4(1-2):95. 1840. ifolius (Nees & Meyen ex Trin.) Nees & Meyen, Gramineae 9-10. 1841. Tyee: Chile. Río Mayno, 1000' alt, Mar 1831, , Meyen s.n. coe B, BAA-3464 fragm. ex B!, LE-TRIN-1679.01b!); Copiapó, Meyen s.n. (SYNTYPE: LE-TRIN-1679.01a!). Agrostis eremophila Speg., Anales Mus. Nac. Hist. Nat. Buenos Aires 7:190. 1902. Agrostis distichophylla Phil., Fl. Atacam. 54. 1860. nom. illeg. hom. Tyre: Chile. prope Tilopozo ad marginem paludos, 23.20 lat., 7000 ft (isotypes: BAA-4153 fragm. ex SGO!, SGO-63100, SGO-37135, US-1939353 fragm. ex SGO-37135!) Rhizomatous perennials, occasionally stoloniferous. Culms 10-60(-100) cm tall, decumbent-ascending, bases somewhat compressed-keeled; internodes glabrous, shiny below the nodes. Leaf sheaths glabrous, margins hyaline; ligules 0.2-1 mm long, firm, truncate, ciliate, without lateral lobes; blades 2-7(-11) cm long, 1-2.8(-4) mm wide, flat, occasionally conduplicate, smooth or scaberulous abaxially, scaberulous adaxially, margins and midveins not conspicuously thickened, greenish, apices acute, not sharp. Panicles 6-21 cm long, 4-16 cm wide, broadly ovoid, open; primary branches 3-12 cm, capillary, lower branches spreading 30—90? from the rachises, never appearing fascicled; pedicels 3-14 mm long, longer than the spikelets. Spikelets 1.2-2.1 mm long, occasionally with 2 or 3 florets; glumes 0.6-1.7 mm long, equal, pur- plish, scaberulous, particularly on the veins, 1-nerved, apices acute; lemmas 1.2-2.1 mm long, lanceolate to oblong-elliptic, somewhat plumbeous, glabrous, usually smooth, occasionally scaberulous near the apices, apices acute, unawned or mucronate, mucros to 0.3 mm long; paleas 1.2-2.1 mm long, lanceolate, glabrous, acute; anthers 1-1.3 mm long, greenish-yellow to purplish at maturity. Caryopses 0.8-1 mm long, fusiform, brownish. 2n = 20, 22, 28. Distribution and habitat.—Muhlenbergia asperifolia grows in moist, often alkaline meadows, beach margins, and sandy washes, on grassy slopes, and around seeps and hot springs, at elevations of 55-3000 m. This species is common in western U.S.A. and Canada (Peterson 2003), and is also found in Argentina, Bolivia, and Chile (Zuloaga et al. 1994). In México M. asperifolia occurs in Baja California, Chihuahua, Durango, Jalisco, and San Luis Potosí (Espejo Serna et al. 2000). Comments.—The caryopses of Muhlenbergia asperifolia are frequently infected by a smut, Tilletia asperifolia Ellis & Everhart, which produces a globose body filled with blackish-brown spores. Specimen examined. MÉXICO. Coahuila: Municipio de Cuatrociénegas, D la Poza de la Becerra en Cuatrociénegas, J.A Villarreal Q. 3200, J. Valdés-Reyna & M.A. Carranza P. (ANSM 8. Muhlenbergia depauperata Scribn., Bot. Gaz. 9:187, f. 1-2. 1884. (Fig. 6, A-C). Tre: U.S.A. Arizona: Cochise Co. or Santa Cruz Co.: M. Mountains, 13 P E n € s.n. mu TOTYPE: US-994221! designated by McVaugh, Fl. Novo- Galiciana 14:236. 1983, b dicated by Peterson & Annable, Syst. Bot. Monogr. 31:35. 1991; IsSOLECTOTYPES: K!, MO!, MSCII, NYI, PI, USE VAN, W-1916- 27671. Muhlenl haffneri E. Fourn., Mexic. Pl. 2:85. 1886. Type: MÉXICO. Tacubaya, W. Schaffner s.n. (Lectotype: P! up Hitchcock, MN US: Natl. Herb. 17:294. 1913; ISOLECTOTYPES: LE (2 sheets)!, MO!, P!, US-3412353!, US-2312354 fragm. ex P Caespitose annuals. Culms 3-15 cm tall; internodes mostly scaberulous or pubescent, pubescent or strigose below the nodes, much branched at the lower nodes. Leaf sheaths often longer than the internodes, some- what inflated, smooth or scabrous, keeled, margins scarious; ligules 1.4-2.5 mm long, membranous, acute, with lateral lobes; blades 1-3 cm long, 0.6-1.5 mm wide, flat or involute, scabrous to strigose, midveins and margins thickened, whitish. Panicles 2.5-8.5 cm long, 0.5-0.7 cm wide, contracted; primary branches 1-2.2 cm long, appressed, spikelet-bearing to the base, spikelets borne in subsessile-pedicellate pairs; lon- ger pedicels 3-6 mm long, scabrous; disarticulation beneath the spikelet pairs at base of pedicel. Spikelets 2.5-4.5 mm long, appressed; glumes 2.3-5.1 mm long, equaling or exceeding the florets; lower glumes 2.3-4 mm long, subulate, 2-nerved, minutely to deeply bifid, teeth aristate or with awns to 1.3 mm long; upper glumes 3-5.1 mm long, lanceolate, 1-nerved, entire, acuminate; lemmas 2.5-4.5 mm long, narrowly Peterson et al., Muhlenl NS $ \ bu 1mm A 227. v RS RATS AES ) K di EZ Bg. ` ame AZ NS NE! y I SONS NS SA A AA E Y N A SN N ` NS A P 951 Fi. 5. Muhlenbergia asperifolia [P.M. Peterson 1703, C.R. Annable, R.F. Thorne & K. Kubitskie (US)]. A. Habit. B. Ligule. C. Glumes. D. Lemma. Muhlenbergia tenuifolia [PM. Peterson 9705 & A. Campos-Vill (US, ANSM)]. E. Habit. F. Ligule. G. Glumes. H. Floret. 952 Journal of the Botanical R h Institute of Texas 1(2) i | i Fic. 6. Muhlenbergia depauperata [PM. Peterson 4082 & C.R. Annable (ARIZ, ENCB, GH, MEXU, MICH, MO, NMC, NY, RSA, TAES, UC, UNLV, UTC, US, WIS, WS)]. A. Habit. B. Glumes. C. Florets. Muhlenbergia fragilis [PM. Peterson 5456 & C.R. Annable (US)]. D. Habit. E. Ligule. F. Glumes. G. Lemma. Muhlenbergia minutissima [PM. Peterson 5601 & C.R. Annable (US)]. H. Habit. I. Ligule. J. Spikelet. K. Floret. Peterson et al., Muhlenl ii f T a Mavi T lanceolate, light greenish-brown to purplish, scabrous, appressed-pubescent on the margins and midveins, apex acuminate, awned, the awns 6-15 mm long, stiff; paleas 2.4—3.6 mm long, lanceolate, intercostal region appressed-pubescent, apex acuminate; anthers 0.5-0.8 mm long, purplish to yellowish. Caryopses 1.5-2.3 mm long, narrowly fusiform, brownish. 2n = 20. Distribution and habitat. —Muhlenbergia depauperata grows in gravelly flats, rock outcrops, exposed bedrock, and sandy banks, in grama grassland associations, usually on soils derived from calcareous parent materials, associated with species of Selaginella, Hechtia, Opuntia, Salvia, Brahea, and Juniperus; 1530-2400 m. The range of M. depauperata extends into the southwestern U.S.A., and in México this species occurs in Chihuahua, Durango, Aguascalientes, México, Distrito Federal, Guanajuato, Jalisco, San Luis Potosí, Puebla, Veracruz, and Oaxaca (Espejo Serna et al. 2000 Comments.—Muhlenbergia depauperata and M. brevis C.O. Goodd., found commonly in the Sierra Madre Occidental, share several features with Lycurus: spikelets borne in pairs, 2-nerved and 2-awned lower glumes, ]-nerved and awned upper glumes, acuminate, awned lemmas with short pubescence along the margins, and pubescent paleas. Specimens examined. MÉXICO. Coahuila: Municipio de Ocampo, W side of Potrero de la Mula, 20 km NW of Ocampo, I.M. Johnston 9250 (MEXU); Municipio de Saltillo, Sierra Madre Oriental, 18 mi S of Saltillo on México 54 and 1.3 mi W on road to microondas, = W of Estación Carneros, P.M. Peterson 8425 & M.A. Carranza P. (ANSM, MEXU, US); Municipio de Torreón, Sierra de Jimulco, 3 f Mina San José and 8 km NE of Estación Otto, M.C. Johnston 9552, F. Chiang C.& T.L. Wendt (ANSM). Nuevo León: 6.5 mi S of en of CHE and Nuevo Leon on Hwy 57 towards Matehuala, P.M. Peterson 13282 & M.B. Knowles (US); 5.2 mi S of Zaragoza on road towards Ejido La Encantada, P.M. Peterson 16741, J. Valdés-Reyna & M. Sosa Morales (US); Municipio de Santiago, La Escondida, I. Cabral- Cord 98 (ANS 9. Muhlenbergia distichophylla (J. Presl) Kunth, Enum. Pl. 1:202. 1833. (Fig. 7, A-C). Podosemum distichophyllum J. Presl, Reliq. Haenk. 1(4—5):231. 1830. Epicampes stricta var. distichophylla J. Presl) M.E. Jones, Contr. W. Bot. 14:6. 1912. Type: MÉXICO: T. Haenke 173 (HoLotype: PR, photo US!; isotypes: MO-1837831!, US-90711 fragm. ex PRI, US fragm. ex W!, WU). Meu Mi ia Swallen, N. Amer. Fl. 17(6):457. 1935. Type: MEXICO. JaLisco: rocky hills near Guadalajara, 11 Nov 1889, C.G. US-822882!; isotypes: LE!, MO-1837815!, US-995828!, US-3274342 fragm!). Caespitose ME Culms 100-180 cm tall, erect, glabrous to pubescent below the nodes; internodes glabrous. Leaf sheaths 8-42 cm long, longer than the internodes below, glabrous, the keels prominent, sometimes coiled to shredded below, basal sheaths compressed-keeled; sheath auricles 0.4-2.6 cm long on lower portions and up to 6.4 cm long above; ligules 4-15 mm long, membranous, apex finely lacerate some- times almost to base; blades 18-90 cm long, 2-7 mm wide, flat or folded, scaberulous to scabrous above and below, the margins and keel saw-toothed. Panicles 35-70 cm long, 4-15 cm wide, densely-flowered, oblong, sometimes lax near apex, greenish-brown, sometimes reddish-purple; primary branches 2-15 cm long, without spikelets near base, appressed to loosely spreading up to 60 from the rachises; pedicels 0.2-4 mm long, glabrous to scaberulous. Spikelets 1.5-2.8 mm long, erect, greenish-brown, to reddish-purple; glumes 1.2-2.8 mm long, longer, as long or a little shorter than the lemma, subequal, oblong to narrowly-oblong, faintly 1-nerved, hyaline, glabrous to scaberulous, usually with faint, widely scattered hairs, the hairs less than 0.1 mm long, apex acute to acuminate; upper glumes rarely mucronate, the mucro up to 0.4 mm long; lemmas 1.4-2.7 mm long, lanceolate to linear-lanceolate, awned, glabrous or sometimes the margins on the lower 1/3 pubescent, the hairs up to 0.2 mm long, rarely the lower 1/3 with scattered hairs, callus usually short pilose, apex acute, minutely bifid, the teeth up to 0.5 mm long, the awn 4-16 mm long, flexuous, often reddish-purple near base; paleas 1.3-2.7 mm long, glabrous or with a few hairs between the nerves on the lower 1/3, apex acute; anthers 1.2-1.5 mm long, yellowish, sometimes reddish tinged. Caryopses not seen. Chromosome number unknown. Phenology.—Flowering late September through December. Distribution and habitat.—Muhlenbergia distichophylla occurs from central México in Jalisco, Guerrero, México, Oaxaca, Chiapas to Guatemala; found in open pine-oak forests and tropical deciduous forests on rocky slopes, canyons, and ravines, with Pinus sp., Quercus spp., Arbutus sp., Dioon sp., Dasylirion sp., and Agave sp.; 400-2000 m 954 I Lnaft+h,D ¿VU Mal OF IN MI d J te of Texas 1(2) 1. King (US)]. D. Habit. E. Panicle. F. Lig ule. G. Glum lla [L.0. Williams 22309, A. Molin T.P. Williams (U /. Gol ae US US)]. A. Ligule. B. Glumes. C. Floret. Peterson et al., Muhlenl ii f T a Mavi T Comments.—Morphological differences among many species of Muhlenbergia subg. Trichochloa sect. Epicampes, of which M. distichophylla is a member, are very minimal. In NE México the following seven spe- cies represent this section: M. distichophylla, M. emersleyi, M. jaime-hintonii, M. lindheimeri, M. pubescens, M. pubigluma, and M. robusta. Preliminary molecular analyses of both nuclear and chloroplast DNA sequences also exhibits little variation among members of this section, perhaps indicating that current species delimi- tation is too fine- juni (Peterson et al. 2001a, 2004). MÉXICO. Tamaulipas: 42 km SW of hwy 85 on road towards Dulces Nombres, P.M. Peterson 15913 & J. Valdés- eee (ANSM, US); 14.6 mi NE of Dulces Nombres, P.M. Peterson 15921 & J. Valdés-Reyna (ANSM, US). 10. Muhlenbergia dubia E. Rn Biol. ae -Amer., Bot. 3(29): 540. 1885. (Fig. 8, A-C). Tre: MÉXICO. Oaxaca: Chinantla, May 1841, EM. Li 68 TYPE: P'; isotypes: C, US-91018 fragm). Muhlenbergia acuminata Vasey, Bot. Gaz. 11(12):337—338. 1886. Type: U.S.A. New Mexico: probably Grant Co. near Santa Fe (fide Allred, Great Basin Naturalist 50:75. 1990), 1851-1852, C. Wright 1993 (notorvee: US-81605!; isotypes: GH, US-995177!). Sporobolus ligulatus Vasey & L.H. Dewey, Contr. U.S. Natl. Herb. 1(8): 268. 1893. Tyre: U.S.A. Texas: Presidio Co.: Limpia Cañon, Sep Nealley 127 (HoLoTYrE: US-82018!; isotype: US-5568871) Muhlenbergia firma Beal, Grass. N. Amer. 2:243. 1896. Type: MÉXICO. Oaxaca: Sierra de San Felipe, summit ledges, 10,500 ft, 18 Sep , C.G. Pringle 4914 (HoLoTYPE: MSC; isorvees: LE!, MO-2974120!, US-746252!, US-822866!, US-250841!; W-1895-4473!). Muhlenbergia densiflora Scribn. & Merr., Bull. Div. Agrostol., U.S.D.A. 24:18, f. 4. 1901. Tree: MÉXICO. Distrito FEDERAL: Serrania de Ajusco, on lava beds, 10,000 ft, 13 Aug 1897, C.G. Pringle 6675 (notorvee: US-316900!; isotypes: CM-279867, LE!, US-822867!; W-1898-2942! Crypsinna brevig umis M.E. Jones, Contr. W. Bot. 14:8. 1912. Type: MÉXICO. Cuinuanua: Guayanopa Canyon, Sierra Madre Mountains, Sep 1903, M.E. Jones s.n. (HOLOTYPE: POM?; IsoTYPE: US-3168555 fragm!). Densely caespitose perennials. Culms 30-100 cm tall, erect, rounded near the base, not rooting at the lower nodes; internodes glabrous for most of their length, minutely pubescent to hirtellous below the nodes. Leaf sheaths longer than the internodes, smooth or scaberulous, not becoming spirally coiled when old, rounded basally; ligules 4-10 mm long, membranous, firm below, acute, lacerate, brownish; blades 10-60 cm long, 1-2 mm wide, usually involute (occasionally flat), scabrous abaxially, hispidulous adaxially. Panicles 10—40 cm long, (0.6-)1-2.4 cm wide, contracted, grayish-green; primary branches 0.6-7 cm long, diverging up to 40° from the rachises, stiff, spikelet-bearing to the base; pedicels 0.1-6 mm long, strongly divergent, hispidulous. Spikelets 3.8-5 mm long, grayish green; glumes (1.8223 mm long, equal, shorter than the florets, glabrous and smooth proximally, scaberulous distally, faintly 1-nerved, acute; lemmas 3.8-5 mm long, narrowly lanceolate, calluses hairy, hairs to 0.5 mm long, lemma bodies glabrous and smooth below, scabrous distally, apices acuminate, unawned, mucronate or awned, the awns 1-6 mm long, straight; pa- leas 3.8-5 mm long, narrowly lanceolate, glabrous below, acuminate; anthers 1.5-2.2 mm long, greenish. Caryopses 2.5-3.5 mm long, fusiform, brownish. 2n = 40, 50. Phenology.—Flowering July to November. Distribution and habitat. —Muhlenbergia dubia grows on steep slopes, ridge tops, limestone rock outcrops, and along draws with Juniperus flaccida, J deppeana, Quercus spp», ugue ponderosa, P. edulis, P. pseudostrobus, P. johannis, Arctostaphylos pungens, Cercocarpus breviflorus, Fall Panicum bulbosum, Piptochaetium fimbriatum, Lycurus phleoides, Buddleja scorpioides, Nassella ru d Juglans sp., Selaginella sp., Dasylirion sp., Hechtia sp., Opuntia spp., Salvia sp., Brahea sp., Agave sp., Yucca sp., Bouteloua hirsuta, B. unioloides, and Aristida spp.; 1500-3200 m. The range of M. dubia extends into the western Texas and southern New Mexico, U.S.A., and in México this species occurs in Chihuahua, Durango, Aguascalientes, Zacatecas, Michoacán, México, Distrito Federal, Tlaxcala, Guanajuato, Querétaro, Jalisco, Hidalgo, San Luis Potosí, Puebla, Vera- cruz, and Oaxaca (Espejo Serna et al. 2000). Comments.—Muhlenbergia dubia is morphologically similar to M. rigens and can be separated from the latter by having a more loosely ascending and wider panicles (up to 2.4 cm wide), longer panicle branches (up to 7 cm long), olivaceous anthers (verses reddish), longer lemmas, and shorter glumes (less than 3 mm long). Traditionally, M. firma was recognized as having longer ligules, a dense spike-like panicle verses a narrow but scarcely spikelike, and glume apices that are sub-aristate verses obtuse. All of these characters 956 Journal of the Botanical R h Institute of Texas 1(2) IY Z^ ze c A AR == <= E, N N s SS LEAR FSS = Oe SS SSS Se < = SEL SISSI => = = L us - Ex > => e << my à = — d f VA » eZ — YL LLY” tz GEE CLARA AE => EXA HÁZ e p ET a EE BEEBE ee T ES c mo A, Z = = = a m : Z <= = EL = SZ LEME : gy, z == = RSE ZA " NSE WT. SS ERR - ASS AO OS s === a m — SS Ko e SC m E SY uo == => => t —À == a WY Z — > WS E AS SSS SON S = MN SS — SS | NS ~ is NS PE ES LA e dE == === -—— — SS = : CES = —— ea == Ss == a z z ==35 > £u Fic. 8. Muhlenbergia dubia [PM. Peterson 8384 & J. Valdés-Reyna (ANSM, US)]. A. Ligule. B. Glumes. C. Floret. Muhlenbergi ida [P. D. Habit, E. Panicle (narrow). [PM. Peterson 10876, C.R. Annable & J. Valdés-Reyna (US, ANSM)]. F. Panicle (wide). G. Ligule. H. Glumes. I. Floret. Peterson et al., Muhlenl ii f +1 í Mavi 35 vary on a single individual and within a population, and there appears to be very little correlation between distribution and morphology. Therefore, it seems best to recognize a single species. Our observations show as one descends in latitude from the U.S.A. and into México a general trend is seen in the coloration of the panicles/spikelets. The northern forms are usually greenish-gray whereas the forms from Chihuahua south and east are often dark-reddish or greenish-gray. The type, from Oaxaca, is the greenish-gray form. Usually the dark-reddish forms tend to have slightly longer florets. Sp i MÉXICO. Coahuila: 2-3.5 mi E of Hwy 54 up San Lorenzo Canyon, P.M. Peterson 13260, 13264, J. Valdés-Reyna e M.B. Knowles (US); Sierra El Pino, 33.5-39.5 km W of Rancho El Cimarron, P.M. Peterson 10659, 10665, 10689 & C.R. Annable (US); 140.3 kn NW ue on Hwy 53 towards Boquilla del Carmen, P.M. Peterson 10593, 10594 & C.R. Annable (US); 4 km E of Los Lirios on road to Laguna hez, 50 km SE of Saltillo, P.M. Peterson 6250 & C.R. Annable (US); 9.7 km E of Los Lirios on road to Laguna de Sanchez, 56.3 km SE of Saltillo, P. M. Peterson 6256 & C.R. Annable (US); 12.9 km E of Los Lirios on road to Laguna de Sanchez, 60 km SE of ui P.M. Peterson 6261 & C.R. Annable (US); Sierra Zapaliname, ca. 5 km east of Saltillo (Las Palapas) up Camino de Quatro; bre, P.M. Peterson 17862, 17866, J. Valdés-Reyna & R.H. Cardenas (ANSM, US); entrance to Monterreal, at 33mi E of TUM P.M. Peterson 15944 & J. Valdés-Reyna (US); Municipio de Acuña, Serranías del Burro, Rancho El Bonito, Cañón El Bonito, 7 mi W of la casa principal, J. Valdés-Reyna 235 & D. H. Riskind (ANSM); Serranías del Burro, Rancho El Bonito, Cañón el Toro, J. Val- ^ Reyna 1173 & D. H. Riskind Nbre eae as Burro, Rancho El Bonito, Cañón Los Ojitos, J. Valdés-Reyna 1259 & D. H. Riskind ANSM M); Municipio de Arteaga, 32 mi SE & E of Jamé, camino towards Las Vigas, P.M. Peterson 10058, C.R. Annable & J. Valdés-Reyna (ANSM, US); 55.3 km SE of Saltillo and 16.6 2 SE of Jame on road to Sierra La Viga, P.M. Peterson 10064, C.R. Annable & J. Valdés-Reyna (US); Jamé -Rayones, G.S. F 7809 (HINT); El Coahuilón, Sierra de la Marta, J. Valdés-Reyna 1835, M.A. Carranza P. & J.A. Villarreal Q. (ANSM); Fraccionamiento Bravo León, E. Pérez-Torres 3 (COCA); Las Vigas, Cañón de la Carbonera, Sierra de Arteaga, J Valdés-Reyna 1792, M.A. Carranza P. & J.A. Villarreal Q. (ANSM); Rancho el Chorro, carretera al Tunal, 3 km E de la desviación al Tunal, sobre la carretera de Saltillo-Matehuala, J. Espinosa-Aburto 168 (ANSM); Sierra de Arteaga, Cañón La Carbonera, camino al Tunal, M.A. Carranza P. 2436 € N. Snow (ANSM); Sierra de Arteaga, El Tunal, carretera estatal 65, P.S. Hoge. 267, M.E. Barkworth & J. Valdés-Reyna (ANSM); Sierra Madre Oriental. SE of San Antonio de las Alazanas and SE of Saltillo at end of road near summit of Coahuilón, P.M. Peterson 8384 & 8391, J.A. Villarreal Q. & J. Valdés-Reyna (ANSM, US); Municipio de Ramos Arizpe, Sierra de la Paila, Ejido el Cedral amino hacia el Valle de Parreños, J.A. Villarreal Q. 5380, M.A. Carranza P. & A. Rodríguez G. (ANSM); Sierra de la Paila, Ejido el Cedral por el camino El Carmen, J. Valdés-Reyna 2184 (ANSM); Municipio de Saltillo, Cañada Salsipuedes, en la Sierra Zapalinamé, 3 km NO de Cuahtémoc, R. López-Aguillóno s.n. (ANSM); Cañón de San Lorenzo, en la Sierra de Zapalinamé, 8 km S de Saltillo, 3.2 km E de la Universidad Autónoma Agraria Antonio Narro, R. López-Aguillón s.n. (ANSM); ca. 6 km S of Saltillo, land owned by Universidad Au- tonoma Agraria Antonia Narro, P.M. Peterson 8344 & J. Valdés-Reyna (US); “Lomas de Lourdes, P.A. Lobato 5 (COCA); highway 54 (road from Saltillo to Concepción del Oro), S of Estación Carneros on telephone line service road, M.E. Barkworth 5122, J. Valdés-Reyna, P. S. Hoge & M.A. Carranza P. (ANSM); 24.2 km S of Saltillo on MEX 54, 16.1 km E to Rancho Experimental Ganadero, P.M. Peterson 8434 & M.A. Carranza P. (US); Rancho Experimental Los Angeles, 48 Km S de Saltillo, carretera 54 Saltillo- Concepción del Oro, Zacatecas, S.L. Hatch Stephen 4551, C.W. Morden, J. Valdés-Reyna (ANSM); 21 km SE of Saltillo on Hwy 57 towards Matehuala, P.M. Peterson 15799 & J. Valdés-Reyna (US); Sierra de ae amé, 1 km S a Cañón de San Lorenzo, R. López-Aguillón s.n. (ANSM); Sierra de Zapalinamé; Lomas de Lourdes, al SE de Saltillo M gi te 2951 (ANSM); Sierra la Concordia, 40 km SO de Saltillo, J.A. Villarreal Q. 6567, M.A. Carranza P., J. Valdés- "i M. VEER, & D. E. Lozano (ANSM); Sierra la Concordia, 6 km NE de la Victoria, J.A. Villarreal Q. 4084, M.A. Carranza P. & A. Rodríguez G. (ANSM); Sierra Madre Oriental, 18 mi S of Saltillo on México 54 and 1.3 Mi W on road to microondas, just W of Estación Carneros, P.M. Peterson 8419 & M.A. Carranza P. (ANSM, US); Sierra Madre Oriental. Cuesta de Palmas Altas, R. Marie Francois 5748 & J. Passini (ANSM); 5 mi W of Chapultepec on cutoff road between Hwy 54 € 57, 23 mi S of Saltillo, P.M. Peterson 13274, 13277 & M.B. Knowles (US). Nuevo León: 3 mi NE of Dulce Nombres, 5.2 mi SE of San Pablo and 4 mi NW of San Francisco Javier, P.M. Peterson 18958 & J. Valdés-Reyna (ANSM, US); Sierra El Pinal Alto, 1.1 mi N of San Pablo, P.M. Peterson 18937, 18941 & J. Valdés-Reyna (ANSM, a Sierra El Pinal Alto, 3.7 mi N of San Pablo, P.M. Peterson 18946 & J. Valdés-Reyna (ANSM, US); 16 W of Laguna de Sanchez and 21 km E of Los Lirios, P.M. Peterson 6272 & C.R. Annable (US); 2.5 mi N of La Siberia on road towards La Encantada, P.M. Peterson 16775, J. Val cue Reyna & M. Sosa Morales (US); 5.2 mi S of dee on road i oe La Encantada , P.M. Peterson 16749, J. Valdés-Reyna & M. Sosa Morales (US); La Lagunita; 9.5 mi SE of Ar ds Agua Fria, P.M Peterson 16695, J. Valdés-Reyna & M. Sosa Morales (US); 13.5 mi SE of A dt ds Agua Fria, P.M. Peterson 16731, J. Valdés- Reyna & M. Sosa Morales (US); 8 km S of La Cruata on Hwy 3 towards Aramberri, P.M. Peterson 15885, 15887 & J. Valdés-Reyna (US); 2 mi W of Dieciocho de Marzo up road towards Cerro Potosi, P.M. Peterson 13319, 13321 & M.B. Knowles (US); 6.7 mi W of Dieciocho de Marzo up road towards Cerro Potosi, P.M. Peterson 13328, 13330 & M.B. Knowles (US); 10.4-12.7 mi W of Dieciocho de Marzo up road towards Cerro Potosi, P.M. Peterson 13331, 13339, 13342, 13356 & M.B. Knowles (US); Municipio d l San José del Río, G.S. Hinton 21508 (HINT); Cañón de los Capulines above San Enrique, Hacienda San José de Raices, H. C. Mueller 2367 (MEXU); Ciénegas del Toro, J.L. Elizondo-Elizondo 204 & R. Banda S. (ANSM); Sierra el Infiernillo, Cañón San Francisco, 15 km NE de Pablillo, J. Valdés-Reyna 2324, M.A. Carranza P. & R. Banda S. (ANSM); p de General Zaragoza, Sierra El Soldado, camino a Puerto Pinos, J.A. Villarreal Q. 4957, M.A. Carranza P., G. Nesom & J. Norris (ANSM, MEXU); Municipio de Iturbide, 10 mi W of Iturbide , J. Brunken. 202 & C. Perino (TAES); Municipio de Linares, Las Palmas-El Pinal, J.J. Ortíz-Díaz 21 (ANSM); N I g ola de Caballo, 958 Journal of the Botanical R h Institute of Texas 1(2) J.A. Villarreal Q. 5556 & M.A. Carranza P..(ANSM); La Escondida, I. Cabral- Corderol 592 (ANSM); Laguna de Sánchez, P. Jauregui-Ramirez 127 (COCA). Tamaulipas: 42 km SW if hwy 85 on road towards Dulce Nombres, P.M. Peterson 15914 & J. Valdés-Reyna (ANSM, US); 14.6 mi NE of Dulce Nombres, P.M. Peterson 15920 & J. Valdés-Reyna (ANSM, US); 101.6 km SW of Ciudad Victoria on MEX 101 towards San Luis Potosi, P.M. Peterson 8326 & R.M. King (US); Municipio de Bustamante, Ejido Felipe Angeles 2 km NO rumbo a Bustamante, R. Díaz-Pérez 187 (UAT); Municipio de Hidalgo, Los Caballos, G.S. Hinton 24809 (HINT); Los Caballos, G.S. Hinton 24809 (MEXU), G. Villegas-Durán 537 (COCA); Municipio de Jaumave, Ejido Avila & Urbina, J.G. Galván-Infante 103 (COCA); Municipio de Palmillas, Ejido Palmillas, R.A. Carranco-Rendon. 197 (COCA); Municipio de Tula, Valle de Tula, G. Villegas-Durán 544 (COCA) 11. Muhlenbergia emersleyi Vasey, Contr. U.S. Natl. Herb. 3(1):66. 1892. (Fig. 9, A-C). Epicampes arid (Vasey) Hitchc., U.S.D.A. Bull. (1915-23) 772:44-145. 1920. Tyre: U.S.A. ARIZONA: 1890, J.D. Emersley 46 (HoLOTYPE: US-73223!). -] 2289 Muhlenbergia vaseyana Scribn., Ann. Rep. Missouri Bot. Gard. 10:52. 1899. Type: U.S.A. Arizona: Jul 1874, J.T. Roth ] US-81633! designated by Soderstrom, Contr. U.S. Natl. Herb 34(4): 166. 1967, without citing a specific sheet in a PC nM barium). Epicampes subpatens Hitchc., U.S.D.A. Bull. (1915-23) 772:144. 1920. Type: U.S.A. New Mexico: Eddy Co.: Ranger Station, Queen, Guadeloupe Mountains, 3-6 Sep 1915, A.S. Hitchcock 13541 [Amer. Gr. Herb. 1382] (HoLotyPE: US-905799!; isotypes: LE!, US- 3215629!, US-3278409!). Muhlenbergia distans Swallen, N. Amer. Fl. 17(6):461. 1935. Type: MEXICO. Oaxaca: Las Sedas, dry hills, 6000 ft, 15 Sep 1894, C.G. Pringle 5575 (HoLotype: US-746297!; isotypes: MO-1837832!, TAES!, US-134322!, US-305676! Muhlenbergia gooddingii Soderstr., Contr. U.S. Natl. Herb. 34(4):115. 1967. Type: U.S.A. Arizona: Pima Co.: Baboquivari Mountains, canyon north of Moristo Canyon, 4000 ft, 19 Nov 1945, L.N. e 462-45 (HoLoTYPE: YU; isotypes: ARIZ!, MO-5073874!, NY, US-2550348!). Densely caespitose perennials. Culms (65—)80-150(-200)cem tall, erect, stout, not conspicuously branched; internodes smooth for most of their length, smooth or scaberulous below the nodes. Leaf sheaths 7-35 cm long, shorter or longer than the internodes, glabrous or puberulent, basal sheaths compressed-keeled, usu- ally keeled; sheath auricles lacking; ligules 10-25 mm long, membranous throughout, acuminate, lacerate, slightly firmer below; blades 20-50 cm long, 2-6 mm wide, flat or folded, scabrous abaxially, smooth or scaberulous adaxially, margins coarsely scabrous. Panicles 20-45 cm long, 3-15 cm wide, loosely contracted to open, light purplish to light brownish; primary branches 1-17 cm long, lax, loosely appressed or diverging up to 70° from the rachises, naked basally; pedicels 0.5-3 mm long, erect, smooth or QUE Spikelets 2.2-3.2 mm long; glumes 2.23.2 mm long, longer or as long as the lemma, subequal, scal b faintly 1-nerved, acute to obtuse, usually unawned, occasionally mucronate, mucro to 0.2 mm long; as 2-3 mm long, oblong-elliptic, pubescent along the midvein and margins on the lower Y2-%, apices acute, usually awned, sometimes unawned and or mucronate, awns generally (1-)6-15 mm, flexuous, purplish; paleas 1.8-2.9 mm long, oblong-elliptic, acute; anthers 1.2-1.6 mm long, yellowish to purplish. Caryopses 1.3-1.6 mm long, fusiform, reddish-brown. 2n = 24, 26, 28, 30, 40, 42, 46, 60, 64 Phenology.—Flowering July to November. Distribution and habitat—Rocky slopes, gravelly washes, canyons, cliffs, and along streams often derived from limestone parent material associated with Quercus spp., Pinus cembroides, P. chihuahuana, P. jefferyi, P. ponderosa, P. edulis, P. reflexa, Juniperus deppeana, Arctostaphylos pungens, A. patula, Ceanothus leucodermis, Agave sp., Prosopis sp., Plantanus wrightii, Rhus trilobata, Cercocarpus breviflorus, Chilopsis linearis, Fallugia paradoxa, and Bouteloua curtipendula, 1200-2600 m; western Texas, New Mexico, Arizona and common throughout LA México. Comments.—Muhlenbergia emersleyi is an extremely variable species. Soderstrom (1967) pointed out that M. emersleyi is the most common species of Muhlenbergia sect. Epicampes and that, “attempts to segregate the forms into more than one species on the basis of morphology are unrewarding because of a thorough intergradation of characters among the specimens." Preliminary molecular evidence seems to support this conclusion because there apparently is very little sequence divergence among closely related species of this section (Peterson et al. 2001a, 2004). Specimens examined. MÉXICO. Coahuila: 26 mi E of Saltillo, J.R. Reeder 3289, C.G. Reeder & T.R. Soderstrom [n=30](US); Municipio de Acuña, del Carmen mountains, E.G. Marsh Jr. 655 (MEXU); Serranías del Burro, Rancho El Bonito, Cañón El Bonito, 7 mi O de la casa principal, en la ultima pila, J. Valdés-Reyna 1237, D.H. Riskind (ANSM); Serranías del Burro, Rancho El Bonito, Cañón el Toro, R. Valdés- Reyna 1174 & D.H. Riskind (ANSM); Municipio de Arteaga, 12 km de Saltillo, hacia Matehuala, M.A. Madrigal-A. s.n. (ANSM); 17 mi SE Peterson et al., Muhl Fic. 9. Muhlenbergi leyi U.R. Reeder 7393 & C.G. Reeder (ARIZ, US)]. A. Ligule. B. Glumes. C. Floret. Muhlenbergia lindheimeri (PM. Peterson 6280 & CR. Annable (US)]. D. Habit. E. Ligule. F. Glumes. G. Floret. 960 Journal of the Botanical R h Institute of Texas 1(2) of Saltillo & 7.4 mi NW of Jamé, P.M. Peterson 10077 (ANSM, US); Sierra de Arteaga, El Tunal, carretera estatal 65, P.S. Hoge 263, M.E. Barkworth & J. Valdés-Reyna (ANSM); Municipio de General Cepeda, km 23 Carretera de General Cepeda a Parras, J. Espinosa-Aburto 154 b oe de Ramos Arizpe, Sierra de la Paila (Lado Norte) Cañada Becerros, J.A. Villarreal Q. 5464, M.A. Carranza P. & L. Arce G. (AN pio d illo, Estación Carneros, carretera a Zacatecas 30 km S de Saltillo, M.A. Carranza P. 515 & P.M. Peterson (ANSM); Rancho experimental Los Angeles, 48 km S de Saltillo, carretera 54 Saltillo- Concepción del Oro, Zacatecas, J.A. Villarreal Q. 3370, J. Valdés-Reyna (ANSM); Sierra la Concordia, 6 km NE de la Victoria, J. A. Villarreal Q. 4083, M.A. Carranza P. & A. Rodríguez G. (ANSM); Sierra Madre Oriental, Sierra Zapalinamé, 1 km S del Cañón de San Lorenzo, R. López-Aguillón s.n. (ANSM). Nuevo León: Municipio de Galeana, Agua Blanco a San Miguel, G.S. Hinton 21337 (HINT); A orillas del p l Gavilán, P. Jauregui-Ramirez 37 (COCA); Municipio de General Zaragoza, 4 km S of Zaragoza at jtn of road to Cerro Viejo-Tepehuanes, P.M. Peterson 15854 & J. Valdés-Reyna (ANSM, US); 6 km S of Zaragoza, P.M. Peterson 15861 & J. Valdés-Reyna (ANSM, US); 8 km S of La Cruata on hwy 3 towards Aramberri, P.M. Peterson 15890 & J. Valdés-Reyna (US); 19 km S de Zaragoza, camino a La Encantada, J.A. Villarreal Q. 7539 & M.A. Carranza P. (ANSM); Muni- cipio de Linares, Las Palmas-El Pinal, J.J. Ortiz-Diaz 53 (ANSM); Municipio de Santiago, Las Gracielas, I. Cabral-Cordero 552 (ANSM); San José de Las Boquillas, I. Cabral- Cordero 563 (ANSM). Tamaulipas: Municipio de Aldama, Ejido Lauro Aguirre, P. Moya-Salgado 246 (COCA); Municipio de Güémez, Camino a Los San Pedros, G. Villegas-Durán 532 (COCA); Los Pedros, G.S. Hinton 25135 (HINT); Los Pedros a Hacienda La Boca, G.S. Hinton 25302 (HINT); Municipio de San Carlos, El Rosario. Vicinity of Marmolejo, H.H. Bartlett 10882 (MEXU); Municipio de Tula, 30 km al N de Tula, I. Nuñez-T li 28 (COCA); Municipio de Victoria, Altas Cumbres, G. Bores-Kulman 81 (COCA); Altas Cumbres, J.F. Iribe-Duarte 160 (COCA); Camino al Molino, J.E. López de la Cruz 28 (COCA); Carretera Victoria-Tula, J.F. Iribe-Duarte J. Fernando 44 (COCA); Ejido Vicente Guerrero, Cisneros M. 187 (COCA); Minas de asbesto, 35 km al O de Ciudad Victoria, F. González-Medrano 3423, P. Torres V. Solís & M. Terrazas (MEXU). 12. piu fragilis Swallen, Contr. U.S. Natl. Herb. os 206. ent (Fig. 6, D-G). Wee: U.S.A. Texas: Brewster Co.: Sunny Glen, W of Alpine, 29 Sep 1935, B.H. Warnoch 23 JLOTYPE: US-1829290!; tsotyre: GH!). Caespitose annuals. Culms 10-38 cm, erect or spreading; scabrous or strigulose below the nodes; internodes mostly glabrous, smooth or scaberulous. Leaf sheaths 2.4-4.2 cm long, often longer than the internodes, scaberulous, margins hyaline; ligules 1-3 mm long, hyaline, obtuse, irregularly toothed to lacerate, with lateral lobes extended into auricles; blades 1-10 cm long, 0.4-2 mm wide, flat, scabrous abaxially, strigulose adaxially, margins and midveins whitish-thickened; panicles 10-24 cm long, 3.5-11 cm wide, diffuse; primary branches 2.2-6.2 cm long, capillary, diverging 80—100? from the rachises, straight; pedicels 6-10 mm long, delicate. Spikelets 1-1.2 mm long, appressed to slightly divergent from branch axes; glumes 0.5-1 mm long, equal to subequal, glabrous throughout or obscurely puberulent, hairs about 0.06 mm long, 1-nerved, apex obtuse or subacute; lemmas 1-1.2 mm long, oblong-elliptic, membranous, purplish to light brownish, not mottled, glabrous or densely appressed-puberulent on the margins and midveins, apices obtuse, unawned; paleas 0.9-1.2 mm long, oblong-elliptic, densely puberulent between the nerves or glabrous, apex obtuse; anthers 0.3-0.5 mm long, purplish. Caryopses 0.7-0.9 mm long, elliptic, reddish-brown. 2n = 20 Phenology. —Flowering August through September. Distribution and habitat.—Muhlenbergia fragilis grows on rocky talus slopes, cliffs, canyon walls, road cuts, and sandy slopes, often on calcareous parent materials, at elevations of 480-2200 m; southeastern California to western Texas, U.S.A., and south in México from Baja California, Sonora, Chihuahua, Sinaloa, Jalisco, Nayarit, Guanajuato, San Luis Potosi, Querétaro, Michoacan, Guerrero, Morelos, Puebla, Veracruz, Oaxaca, and Chiapas. It is usually found in oak-grama savannahs, thorn scrub forests, oak-yellow pine forests, and pinyon-juniper woodlands. Comments.— Occasionally populations may have some individual plants with completely glabrous lem- mas rather than the more common hairy-spikeleted forms. This morphological variation is not correlated with distributional or habitat preference. Specimen examined. MÉXICO. Tamaulipas: San Antonio de los Alamos, E base of Sierra de San Antonio, I.M. Johnston 8244 (US, UT). 13. Muhlenbergia glauca (Nees) B.D. Jacks., Index Kew. 2:269. 1895. (Fig. 10, A-D). Podosemum glaucum Nees, Linnaea 19(6):689. 1847. Tyre: MEXICO. A. Aschenborn 335 (HOLOTYPE: B2). Muhlenbergia huachucana Vasey, Contr. U.S. Natl. Herb. 3(1):69. 1892. Type: U.S.A. Arizona: Huachuca Mountains, Jul 1882, J.G. Lemmon 2915 (HoLoTv?E: US-81617!). Muhlenbergia lemmonii Scribn., Contr. U.S. Natl. Herb. 1(2):56. 1890. Type: U.S.A. Arizona: Huachuca Mountains, Lemon is gi 2918 (LECTOTYPE: US-746111! designated by Hitchcock, N. Amer. Fl. 17(6):449. 1935, witl ISOLECTOTYPES: US-994769!, US-994778!, US-746113!) O 961 Peterson et al., Muhlenl Fic. 10. Muhlenbergia glauca [PM. Peterson 6031 & C.R. Annable (US)]. A. Ligule. B. Glumes. C. Floret. D. Lodicules, stamens, and pistil. Muhlenbergia polycaulis [PM. Peterson 5647 & C.R. Annable (US)]. E. Habit. F. Ligule. G. Glumes. H. Floret. I. Lodicules, stamens, and pistil. 962 Journal of the Botanical R h Institute of Texas 1(2) Perennials with slender, creeping, well-developed rhizomes. Culms 25-60 cm tall, 1-2 mm thick, often decumbent, moderately stiff, sometimes erect; internodes mostly scabrous, retrorsely hispidulous below the nodes. Leaf sheaths longer than the internodes, scaberulous; ligules 0.5-2 mm long, truncate to obtuse, erose or lacerate; blades 4-12 cm long, 1-2.6 mm wide, flat to involute distally, not arcuate, scabrous abaxially, hirsute or scabrous adaxially. Panicles 4—12(-17) cm long, 0.3-2.4 cm wide, contracted, interrupted below; primary branches 0.33 cm long, usually appressed, occasionally diverging up to 30? from the rachises; pedicels 0.1-1.2 mm long, scabrous to hirsute. Spikelets 2.4.5 mm long; glumes 1.5-3.5 mm long, Y to about as long as the lemma, equal, 1-nerved, veins scabrous, apices acute or acuminate, usually Or awned, awns, if present, to 1.5 mm long; lemmas 2.4-3.4 mm long, elliptic, pubescent on the lower the Y of the midveins and margins, hairs to 0.6 mm long, tawny, apices acuminate to acute, awned, awns 0.1-3(-5) mm long, straight; paleas 2.2-3.4 mm, elliptic, intercostal region pubescent on the lower 1⁄2, apices acuminate to acute; anthers 1.8-2.4 mm long, orange. Caryopses 1.7-2 mm long, fusiform, brownish. 2n = 60 Phenology.—Flowering August through October. Distribution and habitat. — Calcareous rocky slopes, cliffs, canyon walls, table rocks, and volcanic rock outcrops in open vegetation associated with Quercus, Pinus cembroides, P. chihuahuana, Juniperus deppeana, Plantanus wrightii, Arctostaphylos pungens, Agave, Dasylirion, Yucca carnerosana, Rhus trilobata, R. glabra, Vitis arizonica, and Ceanothus buxifolius; 1200—2780 m; southeastern Arizona, southern New Mexico, southeastern Texas to northern México, also found in Chihuahua, Durango, Aguascalientes, San Luis Potosí, México, Distrito Federal, Guanajuato, Hidalgo, Jalisco, and Puebla (Espejo Serna 2000; Herrera Arrieta 2001; Peterson 2003). omments.—Muhlenbergia gl is hologically very similar to M. polycaulis and can be distinguished from that species by its E awned imus (the awn usually less than 3 mm long) and slender, well developed rhizomes. More study of these two species, along with the allied, M. arsenei and M. pauciflora, is needed. MÉXICO. Coahuila: Sierra del Paila, Oct 1910, C.A. Purpus 5006 (US); Western Coahuila, Sierra de la Madera, aie of “La Cueva” in Corte Blanco fork of Charretera Canyon, 5300-6500 ft, I.M. Johnston 8906 (US); UAAAN experimental ranch “Los Angeles” 14 km E of Mex pr 54, 30 km S of Saltillo, C. W. Morden 515 (US); Sierra El Pino, 33.5 km W of Rancho El Cimarron, P.M. Peterson & C.R. Annable El Pino, 40.3 km W of Rancho El Cimarron, P.M. Peterson & C.R. Annable 10664 (US); Sierra El Pino, 39.5 km W of Rancho El Cimarron, P.M. Peterson & C.R. Annable 10668 (US); Municipio de Acuña, Serranías del Burro, Rancho El Bonito, 12 km O de la casa del rancho El Bonito, Cañón Los Os J. Valdés-Reyna 1264 & D. H. Riskind (ANSM); Municipio de Arteaga, 17 mi SE of Saltillo & 7.4 mi NW of Jamé, at Bosques de M abin of J. Valdés-Reyna), P.M. Peterson 10072, C.R. Annable & J. Valdés-Reyna (ANSM, US); 6 Km Ed t tera 57 hacia los Lirios, J.A. Villarreal Q.1 5173, J. Valdés-Reyna, M.E. Barkworth & P.S. Hoge (ANSM); Bosques de Montaña, e 6 Km E de carretera 57 camino a Jamé, 30 Km SE e i Valdés-Reyna Jesús 2361, J.A. Villarreal Q. & M.A. Carranza P. (ANSM); SE of San Antonio de las Alazanas and SE of Salti g p Coahuilon, P.M. Peterson 8386, J. Valdés-Reyna & J.A. Villarreal (ANSM, US); Municipio de Múzquiz, Sierra La Encantada, aq ] 170 km NO de Múzquiz, cuesta de Malena, brecha Múzquiz-Boquillas del Carmen, M.A. dd 829 J Val ee Reyna, P.A. Fryxell, R. Vasque A. & O. Meza (ANSM); Municipio de Ramos Arizpe, Sierra de la Paila, Ejido el C Valle de Parreños, J.A. Villarreal " 5284, M.A. Carranza P. & A. Rodríguez G. (ANSM); Municipio de Saltillo, Cañón de San Lorenzo, en la Sierra de Zapalinamé, 8 km S de Saltillo, 3.2 km E de la Universidad Autónoma Agraria Antonio Narro, R. López-Aguillón s.n. (ANSM); Canón de Zapalinamé, en la Sierra Zapalinamé, 8 km S de Saltillo & 3.2 km E de Buenavista, Saltillo, L. Arce-González s.n. (ANSM); Sierra Zapalinamé, 1 k S del Cañón de San Lorenzo, R. López-Aguillón s.n. (ANSM). Nuevo León: Municipio de Aramberri, Carretera Galeana-La Ascensión, a 1.5 km del ieee de San Mns en BOR M.M. Castillo-Badillo 209 & N. Bazaldua B. (COCA); Municipio de Linares, Las Palmas-El Pinal, J.J. l Zaragoza, 6 km S of Zaragoza, P.M. Peterson 15871 G J. Valdés-Reyna (ANSM, US). 14. spas a gypsophila Reeder & C. Reeder, Madroño 18:186-190, f. 1 A1-H1. 1966. (Fig. 11, A-F). ÉXICO. Nuevo Leon: 3 mi E of junction of Linares-Galena road with Hwy. 85, 6400 ft, 30 Oct 1964, J.R. Reeder & C.G. Reeder 3963 (HOLOTYPE: YU; IsoTYPE: US-2524092!). Densely caespitose perennials. Culms (12-)25-80 cm tall, erect, pubescent below the nodes, the basal nodes terete, 2-4 nodes per culm; internodes scabrous. Leaf sheaths (48-22 cm long, shorter or longer than the internodes, pubescent to glabrous basal sheaths rounded becoming brownish, shredded and/or fibrous with age; ligules 1-6(-10) mm long, membranous above and firm and light brown below, decurrent, margins puberulent, apex truncate, acute or acuminate; blades 2-14(-25) cm long, 0.5-1.5 mm wide, tightly involute, 963 Fic. 11. Muhlenbergia gypsophila [PM. Peterson 6235 & C.R. Annable (US)]. A. Habit. B. Panicle. C. Ligule. D. Glumes. E. Floret. F. Lodicules, stamens, and pistil. Muhlenbergia purpusii [PM. Peterson 6227 & CR. Annable (US)]. G. Habit. H. Panicle. I. Ligule. J. Glumes. K. Floret. L. Lodicules, stamens, and pistil. 964 Journal of the Botanical R h Institute of Texas 1(2) somewhat rigid and falcate, scabrous to glabrous below and densely pubescent above, the hairs less than 0.1 mm long. Panicles (3-)5-18(222) cm long, 0.2-1 cm wide, contracted, spikelike, dense, stramineous to purplish; primary branches 0.5-2.7 cm long, ascending and tightly appressed; pedicels 0.6-2.6 mm long, shorter than the spikelets, scabrous. Spikelets 3.5-4.3 mm long, stramineous to purplish; glumes 1.2-2.5 mm long, shorter than the floret, obscurely 1-nerved, puberulent on upper 2/3, apex acute often erose; lemmas 3.5-4.3 mm long, lanceolate, stramineous to purplish, lateral nerves distinct and raised, short pi- lose throughout, the hairs 0.1-0.3 mm long, callus densely short pilose, apex acute to acuminate, entire to minutely bifid, awned, the awn 10-20 mm long, flexuous; paleas 3.5-4.3 mm long, as long as the lemma, short pilose between the nerves throughout, apex acute; anthers 1.5-2.4 mm long, greenish to purplish. Caryopses 1.7-2.1 mm long, fusiform, light brownish. 2n = 20 Phenology.—Flowering September through November. Distribution and habitat. —Muhlenbergia gypsophila occurs on calcareous derived soils primarily on gypsum flats usually on the sides of sink holes, roadsides, and slopes with Flaveria sp., Larrea divaricata, Prosopis sp., Pinus cembroides, Pinus spp., Bouteloua chasei, B. gracilis, B. uniflora, Muhlenbergia purpusii, Acacia sp., Puya sp., Agave lecheguilla, Juniperus flaccida, Rhus sp., Dasylirion longissimum, Condalia sp., Leucophyllum alejandrae, L. hintonii, Gutierrezia microphylla, Aristida spp.Yucca carnerosana, Quercus spp., and Brickellia sp.; 1025-2430 m. 1. MÉXICO. Coahuila: M io d Cuatrociénegas entrada al Casco del Rancho Potrero de Menchaca, R. Vásquez- Ae 17 & L.E. Rodríguez G. (ANSM); along road to Secession de Oro, Zacatecas, 35 mi S of Saltillo, J.R. Reeder & C.G. Reeder 3622. Nuevo Leon: 2 mi W of Dieciocho de Marzo up road towards Cerro Potosi, P.M. Peterson 13318 & M.B. Knowles oe 36 mi NE ix Dr Arroyo on Hwy 61 towards Linares, P.M. Peterson 13367 & M.B. Knowles (US); 8.7 mi NE of San Antonia de Peña N La Liberia, P.M. Peterson 16784, J.Valdés-Reyna & M. Sosa Morales (US); 13.5 mi S of Milagro, P.M. Peterson 17816, J. Valdés-Reyna & G.S. Hinton (US); 5.6 mi E of junction of Hwy 57 on Hwy 58 towards Linares, P.M. Peterson 13289, 13297, 13298 & M.B. Knowles (US); 13.4 mi E of Hwy 57 on Hwy 58 at crossing of Rio Potosí, P.M. Peterson 13299 & M.B. Knowles (US); Municipio d mberri, Aramberri, G.S. Hinton 23999 (ANSM, HINT); La Escondida, Aramberri-Dolores, G.S. Hinton 23597, 23655 (ANSM, HINT); 12 km S of Aramberri, P.M. Peterson 15848 & J. Valdés-Reyna (US); Municipio de Galeana, Near bends 25 Mar 1962, A.A. Beetle M-481 (US); 3 mi SE of Galeana, J.R. Reeder and C.G. Reeder 3659 (US); 10 km E del ent R sal , J. Valdés-Reyna 1721 & R. Banda S. (ANSM); 2 km S del Salero, 1 km E de carretera 257 Matehuala-Saltillo, J. Valdés-Reyna 1619 & J.S. Marroquín de la F. (ANSM); 3 km S of El Salero, P.M. Peterson 17834, J. Valdés-Reyna & G.S. Hinton (US); 7.5 km E of Puentes, P.M. Peterson 17851, J. Valdés-Reyna 6 G.S. Hinton (US); 20 km S of San Roberto along highway 57, S.L. Hatch 4942, J. Valdés-Reyna & J. Kessler (ANSM); 5 km E de Galeana, I. Cabral- Cordero 1082 (ANSM); 4 mi SE of Galeana, J.R. Reeder & C.G. Reeder 3965, 4987 (US); 1 mi S of Galeana, K.W. Allred 5502, T. Columbus & J. Valdés-Reyna (ANSM); Ejido El Tokio, M.L. Avalos-Marín s.n. D n ANSM); Galeana, J.A. Ochoa-Guillemar 1000 (COCA); La Becerra, G.S. Hinton 19868 (ANSM, HINT, MEXU); Santa Rita, 5 | ,J. 4. Villarreal Q. 6319 & M.A. Carranza P. (ANSM); 3mi SE of Galeana on road to Linares, P.M. Peterson 13314 & M. B. Knowles (US); San Jose del Rio, G.S. Hinton 21531 (HINT); Pabillo a La Ascensión, G.S. Hinton 2/496 (HINT); SW of Laguna de Labradores, G.S. Hinton 27891, 28080(HINT); Municipio de General Zaragoza, Zaragoza, G.S. Hinton 23647, 23636 (ANSM, HINT); 12 km S of Aramberri, Cuesta Blanca near La Joya, P.M. Peterson 15840 & J. Valdés- Reyna (ANSM, US) 15. Muhlenbergia jaime-hintonii P.M. Peterson & Valdés-Reyna, Sida 18(3):686, f. 1. 1999. (Fig. 12, A-L). Tyee: MEXICO. Nuevo León: Municipio General ni Ed. Cuesta Blanca, ca. 15 km S of Aramberri on road towards Zaragoza, 1345 m, 31 Oct 1998, J. Valdés-Reyna & M. A. Carranza Péré E: ANSM!; isotypes: US-3377556!, US-3377557)). Loosely caespitose perennials with short, densely leafy rhizomes and extravaginal shoot initiation. Culms 44-82 cm tall, erect, compressed keeled near the base, densely white pubescence below the basal nodes, these hidden beneath the leafy sheaths, the hairs 0.8-1.3 mm long, upper nodes glabrous or puberulent; internodes puberulent below and mostly glabrous above. Leaf sheaths 6-28 cm long, longer than the lower internode, puberulent to glabrous, stiff and brownish below, often curled, margins mostly smooth with a few short hairs near the summit; ligules 0.4-1 mm long, membranous below, apex truncate, ciliolate; blades 5-22 cm long, 1.8-5 mm wide, flat just above ligule to tightly conduplicate above, apically acuminate, somewhat stiff, pubescent above and glabrous below. Panicles 13-34 cm long, 0.7-7 cm wide, narrow to somewhat open, the ascending densely flowered branches tightly appressed or loosely spreading up to 40° from the rachises; primary branches 0.5-6.5 cm long; pedicels 0.5-2 mm long, ascending, scaberulous. Spikelets 1.5-2.1 mm long, appressed to branches, 1-flowered, reddish-gray; glumes 0.7-1.2 mm long, oblong, Peterson et al., Muhlenl ii f T í Mavi T rire ati MAS In A] TG SS Lm zz. E ae ae = LU mas eee a = SS Fic. 12. Muhlenbergia jaime-hintonii U. Valdés-Reyna 2560 & M.A. Carranza P. (ANSM, US)]. A. Habit. B. Ligule. C. Panicle. D. Secondary panicle branch. E. Spikelet. F. Glumes. G. Lemma. H. Lemma, opened. I. Lemma, lateral view. J. Palea. K. Palea with lodicules, stamens, and pistil. L. Lodicules, stamens, and pistil. Scale for Cis shown in A; Scale for F—L is shown in E. 966 Journal of the Botanical R h Institute of Texas 1(2) shorter than the lemma, usually equal in length, faintly 1-nerved, reddish, glabrous, scaberulous along the midvein, apex acute to obtuse, occasionally minutely erose; lemmas 1.5-2.0 mm long, oblong to elliptic, unawned, faintly 3(1)-nerved, greenish mottled with reddish areas, midvein, margins, and proximal Y to 3⁄4 loosely to densely appressed pubescent to villous, often these hairs more numerous along the margins and midvein below, the hairs up to 0.5 mm long, apex acute, rarely minutely mucronate; paleas 1.5-2.0 mm long, oblong, 2-nerved, equal in length to the lemma, the proximal 2/3 to ?4 densely appressed pubescent to villous between the veins and along the margins, apex acute to obtuse; anthers 0.8-1.0 mm long, reddish at maturity, greenish when immature. Caryopsis not seen. Phenology.—Flowering in October through November. Distribution and habitat. —Muhlenbergia jaime-hintonii is known only from southern Nuevo León between 1300-1850 m in the Municipios Aramberri and General Zaragoza, and can be found growing in whitish, alkaline soils derived from gypsum with Leucophyllum hintoniorum G.L. Nesom, Scutellaria lutilabia T.M. Lane € G.L. Nesom, Galium dempsterae B.L. Turner, Lobelia gypsophila TJ. Ayers, Geniostemon gypsophilum B.L. Turner, Callisia hintoniorum B. L. Turner, di striata Zucc., and Hechtia glomerata Zucc. 1 MÉXICO. N I M de A b i. between La Escondida and Aramberri. 23 Oct 1993,G.S. Hinton et t dl. 23707 (ANSM, HINT, US); San Francisco, G.S. Son 22698 (ANSM, HINT, TEX), G.S. Hinton 22725, 28234 (HINT); Municipio General Zaragoza, Aramberri-El Salitre, 26 Oct 1993, G.S. E 23 ANSM, HINT, TEX); La Joya, Cuesta Blanca, approximately 15 km S of Aramberri on road towards Gral. Zaragoza, 29 Jul 1998, M.A. Carranza P. C-2981 & J. Valdés-Reyna (ANSM); near La Joya, 12 km S of Aramberri, 20 Sep 2001, P.M. Peterson 15841 & J. Valdés-Reyna (ANSM, US). 16. Muhlenbergia lindheimeri Hitchc., J. Wash. Acad. Sci. 24(7):291. 1934. (Fig. 9, D-G). Wee: U.S.A. Texas: 1847, E Lindheimer 725 (HOLOTYPE: US-998949t; isotypes: E GH, MOI, UC, US-998947!, WD. Epicampes gracilis Trin., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 6,4(3-4):271. 1841, nom. illeg. hom. Type: U.S.A. Texas oriental, Dec 1870, s.c. s.n. (HOLOTYPE: LE-TRIN-1556.01)). Strongly caespitose perennials. Culms 50-150 cm tall, stout, erect, not rooting at the lower nodes; internodes mostly glabrous, sometimes puberulent below the nodes. Leaf sheaths 10—45 cm long, shorter or longer than the internodes, glabrous, basal sheaths compressed- keeled, not becoming spirally coiled when old, pur- plish-brown or yellowish; ligules 10-35 mm long, decurrent, firm and brown basally, membranous distally, acuminate; blades 25—55 cm long, 2-5 mm wide, flat or folded, firm, scaberulous abaxially, often involute near apex, scabrous and shortly pubescent adaxially. Panicles 15-50 cm long, (0.6-)1-2(3) cm wide, loosely contracted, often purplish-tinged; primary branches 0.5-7 cm long, appressed or strongly ascending, rarely spreading as much as 20? from the rachises; pedicels 0.5-1.2 mm long, scabrous. Spikelets 2.4-3.5 mm long, light grayish; glumes 23.5 mm long, equal, longer than the florets, scabrous or smooth, 1-nerved, obtuse to acute, occasionally bifid and the teeth to 0.3 mm long, unawned, rarely mucronate, mucros less than 0.2 mm long; lemmas 2.4-3.5 mm long, lanceolate, scabrous or smooth, rarely puberulent near the base, apices obtuse to acute, unawned or awned, the awns 1-4 mm long, straight; paleas 2.4-3.5 mm long, lanceolate, glabrous to puberulent between the nerves on the proximal 14, obtuse; anthers 1.1-1.5 mm long, purplish. Caryopses 1.2-1.6 mm long, fusiform, reddish-brown. 2n = 20, 26, 30 (Reeder & Reeder 4577). Phenology.—Flowering August through December. Distribution and habitat.—Muhlenbergia lindheimeri grows in sandy draws to rocky, calcareous soils, generally in open areas, at elevations of 150-700 m. Muhlenbergia lindheimeri is an uncommon species throughout its range, which includes northern México in addition to southern Texas, but it also is grown as an ornamental. Peterson & Valdés-Reyna 18957 from near San Pablo is an unusual collection of M. lindheimeri since it was collected at 2550 m. All other collections of this species are known from below 700 m. Comments.—Muhlenbergia lindheimeri is similar to M. pubigluma, another species with strongly com- pressed-keeled basal sheaths and decurrent ligules that are firm and brown below. Muhlenbergia lindheimeri can be separated from M. pubigluma by having glabrous glumes that are longer than the florets (verses short pubescent glumes, shorter than the florets in M. pubigluma) and lemmas that are glabrous, rarely puberulent (verses lemmas that are villous on lower !? in M. pubigluma). Peterson et al., Muhlenl ii f T a Mavi a cimens examined. MEXICO. Coahuila: 7 mi SSW of Cuatro Cienegas on road to San Pedro, J.R. Reeder & C.G. Reeder 4577 (US); Municipio de Ramos Arizpe, Sierra de la Paila, p > Cedral camino hacia el Valle de Parreños, J.A. Villarreal Q. 5346, M.A. Carranza P. & A. Rodríguez G. (ANSM); Nuevo León: M Carranza P. 232, J.A. Villarreal Q. & J. Valdés-Reyna. (ANSMD. Cafión La Boca, camino a Cola de Caballo-Lag de Sanchez MA ov» 17. Muhlenbergia macroura (Kunth) Hitchc., N. Amer. Fl. 17(6):468. 1935. (Fig. 13, A-F). Crypsis macroura Kunth, Nov. Gen. Sp. (quarto ed.) 1:140-141. 1816. Phleum macrourum (Kunth) Willd. ex Steud., Nomencl. Bot. (ed. 2) 1:365 1840. Epicampes macroura (Kunth) Benth., J. Linn. Soc., Bot. 19:87. 1881. Crypsinna macroura (Kunth) E. Fourn., Mexic. Pl. 2:90. 1886. (Fig. X). Type: MÉXICO. Mexico: near Nevado de Toluca, Sep, EWH.A. Humboldt & A.].A. Bonpland s.n. (HOLOTYPE: P-BONPL!; ISOTYPE: US-A865654 fragm. ex P-BONPL!). Crypsis setifolia J. Presl, Reliq. Haenk. 1(4-5):245. 1830. Cinna setifolia (J. Presl) Kunth, Révis. Gramin. Suppl. xvi. 1830. Tyee: MÉXICO. T. Haenke s.n. (HOLOTYPE: PR; isoTYPE: US-A865655 fragm. ex P-BONPL!) Caespitose perennials. Culms 75-200 cm tall, erect, rounded near base, forming dense clumps of up to 100 culms or more and up to 1 m in diameter, pubescent below the nodes, internodes mostly glabrous. Leaf sheaths 15-40 cm long, shorter than the internodes, glabrous to scaberulous, the basal persistent and keeled with age; ligules (5—)8-40(-50) mm long, strongly decurrent, spliting into broad auricles, membranous to chartaceous above, brownish, firm, the veins evident below and near margins, apex truncate to obtuse; blades 20-60 cm long, 2-5 mm wide, mostly flat and apically involute, scabrous above and below. Panicles (15-)20-40 cm long, 5-12 mm wide, dense, spikelike, erect, exserted and surpassing the blades in height, greenish to greenish-gray; primary branches 0.1-1.2 cm long, ascending and tightly appressed, unexposed, imbricate; pedicels 0.1-1.7 mm long, shorter than the spikelets, scaberulous. Spikelets 3.4-5.6 mm long, erect, strongly laterally compressed, greenish-gray; glumes 3.4-5.6 mm long, linear-elliptic to linear-ovate, usually longer than the lemma, 1-nerved, scabrous along the keel, subequal, unawned, the upper slightly longer, apex acute to acuminate, scabrous; lemmas 3.4-5 mm long, elliptic to linear-elliptic, scabrous, greenish-gray; callus pilose, the hairs 0.1-0.3 mm long, apex acute, rarely mucronate, the mucro less than 0.4 mm long; paleas 3.4—5 mm long, about as long as the lemma, scabrous, apex acute; anthers 1.5-2.2 mm long, dark greenish. Caryopses 2-3 mm long, fusiform, brownish. 2n = 20, 24, 28. Phenology.—Flowering August through December. Distribution and habitat. —Muhlenbergia macroura occurs in the Sierra Madre Occidental in northern México from Chihuahua to Chiapas, Guatemala, and Costa Rica; found on upland slopes, mountain meadows, in pine or pine-oak forests often in deep humid soils associated with Festuca spp., Piptochaetium fimbriatum, P. pringlei, Bromus spp., Trisetum spicatum, Pinus spp., Alnus sp., and Quercus greggii; 1500-3400 m Comments.—Morphologically, M. nigra Hitchc., a species from central and southern México, is dif- ficult to separate from M. macroura. The former, generally has shorter panicles [6—15(-17) cm long], longer glumes [(5.3-)6-8 mm long], longer lemmas (5-6.5 mm long), and narrower leaf blades (2-3 mm wide). Muhlenbergia nigra also tends to be distributed at higher elevations than M. macroura. Specimens examined. MÉXICO. Coahuila: ascent to Sierra Infernillo, 15 mi SW of Galeana, C.H. & M.T. Mueller 832 (US); Municipio de Arteaga, Sierra Zapalinamé, G.S. Hinton 20506, 20868 (HINT); Municipio de Ramos Arizpe, Sierra de la Paila-Ramoz Arizpe, G. Cano > 17 (ENCB); Municipio de Saltillo, Sierra Zapalinamé, E of Saltillo at “El Penitente”, P.M. Peterson 18797 & J. Valdés-Reyna (ANSM, US). 18. Muhlenbergia microsperma (DC.) Kunth, Révis. Gramin. 1:64. 1829. (Fig. 14, A-D). Trichochloa microsper- ma DC., Cat. Pl. Horti Monsp. 151. 1813. Muhlenbergia microsperma (DC.) Trin., Gram. Unifl. Sesquifl. 193. 1824, nom. inval. MÉXICO. cult. at Montpellier, from seeds collected in México and distributed by the Botanical Garden of Madrid, Sésse & Mociño TYPE: s.n. (HOLOTYPE: MPU; IsoTYPES: P!, US fragm. ex P!). Podosemum setosum Kunth, Nov. Gen. Sp. (quarto ed.) 1:129. 1816. Trichochloa setosa (Kunth) Roem. & Schult., Syst. Veg. 2:386. 1817. Agrostis setosa (Kunth) Spreng., Syst. Veg. 1:262. 1825. Muhlenbergia setosa (Kunth) Trin., Gram. Unifl. Sesquifl. 193, t. 5, f. 22. 1824. ees setosa (Kunth) Kunth, Révis. Gramin. 1:63. 1829, isonym. Type: MÉXICO: between Gueguetoque and Tula, Aug, EWH.A. Humboldt & A.J.A. Bonpland 4174 (HoLotype: P-BONPL!; isotypes: B-W, US-91917 fragm. ex P-BONPL!). iin el M oe Gen. Sp. Pl. 2. 1816. Tyre: MEXICO: plants grown at the Botanical Garden of Madrid from seeds collected by xico 2. Sessé s.n.(HOLOTYPE: MA oc ata Trin., Gram. Unifl. Sesquifl. 192, 296, t. 5, f. 21. 1824. Type: North America, boreali-Amer.(HOLOTYPE: LE?). £L Dat PRA D hi PA 968 Journal of i f Texas 1(2) Se eec. Ñ À q N Y ^ WU Nl UP ez SSS SESS ÚS === === e eai - X DA S AAA ÉS PL ES Fic. 13. Muhlenbergia macroura [PM. Peterson 5970 & C.R. Annable (US)]. A. Habit. B. Panicle. C. Ligule. D. Glumes. E. Floret. F. Lodicules, stamens, and pistil. Muhlenbergia rigens [J.R. Reeder 4589 € C.G. Reeder (ARIZ, US)]. G. Ligule. H. Glumes. I. Floret. J. Lodicules, stamens, and pistil. 969 === E NS LLI microsperma [P.M. Peterson 4185 & C.R. Annable (ARIZ, ENCB, GH, MEXU, MICH, MO, NMC, NY, RSA, TAES, UC, UNLV, US, UTC, WIS, WS)]. A. Habit. B. Ligule. C. Glumes. D. Floret. Muhlenbergia spiciformis [PM. Peterson 8361 & J. Valdés-Reyna (ANSM, US)]. E. Habit. F. Ligule. G. Glumes. Fic. 14. Muhlenbergia H. Floret. I. Lodicules, stamens, a nd pistil. 970 Journal of the Botanical R h Institute of Texas 1(2) moe Nutt., J. Acad. Nat. Sci. Philadelphia, ser. 2, 1:186. 1848. Type: U.S.A. CALIFORNIA: Santa Barbara Co.: Santa Barbara and Santa Catalina Island, Gambel s.n. (HOLOTYPE: K!). Malen ramosissima Vasey, Bull. Torrey Bot. Club 13:231. 1886. Tyre: MÉXICO. Crimuanua: SW Chihuahua, Aug-Nov 1885, E. Palmer 158 (LEC E: NY! designated by Hitchcock, N. Amer. Fl. 27:441. 1935, but without indicating the specific specimen; Peterson & pied ps Bot. Monogr. 31:61. 1991 indicated the specific specimen; IsoLECTOTYPES: LE!, MO-2974152!, P!, US-995580!). Caespitose annuals, sometimes appearing as short-lived perennials. Culms 10-80 cm tall, often geniculate at the base, slender, often striate, much branched near the base, scaberulous below the nodes; internodes 1.8-8.6 mm long, mostly scaberulous or smooth. Leaf sheaths 2.2-6.6 mm long, commonly shorter than the internodes, glabrous, smooth or scaberulous; ligules 1-2 mm long, membranous to hyaline, decurrent, margins often extended, apex truncate to obtuse; blades 3-8.5(-10) cm long, 1-2.5 mm wide, flat or loosely involute, scabrous below, strigulose above, often deciduous with age. Panicles 6.5-13.5 cm long, 1-6.5 cm wide, open and not densely flowered, often purplish; primary branches 1.6-4 cm long, ascending or diverging up to 80° from the rachises, spikelet-bearing to the base; pedicels 2-6 mm long, appressed to divaricate, antrorsely scabrous. Cleistogamous panicles with 1-3 spikelets present in the axils of the lower sheaths. Spikelets 2.5-5.5 mm long; glumes 0.4-1.3 mm long, exceeded by the florets, 1-nerved, obtuse, often minutely erose; lower glumes 0.4-1 mm long; upper glumes 0.6-1.3 mm long; lemmas 2.5-3.8(-5.3) mm long, narrowly lanceolate, mostly smooth, scaberulous sally, hairy on the calluses, lower Y of the margins, and midveins, the hairs 0.2-0.5 mm long, api te, awned, awns 10-30 mm long, straight to flexuous; paleas 2.2-4.8 mm long, narrowly lanceolate, acuminate; anthers 0.3-1.2 mm long, purplish. Caryopses 1.7-2.5 mm long, fusiform, reddish-brown. 2n = 20, 40, 60. Phenology.—Flowering March through December. Distribution and habitat. —Muhlenbergia microsperma grows on sandy slopes, drainages, cliffs, rock outcrops, and disturbed roadsides, at elevations of 0-2400 m. It is usually found in creosote scrub with species of Ambrosia, Encelia, Bebbia, Baccharis and Eriogonum, thorn-scrub forest with Acacia and Clethra, sarcocaulescent desert with Acacia, Fouquieria and Bursera, and oak-pinyon woodland associations. Its range extends from the southwestern United States, México in Baja California, Baja California Sur, Sonora, Sinaloa, Chihuahua, Durango, Aguascalientes, Zacatecas, Jalisco, Nayarit, Morelos, Guanajuato, Querétero, Hidalgo, Tlaxcala, México, Michoacán, Guerrero, Oaxaca, Veracruz, and Chiapas, and through Central America and South America along the Andean countries (Espejo Serna et al. 2000; Peterson & Annable 1991). Comments.—Muhlenbergia microsperma can sometimes be confused with M. tenuifolia and differs from it by having cleistogamous panicles in the axils of the lower sheaths and shorter, obtuse glumes, 0.4-1.3 mm nad acute to acuminate, 2-2.8 mm long in M. tenuifolia). ÉXICO. Coahuila: Municipio de Arteaga, 17 mi SE of Saltillo & 7.4 mi NE of Jamé, P.M. Peterson 10074, C.R. T e & J. Valdés- me (ANSM, US); 6 km E del ent tera 57 hacia los Lirios, J.A. Villarreal Q. 5172, J. Valdés-Reyna, M.E. Barkworth & P.S. Hoge (ANSM); Municipio de Cuatrociónegas Ladera baja de la Sierra Cristo, frente al poblado de Cuatrociénegas, A Rodríguez-Gámez 1225, N. Moreno & J.J. López G. (ANSM); Municipio de Múzquiz, Rancho San Manuel, J.A. Santos-L. s.n. (ANSM); Sierra La Encantada, 170 km NO de Múzquiz, cuesta de ira brecha Müzquiz-Boquillas del Carmen, M.A. Carranza P. 828, J. Valdés- Reyna, P.A. Fryxell, R. Vásquez A. & O. Meza (ANSM); Municipio de Ramos Arizpe, Sierra de la Paila, Ejido el Cedral por el camino El Carmen, J.A. Villarreal Q. 5272, M.A. Carranza P. & A. Rodríguez G. (ANSM); Municipio de Saltillo, 3.5 mi E of Hwy 54 up San Lorenzo Canyon, P.M. Peterson 13259, J. Valdés-Reyna & M.B. Knowles (US); Cañón San Lorenzo en La Sierra Zapalinamé, 8 km S de Saltillo, R. López-Aguil- lón s.n. E US); Cañón San Lorenzo, Cañón en la Sierra de Zapalinamé, 1.5 km E del pozo número 2, R. López-Aguillón s.n. (ANSM). Nuevo León: Municipio de Allende, 9 mi N of Allende on Mex 85, O.L. Briones 1841 (ENCB); Municipi n Ejido 1 8 de Marzo, J.A. Ochoa-Guillemar 977 ruben Municipio de San d La ni e 2 km al S de San José, O.L. Bie 1306 (SLPM); Municipio de Santiago, 3 km adel la Cienega camino a Lag Villa de Santiago, P.A. Garcia-Martinez 1865 (COCA). Tamaulipas: Municipio de Bustamante, Camino a la Joya de San Francisco, A. Brito 121 (COCA); Municipio de Gomez Farias, Rumbo al Rancho La Gloria, M.H. Cervera-Rosado 62 Sake evel de San Carlos, Cerro del Diente, J.A. Barrientos-B. s.n. (COCA); Municipio de Tula, 32 km al SO de Tula, cerca del lí uis Potosí & Tamaulipas), F. González-Medrano 4370, R.M. López F. & R. Dirzo M. (MEXU); La Tapona, J.F. Iribe-Duarte 156 (COCA); dM de Victoria, Carretera Victoria-Tula, J.F. Iribe-Duarte 49 (COCA); km 20 carretera Victoria, San Luis Potosí, G. Villegas-Durán 265 (COCA. 19. Muhlenbergia minutissima Un Contr. U. S. Natl. Herb. 29(4):207. 1947. (Fig. 6). Agrostis minutissima Steud., Syn. Pl. Glumac. 1:171. 1854. S} Steud.) Hitchc., Proc. Biol. Soc. Wash. 41:161. 1928. Type: U.S.A. New Mexico: 1847, A. Fendler 986 (HOLOTYPE: not located; isotypes: MO!, NY-327637!, US-825378!, US-997292!). Peterson et al., Muhlenl ii f T a Mavi T Milium microspermum Lag., Gen. de Pl. 2. 1816, non Muhlenbergia microsperma (DC.) Trin. 1824, Panicum microspermum (Lag.) E. Fourn., Mexic. Pl. 2:492. 1886. S Lag.) Hitchc., J. Wash. Acad. Sci. 23(10):453. 1933. Type: MÉXICO: Habitat in Nova Hispania, D. Sessé s.n. SE. MA; Isotype: US-91019 fragm.!). Vilfa confusa E. Fourn., Mexic. Pl. 2:101. 1886. Sporobolus confusus (E. Fourn.) Vasey, Bull. Torrey Bot. Club 15:293. 1888. Muhlenbergia confusa (E. Fourn.) Swallen, Contr. U.S. Natl. Herb. 29(4):207. 1947. Type: MÉXICO: Jalicingo, CJ. W. Schiede & Deppe 913 (SYNTYPES: US-998282 fragm. ex P!, US-3376139!); Orizaba, Botteri 117 (syuTvrES: P!, US fragm.!); Orizaba, Schaffner 93 (syntyes: P!, US fragm. ex P!); Orizaba, e 125 oe us Nevado de Duce Denm Hahn s.n. (syntype: P); U.S.A., Hall & Harbour 643 (syntypE: P!); Jorullo, in d g i, AJ.A. Bonpl part (SYNTYPE: P?). Delicate annuals. Culms 5—40 cm tall, slender, erect or spreading; scaberulous or strigulose below the nodes; internodes mostly glabrous, scaberulous or smooth. Leaf sheaths 0.4—5.2 cm long, shorter or longer than the internodes, smooth or scaberulous; ligules 1-2.6 mm long, hyaline, margins entire, rarely with lateral lobes or auricles not longer than the body of the ligule, apex truncate to obtuse; blades 0.5-4(-10) cm long, 0.8-2 mm wide, flat or involute, scabrous below, puberulent above, margins and midveins not whitish-thickened. Panicles 5-16.2(-21) cm long, 1.5-6.5 cm wide, open; primary branches 8-42 mm long, often capillary, diverging 25-80" from the rachises; pedicels 2-7 mm long, straight or curved, but rarely curved as much as 90°; glumes 0.5-0.9 mm long, subequal, sparsely strigulose, at least near the apices, the hairs 0.1-0.3 mm long, 1-nerved; lower glumes 0.5-0.8 mm long, obtuse to acute; upper glumes 0.6-0.9 mm long, broader than the lower glumes, obtuse; lemmas 0.8-1.5 mm long, lanceolate, brownish to purplish, glabrous or the midveins and margins appressed-pubescent, apex obtuse to subacute, unawned; paleas 0.8-1.4 mm long, puberulent or glabrous; anthers 0.2-0.7 mm long, purplish. Caryopses 0.6-0.9 mm long, fusiform to elliptic, brownish. 2n = 60, 80 Phenology. MEUS. July through Nox Distribution and habita Muhlenl tissi grows in sandy and gravelly drainages, rocky slopes, flats, road cuts, and open sites. It is usually found in yellow pine and oak-pine forests, pinyon-juniper wood- lands, thorn-scrub forests, and oak-grama savannahs, at elevations of 1200-3000 m; western North America from central Washington to Montana south to Texas, U.S.A., and throughout México to Guatemala. Comments.—The shape and length of the lemmas are variable in this species with some individuals having short (0.8-1.1 mm long), obtuse lemmas and others having longer (1.0-1.5 mm long), subacute lemmas. Muhlenbergia texana Buckley has lemmas similar to the longer-flowered M. minutissima individuals and the lemmas are mucronate or short-awned up to 2 mm long (Peterson & Annable 1991). Currently there are no records of M. texana in NE México. Specimens examined. MÉXICO. Tamaulipas: Municipio de Victoria, km 26 carretera Victoria-San Luis Potosí, G. Villegas-Durán 169 (COCA). 20. Muhlenbergia montana (Nutt.) A. S. Hitchc., U.S.D.A. Bull. (1915-23) d Si e 1920. (Fig. 15, A-C). Calycod Nutt., Proc. Acad. Nat. Sci Phil adelphia 4:23. 1848. Type: U.S.A. ^ Fe Co.: near Santa Fe, W. Gambel s.n. (HoLoTYPE: BM!, isotypes GH, PH). Muhlenbergia gracilis var. enervis Scribn. ex Beal, Grass. N. Amer. 2:242. 1896. Muhlenbergia enervis (Scribn. ex Beal) Hitchc., Contr. U.S. Natl. Herb. 17(3):302. 1913. Type: MÉXICO. Cuinuanua: Sierra Madre, dry ledges, 7 Oct 1887, C.G. Pringle 1413 (HoLotyre: MSC; ISOTYPES: GH, US-995814!, VT, W-1916-27712!). Muhlenbergia trifida Hack., Repert. Spec. Nov. Regni Veg. 8:518. 1910. Tyre: MÉXICO. MicHoacáx: vicinity of Morelia, Quinceo, 11 Nov 9, Bro. Arsene 3217 (HOLOTYPE: W-1916-!; isotypes: BM!, MO-843315!, US!, US-86637 fragm!, W-1916-321451). Densely caespitose perennials. Culms 10-80 cm tall, erect, rounded near base, glabrous below the strictly basal nodes; internodes mostly glabrous, occasionally glaucous. Leaf sheaths 2-35 cm long, longer than the lower internode, glabrous to scaberulous, often glaucous, becoming flattened, loose and papery, and occasionally spirally twisted near base; ligules 4—14(-20) mm long, membranous, decurrent, apex acute to acuminate, often lacerate; blades 6—25 cm long, 1-2.5 mm wide, flat becoming loosely involute to subfili- form, somewhat stiff, scabrous below and hirsute above. Panicles 4-25 cm long, (1-)2-6 cm wide, narrow to somewhat open, loosely-flowered, not dense; primary branches 0.5-10 cm long, ascending, appressed or spreading up to 40? from the rachises; pedicels 0.5-6.5 mm long, flattened, scabrous, occasionally stiffly 972 Journal of the Botanical R h Institute of Texas 1(2) Fic. 15. Muhlenbergia montana [PM. Peterson 8191 & R.M. King (US)]. A. Ligule. B. Floret. C. Glumes. Muhlenbergia setifolia [PM. Peterson 8368 & J Valdés-Reyna (ANSM, US)]. D. Habit. E. Ligule. F. Glumes. G. Floret. H. Lodicules, stamens, and pistil. Peterson et al., Muhlenl ii f T a Mavi 35 reflexed. Spikelets 3—4.5(-7) mm long, erect, occasionally reflexed; glumes (1-)1.5—3.2(-4) mm long, 1/3 to 2/3 as long as the lemma, subequal, glabrous to scaberulous above; lower glumes 1-nerved, sometimes mucronate, the mucro less than 1 mm long; upper glumes 3-nerved, 3-toothed and 3-awned, the teeth (including the awns) 1/3 to Y the length of the glume, and the awns up to 1.7 mm long, apex truncate to acute; lemmas 3—4.5(-7) mm long, lanceolate, awned, often greenish or yellowish with dark green or purple mottles, scaberulous above, loosely to densely appressed pubescent to pilose along the midvein, margins, and proximal Ya to 4/5, the hairs up to 0.8 mm long, occasionally glabrous, apex acute to acuminate, the awn (236-25 mm long, flexuous; paleas 3—4.5(-7) mm long, lanceolate, loosely to densely appressed pubescent to pilose between the nerves on the proximal 1/3 to 4/5, apex acute to acuminate, scaberulous; anthers 1.5-2.3 mm long, purplish. Caryopses 1.8-2 mm long, fusiform, light brown. 2n - 20, 40 Phenology.—Flowering July through December. Distribution and habitat. —Mubhlenbergia montana ranges from southwestern USA throughout western Mexico to Guatemala, primarily in upland and mountain habitats. In NE México this species is only known from Madera del Carmen and Sierra El Jardin. This species grows on rocky slopes, dry meadows, ridge tops, and open grasslands usually associated with Pinus pseudostrobis, Pinus spp., Arbutus xalapensis, Pseu- dotsuga menziesii, Juniperus deppeana, Cupressus sp., Abies sp., Populus tremuloides, Quercus spp., Ceanothus sp., Cornus stolonifera, Holodiscus discolor, Rhus trilobata, Ribes cereum, Cercocarpus breviflorus, and Festuca arizonica;1400—3500 m. Comments.—Muhlenbergia montana is a highly variable species and is sometimes difficult to separate from the southern Rocky Mountain endemic, M. filiculmis, which has shorter spikelets with shorter lemmatal awns (1-5 mm long), tightly involute and filiform, sharp-tipped blades (2-6 mm long), and shorter culms (5-40 cm tall). The morphological distinctions between these two sister species are not great and further study is warranted. Specimens examined. MÉXICO. Coahuila: Madera del Carmen, 3 mi N of El Cinco Junction on road to El Dos, P.M. Peterson 18904 & J. Valdés-Reyna (ANSM, US); wooded canyon above Campo El Dos, P.M. Peterson 18919 & J. Valdés-Reyna (ANSM, US); Municipio de Acuña, Sierra El Jardin, steep slopes near ridge top, 2410m, P.M. Peterson 19934 & S. Lara- Contreras (US, ANSM). 21. Muhlenbergia polycaulis Scribn., Bull. Torrey Bot. Club 38:327. 1911. (Fig. 10, E-I). Tre: MÉXICO. CHIHUAHUA: Sierra Madre Mountains, 30 Sep 1887, C.G. Pringle 1414 (notorvpee: US-81636!; isotypes: US-822943!, US-1551731, US-995733!). Loosely caespitose perennials from a firm, knotty, short rhizomatous base. Culms 15-40(—50) cm tall, erect, decumbent at base, often geniculate, strigulose below the nodes; internodes strigulose to glabrous. Leaf sheaths 1-8 cm long, mostly shorter than the internodes, glabrous to scaberulous, without necrotic spots and not becoming spirally coiled when old; ligules 0.5-2.5 mm long, erose or lacerate, apex obtuse to acute, margins hyaline, firmer than the central portion; blades 3-10 mm long, 0.5-2 mm wide, flat or involute, occasionally folded, hirsute or scaberulous above and scaberulous or smooth below. Panicles 2-12 cm long, (0.6-)1-2 cm wide, narrow, contracted and interrupted below; panicle branches 0.5-4 cm long, ascending and appressed occasionally spreading up to 30? from the rachises, spikelet bearing to the base; pedicels 0.1-1.5 mm long, scabrous. Spikelets 2-3.5 mm long, plump near the middle; glumes (1-)1.5-2.6 mm long, more than Y as long as the lemma, subequal, 1-nerved, mucronate, awned or unawned, scabrous along midvein near apex, apex acute sometimes acuminate, the awn up to 1.4 mm long; lemmas 2-3.5 mm long, elliptic, widest near the middle, appressed-pubescent along the midvein and margins on the proximal 15 to 2/3, the hairs up to 0.5 mm long; apex acuminate, scaberulous, awned, the awn 10-20(-25) mm long, flexuous; paleas 23.5 mm long, elliptic, appressed pubescent between the nerves on the proximal !5, apex acuminate; anthers 1.5-2 mm long, orange. Caryopses 1.5-2 mm long, fusiform, brownish. 2n = 20, 40. Phenology.—Flowering August through October. Distribution and habitat.—Steep rocky slopes, canyon walls, cliffs, table rocks, and volcanic rock out- crops in open vegetation associated with Quercus spp., Pinus spp., P. cembroides, P. chihuahuana, Pseudostuga menziesii, Cupressus sp., Abies sp., Ceanothus sp., Juniperus deppeana, Arctostaphylos sp., A. pungens, Cercocar- 974 Journal of the Botanical R h Institute of Texas 1(2) pus sp., Rhus trilobata, Brickellia sp., Agave sp., Tillandsia sp., Platanus wrightii, Fouquieria splendens, Cornus stolonifera, Polypodium sp., and Selaginella sp.; 1200-2410 m; southeastern Arizona, southern New Mexico, southwestern Texas, U.S.A. to northern México in Baja California, Baja California Sur, Sonora, Sinaloa, Chihuahua, Durango, Zacatecas, Aguascalientes, San Luis Potosí, Guanajuato, Hidalgo, Jalisco, México, Nayarit, and Querétaro (Espejo Serna 2000; Herrera Arrieta 2001; Peterson 2003). Specimens examined. MÉXICO. Coahuila: Sierra del Carmen, Ejido San Francisco, oe rock outcrop W of cabin headquarters, P.M. Peterson 18845, J. Valdés-Reyna & C. Sifuentes (ANSM, US); Madera del Carmen, 10 mi NW of Pilares, P.M. Peterson 18883 & J. Val- dés-Reyna (ANSM, US); Madera del Carmen, Wooded canyon above Campo El Dos, P.M. Peterson 18914 & J. Valdés-Reyna (ANSM, US); Municipio de Acuña, W slope of Sierra El Jardin, E of Rancho El Caballo, M.C. Johnston 9292, F. Chiang C. & T.L. Wendt (ANSM); Sierra El Jardin, steep slop ridgetop, 2410m, P.M. Peterson 19937 & S. Lara-Contreras (US, ANSM); Municipio de Múzquiz, Rancho La Encantada, J.A. Santos L. 250F (TAES); Municipio de Saltillo, Rancho Demostrativo “Los Angeles” 48 km S of Saltillo, S.L. Hatch 4548, C.W. Morden & J. Valdés-Reyna (TAES) 22. Muhlenbergia porteri Scribn. ex Beal, Grass. N. Amer. 2: aS Da (Fig. 16, A-E). in enbergia texana Thurb. ex Porter & J.M. Coult., Syn. Fl. Colorado 144. 1874. nom. illeg., non Muhl Buckley. P. i (Scribn. ex Beal) Bush, Amer. Midl. Naturalist 7(2):36. 1921. Type: U.S.A. Texas: Rio San nue 5 Nov 1850, J.M. Bigelow s.n. [AM GH, US fragm. ex GH!); Presidio del Norte, Jul 1852, C.C. Parry s.n. (syntypes: GH, US fragm. ex GH!); Western Texas to El Paso, May-Oct 1849, C. Wright 734 (svxtvees: GH, MO!, US fragm. ex GH). 1 Loosely caespitose perennials with a wirey and knotty base, rhizomes absent, distinctly bushy in appearance. Culms 25-100 em long, 0.5-1.5 mm think, erect, geniculate and widely spreading near base, freely branched, branching at the lower and middle nodes; scaberulous below the nodes; internodes mostly scaberulous. Leaf sheaths 0.7-4 cm long, shorter than the internodes, glabrous; ligules 1-2.5(-4) mm long, toothed or lacerate, apex truncate, margins hyaline, decurrent, sometimes extended to form short auricles; blades 2-8 cm long, 0.5-2 mm wide, flat or folded, scaberulous above and smooth to scaberulous below. Panicles 4—14 cm long, 6-15 cm wide, open, loosely flowered, usually purple, panicle branches 1-7.5 cm long, widely divergent and stiffly spreading 30—90? from the rachises, not floriferous basally; pedicels 2-13(-20) mm long, scabrous. Spikelets 3-4.5 mm long, often purple; glumes 2-3 mm long, subequal, 1-nerved, occasionally mucronate, scabrous along the nerve, apex acuminate, occasionally mucronate, the mucro up to 0.4 mm long; lemmas 3-4.5 mm long, lanceolate, purplish, appressed-pubescent on the margins and midvein on the proximal Ya to %, apex acuminate, awned, the awn 5-13 mm long, straight; paleas 3-4.5 mm long, lanceolate, glabrous or appressed-pubescent between the nerves on the proximal 4/5, apex acuminate; anthers 1.5-2.3 mm long, purple to yellow. Caryopses 2-2.4 mm long, ellipsoid, compressed, yellowish brown. 2n = 20, 23, 24, 40. Phenology.—Flowering June through October. Distribution and habitat. —Rocky slopes among boulders, dry arroyos, desert flats and grasslands, and cliffs, frequently in protection of shrubs associated with Prosopis sp., Larrea tridentata, Dasylirion longissimum, Agave lecheguilla, Jatropha dioica, Opuntia sp., Acacia spp., Yucca sp., Psilostrophe sp., Mimosa sp., Nolina sp., Bouteloua spp., and Eragrostis spp.; 600—1700 m; southeastern California, southern Nevada, southern Utah, Arizona, southern New Mexico, western Texas, and scattered in western Oklahoma and Colorado to México in Baja California, Sonora, Chihuahua, Durango, Zacatecas, and San Luis Potosí. Comments.—This grass is highly palatable to all classes of livestock but is never abundant at any par- ticular site to provide a significant source of forage. d. MÉXICO. Coahuila: 16.9 km NE of San Miguel on road towards Boquillas, P.M. Peterson 10609 & C.R. Annable (US); 7.2 mi SEW of Jaboncillos on road towards Cuesta de Malena, P.M. Peterson 19846 & S. Lara-Contreras (ANSM, US); Municipio de Cuatrociénegas, Cuatrociénegas, A. Miranda 7-a (COCA); Ladera baja de la Sierra Cristo, frente al poblado de Cuatrociénegas, A. Rodríguez-Gámez 1224, N. Moreno & J. J. López G. (ANSM); Laderas de la Sierra de San Marcos 24 mi SWof Cuatrociénegas, P.M. Peterson 10010, C.R. Annable & J. Valdés-Reyna (ANSM, US); Sierra de San Marcos, áreas cercanas a la Poza de La Becerra, A. Zarate-Lupercio 5 (ANSM); Municipio de General Cepeda, 41 mi W of Saltillo, FW. Gould 11535 (TAES); o de Lampazos de Naranjo, Rancho Las Rusias, S. González s.n.(TAES); Municipio de Múzquiz, El S tr retera 57, 32 S de Sabinas, R. Vásquez-Aldape 218 (ANSM); Municipio de Ocampo, Rancho el Barranquito, 50 km e MODE ES a Sierra Pun M.A. Carranza P. 575 & FJ. Carranza P. (ANSM); Sierrra La Encantada, Rancho Puerto de Aire, R. Vásquez-Aldape 238 (ANSM, CIIDIB); Municipio de Ramos Arizpe, Sierra de la Paila (Lado Norte), Cañada Becerros, J.A. Villarreal Q. 5446, M.A. Carranza P. & L. Arce G. (ANSM); Municipio de Saltillo, 15 Mi W of 975 Peterson et al., Muhlenl R.M. King (ANSM, US)]. A. Habit. B. Ligule. C. Glumes. D. Floret. E. Lodicules, stamens, and pistil. Muhlenbergia villiflora var. villiflora [PM. Peterson 10982, CR. Annable & J. Valdés-Reyna (ANSM, US)]. F. Ligule. G. Glumes. H. Lemma. I. Lodicules, stamens, and pistil. Fic. 16 Mishinnh 976 Journal of the Botanical R h Institute of Texas 1(2) pon D highway 40 to Torreón, S.L. Hatch 5052 eo Valdés-Reyna (ANSM, TAES); Carretera Saltillo-Torreón km 33, J.F. Cano-Siller NSM); Municipio de Torreón, T 6 E. Palmer 511 (TAES); Sierra de Jumilco, 1 idad lejido La Trinidad. J.A E 4416 (CIIDIR): N Si Mojada, 351 1SEd R. Aguirre C. (INEGD. Nuevo León: EA de Peña Nevada & 0.4 mi E of Jtn Bt Hwy 2 to Dr. Arroyo, P.M. Peterson 16793, J. Valdés-Reyna & M. Sosa Morales (ANSM, US). Tamaulipas: ez-Medrano 9029 (MEXU). Municipio de Miquihuana, 4 Km al O del Tanque de Eguia, hacia la Presa de San Carlos, F. Gonzá 23. Muhlenbergia pubigl Swallen, Proc. Biol. Soc. Wash. 56:78. 1943. (Fig. 17, A-D). Tre: MÉXICO. Coanuta: Municipio "E eu. E en Sierra de B Madera, Cañon del Agua, 10 Sep 1939, C.H. Muller 3264 (HoLoTYPE: US-2209360!). Densely caespitose perennials. Culms (5065-125 cm tall, stout, erect, puberulent below the nodes, 2 or 3 nodes per culm; internodes glabrous. Leaf sheaths 10—32 cm long, usually longer than the internodes, scaberulous, basal sheaths compressed-keeled, glabrous, dark-brown with age; ligules 5-13 mm long, membranous above and firm and brown below, decurrent; blades 10-35 cm long, 1.5-3 mm wide, folded to involute, attenuate, firm, scabrous with whitish pubescent, the short hairs about 0.1 mm long, these slightly longer on the upper surface and near the collar. Panicles (15-)20-34 cm long, 1-3 cm wide, narrow, loosely contracted; primary branches 1-6 cm long, mostly li 1 appressed, rarely spreading up to 20? from the rachises, grayish-green; pedicels 0.3-2.5 mm long, usually shorter than the spikelets, scabrous. Spike- lets 2.5-3.5 mm long; glumes 2-3.3 mm long, usually shorter than the floret, 1-nerved, lightly pubescent especially near base, apex obtuse to acute, unawned; lemmas 2.8-3.5 mm long, lanceolate, grayish-green to purplish, the lateral nerves indistinct, appressed villous on lower 1⁄2, basal margins densely villous, the hairs less than about 0.5 mm long, apex acute, usually short-awned, the awns 0.5-3.5 mm long; paleas 2.73 mm long, pubescent between the nerves on the proximal 1/3, apex obtuse; anthers 1.4-1.6 mm long, yellowish. Caryopses not seen. Phenology.—Flowering September through November. Distribution and habitat. —Muhlenbergia pubigluma primarily occurs on calcareous A and flats with Bouteloua, Aristida, Quercus, Rhus virescens, Pinus cembroides, Pinus spp., Pseud lecheguill Abies, Holodiscus, ni uu ene Ru m. Comments. i hologically very similar to Muhlenbergia pubescens (Kunth) Hitchc., a species common in be oe o of western México. Individuals of M. pubigluma have broadly decurrent ligules 5-13 mm long that are firm and brown at the base, whereas ligules of M. pubescens are only 1.5-6 mm long and are not decurrent or firm and brown near the base. In general, the culms, leaves, glumes, lemmas, and paleas of M. pubescens are villous with wavy hairs 0.2-1 mm long. In M. pubigluma, these same structures are pubescent with straight hairs usually less than 0.2 mm long. However, there are four collections of M. pubigluma: Peterson 6251, 10068 & Annable, Peterson 8385, Valdés-Reyna & Villarreal G.; Peterson 13355 & Knowles, that have longer hairs on the glumes, lemmas, and paleas that resemble individu- als of M. pubescens. Specimens examined. MÉXICO. Coahuila: 4 km E of Los Lirios on road to Laguna de Sanchez, 50 km SE of Saltillo, P.M. Peterson 6251 & C.R. Annable (US); 12.9 km E of Los Lirios on road to Lag 60 km SE of Saltillo, P.M. Peterson 6258 .R. Annable (US); Municipio de Arteaga, SE of San Antonio de las Alazanas and SE of Saltillo al d oahuilon, P.M. Peterson 8385, J. Valdés-Reyna & J.A. Villarreal G. (ANSM, US); 41.4 km SE of Saltillo and 4.3 km SE of Jame on i to Sierra La Viga, P.M. Peterson 10068, C.R. Annable & J. Valdés-Reyna(US); 26.7 mi SE of Saltillo & 2.7 mi SE of Jamé, on road towards Sierra La Viga, 2640 m, P.M. Peterson 10058, C.R. An- nable & J. Valdés-Reyna (ANSM, US); 5 mi W of uere on cutoff road between Hwy 54 & 57, 23 mi S of Saltillo, P.M. Peterson 13279 > M.B. Knowles (US); Sierra de Arteaga, El Tunal tal 65, P.S. Hoge 280, M.E. Barkworth & J. Valdés R. (ANSM); Municipio de , at top of S-facing scarp, near Cañón del Desiderio, J. Valdés-Reyna 1074 & T.L. Wendt (ANSM); Municipio de Saltillo, Cañada Salsipuedes, en la Sierra Zapalinamé, 3 km NO de Cuahtémoc, R. López-Aguillón s.n. (ANSM); Sierra Zapalinamé, 1 Ocampo, Crest of the range km S del Canón de San Lorenzo, R. López-Aguillón s.n. (ANSM); Sierra Zapalinamé, ca. 5 km east of Saltillo (Las Palapas) up Camino de Quatro, P.M. Peterson 17864, 17867, J. Valdés-Reyna & R.H. Cardenas (ANSM, US); 18 mi S of Saltillo on México 54 and 1.3 mi W on road to microondas, just W of Estación Carneros, P.M. Peterson 8418 8420, 8421 & M.A. Carranza P. (ANSM, US); 30.6 km SE of Saltillo on hwy 57 to San Luis Potosí, P.M. Peterson 6241 & C.R. Annable (US); 15 mi S of Saltillo on México 54 and 10 mi E to Rancho Experimental Los Angeles, P.M. Peterson 8426, 8427 & M.A. Carranza P. (ANSM, US); 3.5 mi E of Hwy 54 up San Lorenzo Canyon, P.M. Peterson 13262, J. Valdés-Reyna & M.B. Knowles (US). Nuevo Leon: 6.7 mi W S cue de Marzo up road towards Cerro Potosí, P.M. Peterson 13329 & M.B. Knowles (ANSM, US)10.4 mi W of Di 1 1s Cerro Potosi, P.M. Peterson 13355 & M.B. Knowles (ANSM, US); LS 977 ES ÁS Ue ren SSS Te er RES DT AR SS T 3 YE. 5 Semet eg PS —— aa = zc c= — TIe a ee POZA SS a ES AE p LE y m yl Ñ ve) T y and pistil A, US)]. A. Ligule. B. Glumes. C. Floret. D. Lodicules, stame Fic. 17. Muhl IDM 24265 € M A f, Muhlenbergia robusta [PM. Peterson 6131 & CR. Annable (US)]. E. Habit. E Panicle. G. Ligule. D. Glumes. I. Floret. J. Lodicules, stamens, a pistil. 978 Journal of the Botanical R h Institute of Texas 1(2) M de Gal , 12 km E of highway 57 on hwy 58, C.W. Morden 519 (ANSM, TAES); San Pablo to San Pedro Sotolar, G.S. Hinton 27299 (HIND: Municipio de General Zaragoza, 9 km S of Aramberri on road towards xr P.M. Peterson 15838 & J. Valdés-Reyna (ANSM, US); Zaragoza, G.S. Hinton 23654 (ANSM, HINT); above El Barro, G.S. Hint 98 IT); Municipio de General Zaragoza, W of Zaragoza, G.S. Hinton 23654 (HINT); 9 km S of Aramberri on road towards Zaragoza, P.M. Peterson 15838 & J. Valdés-Reyna (ANSM, US); Municipio de Santi Santiago, I. Cabral- Cordero 609 (ANSM). o 24. decias purpusii Mez, Repert. Spec. Nov. Regni Veg. 17(13-18):214. 1921. (Fig. 11, G-L). Tr: CO. San Luis Potosi: Minas de San Rafael, Nov 1911, C.A. Purpus 5011 (HoLotype: B; isotypes: MO-2974180!, US-463679!, US-72636 fragm. ex B!). Caespitose perennials. Culms 25-60 cm tall, erect, pubescent below the nodes, the nodes all basal terete, usually 1 node per culm; internodes scabrous and pubescent. Leaf sheaths 1-20 cm long, shorter than the internodes, pubescent to hispidulous, sheaths rounded, brownish with age below; ligules 4-10 mm long, membranous above and firm and brownish below, decurrent, margins puberulent and wider than Y the adjacent blade width, apex acuminate; blades 5-13 cm long, 0.7-1.8 mm wide, flat or folded, falcate, pu- berulent to hispidulous above and below, the hairs about 0.1 mm long, margins scabrous. Panicles 7-25(-30) cm long, 1-10 cm wide, loosely contracted to open, ovate to pyramidal, purplish; primary branches 2-8.5 cm long, capillary, ascending, appressed or spreading up to 50° from the rachises; pedicels 2224-14 mm long, mostly longer than the spikelets, flexuous, glabrous and smooth. Spikelets 2.4—3.2 mm long, purplish; glumes 0.7-1.1 mm long, less than Y as long as the lemma, obscurely 1-nerved, puberulent on upper 2/3, apex obtuse to acute often erose; lemmas 2.3-3.1 mm long, lanceolate, purplish, short pilose, the hairs 0.1-0.2 mm long, callus short pilose, apex acute, entire to minutely bifid, awned, the awn 10-20 mm long, flexuous; paleas 2.4.2 mm long, slightly longer than the lemma, short pilose between the nerves, apex acute; anthers 1.6-2 mm long, purplish. Caryopses 1-1.2 mm long, fusiform to ovoid, brownish. 2n = 20 Phenology.—Flowering September through November. Distribution and habitat.—Muhlenbergia purpusii occurs on calcareous derived soils primarily on gypsum flats usually on the sides of sink holes, rock outcrops, roadsides, and slopes with Muhlenbergia gypsophila, Pinus cembroides, Leucophyllum hintonii, A lecheguillaa, Juniperus flaccida, Gutierrezia microphylla, Bouteloua chasei, and Yucca filifera; 1240—1900 m. This species has also been reported in Chihuahua, Coahuila, and San Luis Potosí (Espejo Serna et al. 2000; Davila et al. 2006). Specimens examined. MÉXICO. Nuevo León: 7.5 km E of Puentes on dirt road, P.M. Peterson 17847, J. Valdés-Reyna & G.S. Hinton (ANSM, US). Tamaulipas: 101.6 km SW of Ciudad Victoria on MEX 101 towards San Luis Potosí, P.M. Peterson 8325 & R.M. King (ANSM, US); Municipio de Jaumave,19 km al S de Avila y Urbina, F. González-Medrano 14248, P. Hiriart, V. Juárez, R. Molczadzhi & L. Hernández (MEXU). 25. Muhlenbergia quadridentata (Kunth) Trin., Gram. Unifl. Sesquifl. 1a t. 5b, E I. i (Fig. 18, A-D). Podosemum quadridentatum Kunth, Nov. Gen. Sp. (quarto ed.) 1:130-131. 1816. A | Révis. Gramin. 1:64. 1829, isonym. Trichochloa ee (Kunth) Roem. & Schult., Syst. Veg. 2: 388. 1817. mue d virescens subsp. quadridentata (Kunth) Y. Herrera, Amer. J. Bot. 81(8):1043. 1994. Type: MEXICO. Mexico: near Toluca, Sep, EWH.A. Humboldt npland s.n. (LECTOTYPE: P-BONPL!, designated by McVaugh 253. 1983; isoLectotyPe: GH, US-2557456!, US-86634 p ex P!, US-86635!). Podosemum gracile Kunth, Nov. Gen. Sp. (quarto ed.) 1:131-132. 1816. Muhlenbergia gracilis (Kunth) Trin., Gram. Unifl. Sesquifl. 193, t. 5a, f. 6. 1824. Muhlenbergia gracilis (Kunth) Kunth, Révis. Gramin. 1:64. 1829, isonym. Type: MÉXICO. Michoacán: Volcán de Jorullo, Sep, EWH.A. Humboldt & A.J.A. Bonpland s.n. (HoLOTYPE: P-BONPL!; isotypes: LE-TRIN-1501.02!, US-86636 fragm. ex P-BONPL!) Caespitose perennials with short, stout rhizomes. Culms 20-70 cm tall, erect, mostly glabrous below the nodes, the nodes basal, flattened, 1 node per culm; internodes mostly scabrous. Leaf sheaths 10—30 cm long, shorter than the internodes, scabrous to smooth; basal sheaths densely pubescent to glabrous abaxially, smooth and shiny adaxially, becoming flattened and usually not spirally twisted with age; ligules 2-8 mm long, membranous to hyaline above, firm and often brownish with evident veins near the margins below, decurrent, apex acuminate often lacerate; blades 5-15 cm long, 0.6-2 mm wide, flat or usually tightly involute, scaberulous below, short-spiculate and often villous above, the hairs 0.2-0.5 mm long, usually 979 Peterson et al., Muhlent ICH, US)]. A. Habit. B. Ligule. C. Glumes. D. Floret. Muhlent 2472/(M [PM. Peterson 6201 J J Fic. 18. Muhl & CR. Annable (US)]. E. Ligule. F. Portion of panicle branch. G. Glumes. H. Floret. 980 Journal of the Botanical R h Institute of Texas 1(2) appressed, the spicules shiny to whitish. Panicles 5-20 cm long, 0.5-2 cm wide, narrow, loosely-contracted, interrupted below, mostly plumbeous; primary branches 0.5—5(-6) cm long, appressed and ascending to spreading up to 30? from the rachises; central axis flattened with 2 ribs, scabrous; pedicels 0.5-2 mm long, shorter than the spikelets, scabrous. Spikelets 3.4—4.7 mm long, mostly plumbeous; glumes 1.8-4 mm long, shorter to almost as long as the floret, unequal, mostly greenish-plumbeous, scabrous, usually with a few short hairs below; lower glumes 1.8-2.5(23) mm long, 1-nerved, apex obtuse to acute, often with 2 small teeth; upper glumes (3-)3.2-4 mm long, 3-nerved, apex truncate, obtuse or acute, often with 3 or 4 small teeth, the teeth less than 1/6 the length of the glumes; lemmas 3-4.7 mm long, lanceolate, terete, usually awned, greenish-plumbeous to mottled-plumbeous, sparsely pilose near base and margins on lower !5, apex acuminate, scabrous, the awn 0-20 mm long, flexuous, scabrous, greenish-plumbeous; paleas 2.8-4.3 mm long, shorter than the lemma, pilose on the proximal ^; anthers 1-2.5 mm long, purple. Caryopses 1.8-2 mm long, fusiform, brownish. 2n = 20. Phenology.—Flowering July through November. Distribution and habitat. —Muhlenbergia quadridentata occurs on open to forested slopes derived from calcareous and volcanic rocks, and is associated with Pinus spp., P. culminicola, Abies sp., Holodiscus discolor, Populus tremuloides, Pseudostuga menziesii, and Quercus spp.; (1900-)2500-4100 m; throughout México in the higher mountains and found in Guatemala. Comments.—The distinction between M. quadridentata and M. virescens is minimal and it is quite pos- sible that they represent different morphological forms corresponding to their distinctive habitat preferences (McVaugh 1983). Generally the greenish-plumbeous spikeleted forms (M. quadridentata) are found above 2500 m whereas the whitish-hyaline to grayish-green forms (M. virescens) are found between 1600-2700 m. Even this color distinction can break down since intermediate individuals are not uncommon. Reeder (1995) mentioned that in M. quadridentata the ligule is shorter, firmer near the base, and frequently with vascular traces, and the blades are strongly ribbed with tiny spicules on the ribs. After studying the types, including that of M. curvula, we find these same characteristics in many specimens of M. virescens as anno- tated by C.G. Reeder. One character that seems to be fairly consistent within each species is the presence of hairs at the base of the glumes (Reeder 1995). In addition to having dull, scabrous glumes, most individu- als of Muhlenbergia quadridentata have a few short hairs near the base, whereas individuals of M. virescens have whitish or stramineous glumes that are glabrous and shiny near the base. The panicles of M. virescens are often wider (0.5-5 cm) with branches spreading up to 45? from the rachises. Muhlenbergia montana, a common and widespread species known only in NE México from Madera del Carmen and Sierra El Jardin, is morphologically similar to M. quadridentata and M. virescens. However, M. montana differs primarily by having upper glumes that are 3-awned (McVaugh 1983). Even though there has been a recent revision of the Muhlenbergia montana complex (Herrera Arrieta 1998) a more thorough study of this group is needed since SUE limits have not been tested. Specimen amined. MÉXICO. Coahuila: Municipio de Arteaga, Sierra La Viga 6 km al E de Jamé Puerto de Maravillas, J.A. Villar- n VR-1987 (MEXU); 51.6 km SE of Saltillo and 13 km SE of Jamé on road to Sierra La Viga, P.M. Peterson 10056, 10057, C.R. Annable & J. Valdés-Reyna (US); 55.3 km SE of Saltillo and 16.6 km SE of Jamé on road to Sierra La Viga, P.M. Peterson 10062, C.R. Annable & J. Valdés-Reyna (US); Sierra La Viga 6 km al E oo Puerto Maravillas, J.A. Villarreal Q. 1987, J. Valdés-Reyna, P.S. Hoge & M.E. Barkworth (ANSM, MEXU, US); Sierra La Viga, J.A. McD 00 (MEXU, TEX); A. Moreno T. 1192 (INEGI), C.E. Zermeño B. 1155 (INEGD; Cima de Sierra La Marta en la ceja de la ladera sur, J.A. McDonald 1235 (COCA). Nuevo León: 12.7 mi W of Dieciocho de Marzo up road towards Cerro Potosi, P.M. Peterson 13340 & M.B. Knowles (US) 26. Muhlenbergia racemosa (Michx.) Britton, Sterns & Poggenb., Prelim. Cat. 67. 1888. (Fig. 19, A-E). Agrostis racemosa Michx., Fl. Bor.-Amer. 1:53. 1803. Vilfa racemosa (Michx.) P. Beauv., Ess. Agrostogr. 16, 148, 182. 1812. oe racemosus (Michx.) Nutt., Gen. N. Amer. Pl. 1:51. 1818. Cinna racemosa (Michx.) Kunth, Révis. Gramin. 1:67. 1829. T ILuNoIs: in ripis sabulosis inundatis fluminis Mississippi, Michaux s.n. (HOLOTYPE: P!; isotype: US-76287 fragm. & photo ex ae Muhlenbergia gom var. ramosa Vasey, Descr. Cat. Grass. U.S. 40. 1885. Muhlenbergia racemosa var. ramosa (Vasey) Vasey ex Beal, Grass. N Tyre: U.S.A. DAkKoTA-WiscowsiN: 188?, G.R. Vasey s.n. (LECTOTYPE: US-994606!, ps designated, Hitchcock designated a specimen “type from Dakota and Wisconsin" in N. Amer. Fl. 17(6):453. 1935, but did not specify an herbarium). Peterson et al., Muhlenl gii f +) vi Mavi " Sa" s id a ara moa ae cud ey & D. Riskind (ANSM, US)]. A. Habit. B. Ligule. C. Glumes. D. Floret. E. Lodicules, stamens, and pistil. g (US)]. F. Habit. G. Ligule. H. Glumes. I. Floret. J. Lodicules, stamens, and pistil. 982 Journal of the Botanical R h Institute of Texas 1(2) Perennials with slender, elongate, scaly rhizomes. Culms 30-110 cm tall, stiffly erect, in clumps, branched above from the middle, glabrous to puberulent below the nodes; internodes smooth and polished, glabrous. Leaf sheaths 1.5-7.5 cm long, slightly keeled, scaberulous; ligules 0.6-1.5(-1.7) mm long, membranous, lacerate-ciliolate, apex truncate; blades 2-17 cm long, 2-5 mm wide, flat, scabrous, scaberulous, or occa- sionally smooth. Panicles 0.8-16 cm long, 0.3-1.8 cm wide, narrow, densely-flowered, condensed, lobed; primary branches 0.2-2.5 cm long, ascending and appressed; pedicels 0-1 mm long, shorter than the spikelets, erect, strigose. Spikelets 3-8 mm long; glumes 3-8 mm long including the awns, 1.3-2 times longer than the lemma, subequal, awn-pointed, 1-nerved, smooth to scaberulous near apex, apex acumi- nate, the awn up to 5 mm long; lemmas 2.23.8 mm long, lanceolate, unawned or mucronate, short pilose along the midvein and margins on the proximal 14, callus short pilose, apex acuminate, scaberulous, the mucro up to 1 mm long; paleas 2.2—3.8(-4.5) mm long, lanceolate, loosely pilose between the nerves on the proximal Y2, apex acuminate; anthers 0.4-0.8 mm long, yellowish. Caryopses (1.2-)1.4-2.3 mm long, fusiform, brown. n = 20. Phenology.—Flowering August through October. Distribution and habitat. —Muhlenbergia racemosa is found on rocky slopes, irrigation ditches, season- ally wet meadows, margins of cultivated fields, railways and roadsides, prairies, sandstone outcrops, stream banks, and along forest ecotones; 30-3400 m. This species is more common in northcentral United States but ranges from southern Canada sporadically throughout the western U.S.A. to Coahuila, México. Sp xamined. MÉXICO. Coahuila: Municipio de Acuña, Serranías del Burro, Rancho El Bonito, 12 km N of headquarters, J. Valdés-Reyna 1227 & D.H. Riskind (ANSM); Serranías del Burro, Rancho El Bonito, Cañón Los Ojitos, J. Valdés-Reyna 1248 & D.H. Riskind (ANSM) 21. pup ae repens (J. Presl) Hitchc., Fl. Calif. 1:111. 1912. (Fig. 20, A-D). sporobolus M Presl, Reliq. 1(4-5):241. 1830. Vilfa repens (J. Presl) Tin., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci Nat. pode 1840. Tyee: MÉXICO. Haenke s.n. (HOLOTYPE: PR; isotypes: LE-TRIN-1732.01!, MO fragm. & vr US (oun Muhlenbergia subtilis Nees, Linnaea 19(6):689. 1847. Type: MEXICO. Aschenborn 206 (HoLotrrE: B; isotype: US fragm. ex B!). Muhlenbergia abata 1.M. Johnst., J. Arnold Arbor. 24:387—388. 1943. Type: U.S.A. New Mexico: Valley of the Rio edel 1851-1852, C. Wright 1982 (HoLoTvrE: GH; Isotype: US-556896!). Perennial with shining, creeping, scaly rhizomes. Culms 5-42 cm tall, decumbent near base, forming dense mats, freely branching above, glabrous below the nodes; internodes mostly glabrous below, lightly nodulose roughened just below the inflorescence. Leaf sheaths 0.6-3.4 cm long, shorter or longer than the internodes, glabrous, margins hyaline; ligules 0.1-1(-1.8) mm long, membranous, decurrent, apex truncate, occasionally lacerate; blades 0.4-6 cm long, 0.5-1.4 mm wide, involute, somewhat arcuate-spreading, mostly glabrous below and scaberulous above. Panicles 1-9 cm long, 0.1-0.6 cm wide, narrow, contracted, loosely-flow- ered, usually included at the base in the uppermost leaf sheath; primary branches 0.2-3 cm long, usually closely appressed, rarely diverging up to 40? from the rachises; pedicels 0.2-3.6 mm long, minutely setose. Spikelets 2.6-4.2 mm long, occasionally 2-flowered; glumes 1.1-3.6 mm long, Y to as long as the lemma, subequal, usually l-nerved, occasionally 2- or 3-nerved, light green, apex acute; lemmas 2.6—3.2(-4.2) mm long, lanceolate, unawned or mucronate, dark greenish or mottled, glabrous or with minute appressed pubescence along the margin and base, the hairs less than 0.1 mm long, apex acute to attenuate and taper- ing, scaberulous, the mucro 0.1-0.8 mm long; paleas 2.1-3.3 mm long, lanceolate, glabrous to scaberulous, apex acute; anthers 0.7-1.4 mm long, yellow to purplish. Caryopses 1.1-1.5 mm long, ellipsoid to ovoid, brownish. 2n = 60, 70-72. Phenology.—Flowering May through November. Distribution and habitat.—Muhlenbergia repens occurs on sandy meadows, canyon bottoms, calcareous rocky flats, gypsum flats, rolling slopes, and roadsides associated with Pinus spp., P. cembroides, P. johannis, Pseudotsuga menziesii, Cupressus sp., Abies sp., Quercus spp., Juniperus deppeana, Cornus stolonifera, Chryso- thamnus nauseosus, Arctostaphylos spp., Prosopis spp., Larrea tridentate, Cercocarpus ledifolius,, Flaveria spp., Ceanothus spp., Fallugia paradoxa, Dasylirion sp., Agaves spp., Yucca sp. , Sporobolus wrightii, Lycurus phalaroi- Peterson et al., Muhlenl gii f +) vi Mov: m W | V X WW W Y RUM SAW R ror N Z y NS X «à N m | Gr / y Y ES | P VAN NI VEN! NY PAN 14 V NI UN | Wr Fic. 20. Muhlenbergia repens U.R. Reeder & C.G. Reeder 4045 (ARIZ, US)]. A. Habit. B. Ligule. C. Glumes. D. Floret. Muhlenbergia utilis [A.S. Hitchcock 5652 (US)]. E. Habit. F. Rhizome. G. Ligule. H. Glumes. I. Floret. £L Dag PRA D hi PEN 984 Journal of f Texas 1(2) des, Aristida sp., and Stipa robusta; 100-3120 m; southern Utah, southern Colorado, eastern Arizona, New Mexico, western Texas, and throughout México. Specimens examined. MÉXICO. Coahuila: 32.21 SE of Saltill d to Los Lirios, P.M. Peterson 6249 & C.R. Annable (US); Madera del Carmen, above campo El Dos, P.M. Peterson 18906 & J. Valdés-Reyna (ANSM, US); Municipio de Arteaga, N de La Cañada el Carbón, 3 km NO del Ejido Sierra Hermosa, R. López-Aguillón s.n. (ANSM); Columpio, 2 km NO del Ejido Sierra Hermosa, R. López-Aguillón s.n. (ANSM); El Coahuilón, Sierra de la Marta, R. Valdés-Reyna 1836 & M.A. Carranza P. (ANSM); SE of San Antonio de las Alazanas and SE of Saltillo at end of road near summit of Coahuilon, P.M. Peterson 8396, 8406, J. Valdés-Reyna & J.A. Villareal (ANSM, US); Puerto Los Tejocotes en la Sierra de Zapalinamé, R. López-Aguillón s.n. (ANSM); Sierra La Viga 6 km E de Jamé, Puerto Maravillas, J. Valdés-Reyna 1981 J A. Villarreal Q., P.S. Hoge & M.E. Barkworth (ANSM); dnd de Miquihuana, Ejido Valle Hermoso, M.H. Cervera 032 (COCA); Station No. 24, J.A. Santos I sd ES); N Saltillo,12 km N of Zacatecas border on México highway 54 CW. Morden 509 (ANSM, TAES); 20 Saltillo-Torreón, R. Almeida 1638 (ANSM); 28.2 mi S de Saltillo, carretera 54 a Concepción del on Zacatecas, P.M. Peterson 10025, C.R. Annable & J. Valdés-Reyna (ANSM, US); 5 km SO de Buenavista, carretera Saltillo- Concepción del Oro, Zacatecas, P.E. García-Solís s.n. (ANSM); Sierra de Zapalinamé, 5 km S Saltillo in San Lorenzo Canyon, S.L. Hatch 4482, C.W. Morden & J. Valdés-Reyna 4482 (TAES); 21 km SE of Saltillo on Hwy 57 towards Matehuala, P.M. Peterson 15801 & J. Valdés-Reyna (ANSM, US); Buenavista, 7 km S de Saltillo, carretera 54 Saltillo- Concepción del Oro, Zacatecas, H. González-Domíngu s.n. (ANSM), C.W. Morden 505, S.L. Hatch, A. Ruíz-Aznar, V.M. Valdés-Rodríguez & J.S. Marroquín-de la Fuente (ANSM); Escuela Superior Agraria Antonio Narro, Buenavista, H.M. Garza-Cantü s.n. (MEXU); Poblado Carneros, a 40 km por la carretera Federal Número 54, Saltillo- Concepción del Oro, Zacatecas, Saldivar 2 (COCA); Rancho Experimental Los Angeles, 48 Km S of Saltillo, carretera 54 Saltillo- Concepción del Oro, Zacatecas, C.W. Morden 505, 513, J. Valdés-Reyna & J.S. Sierra-Tristán (ANSM, TAES), D.H. Rodrígue n. (ENCB); 24.2 km S of Saltillo on MEX 54, 16.1 km E of Rancho Experimental Los Angeles, P.M. Peterson 8437 & M.A. Carranza P. (ANSM, US). Nuevo León: El Salero, Sierra Madre Oriental, P.M. Peterson 15820 & J. Valdés-Reyna (ANSM, US); 2.5 mi N of La Siberia on road towards La Encantada, P.M. Peterson 16776, J. Valdés-Reyna & M. Sosa Morales (ANSM, US); Ejido Tepehuanes, P.M. Peterson 15874 & J. Valdés-Reyna (ANSM, US); El Salero, P.M. Peterson 15820 & J. Valdés-Reyna (ANSM, US); 10.4 mi W of Dieciocho de Marzo up road towards Cerro Potosí, P.M. Peterson 13352 & M.B. Knowles (US); 13.4 mi E of Hwy 57 on Hwy 58 at crossing of Rio Potosí, P.M. Peterson 13310 & M.B. Knowles (US); Municipio de Aramberri, 70 mi N of Matehuala on bui 61 towards Linares and 1.5 mi N of La Rosita, P.M. Peterson 13364 & M.B. Knowles jn Trinidad — Santa Gertrudis, G.S. Hi 27292 (HINT); M d t te, La Cardona, R. López 042 (COCA); M 32 km N of San Roberto, 18 km S of San Rafael, C.W. Moda 521 (ANSM, TAES); 13 km S of Coahuila border in Nuevo León on highway 57, C.W. Morden 524 (ANSM, TAES); Navidad, carretera 57, km 85 al N de Saltillo, F. ur s.n. & M.A. Bernal (MEXU); Hwy 58,8 Km W of “Y intersection, 23 km E of Hwy 57, C.W. Morden 518 (ANSM, TAES); N p l Treviño, km 55 on Mex 54, E of road, C.W. Morden 507, S.L. Hatch & J. Valdés-Reyna (TAES); Municipio de Santiago, La Escondida, I. Cabral- Cordero 590 nA ida 101.6 km SW of Ciudad Victoria on MEX 101 towards San Luis Potosí, P.M. Peterson 8323 & R.M. King (US); Muni te, Ejido La Cardona, C.R. López-Aguilar 42 (COCA); Municipio de Miquihuana, E Valle Hermoso, J.A. Franco- López 32 (COCA); Municipio de Tula, La Tapona, M.H. Cervera-Rosado 132 (COCA), J.F. Iribe-Duarte 155 (COCA). 28. eric rigens sie : Hitchc., J. Wash. Acad. Sci. 23:453. 1933. (Fig. 13, G-J). Epicampes rigens n. Soc., Bot. 19:88. 1881. Crypsinna rigens (Benth.) M.E. Jones, Contr. W. Bot. 14:8. 1912. Type: U.S.A. CALIFORNIA: dein i 1866, H.N. hou 6124 (HOLOTYPE: Kl; ISOTYPES: US-3239741, US-3239759). Crypsinna setiglumis M.E. Jones, Contr. W. Bot. 14:8. 1912. Tyre: MÉXICO. Cutnuanua: Marsh Lake, Sierra Madre Mountains near Garcia in dry pine woods, Sep 1903, M.E. Jones s.n. (HOLOTYPE: POM; Isotype: US-31685561). Ao o Piper, Proc. Biol. Soc. Wash. 18:143. 1905. Muhlenbergia leptoura (Piper) Hitchc., N. Amer. Fl. 17(6):468. 1935. ICO. CHIHUAHUA: Sierra Madre Mountains near Colonia Garcia, 2134m, 21 Sep 1899, C.H.T. Townsend & C.M. Barber 341 HOLOTYPE: US-347144!; isotypes: MO-1837827!, US-347144!, US-998992!). Muhlenbergia marshii I.M. Johnst., J. Arnold Arbor. 24:392. 1943. Tyre: MÉXICO. Coanuila: Sierra del Carmen, 8 Sep 1936, E. G. Marsh Jr. 746 (HOLOTYPE: GH; isotypes: LL, US-1647094!). Muhlenbergia mundula I.M. Johnst., J. Arnold Arbor. 24:392-393. 1943. Type: MÉXICO. Chinuanua: near Chihuahua, 13 Oct 1885, C.G. Pringle 417 (HoLorYrE: GH; isotypes: MO-1837822!, US-999023!). Densely caespitose perennials. Culms (35-)50-150 cm tall, erect, rounded and up to 5 mm thick near base, mostly glabrous below the nodes; internodes glabrous. Leaf sheaths 3-45 cm long, longer than the internodes, glabrous to scaberulous, rounded to keeled and chartaceous near base; ligules 0.5-2(-3) mm long, somewhat firm, decurrent, apex truncate, usually minutely ciliolate; blades 10-50 cm long, 1.5-6 mm wide, flat or involute, stiff, glabrous below and scabrous between the prominent ridges above. Panicles 15-60 cm long, 0.5-1.2 cm wide, narrow, elongate, contracted and spikelike, densely flowered, often interrupted below, grayish-green; primary branches 0.2-4 cm long, ascending and tightly appressed; pedicels 0.2-3 mm long, mostly shorter than the spikelets, hispidulous. Spikelets 2.4-4 mm long, erect, grayish-green; glumes 1.8-3.2 mm long, subequal, shorter than the lemma, scabrous to scaberulous, 1-nerved, apex acute Peterson et al., Muhlenl ii f T í Mavi T or obtuse, occasionally acuminate or notched, mucronate or short awned, the awn up to 1.7 mm long; lem- mas 2.4-4 mm long, lanceolate, unawned, rarely mucronate, glabrous below and scabrous above, short ap- pressed pubescence on the callus, midvein, and margins on the proximal 1/6, the hairs up to 0.4 mm long, apex acute or obtuse, the mucro up to 0.9 mm long; paleas 2.3-3.8 mm long, lanceolate, glabrous below and scabrous above, apex mostly acute; anthers 1.3-1.8 mm long, yellow to purple. Caryopses 1.8-2.2 mm long, fusiform, brownish. 2n = 40. Phenology.—Flowering August through November. Distribution and habitat.—Sandy washes, gravelly canyon bottoms, rocky drainages, and moist, sandy slopes often along small streams with Pinus spp., P. cembroides, P. chihuahuana, P. reflexa, Juniperus deppeana, J. flaccida, Quercua spp., Q. greggii, Salvia regla, Abies concolor, Arctostaphylos pungens, A. patula, Ceanothus lei dermis, Garrya grisea, Prosopis, Cowania plicata, Berberis sp., Bouteloua curtipendula, Rhus trilobata, Cercocarpus breviflorus, and Mimosa; 90—2500 m; California, Arizona, southwestern New Mexico, southwestern Texas and scattered in Nevada to northcentral México in Baja California, Baja California Sur, Sonora, Chihuahua, Durango, Aguascalientes, Zacatecas, Jalisco, Michoacán, and Puebla (Espejo Serna et al. 2000). Comments.—A highly variable taxon, M. rigens is morphologically similar to M. macroura and M. dubia. Muhlenbergia rigens can be separated from M. macroura by having glumes shorter than the florets (glumes longer than the florets in M. macroura) and shorter ligules 0.5-2(3) mm long [ligules (5—)8-—40(-50) mm long in M. macroura]. Muhlenbergia rigens can be separated from M. dubia by having yellow to purple anthers, 1.3-1.8 mm long (anthers greenish, 1.5-2.2 mm long in M. dubia), shorter lemmas, 2.4-4 mm long (lemmas 3.8-5 mm long in M. dubia), and shorter ligules (ligules 4-10 mm long in M. dubia). Muhlenbergia rigens is available commercially as an ornamental. Speciments examined. MÉXICO. Coahuila: Municipio de Arteaga, Las Vigas, Cañón de la Carbonera, Sierra de Arteaga, J. Valdés-Reyna 1793, J.A. Villarreal Q. & M.A. Carranza P. b Municipio de nu Carneros ] d d Bu O de la estación, 30 km S de Saltillo, J. Espinosa-Aburto 117 (AN 5 km S of Saltillo in San Lorenzo Canyon, S.L. Hatch 4487, C. W. Morden & J. Valdés-Reyna ao TAES); Sierra zm amé, 2 km east of Saltillo up Camino de Quatro (Las Palapas), P.M. Peterson 17855, J. Valdés-Reyna & R.H. Cardenas (ANSM, US); P.M. Peterson 17855 & J. Valdés-Reyna 18806 (ANSM, US). Nuevo León: 13.4 mi E of Hwy 57 on Hwy 58 at crossing of Rio Potosí, P.M. Peterson 13311 & M.B. Knowles (US) 29. Muhlenbergia rigida (Kunth) Kunth, Révis. Gramin. 1:63. 1829. (Fig. 8, D—D). Podosemum rigidum Kunth, Nov. Gen. Sp. (quarto ed.) 1:129. 1816. Trichochloa rigida (Kunth) Roem. & Schult., Syst. Veg. 2:386. 1817. Agrostis rigida (Kunth) Spreng., Syst. Veg. 1:262. 1825. Type: MÉXICO. GUANAJUATO: near EWH.A. Humboldt & A.J.A. Bonpland s.n. (HOLOTYPE: P!; isotype: US-91920 fragm. ex P!). J Podosemum glabratum Kunth, Nov. Gen. Sp. (quarto ed.) 1:130. 1816. Trichochloa glabrata (Kunth) Roem. & 2 ay ME 2:387. 1817. Agrostis glabrata (Kunth) Spreng., Syst. Veg. 1:262. 1825. Type: MÉXICO. Santa Rosa de la Sierra and Ca EWH.A. Humboldt & A.J.A. Bonpland s.n. (HoLoTYPE: P-Bonpl.!; isorvee: US-91921 fragm. ex P-Bonpl.!). Muhl a berlandieri Trin., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 6,4(3-4):299. 1841. TYPE: XICO. Distrito FEDERAL: Mountains near México, Aug 1827, J.L. Berlandier 676, 684 (syntYPE: LE-TRIN-1487.01!, both collec- tion numbers appear on the same sheet, with a single specimen and figure); México, 26 Aug 1827, J.L. Berlandier 676 (ISOSYNTYPES: US-2557457!, US-87241 fragm.!, W-2396040; México, ca. México in montibus, J.L. Berlandier 684 (isosyNTYPE: MO-2974285!, W-1889-239603!). Muhlenbergia affinis Trin., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 6,4(3-4):301. 1841. Podosemum affine (Trin.) Bush, Amer. Midl. Naturalist 7(2):40. 1921. Type: MÉXICO. Mexico: Toluca, J.L. Berlandier 1083 (isotypes: LE-TRIN- 1485.01 fragm.!, US-87237 fragm. !). Muhlenbergia metcalfei M.E. Jones, Contr. W. Bot. 14:12. 1912. Tyre: U.S.A. New Mexico: Grant Co.: Santa Rita Mountains, 1904, O.B. Metcalf 1485 (isotype: US!). ep E Densely caespitose perennials. Culms 40-100 cm tall, stiffly erect, glabrous to scaberulous below the basal, terete nodes; usually 1 node per culm; internodes mostly glabrous. Leaf sheaths 2-30 cm long, longer than the internodes, glabrous to scaberulous, rounded near base; ligules (12)3-12(-15) mm long, often lacerate, firmer below, strongly decurrent; apex obtuse to acute; blades 12-35 cm long, 1-3 mm wide, flat or invo- lute, not falcate, glabrous to scaberulous below and scaberulous to hirsutulous above. Panicles (4-)10-35 cm long, (22)3—5(-15) cm wide, loosely contracted to open and lax, purplish; primary branches 0.4-10 cm long, sometimes capillary, ascending and spreading up to 80? from the rachises; pedicels 1-10 mm long, 986 Journal of the Botanical R h Institute of Texas 1(2) mostly longer than the spikelets. OM 3.5-5 mm long, purplish; glumes 1-1.7(-2) mm long, about equal, 1-nerved, unawned, apex obtuse to te, sometimes hirsutulous, rarely mucronate; lemmas 3.5-5 mm long, narrow lanceolate, scaberulous to scabrous, purple, awned, callus with hairs up to 0.5 mm long, apex acuminate, the awn (5-)10-22 mm long, flexuous; paleas 3.5-5 mm long, narrow lanceolate, purple, scaberulous, apex acuminate; anthers 1.7-2.3 mm long, purplish. Caryopses 2-3.5 mm long, fusiform, brownish. 2n 2 40, 44. Phenology.—Flowering August through November. Distribution and habitat. —Rocky slopes, ravines, and sandy, gravelly slopes derived from granitic and calcareous substrates associated with Pinus spp., P. cembroides, Juniperus deppeana, Juniperus sp., Quercus spp., Arbutus sp., Acacia sp., Arctostaphylos pungens, Brahea sp., Hechtia sp., Cercocarpus breviflorus, Ceanothus sp., Salvia sp., Arbutus sp., Agave sp., Opuntia sp., and Yucca sp.; 1200-2500 m; southeastern Arizona, south- western New Mexico to southwestern Texas, and throughout México and South America along the Andes in Ecuador, Peru, Bolivia, and Argentina. Comments.—This species is highly variable and is one of the most common upland bunchgrasses form- ing almost pure stands in northern México. Muhlenbergia rigida can be distinguished morphologically from M. setifolia by having purplish spikelets (stramineous in M. setifolia), scabrous lemmas (smooth and shiny in M. setifolia), only one node per flowering culm visible (2-4 nodes per culm in M. setifolia), and an evident demarcation between the lemma body and the awn (this not evident in M. setifolia). Another species similar to M. rigida with only a single node per flowering culm is M. purpusii. However, M. rigida has longer lemmas (3.5-5 mm long verses 2.33.1 mm long in M. purpusii) that are scaberulous to scabrous (short pilose in M. purpusii). Specimens examined. MÉXICO. Coahuila: Sierra El Pino, 26.7 km W of Rancho El Cimarron, P.M. Peterson 10648 & C.R. Annable (US), 33.5 km W of Rancho El Cimarron, P.M. Peterson 10661 & C.R. Annable (US), 39.5 km W of Rancho El Cimarron, P.M. Peterson 10673 & C.R. Annable (US); 5 mi W of Chapultepec on cutoff road between Hwy 54 & 57, 23 mi S of Saltillo, P.M. Peterson 13273 & M.B. Knowles n: 30.6 o SE of ue on HWY 57 to San Luis Potosi, P.M. Peterson 6238, 6242 & C.R. Annable (US); 4 km E of Los Lirios on road to Saltillo, P.M. P 252 & C.R. Annable (US); 12.9 km E of Los Lirios on road to I mn SE of Saltillo, P.M. Peterson 6257, 6263 & C.R. Annable (US); Municipio de Acuña, del Carmen Mountains, E.G. Ma TE 719 (MEXU); Madera del Carmen, 12.2 mi NW of Pilares near *old cabin", P.M. Peterson 18884 & J. Valdés-Reyna (ANSM, US); Municipio de Arteaga, AlN de La Cañada el Columpio, 2 km NO del Ejido Sierra Hermosa, R. López-Aguillón s.n. (ANSM); Las Vigas, Cañón de la Carbonera, Sierra de Arteaga, J. Valdés-Reyna 1796, J.A. Villarreal Q. & M.A. Carranza P. (ANSM); Sierra de Arteaga, El Tunal, carretera estatal 65, P.S. Hoge 266, M.E. Barkworth & J. Valdés-Reyna (ANSM); Municipio de Cuatrociénegas, Sierra de la Madera, Cañón Charreteras, Rancho Charreteras, J.A. Villarreal Q. 7344, M.A. Carranza P. & R. Rodríguez L. (ANSM); 25 km E of San Roberto along Hwy 58, S.L. Hatch 4574, 4577, C.W. Morden & J. Valdés-Reyna (TAES); carretera 57, 4.1 km al SW de Pablillo, S.L. Hatch 4999 (TAES); Municipio de Müzquiz, Sierra Maderas del Carmén, E.A. Estrada-Castillón 1784 (ANSM); Sierra del Carmen, Ejido San Francisco, P.M. Peterson 18848, J. Valdés- Reyna & C. Sifuentes (ANSM, US); Madera del Carmen, 12.2 mi NW of Pilares near “old cabin", P.M. Peterson 18890 & J. Valdés-Reyna (ANSM, US); Municipio de Ocampo, Rancho La Rueda, 87 km al NW de Ocampo, D. Barra s.n. (SLPM); Municipio de Ramos Arizpe, 242 Sierra de la Paila, J.S. Marroquín-de la F ANSM); Sierra de la Paila, Ejido el Cedral por el camino El Carmen, J.A. Villarreal Q. 5379, M.A. Carranza P. & A. a (ANSM) Municipio de Saltillo, Buenavista, 7 km S de Saltillo, carretera 54 Saltillo-Concepción del Oro, Zacatecas, J.A. Villarreal Q. 1413 & M.A. Carranza P. (ANSM); 2.4 km al S de Chapula, C.E. Zermeño B. 1078 (INEGI); Camino ] de los Burros, hacia los Aguajes N del P en la Sierra de Zapalinamé, R. López-Aguillón s.n. (ANSM); Cañada Salsipuedes en la Sierra Zapalinamé, 3 km NO de Cuahtémoc, R. López-Aguillón s.n. (ANSM); 2 mi E of Hwy 54 up San Lorenzo Canyon at mouth, P.M. Peterson 13263, J. Valdés-Reyna & M.B. Knowles (US); Cañón de San Lorenzo, P.A. Lobato 21 (COCA); Estación Carneros, carretera a Zacatecas 30 km S de Saltillo, M.A. Carranza P. 519 & P.M. Peterson (ANSM); 18 mi S of Saltillo on México 54 and 1.3 mi W on roa to microondas, just W of Estación Carneros, P.M. Peterson 8422 & M.A. Carranza P. (ANSM, US); Hwy 54 S of Estación Carneros on telephone line service road, M.E. Barkworth 5121, J. Valdés-Reyna, P. S. Hoge & M.A. Carranza P. (ANSM); Rancho Experimental Los Angeles, 48 km S de Saltillo, carretera 54 Saltillo-Concepción del Oro, Zacatecas, J. Espinosa-Aburto 78 (ANSM), S.L. Hatch 4553 & C.W. Morden (ANSM), J. Valdés-Reyna 1903, M.E. Barkworth, P. S. Hoge & M.A. Carranza P. (ANSM), J.S. Sierra-Tristán (ANSM, MEXU, COCA); 18 mi S of Saltillo on México 54 and 1.3 Mi W on road to microondas, just W of Estación Carneros, P.M. Peterson 8423 & M.A. Carranza P. (ANSM, US); 15 mi S of Saltillo on México 54 and 10 mi E to Rancho Experimental Los Angeles, P.M. Peterson 8435, 8436 and M.A. Carranza P. (ANSM); Terrenos de Tranquitas, R.M. F 830 & J. Passini (ANSM); Municipio de Torreón, Sierra de Jimulco, 11.7 km E of Flor de Jimulco at an abandoned mine, P.M. Peterson 8459, P.A. Fryxel l, J. Valdés-Reyna & J.A. Villarreal Q. (ANSM, US). Nuevo León: 5.6 mi E of junction of Hwy 57 on Hwy 58 towards Linares, P.M. Peterson 13290 & M.B. Knowles (US); 13.4 mi E of Hwy 57 on Hwy 58 at crossing of Rio Potosi, P.M. Peterson 13300, 13301 & M.B. Knowles (US); 2 mi W of Dieciocho de Marzo up road towards Cerro Potosi, MA ll L H4 £, 41 + M g i 987 Peterson etal., P.M. Peterson 13323 & M.B. Knowles (US); 36 mi NE of Dr. Arroyo on Hwy 61 towards Linares, P.M. Peterson 13369 & M.B. Knowles (US); 5.2mi S of Zaragoza on road towards Ejido La Encantada, P.M. Peterson 16732, J. Valdés-Reyna & M. Sosa Morales (ANSM, US); 14 mi S of the junction of Hwy 58 and ape eles oe Pablillo, S.L. Hatch. 4999 & J. Valdés-Reyna (ANSM); Sed de Dr. E Cruz de Elorza, G.S. Hinton 20631 (HINT); M ] , La Cuesta, G.S. Hinton 19887 (HINT, MEXU); 2 ] G.S. Hinton 27308 27310 (HINT); SW of Laguna de I RENE G.S. Hinton 27922 (HINT); km 63.6 carretera Galeana-San Roberto, I. Cabral- Cordero 1089 (ANSM); Puerto México, carretera 57 km 67 al S de Saltillo, F. Cárdenas s.n. (MEXU); Muni d l Zaragoza, Sierra El Soldado camino a Puerto Pinos, J.A. Villarreal Q. 4955, M.A. Carranza P., G. Nesom & J. Norris (ANSMD; Municipio de Santiago, El Guardarralla, I. Cabral-Cordero s.n. (ANSM); Las Gomitas, I. Cabral- Cordero 623 (ANSM); carretera Villa de Santiago-Laguna de Sánchez, km 30.7, P Jauregui R. 112 (COCA). Tamaulipas: Municipio de Bustamante, Ejido Felipe Angeles, 2 km al NW rumbo a Bustamente, R. Díaz 187 (MEXU); Municipio de Miquihuana, Colonia La Peña, J.A. Barrientos-B. 122 (COCA); Municipio de Palmillas, 101.6 km SW of Ciudad Victoria on MEX 101 towards San Luis Potosí, P.M. Peterson 8316 & R.M. King (US); Ejido Palmillas, R.A. Carranza 228 (COCA) 30. Muhlenbergia robusta (E. Fourn.) Hitchc., N. Amer. Fl. 17(6):462. 1935. (Fig. 17, E-J). Epicampes robusta Fourn., Mexic. Pl. 2:89. 1886. Type: MÉXICO. Distrito FEDERAL: Santa Fe, 2 Oct 1865, M. Bou rgeau 1153 (LEcTOTYPE: P!, designated by Hitchcock, N. Amer. Fl. 17(6):462. 1935; IsSOLECTOTYPES: K!, US-999036!, US-999031 fragm!, US-90734 fragm). ipid stricta J. Presl, Reliq. Haenk. 1(4—5):235, t. 39. 1830. Muhlenbergia presliana Hitchc., N. Amer. Fl. 17(6):462. 1935, nom. nov. Tre: MÉXICO. T. Haenke s.n. (HOLOTYPE: 2 isotypes: LE-TRIN-1558.01 fragm!, US-865970 fragm). Med PE E. Fourn., Mexic. Pl. 2:89. 1886. Muhlenbergia fournieriana Hitchc., J. Wash. Acad. Sci. 23:453. 1933. Tree: XICO. México: Feb 1839, J.L. Berlandier 670 (Lectotype: P designated by Hitchcock, N. Amer. Fl. 17(6):462. 1935; ISOLECTOTYPE: US-1127013)) Epicampes macrotis Piper, Proc. Biol. Soc. Wash. 18:144. 1905. Muhlenbergia macrotis (Piper) Hitchc., N. Amer. Fl. 17(6):463. 1935. Type: MÉXICO. Zacatecas: Sierra Madre Mountains, ca. 40 km W of San Juan Capistrano, 7 Aug 1897, J.N. Rose 3528 (HOLOTYPE: US-302505!). Epicampes minutiflora Mez, Repert. Spec. Nov. Regni Veg. 17:212. 1921. Muhlenbergia meziana Hitchc., N. Amer. Fl. 17(6):461. 1935, nom. nov. Type: MÉXICO. Michoacán: near El Canizal, 600m, 15 Jan 1899, E. Langlassé 750 (isotype: US-386160!). Caespitose perennials. Culms 100-230(-300) cm tall, erect, compressed-keeled near base, glabrous to sometimes pubescent below the nodes; internodes glabrous. Leaf sheaths 15-70 cm long, longer than the internodes below, glabrous, becoming brownish below, sometimes shredded; sheath auricles present, (1-)2—4(-10) cm long, linear subulate to broadly triangular, longer above, straight or twisted, firm below; ligules 2-10-12) mm long, membranous, lacerate throughout; blades 40-100 cm long, 4-7 mm wide, folded sometimes involute towards tip, scaberulous above and below, the margins and keel saw-toothed. Panicles 30-80 cm long, (2-)3-8 cm wide, narrow to loosely contracted, greenish gray to silvery gray or purplish; primary branches 1-15-17) cm long, naked on the lower 4, ascending and closely appressed to spreading up to 40° from the rachises; pedicels 0.3-1.1 mm long, shorter than the spikelets, erect, scaberulous; central axis prominently ribbed, scabrous. Spikelets (1.8—)2-3(-3.2) mm long, erect, greenish gray or purplish; glumes 1.8-3.2 mm long, usually longer than the floret, subequal, narrowly oblong to elliptic, unnerved to indistinctly 1-nerved, hyaline to greenish-gray, glabrous to scaberulous, apex acute to obtuse occasionally erose; lemmas 1.7-2.6 mm long, linear oblong, unawned or rarely mucronate, greenish to yellowish-brown, glabrous or pubescent with scattered hairs on lower 42, the hairs up to 0.3 mm long, callus glabrous or with a few hairs, apex acute, the mucro when present up to 1 mm long; paleas 1.7-2.6 mm long, glabrous to sparingly pilose between the nerves on the lower 4, apex acute; anthers 1.1-2 mm long, purplish. Caryopses 1.2-1.7 mm long, fusiform, brownish. 2n = 40. Phenology.—Flowering July through April. Distribution and habitat. —Muhlenbergia robusta occurs in mountainous areas from Sinaloa and Chihua- hua south to Chiapas and Central America; found on rocky slopes, along barrancas, pine, pine-oak, and/or pine/fir forests, and tropical deciduous forests with Carya sp., Pinus spp., Arbutus sp., and Quercus spp.; 850-3000 m. Comments.—This species is highly variable. Historically, M. macrotis was separated from M. robusta by having sheath-auricles from 1-10 cm long whereas M. robusta has sheath-auricles 2-5 mm long or lacking. The length of sheath-auricles seems to be under environmental control since we can find no discrete length classes to separate these forms. This character varies widely on individuals within a population and there appears to be no other character that can be used to separate these two forms. 988 Journal of the Botanical R h Institute of Texas 1(2) Ds examined. MÉXICO. Coahuila: ipio de Saltillo, Lomas de Lourdes, Saltillo, P. Lobato 040 (COCA). Nuevo León: 9.2 NE of Dulces Nombres, P.M. Peterson 15928 & J. val dés-Reyna (ANSM, US). 31. Muhlenbergia setifolia Vasey, Bot. Gaz. 7(8-9):92. 1882. Ta 15, D-G). Tee: U.S.A. Texas: western Texas, Guadalupe Mountains, V. Havard s.n. (HOLOTYPE: US-81634!, W-1916-2906 Caespitose perennials. Culms 30-85 cm tall, erect, slender, slightly decumbent near base, hirsutulous below the terete nodes, 2-4 nodes per culm; internodes glabrous to hirsutulous. Leaf sheaths 2-13 cm long, shorter than the internodes, glabrous to puberulent, basal sheaths rounded, margins whitish; ligules 4-7(-10) mm long, membranous, firm below, decurrent, margins glabrous to puberulent and wider than 1⁄2 the adjacent blade width, apex acuminate, lacerate; blades 5-20(-25) cm long, 0.2-1.2 mm wide, tightly involute, fal- cate, scaberulous below and scaberulous to hirsutulous above, the hairs less than 0.1 mm long. Panicles 8-20(-25) cm long, (2-)3-5 cm wide, loosely contracted, ovate; primary branches 0.5-7 cm long, capillary, ascending, appressed or spreading up to 70° from the rachises; pedicels 3-20 mm long, mostly longer than the spikelets, delicate, flexuous, glabrous and smooth. Spikelets 3.5-5.1 mm long; glumes 1.5-2.5 mm long, subequal, thin and hyaline, often purplish near base; lower glumes unnerved, apex truncate or obtuse, often toothed or notched; upper glumes 1-nerved or unnerved, apex obtuse to acute often mucronate, the mucro less than 0.7 mm long; lemmas 3.5-5.1 mm long, narrow lanceolate, glabrous, smooth and shining, stramineous, awned, callus with hairs up to 0.6 mm long, apex acuminate, demarcation of lemma body and awn not evident, the awn 10—30 mm long, flexuous; paleas 3.5—5.1 mm long, as long as the lemma, narrow lanceolate, glabrous, apex acuminate; anthers 2-2.6 mm long, greenish. Caryopses 2.4-3.2 mm long, fusiform, brownish. 2n = 40 Phenology.—Flowering June through November. Distribution and habitat. —Muhlenbergia setifolia grows on calcareous rocky slopes, rock outcrops, and in desert grasslands associated with Quercus spp., Q. saltensis, Juniperus deppeana, J. flaccida, Pinus cembroides, Larrea tridentata, Parthenium argentatum Agave lecheguilla, Fallugia paradoxa, Rhus virescens, Acacia berlandieri, Dasylirion longissimum, Karvinskia humboldtiana, Salvia regla, Cowania plicata, Mimosa sp., Berberis sp., Prunus sp., Bouteloua spp., B. curtipendula, Aristida sp., and Sorghastrum sp.; 1000-2250 m. Muhlenbergia setifolia also ranges to western Texas and New Mexico, U.S.A Comments.—Muhlenbergia setifolia is very similar to M. rigida but differs from the latter by having lem- mas that are smooth, shining, and stramineous; leaf blades that are tightly E. falcate, and narrow (0.2-1.2 mm wide); culms usually with 2-4 nodes; and greenish anthers. Muhl gia setifolia can be easily confused with M. purpusii, another species commonly found on calcareous habitats. However, Muhlenbergia purpusii has smaller spikelets (2.4-3.2 mm long, pilose lemmas and paleas, shorter anthers (1.6-2 mm long), shorter caryopses (1-1.2 mm long), and only a single, basal node per culm. d. MÉXICO. Coahuila: 35.4 km W of Ocampo at Cuesta Zozaya, P.M. Peterson 8368 & J. Valdés-Reyna (US); 140.3 m NW of nsus on Hwy 53 towards Boquilla del Carmen, P.M. Peterson 10599 & C.R. Annable (US); 135.4 km NW of Muzquiz on Hwy 53 Boquilla del Carmen, P.M. Peterson 10587 & C.R. Annable (US); Sierra El Pino, 9.6 km SW of Rancho El Cimarron, P.M. Peterson 10637 & C.R. Annable (US); Sierra El Pino, 39.5 km W of Rancho El Cimarron, P.M. Peterson 10669 & C.R. Annable (US); Sierra La Lagunita; 6.4 mi SE of Aramberri on road towards Agua Fria, P.M. Peterson 16687, J. Valdés-Reyna & M. Sosa Morales (ANSM, US); Municipio de Acuña, Serranías del Burro, Rancho El Bonito, Cañón el Toro, J. Valdés- En 1182 & D.H. Riskind (ANSM); Cañón Los Ojitos, J. Valdés-Reyna 1250, 1260 & D.H. Riskind (ANSM); Municipio de Arteaga, P las Fl R.M. Francois 183 (ANSM); 18 mi E of Saltillo, J.R. Reeder 5000 & C.G. Reeder (ENCB); Municipio de Castaños, Paso de San Lázaro, Si del ia, 37.6 mi S of Monclava on Mex 57 Rn. E P.M. Peterson 9977, 9979, C.R. odd € 2 Valdés-Reyna (ANSM, US); Paso de San Lázaro, Si M Cepeda, 2 peda-Parras, J. Espinosa-Aburto 147 (ANSM); Municipio de TN RENS Santo Domingo, J.A. Santos-L s.n. (ANSM); 26 mi NW of AM Margareta, 90 mi NW of Sabinas, F.W. Gould 10688 (TAES); La Cuesta del Plomo on the Müzquiz-Boquillas hwy, J. Cano 19 (TAES); Sierra La Encantada, 140 km al N de Müzquiz, entronque Boquillas del Carmén brecha & túnel, Flourita de México Unidad Minera, 6 km al SO pasando el túnel, M.A. Carranza P. 710, J. Valdés-Reyna, P.A. Fryxell & R. Vásquez A. (ANSM, MEXU); Municipio de Parras, Rancho El Tunal, 25 km al ESE de Parras de la Fuente, C.P. Cowan 3669 (TAES); Municipio de Ramos Arizpe, Cañada el Diente, Sierra de la Paila, J.A. Villarreal Q. 4811 & M.A. Carranza P. (ANSM); Sierra de la Paila (Lado Norte) Cañada Becerros, J.A. Villarreal Q. 5468, M.A. Carranza P. & L. Arce G. (ANSM); Sierra de la Paila, Ejido el Cedral por el camino El Carmen, J. Valdés-Reyna 2183, J.A. Villarreal Q. 5378, M.A. Carranza P. & A. Rodríguez G. (ANSM); Sierra de la Paila, Valle de Parreños, J. A. Villarreal Q. 4171, J.L. Elizondo E. & J.J. López (ANSM de Saltillo, Cerro del Pueblo, O de la Ciudad de Peterson et al., Muhlenl ii f T í Mavi T Saltillo, J. Valdés-Reyna 2054, M. Martínez M. & M.A. Carranza P. (ANSM, MEXU); San Lorenzo Canyon, 6 mi SE of Saltillo, E. Palmer 400 US); 6 km S of Saltillo, land owned by Universidad Autonoma Agraria “Antonia Narro”, P.M. Peterson 8346 & J. Valdés-Reyna (ANSM, US); Carneros, camino a la torre de microondas, 30 km al S de Saltillo, J. Espinosa 103 (TAES); alrededores de la Escuela de Agricultura “Antonio Narro”, J.G. García G. s.n. (IBUG); Rancho Demostrativo “Los Angeles” 48 km al S de Saltillo, J. S. Sierra PRO4 (ENCB); Estación Carneros, carretera a Zacatecas 30 km S de Saltillo, M.A. Carranza P. 517 & P.M. Peterson (ANSM); Paso de Carneros, al S de Saltillo, J.S. Marroquín-de la Fuente 3056 (ANSM); Ranch tal Los Angeles, 48 km S de Saltillo, carretera 54 Saltillo- Concepción del Oro, Zacatecas, J.S. Sierra-Tristán s.n. & J. Valdés-Reyna , (ANSM); 29 km S of Saltill Mex 54 and 2.1 km W on road to Microdoonas, just W of Estación Carneros, P.M. Peterson 8417 & M.A. Carranza P. (ANSM, US); Municipio de San Buenaventura, Sierra La Encantada, Rancho Carrizalejo, entrada S al rancho Puerto del Aire, M.A. Carranza P. 725, J. Valdés- 3 du PA o R. Mad & O. Meza (ANSM); 9 km al SW del Rancho El Cimarrón, J. Blando Navarrete 8109181 (INEGI). Nuevo León: , Near Puerto México along highway 57, S.L. Hatch. 5009, C.W. Morden & J. Valdés-Reyna (TAES); Municipio Mier & Moers N MN Morena, G.S. Hinton 27335 (HINT); Municipio de San Pedro, 16 km al NE de San Fernando, Sierra La Fragua, H. García G. s.n (INEGD; N Gomitas, I. Cabral- Cordero 617 (ANSM); Sierra Zapalinamé, at 2 km east of Saltillo up Camino de Quatro (Las Palapas), P.M. Pee 17854, J. Valdés-Reyna & R.H. Cardenas (ANSM, US); Sierra Zapalinamé, along camino “El Cuatro”, E of Saltillo, P.M. Peterson 18811, J. Valdés-Reyna & R.H. Cardenas (ANSM, US) Ls 32. Muhlenbergia spiciformis Trin., Mém. Acad. Imp. Sci. Saint-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 6,4(3-4):288. 1841. (Fig. 14, E-I). Tee: MÉXICO. “Southern México,” Karwinsky s.n. (Lecrorvpe: W! here designated, sheet is annotated by C.B. Trinius; isotecrorvre: LE fragm!) Muhlenbergia acutifolia E. Fourn., Mexic. Pl. 2:86. 1886. Type: MÉXICO Veracruz: Orizaba, 8 Nov 1866, M. Bourgeau 3327 (HOLOTYPE: P!; ISOTYPES: MO-2974301!, US-87235 fragm!, US-2561240!). Muhlenbergia parviglumis Vasey, Contr. U.S. Natl. Herb. 3(1):71. 1892. Tyre: U.S.A. Texas: 1887, G.C. Nealley s.n. (HOLOTYPE: US-81638!; ISOTYPE: US-994967!). Caespitose perennials, often short-lived and appearing as annuals. Culms 25-80 cm tall, erect, slender and wiry, freely branching at the base, strigose to glabrous below the nodes; internodes mostly glabrous, usu- ally 4-8 nodes per culm. Leaf sheaths 3.5-12 cm long, shorter than the internodes, scaberulous; ligules 1-3 mm long, deeply lacerate, margins hyaline, apex acuminate; blades 2-12 cm long, 1-3 mm wide, flat to involute, hirsutulous to scabrous above and scaberulous below. Panicles 4-18 (220) cm long, (0.6-)1-2.8 cm wide, narrow, contracted, sometimes interrupted below, loosely flowered; primary branches 0.6-5 cm long, ascending and appressed occasionally spreading up to 30° from the rachises; pedicels 0.1-3.0 mm long. Spikelets 2.8-4 mm long, erect; glumes 0.3-1.0 mm long, less than Y as long as the lemma, 1-nerved, unequal, apex obtuse to acute, sometimes erose; lower glumes shorter than the upper glumes; lemmas 2.8-4 mm long, narrowly lanceolate, awned, purplish, scabrous roughened, sparsely appressed-pubescent on the calluses and lower 1⁄4 of the midveins and margins, the hairs less than 0.3 mm long, apex acuminate, the awn (10-)20-40 mm long, straight to flexuous; paleas 2.6-3.9 mm long, narrowly lanceolate, sparsely pu- bescent between the nerves on the basal 1/3, apex acuminate, scabrous; anthers 0.9-1.6 mm long, purplish. Caryopses 2-2.6 mm long, fusiform, brownish. 2n = 40 Phenology.—Flowering July to November. Distribution and habitat—Muhlenbergia spiciformis grows on rocky slopes, cliffs, and calcareous rock outcrops, often in thorn-scrub and open woodland communities associated with Quercus spp., Pinus spp., P. cembroides, Juniperus deppeana, Pseudotsuga menziesii, Abies sp., Cupressus sp., Agave sp., Ceanothus sp., Acacia sp., Cornus stolonifera, Salvia sp., Juglans sp., Arbutus sp., Opuntia, sp., Fraxinus sp., Bouteloua sp., and Aristida sp.; 450-2800 m. This species ranges from southern New Mexico and southwestern Texas, and is found in México in Chihuahua, San Luis Potosí, Hidalgo, Michoacán, Jalisco, Oaxaca, Querétaro, Veracruz, and nm to NC Mexico. Flowering July to October. im xamined. MÉXICO. Coahuila: Madera del Carmen, 9.5 mi NW of Pilares, P.M. Peterson 18880 & J. Valdés-Reyna (ANSM, US): 27.4 km SE of Saltillo and 12 km NW of Jame at Bosque de Montaña, P.M. Peterson 10074, C.R. Annable & J. Valdés-Reyna (US); Sierra El Pino, 33.5 km W of Rancho El Cimarron, P.M. Peterson 10658 & C.R. Annable (US); 9.7 km E of Los Lirios on road to Laguna de Sanchez, P.M. Peterson 6255 & C.R. Annable (US); Municipio de Acuña, Del Carmen mountains, E.G. Marsh Jr. 717 (MEXU); Municipio de Arteaga, 32.2 km SE of Saltill d to Los Lirios, P.M. Peterson 6244 & C.R. Annable (US); Sierra de Arteaga p Tunal, P.S. Hoge 280 M.E. Barkworth & J. Valdés-Reyna (ANSM); Sierra Madre Oriental. SE of San Antonio de las Alazanas and f road summit of Coahuilón, P.M. Peterson 8402, J.A. Villarreal Q. & J. Valdés-Reyna (US, ANSM); Municipio de Monclova, Rancho Agua Bueno, 43 mi N of Monclova, F.W. Gould 6410 (ANSM, TAES); Sierra de la Gloria, Cañón El Cono, near El Chilpitín, T.L. Wendt 1604 & D.H. 99 Journal of the Botanical R h Institute of Texas 1(2) Riskind (ANSM, UT); Municipio de Bep. 137.9 kn NW Md Ue on Hwy 53 1s Boquilla del Carmen, P.M. Peterson 10590 & C.R. Annable (US); Madera del Car Campo El Dos, P.M. Peterson 18915 & J. Valdés-Reyna (ANSM, US); Sierra Santa Rosa, R. Vásquez-Aldape s.n. pne Sierra del Carmen, Ejido San Francisco, P.M. Peterson 18859, J. Valdés-Reyna & C. Sifuentes (ANSM, US); Municipio de Ocampo, 10 km Saltillo towards General Cepeda, J. Espinosa-Aburto 132 (ANSM); 7.5 km al W del Rancho La Palma, Sierra La Encantada, J. Blando Navarrete 810971 (INEGD; Municipio de Ramos Arizpe, 36 mi S of Castaños, J.R. Reeder 4813 & C. G. Reeder (MEXU); Paso de San Lázaro, al N de Ramos Arizpe, carretera 57, 3 mi S of restaurant La Muralla, P.M. Peterson 8361 & J. Valdés-Reyna (ANSM, US); Municipio de Saltillo, 3.5 mi E of Hwy 54 up San Lorenzo Canyon, P.M. Peterson 13259, J. Valdés-Reyna & M.B. Knowles (US); 5 mi W of Chapultepec on cutoff road between Hwy 54 & 57, 23 mi S of Saltillo, P.M. Peterson 13272 & M.B. Knowles (US); km 25 carretera Saltillo-Monclova, R. Almeida 1593 & M. Torres H. (ANSM); Lomas de Lourdes, M. Saldivar 6 (COCA); N del cañón el Carbón 3 km NW of Ejido Sierra Hermosa, R. López-Aguillón s.n. & T.L. Wendt (ANSM); Ranch perim l Los Angeles, 48 km S of Saltillo, carretera 54 Saltillo- Concepción del Oro S.L. Hatch 4548, C.W. Morden & J. Valdés-Reyna (ANSM, TAES), J.J. Villarrea Q. 6704, J.S. Sierra-Tristán, M.A. Carranza P. & D.E. Lozano (ANSM); 5 km S of Saltillo, in San Lorenzo Canyon, S.L. Hatch, 4496, C.W. Morden & J. Valdés- Reyna (TAES); Sierra Madre Oriental, 15 mi S of Saltillo on México 54 and 10 mi E to Rancho Experimental Los Angeles of the Universidad Autónoma Agraria Antonio Narro, P.M. Peterson 8433 & M.A. Carranza P. (ANSM, US). Nuevo León: 6.1 km S of Allende on MEX 85 towards Montemorelos, P.M. Peterson 8334 & R.M. King (US); 16 km E of Los Lirios and 19.3 km W of Laguna de Sanche P.M. Peterson 6270 & C.R. Annable (US); 8.8 km W of Laguna de Sanchez and 29 km E of Los Lirios, P.M. Peterson 6271 & C.R. Annable (US); Muni | 25 km E of San Roberto along highway 58, S.L. Hatch 4562, C.W. Morden & J. Valdés-Reyna (ANSM, TAES); 5.6miE m of Hwy 57 on Hwy 58 towards Linares, P.M. Peterson 13295 & M.B Knowles un 13.4 mi E of Hwy 57 on Hwy 58 at crossing of Rio Potosí, P.M. Peterson 13309 & M.B Knowles (US); 3 mi W of Di 1s Cero Potosí, P.M. Peterson 13327 & M.B. Knowles (US); Highway-Potrero Prieto, G.B. Hinton 28109 (HINT); carretera eM La Asención, a 1.5 km de San Felipe, M. Castillo 211& N. Bazaldu (COCA); Municipio de García, Villa García, C. Díaz L. s.n. (GUADA); Municipio d l Zaragoza, 4km S of Zaragosa at junction of road to Cerro Viejo-Tepehuanes, P.M. Peterson 15847 & J. Valdés-Reyna (ANSM, US); Sierra La Lagunita; 6.4 mi SE of Aramberri on road towards Agua Fria, P.M. Peterson 16685, J. Valdés-Reyna & M. Sosa-Morales (ANSM, US); 5.2mi S of Zaragoza on road towards Ejido La Encantada, P.M. Peterson 16737, J. Valdés-Reyna & M. Sosa-Morales (ANSM, US); Sierra El Soldado, camino a Puerto Pinos, J.J. Villarreal Q. 4956, M.A. Carranza P., G. Nesom & J. Norris (ANSM, MEXU); Municipio de Iturbide, Bosque-escuela, Universidad Autónoma de Nuevo León, 12 km E de Iturbide, I. Cabral-Cordero 1078 (ANSM, UAT); Municipio de Linares, Las Palmas-El Pinal, J.J. Ortíz-Díaz 23 (ANSM); El Rancho El Nogalar, carretera Linares-San Roberto, km 12, M.M. Castillo-Badillo (COCA); km 169 carretera Monterrey-Linares en El Chocolate, J. Garza 166 & M.M. Castillo-Badillo (COCA); Los Pinos, J.J. Ortíz-Díaz s.n. (ANSM, ENCB); Municipio de Montemorelos, 20 km NW of Montemorelos, J.N. Weaver 591 (TAES); Municipio de Monterrey, Sierra Madre, H.C. Muel- ler 395 & M. T. Mueller (MEXU); 9 km S of El Ca.do on Monterrey-México City hwy, F.W. Gould 6340 & J. Ortega (TAES); Municipio de Rayones, Galeana-Rayones, G.B. Hinton 20835, 20859 (HIND): acne de Santiago, Camino a la Cola de Caballo, P. Jauregui-Ramirez 52 (COCA); La Escondida, I. Cabral-Cordero 597 (ANSM). Tamaulipas: 14.6 mi NE of Dulces Nombres, P.M. Peterson 15919 & J. Valdés- Reyna (ANSM, US); Municipio de Hidalgo, Paso de la Muerte, A. Brito 109 (COCA); Municipio de Jaumave, Ejido Avila y Urbina, J.F. Iribe-Duarte 240 (COCA); Sierra Madre rumbo a Jaumave, A. Brito 40 (COCA); Municipio de Palmillas, 88.7 km SW of Ciudad Victoria n MEX 101 towards San Luis Potosí, P.M. Peterson 8332 6 R.M. King (US); Municipio de San Carlos, La Begonia, 2 km S de San José, Sierra San Carlos, O.L. Briones-Villarreal 1306 (ANSM); Municipio de Victoria, Altas Cumbres, G. Bores-Kulman 88 (COCA). 33. Muhlenbergia stricta (J. Presl) Kunth, Enum. Pl. 1:202. 1833. (Fig. 7, D-H). Podosemum strictum J. Presl, Reliq. Haenk. 1(4-5):230. 1830. Type: MEXICO. T. Haenke s.n. (HOLOTYPE: PR). Muhlenbergia elata Vasey, Contr. U.S. Natl. Herb. 1(8):282. 1893. Tyre: MÉXICO. Jausco: near Guadalajara, Jul-Oct 1886, E. Palmer 770 (HOLOTYPE: US-81624!; isotypes: LE!, MO-2974305!, MO-2974306!, US-822890!). Muhlenbergia longifolia Vasey, Contr. U.S. Natl. Herb. 1(8):283. 1893. Type: MÉXICO. JAusco: Rio Blanco, 1886, E. Palmer 523 (HOLOTYPE: -995747!; isotypes: MO-2947307!, US-822923)). Caespitose perennials. Culms 74-150 cm tall, erect, glabrous to pubescent below the terete nodes; usually 2 or 3 nodes per culm; internodes glabrous to sparingly pubescent. Leaf sheaths 15-35 cm long, glabrous to pubescent, closely overlapping at the base and rounded, the lower usually pubescent; ligules 2-5 mm long, chartaceous, firm and brownish below, strongly decurrent, apex obtuse to truncate; blades 20-65 cm long, 1-3 mm wide, flat or involute, not falcate, glabrous below and scaberulous to hirsutulous above. Panicles 15-60 cm long, 5-12(=15) cm wide, open, loose, diffuse, ovoid to pyramidal, dark greenish-purple; primary branches mostly 2-12 cm long, capillary, ascending and spreading up to 45° from the rachises, whorled; pedicels 1-7 mm long, mostly longer than the spikelets but some of the lateral pedicels shorter than the spikelets. Spikelets (22)2.4—3.6 mm long, dark green; glumes 0.8-2 mm long, shorter than the lemma, subequal, 1-nerved, apex obtuse to subacute or erose, apex often mucronate or short-awned; lower glumes 0.8-1.5 mm long; upper glumes 1-2 mm long, usually mucronate or short-awned, the mucros or awns 0.4-1.2 mm long; lemmas (2—)2.4-3.6 mm long, narrow lanceolate, glabrous, dark green, awned, callus Peterson et al., Muhlenl ii f T í Mavi 9 with hairs up to 0.3 mm long, apex acuminate, sometimes scaberulous, the awns 5-15 mm long, flexuous; paleas (2—)2.4-3.6 mm long, as long as the lemma, narrow lanceolate, glabrous, apex acuminate; anthers 1.1-1.6 mm long, greenish to purplish. Caryopses 1.5-1.7 mm long, fusiform, brownish. 2n = 32 (J.R. Reeder 4721 & C.G. Reeder, US-2541318!), ca. 40 (J.R. Reeder 6389 & C.G. Reeder, US-2769903)). Phenology.—Flowering September through November. Distribution and habitat.—In Tamaulipas, M. stricta was found growing on calcareous derived rocky slopes in thorn-scrub with species of Agave, Yucca, Dasylirion, and Juniperus. Muhlenbergia stricta is endemic to México and has been reported from the states of Colima, Durango, Guanajuato, Jalisco, México, Micho- acán, Morelos, Nayarit, Oaxaca, Veracruz and Zacatecas (Espejo Serna et al. 2000); 1100-2100 m. This is the first report of M. stricta for Tamaulipas. Comments.—Muhlenbergia stricta is morphologically very similar to M. capillaris (Lam.) Trin., a species known from lower elevations (0—500 m) of the southeastern USA, the Caribbean Islands, the Atlantic slopes of Veracruz, and the Yucatán (McVaugh 1983; Peterson 2003). In M. capillaris the pedicels are always longer than the florets (verses some of the lateral pedicels shorter than the spikelets in M. stricta), the spikelets range from (3-)3.5-5 mm long [verses (2—)2.4-3.6 mm], and the lower sheaths are mostly glabrous (verses usually pubescent). Muhlenbergia rigida is also morphologically similar to M. stricta, but the former species has larger spikelets 3.5—5 mm long, unawned glumes, and glabrous basal sheaths. It is somewhat difficult to make correct determinations of some individual specimens especially if you have a glabrous form of M. stricta and the exact collection locality is not known. Specimens examined. MÉXICO. Tamaulipas: Municipio de Palmillas, 63 mi SW of Ciudad Victoria on Mex 101 towards San Luis Potosí, P.M. Peterson 8324 & R.M. King (US). 34. Muhlenbergia tenuifolia (Kunth) Kunth, Révis. Gramin. 1(4):63. 1829. (Fig. 5, E-H). Calamagrostis oe uarto ed.) 1:134. 1816. Arundo tenuifolia (Kunth) Poir., Encycl. 4:704. 1816. Podosemum tenuifoliu Nees ex Nees & ee Linnaea 19(6):690. 1847. Type: MEXICO. Distrito FEDERAL: México City, Apr, EWH.A. Humbol 4 REM Bonpland s.n. (HOLOTYPE: P-BONPL!; isotypes: photo K!, microfiche US!, P!) Calamagrostis quitensis Kunth, Nov. Gen. Sp. ee ed.) 1:133-134. 1816. Arundo quitensis (Kunth) Poir., Encycl. 4:704. 1816, nom illeg. hom., non Arundo quitensis Spreng., 1815. Muhl a quitensis (Kunth) Hitchc., Contr. U.S. Natl. Herb. 17(3):292. 1913. Tre: MEXICO. GUERRERO: Sochial et Valle Zopilote, Apr, EWH.A. Humboldt & A.J.A. Bonpland s.n. (HOLOTYPE: P-BONPL!; ISOTYPES: photo K!, microfiche US!, P Muhlenbergia longiseta Benth., Pl. Hartw. 28. 1840. Type: MÉXICO. AGUASCALIENTES: Aguas Calientes, H g 248 (HOLOTYPE: K!; ISOTYPES: K!, NY!, P!, US-91927 fragm. ex W!). ful la Buckley, Proc. Acad. Nat. Sci. Philadelphia 14:91. 1862. Tyre: U.S.A. Texas: side of hills in the Pass of the Limpia, 24 Miey 1849, C. Wright 731 (Lectotype: PH! designated by Dorr & Peterson, Sida 15:591. 1993; ¡soLecToTYPES: MO!, US-81641!). Muhlenbergia sylvatica var. flexuosa Vasey, Rep. U.S. Geogr. Surv., Wheeler 6:284—285. 1879. Type: U.S.A. Arizona: Camp Crittenden, 1874, Rothrock 681 (LectotyPE: US-995253! designated by Hitchcock, N. Amer. Fl. 17:474. 1935). Loosely caespitose to densely tufted, annuals to short-lived perennials with delicate bases, flowering the first year. Culms 20-70 cm tall, erect or decumbent at the base, branching at the lower and middle nodes, scaberulous below the terete nodes; internodes generally 2.0-9.5 cm long. Leaf sheaths 4.0—7.5 cm long, glabrous or scaberulous, commonly shorter than the internodes; ligules 1.2-3.0(-5.0) mm long, membra- nous, apex acute, often lacerate with age; blades 2-13 cm long, 1.2-2.5 mm wide, flat or loosely involute, scaberulous to glabrous below and scabrous above. Panicles 7-20 cm long, 1.0-6.5 cm wide, narrow and contracted to loosely spreading, interrupted below, terminal and axillary, 15-23 nodes per panicle; primary branches 3.5—7.5 cm long, usually one per node, when immature the branches mostly appressed and ascend- ing, when mature the branches sometimes widely spreading up to 70? from the rachises; pedicels 1-3 mm long, usually shorter than the spikelets, antrorsely scabrous, stout, appressed or spreading. Spikelets 2-4 mm long, erect, often purplish; glumes 1.2-2.8 mm long, unequal, 1-nerved, scabrous along the nerves, apex acute to acuminate, often mucronate or erose, the mucro up to 0.5 mm long; lower glumes 1.2-2.0 mm long; upper glumes 1.5-2.8 mm long, more than Y as long as the lemma; lemmas 2.0—3.5(-4.0) mm long, lanceolate, widest near base, awned, scaberulous above and villous on proximal Y along the margins and 992 Journal of the Botanical R h Institute of Texas 1(2) the midvein, the hairs 0.5-1.5 mm long, callus short-pubescent, the awn 10—30(-40) mm long, scabrous, flexuous; paleas 1.8—3.4(-3.8) mm long, lanceolate, sparsely appressed pubescent between the nerves on the proximal Y; anthers 0.9-1.5 mm long, yellowish. Caryopses 1.0-2.2 mm long, narrowly fusiform, terete, brownish. 2n = 20, 40. Phenology.—Flowering July through November. Distribution and habitat. —Muhlenbergia tenuifolia grows on rocky slopes, limestone rock outcrops, gravelly roadsides, and sandy drainages in grama grasslands with Pinus cembroides, Yucca, sp., Agave sp., Acacia sp., Bouteloua sp., Quercus sp., and Opuntia sp., and in pine-oak woodlands; 1200-2500 m; southern Arizona, New Mexico, and Texas, U.S.A. south throughout México in the Mountains and in Venezuela, Bolivia, Peru, and Argentina (Peterson & Annable 1991; Zuloaga et al. 1994). Comments.—Muhlenbergia tenuifolia can be separated from M. microsperma by having longer acute to acuminate pue s 2-2. i mm one) and by ene cleistogamous panicles present in the axils of the lower sheaths variation in overall plant size, ranging from small first-year flowering Senne to TAM caespitose or densely tufted short-lived perennials. Muhlenbergia spiciformis is morphologically similar but can be separated from M. tenuifolia by possessing short glumes (0.3-1.0 mm long) that are often obtuse at the apex and lemmas with longer awns [(10-2)220-40 mm long]. Specimens examined. MÉXICO. Coahuila: Municipio de Acuña, Del Carmen mountains, E.G. Marsh Jr. 872 (MEXU); Municipio de Arteaga, 10.5 mi SW of Arteaga, 22.5 mi SW of Saltillo, F.W. Gould 8695 (TAES); Las Vigas, Cañón de la Carbonera, Sierra de Arteaga, J. Valdés-Reyna 1800, J.A. Villarreal Q. & M.A. Carranza P. (ANSM, ENCB); Jamé a Rayones, G.S. Hinton 27808 (HINT); Municipio de Castaños, Paso de San Lázaro, Sierra de la Gavia, 37.6 mi S de Monclova, hwy 57, P.M. Peterson 9980, C.R. Annable & J. Valdés-Reyna (ANSM, US); S end of Puerto San Lázaro at Cuesta La Muralla, alon hwy 57, J. Valdés-Reyna 1103 & T.L. Wendt (ANSM); Sierra de La Gavia, Rancho La Gavia, 3 km S de la casa por el cañón, M.A. Carranza P. 2299 & J.A. Encina D. (ANSM); Municipio de Cuatrociénegas, 1? mi N of mouth of cañón de la Hacienda, near turnoff to Cañón del Agua, J. Valdés-Reyna 991 & T.L. Wendt (ANSM, TAES); 1 km NE of Mina La Reforma, Sierra de la Purisima, M.C. Johnston 10326, T.L. Wendt & F. Chiang C. (MEXU, TAES); Cañón del Agua, mouth of canyon, vicinity of Ranchito, J. Valdés-Reyna 1007 & T.L. Wendt (ANSM, TAES); Sierra de la Madera, Canón Charreteras, Rancho Char- reteras, J.A. Villarreal Q. 7345, M.A. Carranza P. & R. Rodríguez L. (ANSM); Sierra de San Marcos, Cañón Grande, Ejido Estanque de Norias, M.A. Carranza P. 1706, J.A. Encina D., J. Fierro G. & R. Rodríguez L. (ANSM); Municipio de Múzquiz, 153.2 km NW of Múzquiz on Hwy 53 towards Boquilla del Carmen, P.M. Peterson & C.R. Annable 10600 (US); Municipio de Nadadores, Sierra de la Purisima, 3 km O de Lamadrid, carretera 30, J.A. Villarreal Q. 3232, J. Valdés-Reyna. & M.A. Carranza P. (ANSM); Municipio de Ocampo, 22 mi W of mpo at Cuesta Zozaya, P.M. Peterson 8369 & J. Valdés-Reyna (MEXU, US); Municipio de Parras, 9 km S of Parras on Sierras Negras, RL 1 165, K. L. Retherford & R.D. Northcraft (MEXU); Sierra de Parras, 2.4 km al S de ej. Colorado, 8.8 km al E de Parras, C.P. Cowan 3605 (TAES); 3 mi N of Parras de la Fuente, E. Palmer 1348 (TAES); Municipio de Ramos Arizpe, Cañada el Diente, Sierra de la Paila, J.A. Villarreal Q. 5186 & M.A. Carranza P. (ANSM); Cañón Loma Prieta, 12 km NE de Hipólito, camino al Valle Loma Prieta, J.A. Villarreal Q. 3084, J. Elizondo, M.A. Carranza P. & A. Rodíguez G. (ANSM, CIIDIR); Puerto de San Lázaro, Sierra de La Gavia, J.A. Villarreal 3178, J. Valdés-Reyna & M.A. Carranza P. (ANSM); Sierra de la Paila (Lado Norte) Cañada Becerros, J.A. Villarreal Q. 5448, M.A. Car- ranza P. & L. Arce G. (ANSM); Sierra de la Paila, áreas ca.nas a las minas La Casa Colorada & el Aguirreño, J.A. Villarreal Q. 4504, 4557 M.A. Carranza P. (ANSM, CIIDIR); M de Saltillo, Saltillo, A.S. Hitchcock 5624 (MEXU); Buenavista, ca. 6 km al S de Saltillo, E. García s.n. Rue J.A. De la pou AES: 2 a saliendo del libramiento por la brecha Saltillo-General Cepeda, J. Espinosa 120 (IBUG), J. Espinosa 122 (SLPM); al Escuela de Agricultura “Antonio Narro”, J.G. García G. (IBUG); San Lorenzo canyon, 6 mi SE of Saltillo, M. Saldívar 557 (COCA); P. Lobato 11 (COCA); 5 km S of Saltillo, San Lorenzo Canyon, P.M. Peterson & C.R. Annable 10547 (US); 3.2kmE d dps on did 57 ] huala, San Luis Potosí, P.M. Peterson 10079, C.R. Annable & J. Valdés R. (ANSM, US); 4 km W ltillo-General Cepeda, a la orilla del camino, J. Valdés-Reyna 1554, L.E. Rodríguez G. & R. Vásquez A. (ANSM); W of Saltillo on r 40 and 1.5 mi W of Atenco, J. Valdés-Reyna 984 & T.L. Wendt (ANSM, TAES); Buenavista, 7 km S de Saltillo, carretera 54 Saltillo-Concepción del Oro, Zacatecas, E. García-Aguilera s.n. & M.A. González-Galindo (ANSM); Cerro del Pueblo, W o Ciudad Saltillo, J. Valdés-Reyna 2047, M. Martínez M. & M.A. Carranza P. (ANSM); Entrada camino Fosa el Mamut, 2 km N del Ejido La Encantada, Sierra de Arteaga, J. Valdés-Reyna 1517a, L.E. Rodríguez G. (ANSM); Estación Carneros, camino Torre de Microondas, 3 km O de la estación, 30 km S de Saltillo, J. Valdés-Reyna 1501 & L.E. Rodríguez G. (ANSM), P.S. Hoge 297, M.E. Barkworth & J. Valdés-Reyna. (ANSM); 18 mi S of Saltillo on México 54 and 1.4 mi W on road to microondas, just W of Estación Carneros, P.M. Peterson 8411 & M.A Carranza P. (ANSM, MEXU, US); Municipio de Sierra Mojada, vicinity of Santa Elena Mines, R. M. Stewart 284 (MEXU); Municipio de Torreón, S of Torreón; canyon between Jimulco and Juan Eugenio, P.M. Peterson 8471 & J. Valdés-Reyna (ANSM, MEXU, US); Sierra de Jimulco, 150 km E de La Mina de San José, J.A. Villarreal Q. 5522, P.A. Fryxell, J. Valdés-Reyna & P.M. Peterson (ANSM). Nuevo León: Carretera a Trinidad China Met odd slag 1225 (COCA); 16 km NE of Sandia on road to La Ascensión, P.M. Peterson 15832 & J. Valdés-Reyna (ANSM, US); Muni etera Linares-San Roberto a medio km de la Galeana, M. Castillo 200 & N. Bazaldu (COCA); bank of stream, V.H. Chase 7739 & Mexican iu Expedition of students of the University of Illinois (ANSM, US); Hacienda Peterson et al., Muhlenl ii f T vd Mavi 25 Pablillo, M. Taylor-Edwards (MEXU); km 137 de la carretera 57, tramo Matehuala-Saltillo, al E del Cerro El Potosí, J. García-P 500 (ANSM); 13.4 mi E hwy 57 on hwy 58 at crossing Río Potosí, P.M. Peterson 13302 & M.B. Knowles (US); 10 mi S of San José de Raíces, R. McVaugh 18239 (ENCB); La Becerra, G.S. Hinton 19793, 19888 (HINT, MEXU); Municipio de Iturbide, Bosque-escuela, Universidad Autónoma de Nuevo León, 12 km E de Iturbide, I. Cabral- Cordero 1081 (ANSM); Municipio de Monterrey, carretera Monterrey- Saltillo, N. Bazaldua B. 115 (COCA); Municipio de San Pedro Garza García, carretera Villa de García a Grutas (W of Monterrey), T. Tateoka, 1125 (US); Mu- nicipio de Santa Catarina, Ladera E del Cerro del Potosí, J. García 500 (TAES); Municipio de Zaragoza, La Joya, Cuesta Blanca, 15 km de Aramberri rumbo a Zaragoza, J. Valdés-Reyna 2564, J.A. Villarreal M.A. Carranza P. (ENCB). Tamaulipas: Municipio de Palmillas, 55 mi SW of Ciudad Victoria on México 101 towards San Luis Potosí, P.M. Peterson 8330 & R.M. King (MEXU, US); Municipio de Palmillas, Ejido Palmillas, M.H. Cervera-Rosado 316 (COCA); Municipio de Victoria, 1 km S del Rancho El Novillo, 20 km O de Ciudad Victoria, ez-Medrano 3200 & E. Martínez (MEXU). F. Gonzá 35. Muhlenbergia torreyi (Kunth) Hitchc. ex Bush, Amer. Midl. Naturalist 6:84. 1919. (Fig. 4, F-D). Agrostis torreyi Kunth, Enum. Pl. 1:226. 1833. Agrostis caespitosa Torr., Ann. Lyceum Nat. Hist. New York 1(1):152-153. 1824, nom. illeg hom., non Agrostis caespitosa (L.) Salisb. Prodr. Stirp. Chap. Allerton 25. 1796. Tyre: U.S.A. Prairies of the Missouri and Plate Rivers, E. James s.n. (HOLOTYPE: NY-327621! Muhlenbergia gracillima Torr., Pacif. Railr. Rep. 4:155. 1857. Podosemum gracillimum (Torr.) Bush, Amer. Midl. Naturalist 7:33. 1921. Type: A. Texas: Llano Estacado, 1853-1854, J.M. Bigelow s.n. (svxrvee: NY; isosyNtyPE: US fragm. ex NY); Antelope Hills, 1853-1854, J.M. Bigelow s.n. (svxrveE: NY; ISOSYNTYPE: US fragm. ex NY!). Caespitose perennials. Culms 10—40(-50) cm tall, decumbent at base, hispidulous below the nodes, usually no culm nodes exposed above the tightly clustered leafy base, the leafy portion not reaching more than 1/5 the plant height; internodes mostly scabrous to glabrous. Leaf sheaths 0.1-2.5(-3.2) cm long, shorter than the internodes, scaberulous to glabrous, margins hyaline; ligules 2-5(-7) mm long, hyaline, often splitting down the middle and appearing as auricles, apex acuminate, lacerate, margins entire; blades 1-3(-5) cm, long, 0.3-0.9 mm wide, tightly involute or folded, arcuate, somewhat sharp pointed, scaberulous. Panicles 7—21 cm long, 3-15 cm wide, open, diffuse; primary branches 1-8 cm long, ascending to stiffly spreading at maturity 30-90” from the rachises; pedicels 1-8 mm long, erect, sometimes appressed to the branches. Spikelets 2-3.5 mm long, erect; glumes 1.3-2.5 mm long, about equal in length, 1-nerved, unawned, mu- cronate, or awned, scaberulous towards apex, apex acute to acuminate, minutely erose, the mucro or awn up to 1.1 mm long; lemmas 2-3.2(3.5) mm long, narrow elliptic to lanceolate, mucronate or awned, ap- pressed pubescent on the margins and midvein on the proximal Y2 to ?4, scabrous above, apex acuminate, the mucro or awn 0.5-4 mm long; paleas 2-3.2(-3.5) mm long, narrow elliptic, sparsely pubescent between the nerves, apex acuminate sometimes with two mucros, the mucros up to 0.2 mm long; anthers 1.2-2.1 mm long, greenish. Caryopses 1.7-2.0 mm long, fusiform, brownish. 2n = 20, 21. Phenology.—Flowering May through October. Distribution and habitat. —Muhlenbergia torreyi grows in desert grasslands, sandy mesas, calcareous rock outcrops, rocky slopes, and open woodlands with Prosopis spp., Yucca sp., Larrea tridentata, Lycium spp., Parthenium argentatum, Agave lecheguilla, Artemisia tridentata, Psilostrophe sp., Ceratoides lanata, Bouteloua gracilis, Juniperus osteosperma, and Pinus edulis; 1000—2450 m; southeastern Arizona to eastern Colorado, and extreme southeastern Wyoming, western Kansas to western Texas, New Mexico, U.S.A., northern México in Sonora and Chihuahua, and disjunct in northwestern Argentina and Bolivia. Specimens examined. MÉXICO. Coahuila: 35.4 km W of Ocampo at Cuesta Zozaya, P.M. P 8364 & J. Valdés-Reyna (US); Municipio de Arteaga, 18 mi E of Saltillo, J.R. Reeder 5002 & C.G. Reeder ee Municipio de Saltillo, Rancho Demostrativo “Los Angeles” 48 km al S de Saltillo, L.H. Harvey 8738 (ENCB). Nuevo León: Municipi Galeana, Rancho Aquililla, G.B. Hinton 27026, 27073 (HINT). 36. Muhlenbergia utilis (Torr.) Hitchc., J. Wash. Acad. Sci. 23(10):453. 1933. (Fig. 20, E-I). Vilfa utilis Torr., Pacif. Railr. Rep. 5(2):365-366. 1857. Sporobolus utilis (Torr.) Scribn., Bull. Div. Agrostol., U.S.D.A. 17:171, f. 467. 1899. Type: U.S.A. CALIFORNIA: Lost Mountain Spring, from Tejon to the Lost Hills, in stony places, WP Blake s.n. (HoLoTYPE: NY; isotypes: GH, MO!, US fragm. ex NY!). Perennials with slender, scaly rhizomes. Culms 7-30 cm tall, erect to decumbent, older plants trailing, up to 1 m long, minutely pubescent to glabrous below the nodes; internodes mostly smooth to lightly nodulose- roughened. Leaf sheaths 0.3-2.4 cm long, shorter or longer than the internodes, glabrous, margins hyaline; ligules 0.2-0.8 mm long, membranous, decurrent, apex truncate; blades 0.5-4.7 cm long, 0.2-1.8 mm wide, 994 Journal of the Botanical R h Institute of Texas 1(2) involute, sometimes flat, straight or arcuate-spreading, blades often at right angles to culm, mostly glabrous below and hirsutulous above. Panicles 1-5 cm long, 0.1-0.4 cm wide, narrow, contracted, interrupted between each branch, partially included in the upper sheaths; primary branches 0.2-1.2 cm long, appressed, rarely ascending up to 30? from the rachises; rachises usually visible between the branches; pedicels 0.1-1.1 mm long, glabrous. Spikelets 1.4-2.4 mm long, erect; glumes 0.5-1.4 mm long, 1/3 to Y as long as the lemma, subequal, unawned, glabrous, usually 1-nerved, occasionally 2- or 3-nerved, yellowish to light green, apex acute; lemmas 1.3-2.4 mm long, lanceolate, unawned, glabrous or with minute appressed pubescence along the margins and base, the hairs about 0.1 mm long, green or purplish, apex acute; paleas 1-2 mm long, lanceolate, glabrous, apex acute; anthers 0.7-1.4 mm long, yellow to purplish. Caryopses 0.7-1.2 mm long, ellipsoid to ovoid, brown. 2n = 20 Phenology.—Flowering July to December. Distribution and habitat.—Muhlenbergia utilis occurs in wet soils along streams, ponds, depressions in grasslands, and alkaline or gypsiferous plains associated with Quercus spp., Q. agrifolia, Acacia sp., Prosopis sp., Larrea tridentata, Pinus spp., and Cupressus sp.; 200-2500 m; southwestern California, southern Nevada, southern Arizona, southcentral Texas, U.S.A. to México in Estados Chihuahua, Sonora, Durango, Guanajuato, Hidalgo, Jalisco, Zacatecas, México, Distrito Federal, Michoacán, Querétaro, Puebla, Veracrúz, and Chiapis to Costa Rica. Specimens examined. MÉXICO. Coahuila: Municipio de Arteaga, Rancho el Chorro, carretera al Tunal, 3 km E de la desviación al Tunal, sobre la carretera de Saltillo-Matehuala, San Luis Potosí, J. Espinosa-Aburto 170. (ANSM, SLPM); Ojo Caliente ca. 50 km Monterrey, F.A. Barkley 16281 (US); Municipio de Parras, 3 mi N of Parras de la Fuente, F.W. Gould 11554 (TAES); Municipio de Saltillo, Saltillo, A.S. ur 1378 (TAES), A.S. Hitchcock 5652 (US); 2 mi S of San Lorenzo, J.R. Reeder 4830 & C.G. Reeder (ENCB, MEXU); 3 mi N of Saltillo on Piedras Negras Hwy, F.W. Gould 11212 (TAES). Nuevo León: Municipio de Galeana, Galeana, J.A. Ochoa-Guillemar 1003 (COCA); un 57, 32 km N of San Roberto, 18 km S of San Rafael, C.W. Morden 522, 523, S.L. Hatch & J. Valdés-Reyna (ANSM, TAES); 4 mi SE of Galeana, J.R. Reeder 4986 & C.G. Reeder (US); 3 mi E of jet of hwy 60 & 57, C.W. Morden 519, S.L. Hatch & J. Valdés- Reyna (ANSM, SLPM); km 63.6 carretera Galeana—San Roberto, I. Cabral- Cordero 1085 (ANSM, ENCB); Municipio de Linares, Baño de San Ignacio, 22 km NE de Linares, I. Cabral-Cordero 771 (ANSM); 12.9 km E of Los Lirios on road to Laguna de Sánchez, 60 km SE of Saltillo, P.M. Peterson 6259 & C.R. Annable (US). 37. Muhlenbergia villiflora Hitchc. var. villiflora, N. Amer. Fl. 17(6):470. 1935. (Fig. 16, F-D. vilfa pubes- cens E. Fourn., Mexic. Pl. 2:102. 1886, non Muhlenbergia pubescens (Kunth) Hitchc. Tyee: MEXICO. TamauLiras: inter Michiguana et Tanquecillos, Cañon de las Minas et Victoria, W. Karwinsky 1012 (HoLotype: P!; isotype: US fragm. ex P!). Perennials with scaly rhizomes. Culms 4-20(227) cm tall, to 2 mm thick, erect, wiry, smooth to nodulose roughened below the nodes; internodes smooth or nodulose. Leaf sheaths 5-15 cm long, shorter than the internodes, about Y the length of the internodes, glabrous to nodulose roughened, margins hyaline; ligules 0.4-1.5 mm long, membranous, decurrent, apex acute, erose and toothed; blades 0.7-2 cm long, 0.2-1.2 zl mm wide, involute, arcuate-spreading, glabrous below and hi above. Panicles 1-5 cm long, 0.1-0.5 cm wide, narrow, contracted, loosely-flowered, usually on an exserted peduncle; primary branches 0.2-1.1 cm long, appressed, ascending; pedicels 0.1-0.8(-1.2) mm long, shorter than the spikelets, minutely se- tose. Spikelets 1.4-2.4 mm long; glumes 0.6-1.8 mm long, equal, unawned, glabrous, usually 1-nerved, occasionally 2- or 3-nerved, 1⁄2 to 2/3 the length of the floret, green or purple, apex acute; lemmas 1.4-2.4 mm long, lanceolate, indistinctly 3-nerved, mucronate or not, green or purplish, densely villous along the midvein and margins on the proximal 4/5, the hairs 0.4-1 mm long, apex acute, the mucro 0.1-0.6 mm long; paleas 1.2-2.1 mm long, lanceolate, densely villous between the nerves on the proximal 2/3, apex acute; anthers 0.8-1.5 mm long, yellow, dark green, or purple. Caryopses 1-1.4 mm long, ellipsoid to fusiform, dark brown. 2n = 20, 22. Phenology.—Flowering July through October. Distribution and habitat. —Muhlenbergia villiflora occurs in openings primarily on alkaline soils derived from gypsum, rocky flats, and desert g 1s usually in small, isolated populations associated with Larrea tridentata, Juniperus monosperma, Flaveria sp., Isocoma sp., Sarcobatus vermiculatus, Parthenium sp., Prosopis sp., Dasyochloa pulchella, Enneapogon desvauxii, Sporoblolus airoides, Opuntia imbricata, Buddleja sp., Muhlenbergia Peterson et al., Muhlenl ii f T í Mavi 25 tenuifolia, Erioneuron avenaceum, and Bouteloua spp.; 1500-2100 m. In México, M. villiflora var. villiflora also occurs in Chihuahua, Durango, Zacatecas, and San Luis Potosí. Comments.—Plants that occur in New Mexico and Texas, U.S.A. belong to M. villiflora var. villosa (Swal- len) Morden. This variety differs from var. villiflora by having longer spikelets (1.8-2.5 mm versus 1.4-2.3 mm long) and a preference for alkaline to calcareous, rather than gypsiferous soils. Specimen mined. MÉXICO. Coahuila: Municipio de Dr. Arroyo, Ejido El Jarro, J. Garza 192 & M. Castillo (COCA); pesa) de RAM Rancho Experimental Santa Teresa de La Rueda, 87 km NE de Ocampo, brecha Ocampo-Boquillas del Carmen, M.T. Ruíz-de León s.n., M.A. Carranza P. & M. Vásquez R. (ANSM); Municipio de Saltillo, 28.3 mi S de Saltillo on Hwy 54 to Concepción del Oro, Za- catecas P.M. Peterson 10029, C.R. Annable & J. Valdés-Reyna (ANSM, US); 53.2 km S of Saltillo on MEX Hwy 54 and 28.5 km E on road to La Ventura, P.M. Peterson 10040, C.R. Annable & J. Valdés-Reyna (US); 50 km S de Saltillo, carretera 54 a Concepción del Oro, Zacatecas, J. Valdés-Reyna 2059, J.A. Villarreal Q. & M.A. Carranza P. (ANSM); 6 mi N of La Ventura, I.M. Johnston 7642 (US); 10 mi E of Fraile, road from Saltillo to Concepción del Oro, I.M. Johnston 7305 (US); Estación Carneros, carretera a Zacatecas 30 km S de Saltillo, M.A. Carranza P. 518 & P.M. Peterson (ANSM); La Ventura, 80 km S de Saltillo, J.A. Villarreal Q. 1992, 3501, M.A. Carranza P. & J. Valdés-Reyna (ANSM); Rancho Experimental Los Angeles, 30 km S of Saltillo, Hwy 54 Saltillo- Concepción del Oro, Zacatecas, C.W. Morden 510, 514 (ANSM, TAES); S of Saltillo, 55 km on Hwy 54, C.W. Morden 506 (ANSM, TAES); 46 mi SE of Saltillo along hwy, F.W. Gould 10547 & D. Watson (TAES); Saltillo, A.S. Hitchcock 5613 (MEXU). Nuevo León: 5.6 mi E of junction of Hwy 57 on Hwy 58 towards Linares, P.M. Peterson 13291 & M.B. Knowles (US); near San Rafael (Hwy 57), P.M. Peterson 158116 J. Valdés-Reyna (US); Sandia where small laguna (Barrano) used to be, P.M. Peterson 17842, J. Valdés-Reyna & G.S. Hinton (ANSM, US); 5.1 mi S of Milagro, P.M. Peterson 17817, J. Valdés-Reyna & G.S. Hinton (ANSM, US); Municipio de Aramberri, San Juan Puente & Avilés, G.S. Hinton 18789 (HINT); Trinidad a Santa Gertrudis, G.S. Hinton 27296 (HINT); Municipio de Galeana, 2 km S del Salero, 1 km E de carretera 57 Matehuala-Saltillo, J. Valdés-Reyna 1616 & J.S. Marroquín de la F. (ANSM); Rancho Aquililla, G.S. Hinton 19532, 27044 (HINT); 20 km E of border with Nuevo León, Hwy 57 SE of La Providencia, C.W. Morden 516 (ANSM, TAES); 6.5 mi S of border of Coahuila and Nuevo Leon on Hwy 57 towards Matehuala, P.M. Peterson 13280 & M.B. Knowles (US); 20 km S of San Roberto along Hwy 57, S.L. Hatch 4939, J. Valdés-Reyna & J. Kessler (ANSM, SLPM, TAES); 5 km SE of “Y intersection in Galeana on Hwy 58, 34 km SE of San Roberto intersection with Hwy 57, C.W. Morden 517 (ANSM, TAES); 18 mi S of San Roberto, S.L. Hatch 4939b, J. Valdés-Reyna & J. Kessler (TAES); 34 mi NNW of San Roberto, 2 mi S of Providencia, J.R. Reeder 4258 & C.G. Reeder (US); 18 mi S of San Roberto, J.R. Reeder 4261 & C.G. Reeder (US); 20 km E of Coahuila border on Hwy 57, C.W. Morden 516, S.L. Hatch & J. Valdés-Reyna (TAES); 107.7 km Matehuala-Saltillo, E. Aldrete M. s.n. (INEGD; 73 mi SE of Saltillo, FW. Gould 10114 (TAES); 2 km al NW de San Rafael, sobre carretera Matehuala-Saltillo, H. López S. s.n. (INEGD; 3 mi E of jct of Hwy 60 & 57, J.R. Reeder 3961 & C.G. Reeder (US); 5 mi S of La Trinidad, 93 mi N of Matehuala, J.R. Reeder 2914 & C.G. Reeder (MEXU); 50 km S de San José de Raíces, sobre la carretera a Matehuala, J. Rzedowski-Rotter 32273 (ANSM, IBUG); 1 mi S of Galeana, K.W. Allred 5503, T. Columbus & J. Valdés-Reyna (ANSM); 6 mi N of San Roberto on Hwy 57, K.W. Allred 5511, J. Valdés-Reyna & T. Columbus (ANSM), Ejido ‘El Tokio’, I. Cabral-Cordero 1094 (ANSM); 32 km N of San Roberto, 18 km S of San Rafael, C.W. Morden 520 (ANSM, TAES); Municipio de Los Ramones, 1 mi $ of La Providencia, J.R. Reeder 5172, 5173 & C.G. Reeder (ENCB). Tamaulipas: Municipio de Bustamante, Ejido La Cardona, C.R. López-Aguilar 42a & A. Brito (COCA). 38. Muhlenbergia virescens (Kunth) Trin., Gram. Unifl. Sesquifl. 193, t. 5a, f. 7. 1824. (Fig. 18, E-H). Podosemum virescens Kunth, Nov. Gen. Sp. C ed.) 1:132. 1816. Trichochloa virescens (Kunth) Roem. & Schult., Syst. Veg. 2:389. 1817. Muhlenbergia virescens (Kunth) Kunth, Révis. Gramin. 1:64. 1829, isonym.Tvre: MÉXICO. Guanajuato: near Santa Rosa de la Sierra, Sep, EWH.A. Humboldt & A.J.A. Bonpland s.n. (HOLOTYPE: P!; IsoTYPE: US-86633 fragm. ex P-BONPL!) Muhlenbergi la Swallen, Contr. U.S. Natl. Herb. 29(9):410. 1950. Tree: MÉXICO. Guanajuato: dry oak-wooded slopes ca. 12 mi o from juat d to Santa Rosa, 30 Sep 1946, H.E. Moore, Jr. 1353 (HoLoTYrE: US-1963089!). Caespitose perennials with short, stout rhizomes. Culms 30-80 cm tall, erect, mostly glabrous below the nodes, the nodes basal, flattened, usuallyl node per culm; internodes mostly scabrous. Leaf sheaths 10—30 cm long, shorter than the internodes, scabrous to smooth; basal sheaths densely pubescent to glabrous abaxially, smooth and shiny adaxially, becoming flattened and spirally twisted with age; ligules 2-12 mm long, membranous to hyaline, somewhat firmer below, decurrent, apex acuminate often lacerate; blades 8-30 cm long, 1-3 mm wide, flat or usually tightly involute, scaberulous below, short-spiculate and often villous above to nearly glabrous, the hairs 0.2-0.4 mm long, usually appressed, the spicules shiny to whitish. Panicles 5-20 cm long, 0.5-6 cm wide, narrow, loosely-contracted to weakly spreading, interrupted below, mostly whitish to strami or grayish-green; primary branches 0.5-7 cm long, appressed and ascending to spreading up to 45? from the rachises; central axis flattened with 2 ribs, scabrous; pedicels 0.5-3(-5) mm long, usually shorter than the spikelets, scabrous. Spikelets 3.4—5 mm long, mostly stramineous to whitish or grayish-green; glumes (2.2-)2.5-5 mm long, shorter to longer than the floret, unequal, mostly whitish to stramineous or grayish-green sometimes with plumbeous mottles, shiny and usually glabrous to rarely 996 Journal of the Botanical R h Institute of Texas 1(2) scabrous below, usually without short hairs; lower glumes (2.2-)2.5-3.5 mm long, 1-nerved, apex obtuse to acute; upper glumes (3-)3.4-5 mm long, 3-nerved, apex truncate, obtuse or acute, often with 2-4 small teeth, the teeth less than 1/6 the length of the glumes; lemmas 3-5 mm long, lanceolate, terete, awned, gray- ish-green sometimes with plumbeous mottles, pilose nearly throughout and scabrous along the acuminate apex, the hairs 0.2-1.2 mm long, the awn 8-17 mm long, flexuous, scabrous, greenish-plumbeous; paleas 2.8-4.5 mm long, shorter than the lemma, pilose throughout, apex obtuse; anthers 222.2 mm long, purple. Caryopses 1.8-2 mm long, fusiform, brownish. 2n = 20, 40 (Reeder 1995). Phenology.—Flowering August through November. Distribution and habitat. —Muhlenbergia virescens is found on rocky slopes, volcanic tuffs, calcareous flats, canyon bottoms, and ridges usually beneath closed pine forests associated with Pinus spp., P. reflexa, P. chihuahuana, Quercus spp., Q. gambelii, Juniperus deppeana, Arctostaphylos pungens, Ceanothus spp., Arbutus sp., Alnus sp., and Festuca sp.; 1600—2700 m. This species occurs in central México from Estados Guanajuato, Hidalgo, San Luis Potosí, Zacatecas, Queretaro, México, Nayarit, Michoacán, Jalisco, Aguascalientes, and ied d. MÉXICO. Nuevo León: BE de Galeana, SW of Laguna de Labradores, G.S. Hinton 27883, 27886, 27945 (HINT. Tamaulipas: Peña Nevada, J.A. McDonald 2045 (MEXU, TEX); Municipio de Miquihuana, Arroyo Las Huertas, col. La Peña, J. A. Barrientos & M. H. Cervera 120 (COCA); Est. Ant. TELMEX, "s Valle Hermoso, Cisneros M. 199 (COCA). 39. Muhlenbergia wrightii Vasey ex J.M. Coult., Man. Bot. Rocky Mt. 409. 1885. (Fig. 19, F-D. Wee: U.S.A. New Mexico: 1851-1852, C. Wright 1986 (HoLotvrE: US-556872!; IsoTYPE: P!). Muhlenb is Mez, Repert. Spec. Nov. Regni Veg. 17:213. 1921. Tyre: U.S.A. CoLoraDo: Chiann (Cheyenne) Canyon, 5 Oct 1878, M. E. Jones 806 ME US-994951!). Caespitose perennials. Culms 15-60 cm tall, erect with knotty, thickened bases, flattened, strigose to his- pidulous below the nodes; internodes hispidulous to almost glabrous. Leaf sheaths 1.2-7.5 cm long, mostly shorter than the internodes, glabrous to scaberulous, compressed-keeled near base; ligules 1-3(-5) mm long, membranous, apex truncate; blades 1.4-12 cm long, 1-3 mm wide, flat to folded, slightly keeled, strigose above and glabrous to scaberulous below. Panicles 5-16 cm long, 0.2-1.2 cm wide, narrow, densely flowered, contracted, and spikelike; panicle branches 0.3-2 cm long, ascending and tightly appressed; pedicels 0.1-1.4 mm long, shorter than the spikelets. Spikelets 2-3 mm long, dark green or plumbeous; glumes 0.5-1.6 mm long, about equal, 1-nerved, mucronate, scaberulous along midvein, apex abruptly narrowed, acute or obtuse, the mucros 0.5-1 mm long; lemmas 2-3 mm long, lanceolate, mucronate, plumbeous, sometimes mottled with greenish-black areas and lighter areas, short appressed pubescence along the margins and midvein on the proximal Y to %, the hairs about 0.5 mm long, apex acute to acuminate, scaberulous, the mucro 0.3-1 mm long; paleas 1.9-3 mm long, lanceolate, short appressed pubescence between the nerves on the proximal %, apex acute to acuminate, the nerves sometimes mucronate, the mucros up to 0.3 mm long; anthers 1.3-1.8 mm long, greenish. Caryopses 1.2-2 mm long, fusiform, brownish. Phenology.—Flowering July through September. Distribution and habitat. —Muhlenbergia wrightii is found on gravelly prairies, grassy flats, rocky slopes, open wet and dry meadows on granitic, sandstone, or limestone derived soils associated with Pinus edulis, P. ponderosa, P. reflexa, Juniperus spp., J. monosperma, Picea engelmanni, Pseudostuga menziesii, Arctostaphylos pungens, Quercus gambelii, Q. turbinella, Cercocarpus breviflorus, Opuntia sp., Bouteloua gracilis, Pascopyrum smithii, Nassella tenuissima, Bromus carinatus, and Chrysothamnus nauseosus; 1100-3000 m. This species occurs in southeastern Utah, eastern Arizona, New Mexico, western Colorado to northern México, also in Estados Baja California, Chihuahua, Sonora, and Durango. Specimens examined. MÉXICO. Coahuila: Madera del Carmen, 13.3 mi NW of Pilares, P.M. Peterson 18893 & J. Valdés-Reyna (ANSM, US). Tamaulipas: burned over area between Marcella and Hermosa, Stanford 2672, Lauber & Taylor (US EXCLUDED SPECIES — Muhlenbergia ramulosa (Kunth) Swallen has been reported from the flora region (Beetle et al. 1995; Espejo- Peterson et al., Muhl Serna et al. 2000) but no specimens supporting its presence have been located. Muhlenbergia schreberi Gmel. was erroneously identified as M. microsperma (DC) Kunth on a few specimens examined by the authors. ACKNOWLEDGMENTS We wish to thank the curators at the following herbaria for help locating specimens: ANSM, ARIZ, B, BA, BAA, BM, CIDIIR, COCA, ENCB, GH, HINT, IBUG, INEGI, K, LE, MEXU, MICH, MO, MSC, NMC, NMSU, NY, P, RSA, SI, SLPM, TAES, TEX, UAT, US, UT, UTC, VT, and W; Smithsonian Institutions, Fellowships and Grants for supporting research visits by YHA and JVR to the United States National Herbarium; the Restricted Endowments Fund, the Scholarly Studies Program, Research Opportunities, Atherton Seidell Foundation, and Biodiversity Surveys and Inventories Program all at the Smithsonian Institution for fi- nancial support; Utah State University for permission to use illustrations that appeared in Flora of North America, Volume 25, prepared by Linda A. Vorobik and Annaliese Miller; Susan C. Escher, Midge Gillete, Cathy Pasquale, Lisa E. Roberts and Alice R. Tangerini for preparing the illustrations; Alice R. Tangerini for preparing all the plates; and Juan Javier Ortíz Diaz, Maria Teresa Mejía Saulés, and Barney Lipscomb for reviewing the manuscript. REFERENCES BEETLE, A.A., J.A. MIRANDA S, V. JARAMILLO L., A.M. RobrícuEz R., L. ARAGON M., M.A. VERGARA B., A. CHIMAL, and O. Dom- GUEZ S. 1995. Muhlenbergia Schreb. In: Las Gramíneas de México, Tomo IV. COTECOCA-SAGAR,, México, D. F, México. Pp. 32-224. CoLumBus, J.T., H.L. Bett, R. CeRROs-TLATILPA, M.P GRIFFITH, and J.M. Porter. 2002. Schaffnerella rediscovered! (Gramineae: Chloridoideae). Aliso 20:45-50. CoLumBus, J.T., R. CERROs-TLATILPA, M.S. Kinney, M.E. Siqueiros-DELGADO, H.L. Bett, M.P. GRIFFITH, and N.F. REFULIO-RODRIGUEZ. 2007. Phylogenetics of Chloridoideae (Gramineae): a preliminary study based on nuclear ribosomal internal transcribed spacer and chloroplast trnL-F sequences. Aliso 23:xxx-xxx. DAVILA-ARANDA, PR. LiRA-SAADE, and J. VALDES-REYNA, 2004. Endemic species of the grasses in Mexico: a phytogeo- graphic approach. Biodivers. & Conservation 13:1101-1121 Davita, P, M.T. MEJfA-SAULÉs, M. Gomez-SANCHEZ, J. VALDÉs-REYNA, J.J. Ortiz, C. Morin, J. CASTREJÓN, and A. Ocampo. 2006. Catálogo de las gramíneas de México. Universidad Nacional Autónoma de México and Comisión Nacional para el conocimiento & uso de la Biodiversidad, México, D.F. Duval, M.R., PM. Peterson, and A.H. Christensen. 1994. Alliances of Muhlenbergia (Poaceae) within New World Eragrostideae are identified by phylogenetic analysis of mapped restriction sites from plastid DNA's. Amer. J. Bot. 81:622-629 ESPEJO SERNA, A., A.R. LÓPEZ-FERRARI, and J. VaLDEs-REYNA. 2000. Poaceae. Pp. 10:8-236. In: A. Espejo Serna & AR. López-Ferrari, eds. Las Monocotyledóneas Mexicanus: una Synopsis Florística, Partes IX-XI. Consejo Nacional de la Flora de México, A.C., Univerisidad Autónoma Metropolitana-Izapalapa, and Comisión Nacional para el conocimiento & uso de la Biodiversidad, México, D.F. HarrensLEY, PW. and L. Watson. 1992. Diversification of photosynthesis. In: G.P. Chapman, ed. Grass evolution and domestication. Cambridge University Press, Cambridge, UK. Pp. 38-116. HERRERA ARRIETA, Y. 1998. A revision of the Muhlenbergia montana (Nutt.) Hitchc. complex (Poaceae: Chloridoideae). Brittonia 50:23-50. HERRERA ARRIETA, Y. 2001. Las Gramíneas de Durango. Instituto Politécnico Nacional and Comisión Nacional para el Conocimiento & Uso de la Biodiversidad, Durango, México. HERRERA ARRIETA, Y. and P.M. Peterson. 2007. Muhlenbergia (Poaceae) de Chihuahua, México. Sida, Bot. Misc. HiLu, KW. and L.A. Auct. 2000. Bund el in subfamily Chloridoideae (Poaceae) based on matK sequences: A preliminary as t. In: $.W.L. Jacobs and J. Everett, eds. Grasses: Systematic | i CSIRO, Melbourne. Pp. 184-188. £L Dag PRA D hi PEA 998 Journal of f Texas 1(2) Hilu, KW. and L.A. Auce. 2001. A phylogeny of Chloridoideae (Poaceae) based on matK sequences. Syst. Bot. 26:386-405. Mez, C. 1921. LXVII. Gramineae novae vel minus cognitae. Repert. Spec. Nov. Regni Veg. 17:204—214. McVauaH, R. 1983. Flora Novo-Galiciana. A descriptive account of the vascular plants of western México. Gra- mineae. 14:1—436. University of Michigan Press, Ann Arbor. PETERSON, PM. 1989. A re-evaluation of Bealia mexicana (Poaceae: Eragrostideae). Madroño 36:260-265. PETERSON, PM. 2000. Systematics of the Muhlenbergiinae (Chloridoideae: Eragrostideae). In: S. W.L. Jacobs and J. Everett, eds. Grasses: Systematics and Evolution. CSIRO, Melbourne. Pp. 195-212. Peterson, PM. 2003. Muhlenbergia Schreb. In: M.E. Barkworth, K.M. Capels, S. Long, and M.B. Piep, eds. Magno- liophyta: Commelindidae (in part): Poaceae, part 2. Flora of North America North of México, volume 25. Oxford University Press, New York. Pp. 145-201 PETERSON, PM. and C.R. Annable. 1990. A revision of Blepharoneuron (Poaceae: Eragrostideae). Syst. Bot. 15:515- PETERSON, PM. and C.R. ANNABLE. 1991. Systematics of the annual species of Muhlenbergia (Poaceae-Eragrostideae). Syst. Bot. Mongr. 31:1-109. PETERSON, PM. and C.R. AnnaBLE. 1992. A revision of Chaboissaea (Poaceae: Eragrostideae). Madroño 39:8-30. PETERSON, PM and C.R. AnmabLe. 2003. Blepharoneuron. In: M.E. Barkworth, K.M. Capels, S. Long, and M.B. Piep, eds., Magnoliophyta: Commelinidae, (in part): Poaceae, part 2. Flora of North America North of Mexico, volume 25. Oxford University Press, New York. Pp. 47, 48, 50. Peterson, P.M, and J.T. Cotumeus. 1997. Allelic variation in the amphitropical disjunct Scleropogon brevifolius (Poa- ceae: Eragrostideae) Biolannia, Edicion Especial 6:473-490. PETERSON, P.M, J.T. ee E os TES and M.S. Kinney. 2001a. Phylogenetics of Muhlenbergia and relatives (Poaceae: Chloridoic | ribed spacer region sequences (nrDNA). Botany2001 abstract: E EE UI uu m I PETERSON, PM. J.T. Columbus, and S.J. PENNINGTON. 2007. Classification and biogeography of New World grasses: Chloridoideae. Aliso 23:xx-xxx. PETERSON, P.M. J.T. CoLumeus, N.F. Reruuo RODRIGUEZ, R. CERROS TLATILPA, and M.S. Kinney. 2004. A phylogeny of the Muhlen- bergiinae (Poaceae: Chloridoideae: Cynodonteae) based on ITS and trnL-F sequences. Botany2004 abstract: http://www.2004.botanyconferenc os php?func= iia A Peterson, PM., M.R. Duva, and A.H. CENSET 1993. Allozy liff mong B , Muhlenberg argentea, and M. lucida (Poaceae: Eragrostideae). Madroño 40:148-160. PETERSON, P.M. and Y. HERRERA Arrieta. 1995. Allozyme variation in the amphitropical disjunct, Chaboissaea (Poaceae: Eragrostideae). Madroño 42:427-449. PETERSON, P.M. and Y. HERRERA ARRIETA. 2001. A leaf blade anatomical survey of Muhlenbergia (Poaceae: Muhlenber- giinae). Sida 19:469-506 Peterson, PM. and O. Morrone. 1997 Allelic variation in the amphitropical disjunct Lycurus setosus Muhlenbergiinae). Madroño 44:334-346, PETERSON, P.M. and J.J. Ortiz-Diaz. 1998. Allelic variation in the amphitropical disjunct Muhlenbergia torreyi (Poaceae: Muhlenbergiinae). Brittonia 50:381-391. PETERSON, P.M. R.J. SORENG, G. DaviDsE, T.S. FILGUEIRAS, F.O. ZULOAGA, and E.J. Jubziewicz. 2001b. Catalogue of New World grasses (Poaceae): Il. Subfamily Chloridoideae. Contr. U.S. Natl. Herb. 41:1-255 Peterson, P.M., R.D. Wesster, and J. Vatpés-Reyna. 1995. Subtribal classification of the New World Eragrostideae (Poaceae: Chloridoideae). Sida 16:529-544. PETERSON, P.M., R.D. Wesster, and J. VALDEs-REYNA. 1997. Genera of New World Eragrostideae (Poaceae: Chloridoideae). Smithsonian Contr. Bot. 87:1—50. PowtLL, A.M. 1994. Grasses of the Trans-Pecos and adjacent areas. University of Texas Press, Austin. REEDER, C.G. 1985. The genus Lycurus (Gramineae) in North America. Phytologia 57:283-291. Reever, C.G. 1995. The resurrection of a species: Muhlenbergia straminea (Gramineae). Phytologia 78:417-427. Poaceae: — Peterson et al., Muhlenl ii f T í Mavi 35 SANCHEZ, E. and Z.E. RUGOLO DE AGRASAR. 1986. Estudio taxonomica sobre el genero Lycurus (Gramineae). Parodiana SobtERSTROM, T.R. 1967. Taxonomic study of subgenus Podosemum and section Epicampes of Muhlenbergia (Gra- mineae). Contr. U.S. Natl. Herb. 34:75-189. Sykes, G.R., A.H. CHRISTENSEN, and P.M. Peterson. 1997. A chloroplast DNA analysis of Chaboissaea (Poaceae: Eragros- tideae). Syst. Bot. 22:291-302. ZULOAGA, F.O., E.G. Nicora, Z.E. RÚGOLO DE AGRASAR, O. Morrone, J. PiENSIERO, and A.M. CIADILLA. 1994. Catálogo de la familia Poaceae en la Republica Argentina. Monogr. Syst. Bot. Missouri Bot. Gard. 47:1-178 APPENDIX 1 Index of names treated; accepted names are given in Roman type (bold for this treatment) and synonyms are italicized. Aegopogon, 938 ideo 938 ar. abortivus, 939 gritos 939 ortivus, 939 nt purpureus, 939 his unisetus, 939 gracilis, 939 dean 939 tenellus, 9 var. abortivus, 939 unisetus, Agrostis caespitosa, 993 di Pond lla, 950 eremophila, 950 glabrata, 985 microsperma, 970 minutissima, 970 e 993 Arundo quitensis, 991 tenuifolia, 991 Blepharoneuron, 939 tricholepis, 941 Calycodon montanum, 971 Calamagrostis quitensis, 991 tenuifolia, 991 Cinna racemosa, 980 setifolia, 967 Crypsinna breviglumis, 955 macr , 967 rigens, 984 setiglumis, 984 Crypsis macroura, 967 setifolia, 967 Epicam berlandieri 987 minutiflora, 987 stricta, 987 var. distichophylla, 953 subpatens, 958 ri robusta, 987 Hymenothecium unisetum, 939 Lamarckia tenella, 939 Lycurus, 941 a phleoides var. asco 945 setosus, Milium microspermum, 971 Muhlenbergia, 946 a 1 acuminata, 955 dA i affini o 953 coloradensis, 996 confusa, 971 curvula, 995 densiflora, 955 depauperata, 950 distans, 958 distichophylla, 953 emersleyi, 958 enervis, 971 fasciculata, 970 gooddingii, 958 gracillima, 993 gracilis, 978 EE 960 porter presliana, 987 ubescens, 976 purpusii, 978 1000 quadridentata, 978 quitensis, 991 racemosa, 980 var. ramosa, 980 ramosissima, 970 var. fuos 991 tenuifolia, 97 vaseyana, 9 villiflora, 994 var. villiflora, 994 £L Dag PRA D hi Journal of virescens, 995 Sporobolus subsp. quadridentata, 974 arenaceus, 947 wrightii, 996 asperifolius, 950 Panicum microspermum, 971 auriculatus, 947 confusus, 971 Phleum macrourum, 967 ligulatus, 95 Pleopogon setosum, 945 microspermus, 971 T minutissimus, 970 e pm repens, 982 GTP. tricholepis, 941 [SIG de utilis, 993 distichophyllum, 953 glabratum, 985 Trichochloa glau glabrata, 985 gracile, 978 microsperma, 967 gracillimum, 993 quadridentata, 978 p j setosa, 967 quadridentatum, 978 rigida, 985 rigidum, virescens, 995 setosum, 96 ilfa strictum, 990 asperifolia, 950 tenuifolium, 991 confusa, 971 virescens, 99 pubescens, 994 Polypogon racemosus, 980 racemosa, 980 repens, 982 tricholepis, 941 utilis, 993 4 f Texas 1(2) A SECOND SPECIES OF CYCLOPOGON S.S. (ORCHIDACEAE: SPIRANTHINAE) Eric A. Christenson 4503 21% Ave. West Bradenton, Florida 34209, U.S.A. orchideric@juno.com ABSTRACT A new species, Cyclopogon secundum sp. nov., is described from Ecuador. RESUMEN Se describe especie, Cyclopogon secundum sp. nov., de Ecuador. Comparison of modern specimens with the holotype of Cyclopogon ovalifolium Presl from Peru shows that three distinct species have been confused in Ecuadorian literature under this name. The two published drawings based on Ecuadorian collections as C. ovalifolium (Dodson & Dodson 1989; Dodson & Escobar, s.d.) are based on misidentifications and neither one agree with the holotype. Both appear to be undescribed species and I take this opportunity to describe one of them here. P2 oco ovalifolium mud REL re 1:93. 1827. Gyrostachys ovalifolia (Presl) Kuntze, Rev. Gen. Pl. 2:664. 1891. E: PERU: Mountain near Huanocco (=Huanu ), Haenhe s.n. (PR! 305737/1780). Spiranthes preslii Lindl., Gen. & Sp. Orch. Pl. 470. 1840. a secundum Christenson, sp. nov. (Fig. 1). Te: ECUADOR. Prov. Pastaza: Colonia Játiva, ca. 12 km from Mera, Lugo 98 (HOLOTYPE: S!; IsoTypes: AMES, GB). 1 lifolium Presl sensu Dodson & Dodson, Icon. Pl. Trop., ser. 2, pl. 430. 1989, non Presl (1827). To Species haec C. ovalifolium Presl similis sed florib g dis, lobo apicali labelli pubescenti differt. Rosulate terrestrials arising from fleshy fasciculate villose roots. Leaves 2—4, petiolate, elliptic, attenuate, subacute, the petioles to 4 cm long, the blades to 5.5 x 3 cm. Inflorescences erect pedunculate secund ra- cemes, the peduncle slender, to 19 cm long, with several approximate bracts, the bracts lanceolate, aciculate at the apex, the rachis 3.5 cm long, the floral bracts narrowly linear-lanceolate, aciculate, conspicuous, to 1.3 cm long. Flowers 10-15, tubular, the sepals and petals greenish white suffused with purple toward the base, the lip white. Sepals fused for more than 1/3 their length into a tube 3 mm long, oblong, obtuse, the free limbs 5.5 mm x 1.5 mm. Petals oblong from a narrower base, obtuse, asymmetric, appressed to the dorsal sepal, 8 x 1 mm, fused at the base for 1 mm. Lip weakly lobed by a subapical constriction, clawed, auriculate, oblong in overall outline, 8.5 x 3 mm, the auricles retrorse, pubescent, the midlobe transversely reniform, obscurely lobulate, the center of the lip sparsely pubescent, the midlobe with a densely pubescent central patch. Column typical for the genus, pubescent, 5 x 1 mm. Etymology.—From the Latin secundus, referring to the strongly secund raceme. Additional specimens: ECUADOR. Prov. Imbarbura: along trail to Río Chalguayaco, below Magnolia, lower Intag Valley, Drew E-587 (AMES). Prov. Pastaza: Río Topo, road from Baños to Puyo, 1400 m, 4 Jun 1985, B. Stein 2982 (MO, RPSC Cyclopogon secundum is easily separated from C. ovalifolium by its densely flowered secund inflorescence and pubescent lip apex. In C. ovalifolium the flowers are borne in a loose cylindric raceme and the lip apex is glabrous. Both of these species share the character of having the sepals fused into a tube for more than one third their length, the generic character for Cyclopogon in the strict sense when it is kept separate from Beadlea Small. J. Bot. Res. Inst. Texas 1(2): 1001 — 1002. 2007 1002 Journal of the Botanical R h Institute of Texas 1(2) J J hri | B. dorsal sepal: C, petal: D. lateral sepals: E. lip from above: F. col fi | G J pog Ji F L I L F 1 F z £ hal LA Garay. Finally, note should be made that the plant illustrated by Dodson and Escobar (s.d.: No. 267, p. 161) as C. ovalifolium appears to be the same species shown below it as a color photograph (based on Hirtz 3001). The color photograph they published as C. olivaceus (Rolfe) Schltr. (No. 276, p. 163) is clearly not that spe- cies (q.v., Mori et al., 1997: p. 299) and appears to be C. secundum. ACKNOWLEDGMENTS I thank the curators and staff in Prague (PR) and Stockholm (S) for courtesies extended during my visits. I also thank Leslie Garay for generously sharing his research archives and allowing me to publish his drawing. REFERENCES DopsoN, C.H. and P.M. Dopson. 1989. Cyclopogon ovalifolium Presl. Icon. Pl. Trop., ser. 2, pl. 430. Dobson, C.H. and R. Escogar R. s.d. Native Ecuadorian orchids, vol. 1. Editorial Colina, Medellin. MORI, S.A., G. Cremers, C. GRACE, J.-J. DE GRANVILLE, M. Horr, and J.D. MrrcheLL. 1997. Guide to the vascular plants of central French Guiana, Part 1. Mem. New York Bot. Gard. 76:1-422. LECTOTYPIFICATION OF MAXILLARIA HETEROCLITA (ORCHIDACEAE) Eric A. Christenson 4503 21% Ave. West Bradenton, Florida 34209, U.S.A. orchideric@juno.com Frequently there are problems with early botanical works that involve mixed collections of more than one element. Sometimes this is caused by specimens getting physically mixed and sometimes this is caused by combining unrelated elements when producing illustrations. Examination of the type illustration of Maxil- laria heteroclita Poepp. & Endl., now accepted as a species of Chaubardia, shows it to be a combination of two quite different genera requiring lectotypification. Chaubardia heteroclita (Poepp. & Endl.) Dods. & D.E. Benn., Icon. Pl. Trop., ser. 2, pl. 23. 1989. (Fig. . Maxillaria heteroclita Poepp. & Endl., Nov. Gen. Sp. Pl. 1:37, t. 63. 1836; Warscewiczella heteroclita (Poepp. & Endl.) Hoehne, Arq. Bot. Estado Sao Paulo, nov. ser., 2(6):126. 1952. Cochleanthes heteroclita (Poepp. & Endl.) Schultes & Garay, Bot. Mus. Leafl. 18:325. 1959. eee heteroclita r & Endl.) Garay, Orquideología 4:146. 1969. Tver: original drawings of a flower and floral dissections, Poeppig s.n 2 ted here: W!, Herb. Reichenbach 51473). Tabula 63 comprises a plant "S two unifoliate pseudobulbs, a scape with an open flower, a scape with a bud, a separate view of the ovary, column and lip in profile, and a separate view of the undersurface of the column. The scapes and the separate views represent Chaubardia heteroclita. The vegetative parts shown in t. 63 are of an unknowable species of Maxillaria Ruiz & Pav. In Chaubardia there are no pseudobulbs and the plants produce fan-shaped growths of several leaves. I am unaware of any authentic specimen with the name Maxillaria heteroclita conserved at W. The original drawings are unambiguously identifiable to the species level and thus serve as lectotype. The lec- totype is one of six separate elements mounted as Herb. Reichenbach 51473. In the upper left corner is a drawing of a Maxillaria from 1866. On the upper right is a packet without provenance. On the lower right is a drawing of a Maxillaria from Mexico collected by Schiede. On the lower left is a drawing of a Maxillaria lip and column with an illegible annotation. The lectotype itself is one of two pieces of paper mounted together on a slightly larger piece of paper at left center. These comprise part of the Endlicher Herbarium. The lefthand drawing is a small sketch of a plant that became the basis for the Maxillaria element of t. 63. The righthand set of drawings is the lectotype and was faithfully reproduced in part as t. 63. Lectotypification in this manner maintains the well established use of this name (Brako & Zarucchi 1993; Dodson & Bennett, 1989) for an attractive species that has been widespread in cultivation for several decades (e.g., Teuscher 1966). ACKNOWLEDGMENTS I thank the curators and staff of the Reichenbach Herbarium (W) for courtesies extended during my visit. REFERENCES Brako, L. and J.L. ZanuccHi. 1993. Catalogue of the flowering plants and gymnosperms of Peru. Monogr. Syst. Bot. 45:1-1286. Dobson, C.H. and D.E. Bennett. 1989. Chaubardia heteroclita (Poepp. & Endl.) Dodson €: Bennett. Icon. Pl. Trop., ser. 2, pl. 23. TEUSCHER, H. 1966. Zygopetalum rhombilabium viz. Cochleanthes rhombilabia. Amer. Orchid Soc. Bull. 35: 8 J. Bot. Res. Inst. Texas 1(2): 1003 — 1004. 2007 1004 Journal of the Botanical R h Institute of Texas 1(2) Fic. 1. The type ill E lita Poepp. & Endl. s Chaubardia. MISCELLANEOUS TYPIFICATIONS, NEW COMBINATIONS AND ONE NEW VARIETY IN NORTH AMERICAN CRATAEGUS (ROSACEAE) J.B. Phipps The University of Western Ontario L4ACPUE CLEFT TE MZ LIVIUY y 51 Richmond St. N. London, Ontario, NGA 5B7, CANADA jphipps@uwo.ca ABSTRACT Y Various nomenclatural actions to validate certain names of North American Crataegus are taken. Firstly, the following new variety is eee C. chrysocarpa vat. eA B. Phipps. In addition, eight new names or combinations are provided here, as fol- C. brazoria Sarg. var. viburnifolia, C. chrysocarpa Ashe var. blanchardii, C. chrysocarpa Ashe var. praecox, C. coccinea L. var. pringlei, C. pruinosa (H.L. ap K. Koch var. dissona, C. reverchonii Sarg. var. mohrii, C. succulenta Schrader ex Link var. TE C. viridis L. var. velutina. Finally, the following 10 Crataegus names are typified: C. blanchardii Sarg., C. dissona Sarg., C. gattingeri Ashe, C. georgiana Sarg., C. jackii Sarg., C. mohrii Beadle, C. neofluvialis Ashe, C. praecox Sarg., C. velutina Sarg., and C. viburnifolia Sarg Key Wonps: Crataegus, C. chrysocarpa var. vigintistamina J.B. Phipps, var. nov., new combinations, various typifications, Flora North America, volume 9 RESUMEN ali E 1 i| Sere para validar ciertos nombres de Crataegus norteamericas. Primero, se describe la siguiente cane C. chrysocarpa var. vigintistamina J.B. Phipps. Además, se aportan ocho nuevos nombres o combinaciones: C. brazoria Sarg. var. poes C. chrysocarpa Ashe var. blanchardii, C. chrysocarpa Ashe var. praecox, C. coccinea L. var. pringlei, C. pruinosa (H.L. Wendl.) K. Koch var. dissona, C. pd PUE var. mohrii, C. succulenta Schrader ex Link var. neofluvialis, C. viridis L. var. velutina. Finalmente rataegus: C. blanchardii Sarg., C. dissona Sarg., C. gattingeri Ashe, C. georgiana Sarg., C. jackii Sarg., c mohrii Beadle: C. neofluvialis Ashe, C. praecox Sarg., C. velutina Sarg., y C. viburnifolia Sarg. When preparing the treatment of so large a genus as Crataegus for the Flora of North America it is inevitable that one encounters a number of situations where taxa are quite well-known but do not fit comfortably into the taxonomic system without an alteration in rank. Such is the case for the eight varieties that had each originally been described as species and are proposed as new combinations here. This is primarily a ‘housekeeping’ document and represents the best judgment of the author without entering into the complex and detailed study required in each case to more fully defend it. An exception is in the case of C. coccinea var. pringlei where a much more elaborated paper (with J.A. Macklin) is being pre-empted for the sake of meeting the publication schedule for Flora of North America vol. 9 (Rosaceae). The basionyms to be treated here usually need lectotypification and all taxa are assigned to the taxo- nomic series to be used by the author in Flora of North America vol. 9. In some cases the author cited a type number from among a range of paratypes that does not really provide the discrimination needed for that taxon. In such cases I have also provided an epitype. Summary comments may also be given on distribution and differentiation from similar taxa.. The taxa treated are arranged alphabetically by series, as follows: series Coccineae (Loud.) Rehd. 1. es coccinea L. var. pringlei (Sarg.) J.A. Macklin & J.B. Phipps, a et stat. NOV. Crataegus pringlei ., Rhodora 3:21. 1901. Type: VERMONT: CHITTENDEN Co.: Charlotte, 27 May 1877 (LECTOT d by Macklin & Phipps ms 26). Leaf blades broadly elliptic to elliptic-ovate (typically 1.25 x as long as broad), concavo-convex, 4—7 cm, lobes often shallow, bases rounded to broad-cuneate. Stamens 8-10. Pomes often oblong. J. Bot. Res. Inst. Texas 1(2): 1005 — 1010. 2007 1006 Journal of the Botanical R h Institute of Texas 1(2) Flowering May; fruiting Sept-Oct. Brushy places, woodland margins, fencelines, overgrown pastures; 10300 m; Ont., Que.; Conn., Ill., Ind., Mass., ?Me., Mich., Minn., N.H., NY., Oh., Penna., R.I., Vt., Wisc. Crataegus coccinea var. pringlei in its most characteristic form | kedl A shaped leaves from pure coccinea as well as a tendency to oblong, rather than subspherical fruit (see key below). The extreme forms of variety pringlei are very distinct in appearance with their convex, broad-elliptic, usually only shal- lowly lobed leaves. Populations of pure examples of both varieties are quite frequently encountered. However, intermediates with var. coccinea are also quite common. Crataegus coccinea var. fulleriana (20 stamens), the third variety of C. coccinea, is in other respects, rather similar to var. pringlei. This matter will be elaborated in Macklin and Phipps (in prep.). Variety pringlei is found throughout the range of the species except not in Virginia and North Carolina. KEY TO VARIETIES OF CRATAEGUS COCCINEA . Leaf blades plane, ovate (1.4-1.5 x as long as broad), 5-8 cm, broadest in lower 1/3; lobes well-marked, bases wide- a to truncate; pomes usually suborbicular Crataegus coccinea var. coccinea 1. Leaf! concavo-convex, broadly elliptic to elliptic-ovate (typically 1.25 x as long as broad), 4-7 cm, lobes a /, bases rounded to broad-cuneate; pomes oblong to suborbicular. 2. Stamens 8-10; leaf blades conspicuously concavo-convex, this usually evident even from pressed spec- mens; pomes often oblong raat ia coccinea var. pringlei 2. Stamens ca. 20; leaf blades plane; pomes usually suborbicular g ar. fulleriana series Crus-galli (Loud.) Rehd. 2. Crataegus reverchonii Sarg. var. mohrii (Beadle) J.B. Phipps, comb. et stat. nov. Crataegus mohrii Beadle, Bot. Gaz. 28:416. 1899. Type: GEORGIA: Frovp Co.: Rome, flatwoods, 4 May 1899, C.D. Beadle 236 (Lectotype designated here: A; ISOLECTYPE: US). Comments.—The type number is indicated in the protologue; a fruiting specimen of the same collection number but dated 16 Sep 1898 is on the quoted sheet at A, duplicate US. Crataegus denaria Beadle, Biltmore Bot. Stud. 1:131. 1902. The co-type, Beadle 236A, fits the protologue precisely except that the inflorescences are variably perfectly glabrous to quite densely pilose! This variety is notable for its very strongly and sharply toothed foliage. Crataegus reverchonii var. mohrii grows primarily east of the Mississippi River with definite records for Mississippi, Alabama, Georgia and Tennessee. However, in reviewing over 1500 specimens of ser. Crus- galli for the southeastern United States, I have only encountered a few specimens clearly matching the type description. Likewise 1 have not knowingly encountered this variety in the field during over 15 field trips to the region so perhaps it is very rare. Crataegus reverchonii var. mohrii clearly belongs to C. reverchonii on account of its small, short leaves, 2.5-3.5 cm long at maturity, smallish flowers, 3-5 styles, and broad-elliptic to suborbiculate or rounded extension shoot leaves, characters that also very well differentiate the much more abundant var. reverchonii from C. crus-galli. Variety reverchonii in addition has a very distinct range. It is possible both that var. mohrii represents scarce introgressants of the type variety and a form of C. crus-galli towards the edge of the range of var. reverchonii or it may have arisen independently. As var. mohrii is hardly ever seen today it may be some time before this can be answered. The following key illustrates the rather small differences from the remainder of C. reverchonii. 1. Short-shoot leaves 2-3.5 cm; flowers 8-12 mm diam ; styles 3-5. 2. Flowers 8-10 mm diam.; inflorescence usually glabrous; west of Mississippi River Crataegus reverchonii var. reve rchonii 2. Flowers ca. 12 mm diam; inflorescence + pilose; east of Mississippi River — — Crataegus reverchonii var. mohrii 1. Short-shoot leaves 4-6 cm; flowers 10-12 mm diam; styles (2-)3 Crataegus reverchonii var. palmeri Phipps, New combinations and typifications in North American Crat 1007 series Macracanthae (Loud.) Rehd. 3. Crataegus succulenta Schrader ex Link var. neofluvialis (Ashe) E.J. Palmer, in E.J. Dole, ed., Fl. Vermont, 3 ed 155. 1937. Crataegus neofluvialis Ashe, J. Elisha Mitchell Sci. Soc. 16:71. 1900. Tyre: NORTH CAROLINA. AsuHE Co.: along New River, Jul 14 1897 or ‘09, WW. Ashe s.n. (Lectotype designated here: NCU). A damaged original label accounts for the > lack of certainty about the year of collection. Comments.—The designated specimen is a l th th nes with type in Ashes hand, another of which is represented by a photograph at US. Ashe (op. cit., 1900) ol d tl j ] the North Fork of the New River in Ashe Co., North Carolina and the adjacent part of Virginia. This variety is described as having particularly narrow leaves softly and abundantly pubescent on unfolding, later with scanty and roughish hairs or with age nearly glabrous, 2-3 pairs of shallow lateral lobes, 5—7 pairs of lateral veins; petiole eglandular; inflorescences 5-15 flowered, branches nearly glabrous; sepal margins strongly glandular-serrate in flower, sometimes + eroded in fruit and may appear subentire; stamens 10-15 (Kruschke 1965, counted 14-20); fruit 6-9 mm diam., greenish, orange or rosy-cheeked. series Molles (Loud.) Rehd. 4. Crataegus brazoria Sarg. var. viburnifolia (Sarg.) J.B. Phipps, comb. nov. Crataegus viburnifolia Sarg., Trees & Shrubs 2(3):145, pl. 159. 1911. Tyee: bM Brazoria Co.: Columbia, 23 Mar 1909, B.E Bush & C.S. uidi 11 (HOLOTYPE: A). Comments.—A g th ber and date, I have selected the one with the g t ber of flowers in good condition. It has a more or ES horizontal flowering spray across the top of the sheet. TEXAS. Brazoria Co.: Columbia, common in woods, fruit canary d 25 a 1901, B.E en tree 11, = pas 912 dud. een here: A). T : 1 omm nts nents. p I B & €t E E nere it t th t frit 1 [e] without which identity is ambiguous. Crataegus brazoria var. viburnifolia is a large bush or small tree from southeastern Texas very similar to C. mollis (Torr. & A. Gray) or C. texana Buckl. differing from them primarily in fruit color. The pure form is thus effectively a bright yellow-fruited C. mollis. It is only occasionally encountered today. It flowers early. Well grown, var. viburnifolia is one of the most handsome and spectacular of all hawthorns. KEY TO EXTREME FORMS OF CRATAEGUS BRAZORIA . Most! E | lovatet | elliptic, barely but sharply lobed, rough-hairy abaxially at maturity; anthers ivory; pomes 15-20 mm dia Crataegus brazoria var. viburnifolia . Most leaves + elliptic, not cr glabrous abaxially at maturity; anthers rose; pomes 8-10 mm diam Crataegus brazoria var. brazoria Crataegus brazoria is a large bush or small tree restricted to the lower Columbia watershed of southeast Texas. Its best growth is in rich alluvial soil with adequate moisture but it also occurs in a variety of brushy habitats. It is var. brazoria that is most different from C. mollis-texana on the basis of both leaf shape and indumentum. Pure variety brazoria is similar to the red-fruited C. texana but is rarely seen today. For now it is united with C. viburnifolia both because of the unusual fruit color and its similar local range, suggesting an adaptation to a particular frugivore disperser. Molecular work and intensive modern field studies are required to confirm this interpretation but it is the one that provisionally seems most realistic. series Pruinosae Sarg. ex Rehd. 5. Crataegus pruinosa (H.L. Wendl.) K. Koch var. dissona (Sarg.) J.B. Phipps, comb. et stat. nov. Crataegus dissona Sarg., Rhodora 5:60. 1903. Tyre: CONNECTICUT. New Haven Co.: Oxford, 27 May 1901, E.B. Hager 39 (LECTOTYPE desig- nated here: A Comment.—An llent fruiti i 29 Sep 1901) of the same number is mounted on the same sheet as the lectotype. Crataegus pruinosa (H.L. Wendl.) K. Koch f. dissona (Sarg.) Eggl., Rhodora 10:81. 1908. Crataegus brachypoda Sarg., Annual Rep. Missouri Bot. Gard. 19:100. 1908. Crataegus disjuncta Sarg., Trees & Shrubs 1:109. 1 Crataegus rigida Sarg., Annual Rep. Missouri Bot. A 19:99. 1908. 1008 Journal of the Botanical R h Institute of Texas 1(2) This is the only member of series Pruinosae with 10 stamens and as such is readily separated from other varieties of C. pruinosa. Crataegus pruinosa var. dissona, found from Missouri to Wisconsin and through the Great Lakes to New Eng- land, then south to Georgia, is widely scattered throughout the range of C. pruinosa. The detailed distribution is not well known, however and may be more widespread than recorded. The variation in leaf shape of var. dissona approximately parallels that of var. pruinosa. Crataegus brachypoda is a form with cream anthers while C. disjuncta has particularly large foliage. 6. Crataegus gattingeri Ashe, J. Elisha Mitchell Sci. Soc.17:12. 1900. Ter: TENNESSEE. Davipson Co.: Nashville, Sep 1880, Dr. Gattinger s.n. (LEcTOTYPE designated here: MO) Comment.—MO claims this is an isotype, but I could not locate duplicates at HUH, NY, TENN or US. E X georgiana Sarg., Bot. Gaz. 32:113. 1902. Type: GEORGIA. Foyn Co.: Rome, 22 Apr 1900, C.S. an a (LECTOTYPE designated iba Comment. md n georgiana is also typified here because of the long-standing vi f C. pruinosa or . However, this proves to be incorrect because careful examination of the type of C. georgiana shows that it cannot be mou differentiated from that of C. gattingeri Ashe, an earlier name. There are two sheets at A with this information, one the lectotype. series Rotundifoliae (Eggl. ex Sarg.) Rehd. 7. Crataegus jackii Sarg., Rhodora 5:162. Jun 1903. Tee: QUEBEC. La Prairie Co.: Caughnawaga, 29 May 1901, J.G. Jack 1 (LecToTYPE designated here: A). Comment mostly in flower. Crataegus jackii was restricted to southern Quebec in and near Montreal and was generally believed to be a variant of Crataegus chrysocarpa. However, it differs in its larger flowers, smaller and more broad-elliptic short-shoot leaf-blades, more suborbiculate outline of the extension shoot leaf-blades and very conspicuous bracteoles that are larger and more prominently glandular than in the latter species. Further, its bracteoles prove to be similar to those of the C. dodgei complex and the larger, better-developed bracteoles have stipitate glands. Crataegus jackii is very hairy in the manner of C. lumaria (Ashe, J. Elisha Mitchell Sci. Soc. 19:25, April, 1903) of the dodgei complex and differs significantly from that species only in its larger flowers and disjunct distribution. It is thus here made a synonym of C. lumaria. 8. Crataegus chrysocarpa Ashe var. vigintistamina J.B. Phipps, var. nov. Tr: MINNESOTA. Wabexa Co.: 1 mi NNW of Blue Grass, 6 Jun 1996, Welby R. Smith 26113 (nototvee: UWO). Comment.—The matching fruiting specimen, Smith 26424, is held in the unlisted herbarium of the Minnesota Dept. of Natural Resources, Minneapolis-St. Paul. A var. chrysocarpa differt in numero staminum, constanter ca. 20. Variety vigintistamina is the name to be given to specimens of Crataegus chrysocarpa only differentiable from the type variety by stamen number. Thus, there are 20 stamens and ivory to cream anthers. ANADA: Man., ?Ont.; U.S.A.: Mich., Minn., NY., Wisc., ?others. The new variety appears to be scat- tered throughout the eastern part of the range of C. chrysocarpa but it is poorly recorded. We have by far the most records from Minnesota. Variety vigintistamina has been confused with C. divergens (Peck) Sarg., sens. auctt. [New York State Mus. Bull. 105:66. 1906] and C. irrasa var. divergens Peck, sens. auctt. [New York State Mus. Bull. 75:51. 1904] which has a type from New York, Rensselaer County, North Greenbush but which, according to Peck and Sargent, has 10-18 stamens. The type of this variety (C.H. Peck 70) actually has 10 stamens and thus its name is not usable for the novelty named here. Nevertheless, divergens has been incorrectly utilized for the 20-stamen variety proposed here. Variety vigintistamina is also neither the same as the 20-stamen British Columbian C. sheila-phippsiae J.B. Phipps & O'Kennon with glabrous inflorescences and pink anthers nor the eastern species C. irrasa Sarg. and C. oakesiana Eggl. which have rather differently formed leaves. Leaves of the latter two are generally larger and more venous than in C. chrysocarpa varieties. Phipps, New combinations and typifications in North American Crat 1009 9. Crataegus chrysocarpa Ashe var. praecox (Sarg.) J.B. Phipps, comb. et stat. nov. Tver: NEW YORK. Essex Co.: Crown Point, 9 Sep 1900, E. Brainerd and C.S. Sargent s.n. (Lectotype designated here: A). Comments. —Brainerds name is not on the E ni and my a 6S. M um 17 Hee with hs correct label data can be dh located. However, it is assumed that the latter cited here as lectotype E E Crataegus praecox Sarg., Rhodora 3:27. 1901. Crataegus praecoqua Sarg., Rhodora 5:167. 1903, nom. illegit. The diagnostic characteristic of the new variety, pink anthers, has unfortunately not been possible to verify due to lack of a specimen. However, a currently unlocatable Brainerd and Sargent collection from Crown Point, New York, came from a plant said to have pink anthers by Sargent (op. cit. 1903). That it has pink anthers, while it is, therefore, just Sargent's memory, is specifically given as a correction in 1903 and must therefore be given credence. If it can ever be located, it would be worth consideration for an epitype as the fruiting lectoype lacks diagnostic information. Crataegus praecoqua was created as a replacement name for C. praecox when Sargent was following the American rules which would have treated his 1901 name as a homonym of C.oxyacantha var. praecox Loudon, 1838. The rules of the present code are, however, retroactive. Variety praecox is essentially the same as Crataegus chrysocarpa var. chrysocarpa, except with pink an- thers. Although the type of this variety comes from New York, forms of C. chrysocarpa with pink anthers are scattered throughout the range of the species, being especially frequent in Minnesota, and need a home. Minnesota material may have thorns 7-8 cm long. 10. Ew chrysocarpa Ashe var. blanchardii (Sarg.) J.B. Phipps, comb. et stat. nov. Crataegus blanchardii ., Rhodora 7:218. 1905. Type: VERMONT. WiNpHAM Co.: Deerfield River Valley, Wilmington, 26 Sep 1903, WW Eggleston 3452 cM A). t.—This number was designated as type in the protologue. VERMONT. WinbHam Co.: Deerfield River Valley, Wilmington, 20 May 1903, WW. E d dd 3451 Md: aee here: E 1 1 ymment.—Because a , itis essential on 1 J; F4] 1 zs zn JUR designate as epitype a UM specimen which has this character explicitly mentioned in gly still visible. Crataegus irrasa Sarg. var. blanchardii (Sarg.) Eggl., Rhodora 10:79. 1908. Variety blanchardii is like the typical variety, but with 20 pink anthers. The nutlets of the lectoype are plane- sided. Its range is from Quebec to Vermont and New York in the east and in Minnesota in the Midwest, where it is most abundant. series Virides (Loud.) Rehd. 11. Crataegus viridis L. var. velutina (Sarg.) J.B. Phipps, comb. et stat. nov. Crataegus velutina Sarg., Trees & Shrubs 1913. Type: ARKANSAS. HEMPSTEAD Co.: near Fulton, upland woods in dry soil, 25 Apr 1905, B.E Bush 10C om A). omment.—Type number is pP in protologue and duplicates with this label data are not known. Variety velutina is concentrated in western Louisiana, western Arkansas, and eastern Texas and is the only variety of Crataegus viridis y i ifi pubescence beyond the tufts of hair in the adaxial vein axils. This is sufficient for diss MN similar forms but less velutinous young are found in Florida where they have been named Crataegus subviridis. Apart from this Florida situation, var. velutina is easily recognized in young foliage and at anthesis, has a compact range, and has virtually no intermediates with the glabrous form of Crataegus viridis. Crataegus poliophylla Sarg. represents a red-anthered form of the variety. A strong tendency to glabrescence may explain the small percentage of fruiting specimens recognized for this variety. It is not yet clear whether the Florida plants referred to should be referred to variety velutina, var. viridis, or perhaps subviridis at a varietal rank. Crataegus viridis var. velutina might eventually be construed as a hairy form of C. viridis var. lanceolata. Suspicions that var. velutina might represent C. viridis xC. collina or C. viridis xC. berberifolia hybrids are annulled by their consistently small fruit size and entire-margined sepals. 1010 Journal of the Botanical R h Institute of Texas 1(2) ACKNOWLEDGMENTS Jessica Dolan, formerly on staff at HUH, is thanked for diligently searching out most of the syntype mate- rial on which this paper is based. Alan Weakley, NCU, efficiently sought out the type and other important material of C. neofluvialis. James Macklin, Director of Collections, HUH, enabled a valuable visit to HUH where the early observations were made. Arthur Haines, New England Botanical Club and Delta Institute, is thanked for information on certain critical points. This study was funded by a Discovery Grant from the National Scientific and Engineering Research Council of Canada and a microgrant for travel to Harvard University from the Flora of North America Association. REFERENCES KauscHkE, E.P. 1965. Contributions to the taxonomy of Crataegus. Milwaukee Public Mus. Publ. Bot. 3:1-273. Mackuin, J.A. and J.B. Phipps. 2002. Sargent's names in Crataegus series Coccineae (Rosaceae). Harvard Pap. Bot. 7125-53 KRUSCHKE NAMES IN NORTH AMERICAN CRATAEGUS (ROSACEAE) J.B. Phipps Ihe piedi of Western Ontario 1151 ras N London, Ontario, NGA 587, CANADA johippseuwo.ca ABSTRACT TES : lwitl q f E.P. Kruschke are typified: these are: C. apiomorpha Sarg., C. aboriginum Sarg., C cyanophylla Sarg., C. divida ae e T Ashe, C. gemmosa Sarg., C. integriloba Sarg., C. mollis (Torr. & A. Gray) Scheele var. incisifolia Kruschke, C. pertomentosa Ashe, C. pisifera Sarg., C. rugosa Ashe, C. schuettei Ashe var. gigantea Kruschke, C. virella Ashe RESUMEN Se tipifican t b d bi de E.P. Kruschke: éstas son: C. p d Sarg., C. aboriginum Sarg., C cyanophylla Sarg., C. divida Sarg., C. ferrissii Ashe, C. gemmosa Sarg., C. integriloba Sarg., is (Torr. & A. Gray) Scheele var. incisifolia Kruschke, C. pertomentosa Ashe, C. pisifera Sarg., C. rugosa Ashe, C. schuettei Ashe var. gigantea irri C. virella Ashe The late Wisconsin botanist Emil P. Kruschke (d. 1974) was very interested in Crataegus taxonomy in the middle of the last century and published a number of novelties in his “Contributions to the Taxonomy of Crataegus’ (Kruschke 1965). Kruschke's opus is a large and primarily nomenclatural work. It is his only taxonomic paper on Crataegus, but one that is widely cited. Kruschke's new taxa are mostly very local and from Wisconsin and Illinois some of which are probably interserial hybrids. The latter have been largely ignored in later works on the genus and none figure in this paper. He also described a number of local ‘for- mae’ which I am not using. However, he rationalized a lot of the existing taxonomy by reducing to varietal status about 20 early 20'^ century names mainly by Sargent and Ashe. These names, being of mostly rather wide-ranging and commonly accepted taxa, in many cases fairly easy to identify, consequently have ample currency in later literature. Unfortunately, Kruschke did everything correctly except precisely cite their types as he generally made no choice between his flowering and fruiting syntypes. Itis this oversight which is corrected here so as to validate those of his new combinations that will be used in FNA vol. 9 (Rosaceae). In one case, that of C. apiomorpha Sarg., it became necessary to lectotypify a species that is not a varietal basionym. The 13 validations made in this paper, which usually required lectotypification, follow hereunder and are arranged alphabetically by the name that Kruschke or Sargent used. SARGENT NAME 1. Crataegus apiomorpha Sarg., Bot. Gaz. 35:38. 1903. Wee: U.S.A. Ituwors. Cook Co.: Tinley Park, near Chicago, 3 Sep 1902, EJ. Hill 49C (LECTOTYPE i ien here: A). Comment.—The syntypes were a mixed a gathering and I accept Kruschkes (1965) suggestion about the type in here validating an entity with Eripe inflorescences. Crataegus apiomorpha is now definitively treated as a pubescent entity that is similar to C. macrosperma Ashe. According to Sargent (1903) it was common near Chicago. KRUSCHKE NAMES 2. Crataegus apiomorpha var. cyanophylla (Sarg.) Kruschke, Milwaukee Public Mus. Publ. Bot. 3:174. 965. Basionvm: Crataegus cyanophylla Sarg., Bot. Gaz. 35:387. 1903. Tyre: U.S.A. ILLINoIs. Will Co.: Joliet, 9 May 1902, H.C. Steele s.n. (LECTOTYPE designated here: A omment a r As with Crataegus apiomorpha, Sargent's syntypes of C. cyanophylla comprised both pubescent and glabrous J. Bot. Res. Inst. Texas 1(2): 1011 — 1014. 2007 1012 Journal of the Botanical R h Institute of Texas 1(2) inflorescence specimens. Most of the syntypes are of the glabrous kind and it is one of these that is se- lected for the cyanophylla lectotype. The leaf shape of this is, in any case, different from the pubescent form [represented by C. apiomorpha] and its young leaves are much less densely scabrous-pubescent. Crataegus cyanophylla was considered to be worth describing by Sargent because of its strongly blued-tinted (i.e., pre- sumably rather glaucous) leaves. However, this does not show on herbarium specimens and as such they cannot be differentiated from C. macrosperma. 3. Crataegus ion Ashe var. aboriginum (Sarg.) Kruschke, Milwaukee Public Mus. Publ. Bot. 3:164. 1965. Bastonyw: Crataegus aboriginum Md Rhodora 5:163. 1903. Type: CANADA. Querrec. La Prairie Co.: Caughnawaga, 27 May 1900, J.G. Jack 45 (LectotYPE designated here: A). omments.—The choice of lectotype is, cm a flowering specimen still in bud and is selected to illustrate the very conspicu- ous and abundant large bracteoles which have stipitate marginal glands. CANADA: Quebec: La Prairie Co.: Caughnawaga, 30 May 1900, J.G. Jack 45 (epityPe designated here: A). Comment.—An epitype is needed to show well-developed floral features that are more familiar to identifiers of hawthorns; however by full anthesis it is very difficult to locate any of the distinctive caducous bracteoles. Crataegus aboriginum, now that it has been critically examined for typification, is seen to have a completely glabrous inflorescence (in contrast to some later usage) and conspicuously glandular bracteole margins and thus can no longer be treated as part of the C. chrysocarpa complex in FNA. Rather, it belongs to the C. dodgei complex. Crataegus aboriginum has been generally construed as the glabrous hypanthium form of eastern C. chrysocarpa with thinly pubescent pedicels but this is clearly wrong. 4. eser Mp Is Lodd. ex Loud. var. pertomentosa (Ashe) Kruschke, Milwaukee Public Mus. Publ. Bot. 3:161. 1965. Basioxvw: gee per Md nae Pi Elisha Mitchell Sci. Soc. 16:70. 1900. Tyre: U.S.A. MISSOURI. Jackson Co.: al 4 Oct 1906, B.E d here: MO) Comments. —This variety was abe p Co Ta but I t locat Ashe putati type. Dodson is now part of Kansas City, M Variety pertomentosa is distinguished from other forms of C. macracantha by its strong abaxial leaf pubescence and its distribution is mapped, in part, in McGregor and Barkley (1977). Recently (Sep 2007), I have col- lected entirely typical material of this in the Judith Mtns., MT. I was impressed by its thin and also smallish leaves. It seems to me to be one of the better-marked forms of C. macracantha. 5. Crataegus macracantha um ex idm var. divida (Sarg.) Kruschke, Milwaukee Public Mus. Publ. Bot. 3:161. 1965. Basowvw: Crataeg Sarg., Bot. Gaz. 35:401. 1903. Type: U.S.A. Inuwors. Lake Co.: near Barrington, 22 May 1901, EJ. Hill 38A (Lec designated here: D Comment.—a a fruiting synype reads nes northeast of Barrington” which is wl its the deducti fth ty Gwhich is hel 1s “clay hills, nr. Barrington,” eee certainly, however, the same area. Otherwise LOL Or thle laDEL J the lectotype would be in Cook Co. Crataegus divida seems to be similar to characteristic forms of C. macracantha except that it has glabrous inflorescences, thin leaves and ‘hard’ fruit. 6. Crataegus macracantha Lodd. ex pou var. a (Sarg.) Kruschke, Milwaukee Public Mus. Publ. Bot. 3:161. 1965. Bastonym: C i g., Rhodora 3:78. 1901. Type: CANADA. Quesec. La Prairie Co.: Caughnawaga, 29 Aug 1899, J.G. Jack 28 (LECTOTYPE due her Comment.—A flowering lectotype would have been preferable but instead a fruiting specimen has to be used as there is not a suit- able flowering specimen ME De DD CANADA: Quebec: La Prairie Co.: Adirondack Jet., 30 May 1900, J.G. Jack 44 (EPITYPE designated here: A). Comment. E E lu E E o o de [2] characteristic. I. Mcr mollis (Torr. & A. Gray) Scheele var. incisifolia Kruschke, Milwaukee Public Mus. Publ. Bot. 65. Type: U.S.A. Wisconsin. Rock Co.: Avon, along Sugar River, 27 May 1947, E.P Kruschke K-47-8 (Lectotype designated ee um duplicate at A). Comment.—Kruschke cited two syntypes as the type, from May and September 1947, respectively. This variety is notable within C. mollis for its deeply and sharply incised leaves. 8. Crataegus pruinosa (H.L. Wendl.) K. Koch var. rugosa (Ashe) Kruschke, Milwaukee Public Mus. Publ. Phipps T O £V hi *oMartkh A e Font 1013 Pro (YP J Bot. 3:181. 1965. Basowvw: Crataegus rugosa Ashe, J. Elisha Mitchell Sci. Soc. 17:5. 1900. Tver: U.S.A. NorTH CAROLINA. Ashe Co.: 26-28 Sep 1908, WW. Eggleston 4171 (NeotyPE designated here: NCU). Comment.—Ashes protologue says Ashe and Watauga Cos. and northward, NC, but tl ining | i with the name ee rugosa in Ashe’ hand are much less specific. The Eggleston neotype was selected as it was the ae NCU specimen seen from the type area and time frame that was a close match for the protologue. Twenty stamen-bases can be clearly seen.. This variety is distinguished by its usually broad leaves, many + truncate at the base. 9. emn pt (H.L. Wendl.) K. Koch var. virella (Ashe) Kruschke, Milwaukee Public Mus. Publ. 1965. Basionym: Crataegus virella Ashe, Ann. Carnegie Mus. 1:396. 1902. Tyre: U.S.A. PENNSYLVANIA. Berks C Kutztown, id 1902, C.L. Gruber s.n. (Lectotype designated here: PH 548493). Comments.—According to the Ashe ie d the pue i bea bed & Ashe collection but the lectotype d t say this on this label. The lectotypes ] I g ] ite d ] rough-hairy | idvein abaxially. in agreement with the protologue. This taxon is unique among the pruinosa complex by virtue of its conspicuously hairy foliage abaxially, at least young. It is also likely that forms with other, for instance, adaxial, pubescence and, possibly, inflo- rescence branch pubescence should be placed here. Other than this pubescence, C. virella appears to be a typical member of the pruinosa group. 10. Crataegus schuettei Ashe var. ferrissii (Ashe) Kruschke, Milwaukee Public Mus. Publ. Bot. 3:176. 1965. Basionvm: Crataegus ferrissii Ashe, J. Elisha Mitchell Sci. Soc. 17:11. 1901. Tyre: U.S.A. ILLINOIS: Lake Co.: northern Illinois, Forest Grove, May 1902, W.C. Egan s.n. (NEoTyPE designated here: A). This entity is distinguished from other forms of C. schuetttei by its fairly deeply divided leaf-margins wherein the more distal lobes point outwards at an angle and the proximal lobes are often held almost at right angles, rather as in C. stolonifera Sarg., and by the strongly glandular-serrate sepals. It is the most distinct form of this species. 11. Crataegus schuettei Ashe var. gigantea Kruschke, Milwaukee Public Mus. Publ. Bot. 3:75. 1965. Tre: U.S.A. WISCONSIN: City of Milwaukee, north side, 16 Sep 1942, E.P Kruschke K-42-288 (Lectotyee designated here: MIL; ISOLECTOTYPE: A) This variety was distinguished by its particularly large flowers and fruit. 12. copas succulenta Schrad. ex Link var. gemmosa (Sarg.) Kruschke, Milwaukee Public Mus. Publ. t. 3:159. 1965. Basioxvw: Crataegus gemmosa Sarg., Bot. Gaz. 33:119. 1902. Tree: CANADA. Ontario. Middlesex Co.: London, 22 s EOS CS. Sue 15 (ectotyee designated here: A). Comment. gemmosa, several in fruit ] the | is sel lf isplavi hel size fruit, ca. 12-15 mm diam. when n which is what is typical for C. gemmosa E to the uU It is distinguished from other forms of C. succulenta by its larger fruit; otherwise it is a typical C. succulenta type with 20 small rose anthers. 13. Crataegus succulenta Schrad. ex Link var. pisifera (Sarg.) Kruschke, Milwaukee Public Mus. Publ. Bot. 3:159. 1965. Basionwm: Crataegus pisifera Sarg., Rhodora 7:163. 1905. Tyre: U.S.A. Vermont. Addison Co.: Cornwall, 26 Jul 1901, E. Brainerd 15d (Lectotype designated here: A). Comment.—Sargent designated the type number in his protologue. This taxon was separated primarily on the basis of its very small fruit but while this size does obtain in the lectotype that specimen has immature fruit which may well have become larger as in some other syntypes. Thus, perhaps this differentium is not as reliable as earlier believed. However, in spite of this, syntype flowering material of var. pisifera is also differentiated from other forms of C. succulenta by a distinctive and unusual form of marginal lobing to the leaves. ACKNOWLEDGMENTS The curators and staff of HUH, MO, NCU and PH are thanked for their careful searches for relevant speci- mens. Kanchi Gandhi, Harvard University, is thanked for nomenclatural advice. Timothy A. Dickinson and an anonymous reviewer made helpful comments. 1014 Journal of the Botanical R h Institute of Texas 1(2) REFERENCE KRuscHkE, E.P. 1965. Contributions to the taxonomy of Crataegus. Milwaukee Public Mus. Publ. Bot. 3:1-273. McGREGOR, R.L. and T.M. Barkley. 1977. Atlas of the flora of the Great Plains. Ilowa State University Press, Ames. NEW COMBINATIONS IN DUDLEYA (CRASSULACEAE) AND RIBES (GROSSULARIACEAE) Nancy R. Morin Flora of North America P.O. Box 716 Point Arena, California 95468, U.S.A. The following new combinations are required for the treatment of Dudleya by Reid. Moran (in prep) for Flora of North America. The provision of nomenclature here simply implements Dr. Moran's taxonomy and the combination should be credited to him, with citation as Moran *in" Morin. Dudleya abramsii Rose subsp. costifolia [originally costafolia] (Bartel & Shevock) Moran, comb. nov. Ba- sionyM: Dudleya cymosa (Lem.) Britton & Rose subsp. costifolia Bartel & Shevock, Aliso 12:701. 1990. Dudleya costifolia (Bartel & Shevock) PH. Thomson. Common name.—Pierpoint Springs liveforever. Dudleya abramsii subsp. costifolia is known only from one southwest-facing outcrop of pre-Cretaceous limestone just west of Pierpoint Springs, Tulare County, California, in the southern Sierra Nevada, in an area that has been subdivided. It forms clumps to 10 cm wide. The bright yellow corolla and more-branching cyme and sometimes longer pedicels set subsp. costifolia apart from the other subspecies of Dudleya abramsii and show an approach toward D. cymosa, where the original authors placed it. However, the cespitose small rosettes of narrow leaves have much more the aspect of D. abramsii, especially recalling the lowland subsp. bettinae; the higher insertion of the antesepalous stamens is a mark of D. abramsii, (and the related D. parva) in contrast to D. cymosa and other species. The branching cyme is somewhat approached in subsp. calcicola, which grows just to the north. Dudleya abramsii Rose subsp. setchellii (Jeps.) Moran, OE NOV. Basionym: Cotyledon laxa (Lindl.) S. Watson var. setchellii Jeps., Fl. W. Calif. 267. 1901. Dudleya cymosa (Lem.) B bsp. setchellii (Jeps.) Moran. Dudleya setchellii (Jeps.) Britton & Rose. Common name.—Santa Clara Valley dudleya. Dudleya abramsii subsp. setchellii is known from only a score of places in the Coyote Valley area, Santa Clara County, where it is considered threatened. Although it was formerly placed with Dudleya cymosa, it clearly is closer to D. abramsii and especially to subsp. murina, which likewise grows on serpentine and at about the same elevation but some 200 km to the south. Subspecies setchellii tends to be smaller, with petals unmarked and less sharply acute. Infraspecific taxa in Ribes oxyacanthoides L. will be recognized as varieties in the forthcoming treatment of Ribes in Flora of North America (N. Morin, in prep.). The following combination is required. Ribes oxyacanthoides L. var. cognatum (Greene) Morin, comb. et stat. nov. Basionvm: Ribes cognatum Greene, Pittonia 15. 1896. Grossularia cognata (Greene) Coville & Britton. Ribes oxyacanthoides L. subsp. cognatum (Greene) Sinnott. ACKNOWLEDGMENT Thanks to Guy Nesom and Barney Lipscomb for prompt and helpful comments. REFERENCES Moran, R. In prep. Dudleya. Crassulaceae. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America north of Mexico. 14+ vols. New York and Oxford. Vol. 8. J. Bot. Res. Inst. Texas 1(2): 1015 — 1016. 2007 1016 Journal of the Botanical R h Institute of Texas 1(2) MoRN, N.R. In prep. Grossulariaceae. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America north of Mexico. 14+ vols. New York and Oxford. Vol. 8. NEW COMBINATIONS IN LIMNANTHES (LIMNANTHACEAE) Nancy R. Morin Flora of North America P O. Box 716 Point Arena, California 95468, U.S.A. The following nomenclatural changes are required for the treatment of Limnanthes (Limnanthaceae) in Flora of North America (Morin, in prep.) and revision of The Jepson Manual (Morin, in prep.). Limnanthes alba Hartw. ex Benth. eae gracilis (Howell) Morin, comb. et stat. nov. Basionvm: Limnanthes gracilis Howell, Fl. N.W. Amer. 1:108. 1897 Limnanthes alba Hartw. ex Benth. subsp. pem (Jeps.) Morin, comb. et stat. nov. Basioxvw: Limnanthes versicolor (Greene) Rydb. var. parishii Jeps., Fl. Calif. 2:412. 1936 Allozyme studies of Limnanthes section Inflexae by McNeill and Jain (1983), and subsequent analysis of their data by Plotkin (1998), suggested that Limnanthes gracilis subsp. gracilis and L. gracilis subsp. parishii are more closely related to Limnanthes alba than they are to each other. Jepson originally placed parishii as a variety of Limnanthes versicolor (Greene) Rydb., which is currently treated as a subspecies of L. alba. Limnanthes douglasii R. Br. subsp. striata (Jeps.) Morin, comb. et stat. nov. Basionym: Limnanthes striata Jeps., Fl. Calif. 2:411. 1936 Mason (1952) suggested that Li thes striata Jeps. was closely related to L. douglasii R. Br. var. douglasii and that its status as a species might need reevaluation. ITS analysis by Plotkin (1998) placed L. striata within the L. douglasii clade. Limnanthes striata has been distinguished from L douglasii mainly by having stamens 2-4 mm versus 5-8 mm (Ornduff 1993); its distribution is in the central foothills of the Sierra Nevada, California, adjacent to L. douglasii subsp. rosea (Benth.) C.T. Mason in the Central Valley. Limnanthes macounii Trel. was also placed in the L. douglasii clade (Plotkin 1998), but this taxon is morphologically very distinct (flowers 4-merous rather than 5-merous) and has a completely distinct native distribution (southern Van- couver Island and some adjacent islands, British Columbia). It is known from very few populations; more studies should be done before any change in rank or placement is made. ACKNOWLEDGMENT I thank Guy Nesom and Barney Lipscomb looking over these new combinations. REFERENCES Mason, C.T., Jr. 1952. A systematic study of the genus Limnanthes R. Br. Univ. Calif. Pub. Bot. 25:455-507. McNett, C.l. and S.K. Jain. 1983. Genetic differentiation studies and phylogenetic inference in the plant genus Limnanthes (section Inflexae). Theor. Appl. Genet. 66:257-269. MoRN, N. In Prep. Limnanthaceae. In: The Jepson manual: higher plants of California, revised. University of California Press. MoRN, N. In Prep. Limnanthes, Limnanthaceae. In: Flora of North America north of Mexico. 14+ vols. New York and Oxford. Vol. 13 OnNpure, R. 1993. Limnanthaceae. In: J.C. Hickman, ed., The Jepson manual: higher plants of California. University of California Press. Pp. 736-738. PLOTKIN, M. 1998. Phylogeny and biogeography of Limnanthaceae. M.S. Thesis. University of California, Davis. J. Bot. Res. Inst. Texas 1(2): 1017. 2007 1018 Journal of the Botanical R h Institute of Texas 1(2) BOOK NOTICES STEPHEN G. PALLARDY. 2008. Physiology of Woody Plants (ed. 3). (ISBN 978-0-12-088765-1, hbk.). Elsevier- Academic Press, 30 Corporate Drive, Suite 400, Burlington, MA 01803, U.S.A. (Orders: www.elsevier. com) $136.00, 494 pp- illus, Sep Revised from the 1997 a It is “ led | f ] laref f ] e this eds book should 1 ful f scientists f foresters, agronomists, and arl and soil scientists." A pet peared RA n organization of nd layout, and ET xb. ree reading thr Gag the ] first three m it’s hard t ble of Contents. XC The MO. plant body. Vegetative growth. R ducti th. Photosynthesis. Enzymes, en- ergetics, and respiration. Carbohydrates. Lipids, terpenes, and related substances. Men Se ee Mineral nutrition. Absorption of water and ascent of sap. Transpiration and plant water balance. Plant hormones and other growth regulators. Chapter summaries. General references. Scientific name index. Bibliography. Subject index. Gary Lyons. 2007. Desert Plants: A Curator’s Introduction to the Huntington Botanical Gardens. (ISBN 978-0-87328-231-4, pbk.). Huntington Library Press, 1151 Oxford Road, San Marino, CA 91108, U.S.A. (Orders: California Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A. www.ucpress.edu, 609-883-1759, Fax 609-883-7413). $24.95, 127 pages, color and b/w photos, BS 2 TO The HBG includes “12 acres of plants, rockery, and paths,” begun in 1907, to b f the larg llecti f cacti and l in the world. The book is a quick, easy read with lots of color photos. *A Brief History of the Huntington Desert Garden (documented by B&W photos) *Desert Plants of the Old World (The Aloe Family; Euphorbias; The Crassula Family; Cyphostemma and Dioscorea, The Dragon Tree; Stapelias; Ice Plants; Senecios, or Succulent Sunflowers) *Desert Plants of the New World (New World Crassulas; Cacti; Terrestrial Bromeliads; Ocotillo; Succulents of Baja California; Agave and Agave Relatives) *The World of the Desert Garden (Care of the Desert Garden; Aridity and Survival; Protecting Plant Diversity) PHiLiPPA NIKULINSKY and STEPHEN D Hopper. 2005 [reprinted 2007]. Soul of the Desert. (ISBN 978-1-921064- -7, hbk.). Fremantle Arts Centre Press, 25 Quarry Street, Fremantle, PO Box 158, North Fremantle 6159, Western Australia. (Orders: International Specialized Book Services, 920 NE 58th Avenue, Suite 300, Portland, OR 97213-3786; 503-287-3093; 503-280-8832 fax; info@Isbs.com; www.isbs.com). $54.00, 176 pp., beautiful color paintings, 9" x 11 1/4". Primarily a vehicle for Philippa Nikulinsky's exquisite paintings of the Australian desert but accompanied by brief but very interest- o commentaries 2» PPD ES Oh, if only the artist will give the the next version of the book some landscape examples, in her style of I. Deserts Though Ti Time (Origins; Relati logical age of dseerts; Desert people; European impacts; Soul of A evelopment of A Painting (a brief but eerie narrative, with stages of drawings). III. The Artwork (Ranges and outcrops; Plains; Dunes; Wetlands; Widespread plants and animals; Diversity explored). J. Bot. Res. Inst. Texas 1(2): 1018. 2007 NEW COMBINATIONS IN MICRANTHES (A SEGREGATE OF SAXIFRAGA, SAXIFRAGACEAE) IN NORTH AMERICA Luc Brouillet Richard Gornall Institut de recherche ne MT vegetale mos i biology rsite de Montreal Viversity of Leicester fe noe ur pono Road NOS Quebec, CANADA H1X 2B2 Leicester, LE1 ZRH, U.K. ABSTRACT Molecular phyl i ] ] ly sl hat Saxifi ] t be divided into Saxifraga s. str. and Micranthes. Within the c framework of preparing the Micranthes treatment for the Flora of North America, the species of North American Saxifraga that belong to Micranthes and have yet to be transferred to the ee RESUMEN I lisis filog lecul l ] jue Saxifraga s. 1. debe dividirse en Es s. str. y Micranthes. En el d ión del i le Mi } la Flora of North America las especies de Saxifrag i jue pert Micranthes y aún tienen que t a este género. INTRODUCTION Recent phylogenetic studies (summarized in Soltis et al. 2001) have shown that Saxifraga, as currently un- derstood, must be divided into two genera: Saxifraga L. sensu stricto and Micranthes Haw. Micranthes differs from Saxifraga in having usually leafless (vs leaty) flowering stems, reticulate (vs striate or granular) pollen exine, carpels usually connate less than half (vs more than half) their lengths, ovules with a single (vs two) integuments, and | ribbed (vs. smooth, tuberculate or papillate) seeds (Gornall and Ohba, ined.). In these ES Saxifraga tolmiei, one of two species of section Merkianae, falls at the base of the Micranthes clade and appears to belong to this genus. Webb and Gornall (1989) provide a list of Saxifraga sensu lato species by section for Europe, parts of Asia, and North America, in which section Merkianae is segregated from section Micranthes on the basis of a lack of crystals in leaves and bitegminate ovules. In Russia, Zhmylev (1996, 1997) recognizes Micranthes at the subgeneric level, with two sections, Merkianae and Micranthes, the latter with 6 subsections. This author thus recognized the link between sections Merki- anae and Micranthes but did not acknowledge the distinctiveness of Micranthes. Recently, Banfi et al. (2005) recognize Micranthes and made new combinations for the Italian species. In North America, J.K. Small (in Small and Rydberg 1905, 1918) made combinations for a number of the taxa that belong to Micranthes. Not all North American taxa were given names in this genus, however, since Small also recognized other segregate genera such as Spatularia, the species of which belong to Micranthes, or Leptasea, which included some species related to Micranthes, notably Leptasea (Saxifraga) tolmiei. Fur- thermore, a few species have been described since (Elvander & Denton 1976; Elvander 1984; Skelly 1988). The treatment of Saxifraga for the Flora of North America project prepared by the late Patrick E. Elvander (unpublished manuscript) did not segregate Micranthes, but provided a list of recognized North American taxa, except for the taxa recently described by Zhmylev (1990, 1992, 1993), which were not dealt with. AII species not listed below belong to Saxifraga sensu stricto. The sole exception is Cascadia nuttallii (Small) A.M. Johnson (syn. Saxifraga nuttallii Small), shown to be sister to the South American genus Saxifragodes in phylogenetic analyses, both in turn being the sister group to Micranthes (e.g., Soltis et al. 2001). Cascadia is distinguished by its trailing, leafy stems that do not arise from persistent basal rosettes, its distinct carpels each independently adnate to the hypanthium, its bitegminate ovules, and its spiny seeds. Our goal is to provide the necessary combinations for the North American taxa that have not yet been J. Bot. Res. Inst. Texas 1(2): 1019 — 1022. 2007 1020 f the Bot Journal UI titute of Texas 1(2) treated within Micranthes. Combinations for Eurasian taxa also present into North America were suggested by Gornall and Ohba (ined.) but have yet to be published. Given the necessity of having these names pub- lished, they are reported here as new combinations and are ascribed to these authors (to be cited as “Gornall & H. Ohba in Brouillet & Gornall"). In their work, they accept the division into two genera and provide new combinations for Eurasian taxa. Table 1 provides the list of names for North American taxa available in the literature. NEW COMBINATIONS Micranthes bryophora (A. Gray) Brouillet & Gornall, comb. nov. Saxifraga bryophora A. Gray, Proc. Amer. Acad. Arts 533. 1865. Type: U.S.A. CALIFORNIA: Ebbetts Pass, WH. Brewer 1984 (Lectotype: GH; ISOLECTOTYPE: US!). Saxifraga bryophora A. Gray var. tobiasae J. W. Grimes & PL. Packard; Spatularia bryophora (A. Gray) Small Micranthes calycina (Sternb.) Gornall & H. Ohba, comb. nov. Saxifraga calycina Sternb., Rev. Saxifr. Suppl. 2:10 1831. Micranthes ferruginea (Graham) Brouillet & Gornall, comb. nov. Saxifraga ferruginea Graham, Edinb. N. Phil. J. 7:349. 29. Type: plants grown at Edinburgh from seed sent by Richardson in 1827 and flowered in 1829 (HoLotyPE: E). xaphoma ferruginea Rat. axifraga ferruginea Graham var. macounii Engl. & Irmsch. Saxifraga newcombei (Small) Engl. & Irmsch. Spatularia newcombei Small Spatularia ferruginea (Graham) Small Micranthes foliolosa (R. Br.) Gornall, comb. nov. Saxifraga foliolosa R. Br., Chlor. Melvill. 17. 1823. Tyee: Parry’ first voyage 1819-20, Cap. Nels., Melville Island, Mr James Ross (HOLOTYPE: BM). Micranthes gormanii (Suksd.) Brouillet & Gornall, comb. nov. Saxifraga gormanii Suksd., Torreya 23:106. 1923. Tver: OREGON. Multnomah Co.: Elk Rock, Gorman 4081 (HoLoTyPE: WS!) Saxifraga occidentalis var. dentata (Engl. & Irmsch.) C.L. Hitchc. Micranthes idahoensis (Piper) Brouillet @ Gornall, comb. nov. Saxifraga idahoensis Piper, Bull. Torrey Bot. Club 27:394. 1900. Type: U.S.A. IDAHO. Latah Co.: Kendrick, Henderson 4588 (HOLOTYPE: GH; IsoTYPE: NY) Saxifraga marshallii Greene ssp. idahoensis (Piper) D.L. Krause & Beamish Saxifraga marshallii var. idahoensis (Piper) Engl. & Irmsch. Saxifraga occidentalis S. Watson var. idahoensis (Piper) C.L. Hitchc. Micranthes mexicana (Engl. € Irmsch.) Brouillet & Gornall, comb. nov. Saxifrag icana Engl. & Irmsch., Pflanzenr 67 (IV. 117):42. 1916. Tyee: MEXICO. CHIHUAHUA: near Colonia Garcia, alt. 7500 ft, Townsend & Barber 88 (LEC designated by Webb & Gornall 1989: K). Mcr es nelsoniana (D. Don) Small var. aestivalis (Fisch. & C.A. Mey) Gornall & H. Ohba, comb. et nov. Saxifraga aestivalis Fisch. & C.A. Mey., Ind. Sem. Hort. Bot. Peteropl. 1:37. 1835. Type: RUSSIA. SisERIA: Altai, 1817, Be Fischer s.n. (LECTOTYPE: LE). Mr nelsoniana (D. Don) Small var. carlottae (Calder & Savile) Gornall & H. Ohba, comb. et stat. axifraga punctata Willd. subsp. carlottae Calder & Savile, Canad 2 Islands, Moresby Island, Ca . J. Bot. 38:423. 1960. Type: CANADA. BRITISH COLUMBIA: Queen or 23093 (HOLOTYPE: DAO). der, Savile and Tay ina nelsoniana (D. Don) Small var. cascadensis (Calder & Savile) Gornall & H. Ohba, comb. et stat. OV. Saxifraga punctata Willd. subsp. cascadensis Calder & Savile, Canad. J. Bot. 38:425. 1960. Type: U.S.A. WASHINGTON. ae Co.: Mt. Paddo (Mt. Adams), Suksdorf 2504 (uoLotyee: NY). Micranthes nelsoniana (D. Don) Small var. insularis (Hultén) Gornall @ H. Ohba, comb. et stat. nov. Saxifraga punctata Willd. subsp. insularis Hultén in Sven. Bot. Tidskr. 30:524. 1936. Type: U.S.A. ALEUTIAN IstANDs: Carlisle Island E. Hultén 6591 (HOLOTYPE: S). Micranthes nelsoniana (D. Don) Small var. pacifica (Hultén) Gornall @ H. Ohba, comb. et stat. nov. Saxifraga punctata Willd. subsp. pacifica Hultén, Fl. Alaska & Yukon 5:928. 1945. Type: U.S.A. Alaska: Juneau, H. Anderson 47 (synTYPE: H). HM [^H $ sn AA: n , TABLE 1021 1. List of Micranthes names already published for North America. Micranthes Small = rufidula Micranthes aestivalis (F. & M.) Small = nelsoniana var. aesti- valis Micranthes allen all identali Micranthes nea (Piper) Small Micranthes aprica (Greene) Small Micranthes arguta (D. Don) Small 2 odontoloma Micranthes arnoglossa Small 2 oregana Micranthes bidens Small = integrifolia Micranthes brachypus Small = oregana Mi nudi TD Small A. Gray) Small a (A. peche Small anby) Small = fragosa Me columbiana [s Small = nidifica mall 2 rhomboidea Micranthes davurica TO Small is misapplied in North America th ; > I M + > Mirranth / in ti phora (S. Watson) Small Micranthes fallax (Greene) Small = californica Micranthes flabellifolia (R. Br.) Small 2 unalaschensis, misap- plied in North America Micranthes fragosa (Suksd.) Small Micranthes franciscana Small 2 rhomboidea Micranthes galacifolia Small = spicata Micranthes gaspensis (Fernald) Small Micranthes geum (L) Small: not in North America Micranthes greenei Blankinship = rhomboidea Micranthes hieraciifolia (Waldst. & Kit.) Haw. Micranthes howellii (Greene) Small Micranthes integrifolia (Hooker) Small Micranthes interrupta Small 2 rhomboidea Micranthes kumlienii Small = nivalis Micranthes lata Small = occidentalis Mr lyallii (Engler) Small Mi es marshallii (Greene) Small d micranthidifolia (Haw.) Small Micranthes montana Small = nidifica Mis itn montanensis (Small) Small 2 oregana is (Small) Small = californica niana (D. Don) Small Micranthes nidifica (Greene) Small Micranthes nivalis (L.) Small Micranthes occidentalis (S. Watson) Small Micranthes odontoloma (Piper) A. A. Heller Micranthes ibd a Small Micranthes palme Micranthes d (cn Small = californica Micranthes pennsylvanica (L.) Haw. Micranthes plantaginea (Small) Small = nidifica Micranthes pilosa (Haw.) Bush = virginiensis Micranthes reflexa (Hooker) Small Micranthes icine (Greene) Small Micranthes rufidula Small Micranthes ii quos (Small Small 2 subapeta nthes saximontana (E. Nelson) m z: m. recap sierrae (Coville) A. Heller 2 oregana Micranthes spicata (D.Don) Small Micranthes stellaris (L.) Galasso, Banfi & Soldano Micranthes subapetala (E. Nelson) Small Micranthes tenuis (Wahl.) Small Micranthes tennesseensis (Small) Small = careyana Micranthes texana (Buckl.) Small Micranthes virginiensis d Small Saxifraga virginiensis Michx. var. subintegra Goodman Micranthes yukonensis tone = reflexa co YE Micranthes nudicaulis (D. Don) Gornall & H. Ohba, comb. nov. Saxifraga nudicaulis D. Don, Trans. Linn. Soc. 13:366. 1822. Tyre: [Alaska]: “ LOTYPE: BM). ad oras occidentales Americae borealis” Northwest coast of America [Alaska], Sledge Island, D. Nelson (Ho- Micranthes petiolaris (Raf.) Brouillet & Gornall, comb. nov. Hexaphoma petiolaris Raf., Fl. Tell. 2:67. 1836. Tre: not found. Saxifraga michauxii Britton Saxifraga leucanthemifolia Michx. 1804, not Lapeyrouse 1803 Spatularia leucanthemifolia Haw. Spatularia petiolaris (Raf.) Small Spatularia michauxii (Britton) Small Micranthes razshivinii (P. Yu. Zhmylev) Brouillet & Gornall, comb. nov. Saxifraga razshivinii Zhmylev, Byull. Mosk. Ob- ch. Ispyt. Prir., Biol. 95(3):87. 1990. Type: CANADA. Yukon: Mt. Sedgwick area, British Mts., J.A. Calder 34478 (HoLotyPE: DAO). Saxifraga davurica Willd. subsp. grandipetala (Engl. & Irmsch.) Hultén Micranthes tempestiva (Elvander & Denton) Brouillet & Gornall, comb. nov. Saxifraga tempestiva Elvander € Den- ton, Madroño 23:346 isotypes: NY!, UC!). . 1976. Tyre: U.S.A. MONTANA. Deerlodge Co.: Anaconda Range, Goat Flat, Elvander 492 (HoLotype: WTU!; Micranthes tischii (Skelly) Brouillet & Gornall, comb. nov. Saxifraga tischii Skelly, Madroño 35:126-131. 1988. Tyre: U.S.A. WASHINGTON. Clallam Co.: Olympic Mountains, Mt. Baldy, Tisch 2307 (HoLoTYPE: UC!). 1022 Journal of the Botanical R h Institute of Texas 1(2) Micranthes tolmiei (Torrey & A. Gray) Brouillet & Gornall, comb. nov. Saxifraga tolmiei Torrey & A. Gray, Fl. N. Amer. 1:582. 1840. Tre: U.S.A.: Northwest coast, 1838, Tolmie s.n. (LECTOTYPE: GH). Micranthes unalaschcensis (Sternb.) Gornall & H. Ohba, comb. nov. Saxifraga unalaschcensis Sternb., Rev. Saxifrag., suppl. 2:9. 1831. Tyre: In insula Unalaschka, Chamisso (HOLOTYPE: LE). ACKNOWLEDGMENTS The authors would like to thank Guy Nesom and Barney Lipscomb for promptly reviewing this paper. REFERENCES BANH, E., G. Galasso, and A. SoLDANO, 2005. Notes on the systematics and taxonomy for the Italian vascular flora. 1. Atti Soc. It. Sci. Nat. Museo Civ. Stor, Nat. Milano 146(2):219-244. ELVANDER, PE. and M.F. Denton. 1976. Saxifraga tempestiva (Saxifragaceae): a new species from the Pacific North- west. Madroño 23:346-354. Gervais, C., N. Dicnarb, and R. TRAHAN. 1995. The chromosome number of Saxifraga gaspensis Fernald. Rhodora 97:171-175. McGREGoR, M. and W. HARDING. 1998. Saxifrages. The complete list of species. The Saxifrage Society. Hutton, U.K. SkeLLY, RJ. 1988. A new species of Saxifraga (Saxifragaceae) from the Olympic Mountains, Washington, and Vancouver Island, British Columbia. Madroño 35:126-131. SMALL, J.K. and PA. RvogERG. 1905. Saxifragaceae. North American flora 22(2):81-158. SMALL, J.K. and PA. Rypserc. 1918. Saxifragaceae. Additions and corrections. North American flora 22(6):549- Souris, D.E., R.K. Kuzorr, M.E. Mort, M. Zawis, M. FisHBEIN, L. HUFFORD, J. Koontz, and M.K. Arroyo. 2001. Elucidating deep-level phylogenetic relationships in Saxifragaceae using sequences for six chloroplastic and nuclear DNA regions. Ann. Missouri Bot. Gard. 88:669-693. Wess, D.A. and R.J. GORNALL. 1989. A manual of Saxifrages. Timber Press, Portland, Oregon. ZHMYLEV, PY. 1990. A new Saxifraga L. from Yukon (in Russian). Biull. Mosk. Obsch. Ispyt. Prir., Biol. 95(3):87-91. ZHmvuev, PY. 1996. Rockfoils of the subgenus Micranthes (Haw.) H.G.L. Reichenbach (Saxifraga L., Saxifragaceae) (in Russian). Biull. Mosk. Obsch. Ispyt. Prir., Biol. 101(6):67—77. ZHMYLEV, PY. 1997. Systematical review of Rockfoils (Saxifraga L.) of Russia and contiguous territories: subgenera Micranthes, Diptera and Hirculus [in Russian]. Biull. Mosk. Obsch. Ispyt. Prir., Biol. 102(3):46-51. M —— NEW COMBINATIONS AND NEW SYNONYMIES IN THE GENUS SPERMACOCE (RUBIACEAE) FOR THE FLORA OF GOIÁS AND TOCANTINS (BRAZIL) AND THE FLORA OF THE GUIANAS Piero G. Delprete Federal MU. of Em Campus Il Institute Jical Sciences - ICB-1 pecaninent A General Biology/Botany 74001-970 Goiânia, Goiás, BRAZIL pdelpreteahotmail.com ABSTRACT The pun Pasos has jud Mese d circumscribed 2) Rubi pecialists, including or excluding Borreria and/or other related taxa. N 1 idely deli POMA CoU ERU EIS oj : Sel g pt fg morphology, anatomy, palynology, and re phylogenies. Accordingly, twelve new binati and several new synonymies in € 1 3 1 1 TTD f, 1 Tus f 4l Flora de Goiás e Tocantins, Flora do Distrito Federal, and the Flora of the Guianas. Key Worbs: Borreria, Spermacoce, Rubiaceae, Goiás, Tocantins, Brazil, Guianas, Neotropics RESUMEN O género Spermacoce foi variavelmente circunscrito por ialistas de Rubiaceae, incluíndo ou excluíndo Borreria e/ou outros taxa relacionados. Estudos E b iis é melhor tratar Spermacoce como amplamente delimitado, incluíndo Borreria, sendo ae este conceito 1p logia, anatomia, palinologia e e pele Por estas razóes, Sper macoce, E Flora de Goiás e Tocantins, Flora do Distrito Federal e da Flora a di Guianas. PALAVRAS CHAVE: Borreria, Spermacoce, Rubiaceae, Goiás, Tocantins, Brasil, Guianas, Neotropicos INTRODUCTION The definition of the genera in the tribe Spermacoceae has been debated since their establishment, mostly due to the diverging opinions among specialists. A complete taxonomic history of this tribe and its generic delimitations was presented by Delprete et al. (2005); however, the main facts regarding the history of Borreria and Spermacoce are here summarized. Linnaeus (1753) established the genera Spermacoce [with S. tenuior L. (later selected as the type of the genus), S. verticillata L., and S. hispida L.] and Diodia L. Meyer (1818) founded Borreria G. Mey. (describing B. suaveolens G. Mey.), distinguishing it by having a capsule with both cocci dehiscent, while the fruit of Spermacoce has one coccus dehiscent and the other indehiscent. In the same work he characterized Diodia by having fruits with two indehiscent locules. Richard (1830), in the first important system of classification of the Rubiaceae, synonymized Borreria, Diodia and Spermacoce, with the following statement (Richard 1830: 150-151, free translation from French): ^A careful exam of a great number of species convinced me of the necessity to reunite Spermacoce, Diodia and Borreria into one genus. It is always the same kind of organization, not only in all floral parts, but also in the fruit, upon which was based the distinction among the three genera. In fact, the two cocci remain closed in Diodia, or dehisce longitudinally in Borreria, and the septum might be completely adherent to one coccus or missing, as in the Spermacoce species; I just want to repeat that it is the same organization, and these characters are not always clearly distinct, as it is possible to find one or two of these three types in the same species. I think that instead of forming types of genera, they can be used to establish subdivisions or sections within Spermacoce, which has a large number of species." Hooker (1873), following Richard, maintained Borreria and Sy as synonymous, describing the J. Bot. Res. Inst. Texas 1(2): 1023 — 1030. 2007 1024 Journal of the Botanical R h Institute of Texas 1(2) fruit of Spermacoce as with two dehiscent cocci, or with one indehiscent and the other indehiscent, with a persistent, membranaceous axis, and a persistent calyx. Schumann (1888), in his contribution to the Flora Brasiliensis, recognized the following genera in the tribe Spermacoce: Diodia, Hemidiodia K. Schum. [= Spermacoce], Psyllocarpus Mart. ex Mart. & Zucc., Spermacoce, Endlichera C. Presl [2 Emmeorhiza Pohl ex Endl.], Borreria (treating Galianthe as a section of this genus), Staelia Cham. & Schltdl., Mitracarpus Zucc. in. Schult. & Schult. f., Perama Aubl., Richardsonia Kunth [= Richardia L.], Schwendenera K. Schum., and Machaonia Bonpl. Bremekamp (1934), although he usually adopted rather narrow generic concepts (resulting in the separation of many taxa from several Rubiaceae genera), stated that the differences between Spermacoce and Borreria are small and of little taxonomic significance; however, he opted to maintain them separate for “purely opportunistic reasons." Steyermark (1972, 1974) in several occasions admitted to agree with B bout the similarity of I Tans 11 Borreria and Spermacoce, and also preferred to maintain them separate recognized in the New World, to avoid many new combinations, even though many African species have already been transferred to Spermacoce by contemporary authors. As a matter of fact, Verdcourt (1975, 1976, 1989) treated Borreria as a section of Spermacoce, and proposed the necessary new combinations for the African taxa. In a series of multidisciplinary studies Cabral, Bacigalupo and collaborators (Cabral 1991, 1993; Cabral et al. 2006; Cabral & Bacigalupo 1997, 2000; Pire 1997; Pire & Cabral 1992) maintained Borreria and Sper- macoce separated, and re-established Galianthe, where they transferred many species previously positioned in Borreria. In a successive work, Bacigalupo and Cabral (1999) transferred several species from Diodia to Borreria, Galianthe and Diodella (although without proposing the new combinations in the last genus), based primarily on the type of fruits dehiscence and inflorescence architecture, restricting Diodia to five species with indehiscent fruits. At the same time, they preferred to maintain them as separate, based on the same generic concepts and fruit characters used by Meyer (1818) and Grisebach (1879). Govaerts (1996) in his contribution to the World Checklist of Seed Plants treated Borreria, Ga Spermacoce as synonymous, publishing a total of 80 new combinations and new names in Spermacoce, mostly para ianthe and of Neotropical taxa. De Vré (2000), after a general pollen overview of the tribe Spermacoceae, concluded that there is no palynological evidence to maintain Borreria separate from Spermacoce. He also showed that the pollen of Spermacoce (incl. Borreria) is highly variable, and ranging from 3- to 20-colporate, with tectum foveolate, microspinulate or non-spinulate, with colpi regularly spaced or in couples, in equatorial position or in two series, or arranged in an 8-shaped line. Terrell and Wunderlin (2002) studied the seed morphology of several genera of the tribe Spermaco- ceeae. They concluded that Borreria and Spermacoce (and Galianthe) are not sufficiently distinct to warrant taxonomic separation. In the molecular phylogenies of Dessein (2003) and Dessein et al. (2006), species with fruit types tradi- tionally attributed to Borreria and Spermacoce are intercalated within the same clades. Therefore, data from morphology, anatomy, palynology, and molecular phylogeny support the wide delimitation of Spermacoce, and the two taxa should be treated as synonymous. In the floristic treatments of Costa Rica, Central French Guiana and Santa Catarina (Brazil), Adams (apud Burger & Taylor 1993), Boom and Delprete (2002) and Delprete et al. (2004, 2005), respectively, opted to treat the two genera as synonymous, and arranged all the species under Spermacoce. Therefore, according to Delprete et al. (2004, 2005), Spermacoce is delimited to include species with septicidal fruits with persistent calyx, commonly dehiscent from the top [basipetally] or exceptionally from the bottom [acropetally; e.g., S. palustris (Cham. & Schltdl.) Delprete; S. spicata (Miq.) Delprete, see below], with both cocci dehiscent, one dehiscent and the other indehiscent, or both of them indehiscent. A complete description of Spermacoce as here delimited was recently presented by Delprete € Cortés-B. (2006” [2007]). Continuing with this line of thinking, twelve new combinations are here proposed in Spermacoce to have these names available for my Rubiaceae treatments of the Flora de Goiás e Tocantins, Flora do Distrito Federal, and Flora of the Guianas. Delprete, New combinations in Spermacoce 1025 Two new combinations necessary for these treatments have already been published in Delprete & Cortés- B. 2006”[2007]), and are also reported here to reach a wider distribution in the international botanical community. Specimens cited were all personally examined, unless otherwise indicated by “n.v.” after the herbarium acronym. SYSTEMATIC TREATMENT 1. Spermacoce burchellii (E.L. Cabral & Bacigalupo) Delprete, comb. nov. Borreria burchellii E.L. Cabral & Baciga- lupo, Bonplandia 10:126. 2000. Borreria tenella (Kunth) Cham. & Schltdl. var. pumila K. Schum., in Mart., Fl. Bras. 6(6):56. 1888; non Borreria pumila DC.,1830. Type: BRAZIL. Tocantins: Porto Nacional [as “Goyaz, ad Porto Real"], s.d. [1828-1829] (fL), Burchell 8679-10 (LecTOTYPE selected by Cabral & Bacigalupo 2000: BR). boll Q 0.22 Additional specimens examined: BRAZIL: Tocantins: Porto Nacional [as “Goyaz, ad Porto Real], s.d. [1828-1829] (fl) Bu (BR) and 8657 (BR); without locality, s.d. [1818-1819], Pohl 2497 (Bt; cited by K. Schumann: 1888, p. 56) Geographic distribution —Known only from the historical the collections of early 1800s by Johann Pohl and William Burchell in the northern portion of the Province of “Goyaz” (now the state of Tocantins). 2. Spermacoce crispata (K. Schum.) Delprete, comb. nov. Borreria tenel ella (Kunth) Cham. & Schltdl. var. crispata K. Schum in Martr., Fl. Bras. 6(6):55. 1888. Borreria crispata (K. Schum.) E.L. Cabral & B ] Bonplandia 10:126. 2000. Syntypes: BRAZII Golás: without locality, s.d. [1840], Gardner 4173 (Bt, photo at NY!) and 4175 (BP. Distribution and ecology.—Restricted to northeastern and central Brazil, known from the states of Bahia, Minas Gerais, Goiás, and Tocantins. Growing in open fields in the Cerrado Biome. 3. Spermacoce delicatula (E.L. Cabral) Delprete, comb. nov. Borreria delicatula E.L. Cabral, Hikenia 3:21. 1999 (nom. nov. based on Borreria filiforme E.L. Cabral, pu andia 9:35. 1996, nom. x non Borreria filiformis d Hutch. & Dalziel, Fl. W. Trop. Afr. 2:135. 1931. Tyre: BRAZIL. “Mato Grosso”[R Paca Nova, extreme North I [this area is part of the state of Rondonia], cabeceira do Rio Cantario, Mar 1917 (fl, fr), “Rondon” [J.G. Kuhlmann] 2337 (notorvee: R! 53522; IsoTYPE: CTES n.v.). Taxonomic observations.—This species is similar to Spermacoce neotenuis Govaert (see below), from which it could be distinguished by the glomerules 1-3 mm in diameter (vs. 3-15 mm in diameter in S. neotenuis), 2-lobed calyx (vs. 4-lobed), corollas 1.5-1.7 mm long (vs. 6-7 mm long), and filaments 0.5-0.7 mm long (vs. anthers sessile or subsessile, filaments to 0.2 mm long). Distribution and ecology.—Species know only from the type, collected in the Serra dos Pacaás Novos, state of Rondonia. 4. Spermacoce dimorpha (J.H. Kirkbr.) Delprete, cu Nov. Borreria i H. Kirkbr., Brittonia 49:373. 1997. Tre: BRAZIL. Golds: Mun. Alto Paraíso, Chapada dos Veadeiros, a idade, rod. GO-118 para Teresina de Goiás, 14?03'02"S, 47°31'26"W, 1520 m, 28 Jul 1994 (fl, fr), A. Silva et al. 2117 (HOLOTYPE: IBGE!; isotype: US!). Distribution and ecology.— Apparently known only from the type. Small, annual herb, growing in open fields, below the grass layer. 5. D incognita (E.L. Cabral) Delprete, comb. nov. Borreria incognita E.L. Cabral, Opera Bot. Belg. 7:322. 1996. E: B . Golás: Mun. Campo Alegre, Rod. BR-050, km 222, 6 Feb 1994 (fl, fr), G. Hatschbach & J. Silva 59881 (HoLotyPE: MBM!; ISOTYPES: CTES n.v., SI n.v.). Selected EM studied: BRAZIL: Goiás: Chapada dos Veadeiros, 40 km N of Veadeiros, 1000 m, 16 Mar 1969 (fD, Irwin et al. 24512 (NY, UB); Mun. Sáo Joáo da Alianca, 3 km S of Sáo Joào da Alianca, 850 m, 14 Mar 1971 us Irwin et al. 31719 (NY, UB). Tocantins: Mun. Arraias, rd. Arraias-Paraná, 22 km NW of Arraias, 12 Feb 1994 (fl), Hatschbach 60 Endemic to central Brazil, and known from the states of Minas Gerais, Goiás, and Distribution ande ology Tocantins. Small herb, seldom collected because of its inconspicuous habit, restricted to open grassy fields (“campos limpos”), on sandy soils. 6. Spermacoce irwiniana (E.L. Cabral), Delprete, comb. nov. Borreria irwiniana E.L. Cabral, Bonplandia 9(1-2):36. 1996. Tre: BRAZIL. Tocantins [As “Golás”]: ca. 27 km S of Paraíso [now Paraíso do Tocantins; ca. 10°22'S, 48°52'W], gallery forest, ca. 500 m, 22 Mar 1968 (fl), Irwin, Maxwe & Wasshausen 21617 (HOLOTYPE: NY!; IsoTYPE: F n.v.). 1026 Journal of the Botanical R h Institute of Texas 1(2) Additional specimen examined: BRAZIL: Tocantins: Mun. Araguaína, BR-153, Km 1095, 5 Km S of Araguaína, 7°13'S, 48°14'W, 210 m, 9 Feb 1982 (fD, Krapovickas et al. 37839 (UB) Distribution and ecology.—Rare species, known from only two collections from the central and northern portion of the state of Tocantins: one from sandy soil among rocky outcrops, and the other from the margin of gallery forest. 7. Spermacoce multiflora (DC.) Delprete, comb. nov. Diodia multiflora DC., Prodr. 4:564. 1830. Borreria multiflora (DC.) Bacigalupo & Cabral, Opera Bot. Belg. 7:307. 1996. Tyre: BRAZIL: Locality, date, and collector unknown s.n. (HoLoTyPE: G-DC!). Sel 1: BRAZIL: Distrito Federal: Brasília, Á d botánico, 10 Jan 1967 (fl, fr), Duarte 10118 (UB); Brasilia, REIBGE, 30 Mar 1981 (fr), Heringer et al. 6678 (IBGE); Sobradinho, próximo ao Rio Torto, 975 m, 7 Jan 1966 (fD, Irwin et al. 11429 (NY, UB); Planaltina, 12 km S de Planaltina, DF-13, 1000 m, 21 Feb 1970 (fl-fr), Irwin et al. 26464 (NY, UB); Atoleiro, 15?40'S, 47°40'W, 900 m, 26 Jan 1972 (fl, D Do d ai 1701 (NY, UB); near Cachoeira da Forquilha, Ribeiráo-Sobradinho, 15?44'S, 47?44"W, 990 m, 6 Apr 1983 (fr), Ki id Brasilia, Ponte do Lago Paranoá, near Seminario D. Bosco, 14 Mar 1990 (fr), Silva et al. 965 doc Goiás: Serra Dourada, 2 Jan 1965 do, Heringer 10940 (UB); Mun. Corumbá de Goiás, Serra dos Pireneus, 12 km N de C bá de Goiás, 1000 m, 17 Jan 1972 (fl, fr), Irwin et al. 34416 (NY, UB); Mun. Mossámendes-Goiás, Serra Dourada, Área da UFG, campo rupestre, 2 Jan 1970 (fl, fr), Rizzo 4630 (UFG); Mun. Jataí, 16 Jan 1973 (fl-fr), Rizzo 8733 (UB, UFG). Te 1] Taxonomic observations.—Bacigalupo and Cabral (1996) transferred several positioned in Diodia and Hemidiodia to Borreria subgenus Dasycephala; e.g., B. gardneri (K. Schum.) Bacigalupo & Cabral, B. hyssopifolia (Willd. ex Roem. & Schult.) Bacigalupo & Cabral, B. ocymifolia (Willd. ex Roem. & Schult.) Bacigalupo & Cabral, and B. spicata (Miq.) Bacigalupo & Cabral. They characterized this subgenus by having “fruits septicidally dehiscing into two mericarps, each mericarp indehiscent and with a median longitudinal preformed line of dehiscence on the ventral surface or exceptionally one of them opening at the base.” This definition is encompassed in the present delimitation of the genus Spermacoce, and therefore the new com- binations Spermacoce multiflora (DC.) Delprete and S. spicata (Miq.) Delprete (see below) are here proposed. Distribution and ecology.—Widespread in South America, from Colombia, Venezuela, the Guianas, Brazil, Bolivia, and Paraguay. Herb or subshrub, 50-100 cm tall, erect, climbing or decumbent, usually found at margins or inside gallery forests. Spermacoce neotenuis Govaerts, World Checklist Seed Pl. 2:18. 1996. Borreria tenuis DC., Prodr. 4:543. 1830; non Sper- macoce tenuis Sessé & Moc., Fl. Mexic.: 25. 1893. Tre: BRAZIL: Locality unknown, s.d. [1818-1819], Pohl s.n. (HoLotTYPE: G-DC). Borreria gracillima DC., Prodr. 4:543. 1830, syn. nov. Spermacoce gracillima (DC.) Delprete, Rev. Biol. Neotrop. 3:72. “2006” [2007]. Tyre: BRAZIL. [Goiás or Tocantins]: Locality unknown, s.d. [1818-1819], Pohl s.n. (HoLoTYPE: G-DC; Isotype: F n.v. Selected specimens examined: BRAZIL: Goiás: Mun. Monte Alegre, 8 km W of Monte Alegre, 600—700 m, 11 Mar 1973 (fl), Anderson 6844 (NY, UB); Mun. Cavalcante, Chapada dos Veadeiros, 8 km S of Cavalcante, 1000 m, 10 Mar 1969 (fD, Irwin et al. 24189 (NY, UB); Mun. Ponte Alta, 15 Mar 1974 (fl), Rizzo 9666 (UFG). Tocantins: Mun. Ipueiras, near Rio Tocantins, 11°14'S, 48°27'W, 6 Dec 2001 (fD, E.A. Soares et al. 1858 (HTO, UFG); Mun. Ipueiras, near Rio Formiga, 11°15'S, 48°26'W, 20 Dec 2001 (fr), E.A. Soares et al. 1966 (HTO, Specimens cited by Schumann (1888): BRAZIL: Tocantins: “São Jodo da Palma” [now Paraná], s.d. [1818-1819], Pohl 1242 (B1); Goiás or Tocantins: “prope Gamelleira”, s.d. [1818-1819], Pohl 1916 (Bt); “Rio Reason” [now Rio Razão], s.d. [1818-1819], Pohl 2292 (B*). Taxonomic observations.—In agreement with Schumann [1888, p. 45-47, tab. 75, fig. I (B. tenuis), fig. II (B. ima DC. and B. tenuis DC. as distinct species. In their dichotomous key, they indicated that B. gracillima has corollas 4.5—4.8 mm long, but in the — gracillima)], Cabral & Bacigalupo (1996) maintained Borreria gracil description they wrote that the corollas are 4-4.8 mm long. Aside from this, in the key they distinguished B. tenuis from B. gracillima by the corollas 2.5-4.5 mm long (vs. 4-4.8 mm long in B. gracillima), corolla lobes longer then the tube (vs. shorter then the tube), and leaf blades 0.3-0.5 mm wide with scabrous margin (vs. 0.5 mm wide and glabrous). After careful comparison of more than 50 herbarium specimens and personal observations of natural populations, 1 concluded that all the characters used by Cabral and Bacigalupo to separate the two taxa are widely overlapping. For example, several populations on white sand areas near the city of Ipueiras (state of Tocantins), were observed to have leaves varying from linear to narrowly lanceolate (from 0.2 to 5 mm wide, ti even on the same individual), completely glabrous or with scabrous mar- gins, and corollas ranging from 3 to 5 mm long. Therefore, the two taxa are here treated as synonymous. Delprete, New combinations in Spermacoce 1027 Distribution and ecology.—Endemic to central Brazil, known from the states of Mato Grosso, Tocantins, and Goiás. Delicate, single-stemmed or few-branched herb, mostly found in seasonally flooded fields, on sandy soils and white sand areas of alluvial origin. 8. Spermacoce paraensis (E.L. Cabral & Bacigalupo) Delprete, comb. nov. Borreria paraensis E.L. Cabral & Baciga- lupo, Darwiniana 37:268. 1999. Tyre: BRAZIL. Pará: Marabá, Alto da Serra, 12 May 1982 (fl, fr), R. Secco, C. Sperling, M. Condon, A. Mesquita, B. Gilberto & L. Marinho 117 (HoLoTYPE: MG!; isotypes: CTES n.v., NY!, SI n.v.). Selected specimens examined: BRAZIL: Bahia: “Ad Cabulla”, Aug 1834 (fl, fr), Martius 601 (NY). Pará: Marabá, Serra dos Carajás, 6°00'S, 50°18'W, 700 m, 22 May 1969 (fl), P. Cavalcante 2107 (NY); Marabá, Serra dos Carajás, 1 Apr 1977 (f1), P. Silva et al. 2994 (NY); 5 km W of AMZA, 6%04'S, 50°10'W, 700-800 m, 15 May 1982 (fl, fr), Sperling et al. 5595 (NY) Distribution and ecology.—1t is known only by a few collections from the states of Pará and Bahia, and it may also occur in northern Tocantins. Spermacoce perangusta (S. Moore) Delprete, Rev. Biol. Neotrop. 3:72. “2006” [2007]. Borreria perangusta S. Moore, J. Bot. 42:100. 1904. Type: BRAZIL. Mato Grosso: Porto Murtinho, s.d., A. Roberts 884 (HOLOTYPE: BM!) Geographic distribution. —Apparently known only from the type, without any ecological observations available. 9. Spermacoce pulchristipula (Bremek.) Delprete, comb. nov. Diodia pulchristipula Bremek., Rec. Trav. Bot. Néerl. 33:713. 1936. Borreria pulchristipula (Bremek.) Bacigalupo & E.L.Cabral, Bol. Soc. Argent. Bot. 34:151. 2000. Type: SURINAME: Sipaliwini Savanna, Camp XI, near the Brazilian border, 10 Dec 1935 (fl, fr), Rombouts 360 (HoLoTYPE: U!; isotype: US!; photo-US at NY). Selected specimens examined: GUYANA: Rupununi River, N of Shea, 2257'N, 59*09"W, 190 m, 20 nn 1994 ME f, nm ues et al. 3286 (NY, U, US). SURINAME: Sipaliwini Savanna, 27 Aug 1966 (fl, fr), Donselaar 3631 (U). BRAZIL: Di REIBGE bn do corujáo, na margem do Córrego Roncador, 28 Feb 1978 (fr), Heringer et al. 370 (IBGE). Goiás: Between Jataí and Caiapónia, 40 m Caiapónia, 28 Jun 1966, Hunt 6259 (UB); Alto Paraíso, 21 Mar 1969, Irwin 24810 (UB). Tocantins: near Natividade, Jan 1840, as 3241 e n: BR-153, km 256, 14?30'S, 49°08'W, 3 Mar 1982, Krapovickas et al. 38142 (CEN); Mun. Pirenópolis, Serra dos | | la Portal do Lazaro, towars Cachoeira do Coqueiro, area permanently flooded, 15%47'29"S, 48°54'01"W, 1020 m, 26 Mar 2006 (fl, fr), P.G. Delprete & L.B. Bosquetti 9672 (CTES, HTO, K, MO, NY, R, RB, UB, UFG); Mun. Presidente Kennedy, road from highway BR-153 to Itaporá, 12 km W of Presidente Kennedy, Faz. Primavera, 3 Feb 1980, Plowman et al. 8349 (UB) Pireneus Distribution and ecology.—Amply distributed in South America, from Suriname throughout Brazil, Bolivia (Beni, Santa Cruz) and Paraguay (Amambay, Caaguazú, Central, Cordillera, Paraguarí, San Pedro). In Brazil itis known from the states of Pará, Tocantins, Mato Grosso, Goiás to Sáo Paulo. Present in grassy fields, sometimes seasonally flooded, borders of water courses, and margins of gallery forests. Spermacoce scabiosoides (Cham. & Schltdl.) Kuntze, Revis. Gen. Pl. 3:123. 1898 (como “scabios- od S ). Borreria scabiosoides Cham. € Schltdl., Linnaea 3:318. 1828. Type: BRAZIL. Rio DE JANEIRO: WITHOUT LOCALITY, S.D., SELLOW S.N. (HOLOTYPE: Bt, photo at NY!). Borreria e var. o Huber, Bull. Soc. Bot. Geneve 6:211. 1914, syn. nov. Tyre: BRAZIL. Pará: “Arumandaba” [Aruman- duba], flooded field, 3 May 1903 (fl), A. Ducke 3556 (HOLOTYPE: MG). andl un m Mus. Nat. Hist., Bot. Ser. 7:245. 1931, syn. nov. Borreria scabiosoides var. anderssonii (Standl.) Steyerm., Acta Bot. Venez. 6:194. “1971” [1972]. Type: ECUADOR: Puna Island, near Guayaquil, 1852 (fl, fr), NJ. Andersson 71 (HOLOTYPE: SP). Borreria flexuosa E.L. Cabral, Bonplandia 9:36, fig. 2. 1999, syn. nov. (non Spermacoce flexuosa Lour., Fl. Cochinch.: 79. 1790). Tyee: RAZIL. Maro Grosso: Cuiabá, Santarem Road, next to BR-163, km 601, artificial wet area, 22 Feb 1977 (fl), J.H. Kirkbride & E. Lleras 2986 (HOLOTYPE: NY!; isotypes: BR!, F n.v.). Taxonomic observations.— Cabral (1999) distinguished Borreria flexuosa E.L. Cabral from Spermacoce sca- biosoides (Cham. & Schltdl.) Kuntze because of the leaves 1-2 mm wide and without secondary veins (vs. 5-8 mm wide and with 4-5 veins each side in S. scabiosoides), stipular setae 0.5-3.5 mm long (vs. 3-4 mm long), and white corollas (vs. pale blue). However, the variation among the three varieties of B. scabiosoides recognized by Steyermark and B. flexuosa is within the degree of variation seen in several natural popula- tions in the state of Tocantins. In large populations of S. scabiosoides (Mun. Lagoa da Confusáo, Delprete et al. 6542 and 6558), growing in standing water 30—70 cm deep, it has been personally observed that the habit and leaf shape of this species is quite variable, even within the same individual. Generally, the submersed 1028 Journal of the Botanical R h Institute of Texas 1(2) leaves tend to be slightly narrower, and with 1-2 or no secondary veins, while the emerged ones are usually slightly wider and with (1-)2-4 secondary veins. Also, in the same large populations, the plants growing at the margins of the flooded area had narrower, sturdier stems, while the submersed plants had succulent to sub-succulent stems. Therefore, all the taxa mentioned above are here treated as synonymous, and no pues rank is recognized within the species. Selected st ined: BRAZIL: Tocantins: Mun. Lagoa da Confusão, 1 km W da cidade, 10?47'S, 49?37"W, 170 m, area se- asonally flooded, 17 Aug 1998 (f1), Del p et al. s (NY, ss Mun. Lagoa da Confusáo, Fazenda Trindade, ca. 35 km de Lagoa da Confusão, 10°39'S, 49°51'W, 210 m, flood fa forest, 18 Aug 1998 (f1), Delprete et al. 6558 (NY, UFG); Mun. Palmas, Córrego Santa Luzia, 22 Mar 2000 (fl, fr), E.A. Soares et al. 607 (HTO). eee schumanniana (Taub. ex Ule) Govaerts, World Checklist Seed Pl. 2:18. 1996. Borreria schumanniana Taub. ex Ule in Cruls, Rapp. Comm. Expl. Plat. Centr. Bresil: 351. 1894. Type: BRAZIL. Gouás: “Auf freien stellen an Paranahyba” A im near Rio Paranaíba], Feb 1893 (fl, fr), Ule 2960 here selected: HBG; HOLOTYPE: Bt). Taxonomic observations.—Cabral & Bacigalupo (1999) reported that Borreria schumanniana was originally published by Ule in “Bot. Jahrb. Syst. 21:453. 1895,” but this name was first published by Ule a year earlier (see above). The same authors, knowing that the HoLotYrE: Ule 2960, deposited at B was destroyed during WWII, assumed that no duplicates of this collection are to be found. Therefore, they proceeded in select- ing Ule 427 (P) as a neotype, which is a collection made from practically the same location of the holotype. However, at the HBG herbarium is kept a complete set of Ule up to collection number 7575, and among them is present a duplicate of Ule 2960, that is here selected as the lectotype of this species. Selected specimens examined: BRAZIL: Goiás: Mun. Calda Novas, Serra de Calda Novas, 28 Feb 1970 (fD, Rizzo & Barbosa 4801 (UFG, UB); Mun. Cristalina, Serra dos Topázios, 20 km antes de Cristalina, rod. Brasília-Belo Horizonte, campo rupestre, local arenoso, 26 Apr 1973 x m Rizzo 9014 (UFG); Chapada dos Veadeiros, 4 May 1972 (fl), Rizzo 8077 (UFG); Mun. Mossámendes, Serra Dourada, da Reserva Biológ órregos Cafundó e Picarráo, 18 Oct 1994 (fl, fr), Rizzo 11088 (UFG). 10. Spermacoce semiamplexicale (E.L. Cabral) Delprete, comb. nov. Borreria semiamplexicale E.L. Cabral, Bonplandia 9:37, fig. 4. 1996. Type: BRAZIL. Pará: 25 km NW of camp at Serra Norte, ca. 5°54'S, 50°37'W, 13 Dec 1981 (fl), D. Daly, R. Callejas, M.G. da Silva, E. Taylor, C. Rosario & M. dos Santos 1993 (HOLOTYPE: NY!; isotype: CTES n.v.). Distribution and ecology.—A species characterized by the basally overlapping, cordate to auriculate involucral bracts. It is known only from the type, collected in flooded grassy fields from the state of Para, and it may occur also in northern Tocantins. 11. Spermacoce tocantinsiana (E.L. Cabral & Bacigalupo) Delprete, comb. nov. Borreria tocantinsiana E.L. Cabral & Bacigalupo, Kew Bull. 59:277. 2004. Tyre: BRAZIL. Tocantins: Mun. Conceição do Tocantins, Faz. Cartão de Visita, 11 May 2000 (£D, Hatschbach, Schinini & Barboza 70928 (HOLOTYPE: MBM!; IsoTYPES: CTES n.v., SI n.v.) Distribution and ecology.—Apparently known only from the type. Annual herb, found in open grassy fields, in sandy soils. 12. Spermacoce spicata (Miq.) Delprete, comb. nov. Diodia spicata Miq., Stirp. Sur. 179, t. 52. 1850. Dasycephala spicata (Miq.) Hook. f. in Benth. & Hook. f., Gen. Pl. 2:144. 1873. Borreria spicata (Miq.) Bacigalupo & Cabral, Opera Bot. Belg. 7:307 1996. Type: SURINAME: Bergendaal, s.d. (fl, fr), Focke 1140 (HOLOTYPE: U!; PROBABLE ISOTYPE: K!; photo-K at NY!). Selected specimens examined: SURINAME: Brakopondo Distr., Brownsberg Nature Park, trail to Mazaruni Valley, 4?56'N, 55°11'W, 400-450 m, 24 Jan 1999 (fl, fr), Delprete et al. 7083 (BBS, MO, NY); along Makambie Creek, ca. 121 km of railro ad, 21 Mar 1951, J. Florschuütz & P.A. Florschuütz 1877 (NY, U); Sipaliwini savanna area on Brazilian frontier, 280 m, 14 Jan 1969 (fl, fr), Oldenburger et al. 933 (NY, U). FRENCH GUIANA: Saúl, rd between Saúl and airport, 3°37'N, 53°12'W, 200—400 m, 2 Sep 1994, B.M. Boom 10728 (CAY, NY). GUYANA: 5 mi N of Tunuwan, E. Kanuku Mountains, ca. 110 m, 6 Sep 1958, Cook 107 (NY); Kamakusa, Upper Mazaruni River, ca. 59°50'W, 23-29 Nov 1922, De La Cruz 2808 (NY); Rupuruni Distr., Eastern Kanuku Mountains, NE of Warimure, 23 Jan 1991 (fl, fr), Jansen-Jacobs et al. 2189 (NY, U); Rupuruni Distr., Kanuku Mountains, Crabwood Creek, Camp 2, 4 Feb 1994 (fl, fr), Jansen-Jacobs et al. 3564 (NY, U); Basin of Shodirak Creek (Essequibo River tributary), 1°18'N, 8-22 Jan 1938, A. C. Smith 2894 (NY). Morphological observations.—The fruit of this species, at maturity, dehisces acropetally (from the pedicel up- wards), up to the medial portion of the capsule. The two cocci remain attached to each other at the medio-distal Delprete, New combinations in Spermacoce 1029 portion. Often the whole fruit, divided at base or still unopened, falls off with the pedicel, functioning as a dispersal unit. Distribution and ecology Apparently endemic to the Guianas, but it might be present also in contiguous areas in southern Venezuela and northern Brazil. Spermacoce wunschmannii (K. Schum.) Kuntze, Revis. Gen. Pl. 3(2): 123. 1898. Borreria wunschmannii K. Schum in Mart., Fl. Bras. o 53. 1888. Type. BRAZIL: Tocantins: “prope Porto Real” [now Porto Nacional], s.d. [Nov 1828-Apr 18291. Burchell 8 LE E [selected by Cabral & Bacigalupo 2005]: BR!). R hol] ROSNIRR Selected specimens examined: BRAZIL: Tocantins: “inter Funil et Sáo Joao ad fluvium Tocantins,” s.d. [1828-1829] and 8977 (BR); Mun. Mateiros, Regiáo do Jalapáo, 10°33'S, 46°45'W, 400 m, 8 May 2001 (fD, C. Proenca et al. 2492 (UB, UFG); Mun. Mateiros, Região do Jalapão, 10°33'S, 46°45'W, rd. between Ponte Alta and Mateiros, near Rio Novo, 8 May 2001 (fl), A.B. Sampaio et al. 515 (UFG) Distribution and e ology Species known to occur in the state of Tocantins, and recently reported by several collections from the lowlands of Bolivia, Santa Cruz (Cabral & Bacigalupo 2005). It is therefore expected to occur also in the state of Mato Grosso. In Tocantins, aside from the historical material, it is also known by two recent collections from the Jalapáo Region, in seasonally flooded fields on white sands of alluvial origin. ACKNOWLEDGMENTS This work was undertaken during a fellowship for Visiting Scientist from the National Counsel of Tech- nological and Scientific Development (Conselho Nacional de Desenvolvimento Científico e Tecnológico - CNPq) of the Brazilian Government (grant 309885/2003-5), at the Federal University of Goiás (UFG), under the coordination of Vera Lúcia Gomes-Klein (UFG). Research at the Nationaal Herbarium Nederland, Utrecht University branch (U), in 2004, was supported by a fellowship from the Netherlands Organiza- tion for Scientific Research NWO (grant B 85-368; Rubiaceae treatment for the Flora of the Guianas), under the coordination of Marion Jansen-Jacobs (U). My gratitude also goes to Rob van Aubel (U) and Hendrik Rypkema (U) for help in finding and reproducing relevant taxonomic literature. The directors and curators of the following herbaria are kindly acknowledged for loan of material, sending digital images, and/or pro- viding working space during my visits: BR, CPAP, CTES, F, G, HTO, IBGE, K, MBM, NY, NX, R, RB, S, U, UB, UFG, UFMT, and US. I am very grateful to Joseph Kirkbride (USDA) and Steven Dessein (BR) for their careful and efficient revision of the manuscript, and to Guy Nesom (BRIT) and Barney Lipscomb (BRIT) for reviewing and editing the final version of this work. REFERENCES Apanms, D. 1993. [Spermacoce]. Family 202. Rubiaceae. 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HAWTHORNS (CRATAEGUS: ROSACEAE) OF THE CYPRESS HILLS, ALBERTA AND SASKATCHEWAN J.B. Phipps RJ. O'Kennon Department of Biology Botanical Research Institute of Texas The University of Western Ontario 09 Pecan Street 1151 Richmond Stree Fort Worth, Texas, 76102-4060, U.S.A. London, Ontario, NGA 5B7, CANADA bokennon@brit.org jphipps@uwo.ca ABSTRACT This paper describes the ecological situation of Crataegus found in the Cypress Hills of Alberta and Saskatcl in tl text of cli mate and vegetation patterns, followed by a detailed account of the taxa present. C i l to b in certain habitats and are by far the most abundant shrubby plant of the Cypress Hills. A n combination of physical d of relief, temperature, precipitation and soil have p rise to a remarkable ecological island. Thirteen species of C to occur compared with three in earlier work. S the species, C. cupressocollina, C. aquacervensis, C. rivuloadamensis, C purpurella, C. rivulopugnensis, C. rubribracteolata and C. ursopedensis are new to science and most of them are at least locally common in the Cypress Hills. In addition, C. sheridana A. Nels. has been resurrected. Also one new variety, saskatchewanensis, of C. sheila-phippsiae is described. Four of the new taxa M to bi ds to the Cypress Hills akkengh more intensive fieldwork in Montana might alter this. The biogeography is discussed in t t Plains Holocene h d the small tion available on their differential ecology. All taxa are keyed and the new taxa are ack fully o and illustrated. This work is based on over 500 collections, over 450 of which are by the authors. Key Wokbs: Crataegus, Rosaceae, systematics, taxonomy, biogeography, ecology, Cypress Hills, new taxa RESUMEN En este artículo se describe la situación ecológica de Crataegus en los Cypress Hills de Alberta y Saskatchewan en el contexto de pa- trones de clima y vegetación, seguido de un informe detallado de los taxa presentes. Las especies de Crataegus son extremadamente numerosas en ciertos hábitats y son con mucho las pou un más abundantes de las Cypress Hills. Una combinación peculiar de factores físicos de relieve, temperatura, precipitación y suelo han dado lugar a una isla ecológica singular. Se demuestra la presencia trabajo previo. Siete de las especies, C. cupressocollina, C. aquacervensis, C. rivuloadamensis, C. purpurella, C. rivulopugnensis, C. rubribracteolata y C. ursopedensis son nuevas para la ciencia y la mayoría de ellas son al menos localmente comunes en las Cypress Hills. Además, C. sheridana A. Nels. ha sido resucitada. También se describe una nueva variedad, saskatchewanensis, de C. shei de trece especies de Crataegus comparadas con las tres de un a-phippsiae. Cuatro de los nuevos taxa parecen restringirse a las Cypress Hills aunque un traba jo d más intenso en Montana pudiera alterar esto. Se discute la biog fí l contexto del cambio de la vegetación en el TEM da las Grandes Llanuras y la poca cantidad de información sobre su MOM diferencial. Todos los taxa se incluyen en la clave y los nuevos taxa se describen ampliamente y se ilustran. Este trabajo está basado en más de 500 colecciones, más de 450 de las cuales fueron realizadas por los autores. The authors decided to add the Cypress Hills (Plate 1) to their botanical explorations of the western parts of the North American continent and in doing so unearthed a botanical goldmine. This paper describes in detail the thirteen species of Crataegus discovered, six species and one variety proving new to science. Not only was there a large number of hawthorn taxa present, as in the British Columbia Okanagan, but the number of individuals, over 500 collections studied as emphasized in Figure 2 (collated points for all species), was very large and thus clearly indicates one of the most favorable areas of similar size for Crataegus in the west of the continent. Hawthorns, discovered to be by far the most abundant shrub genus in the Cypress Hills and of great biological significance due not only to this fact, but also, inter alia, to their considerable fruit production providing food for numerous species of bird and mammals, nevertheless do not achieve more than passing mention in significant ecological treatments of the area, a situation which is remedied here. Questions were prompted about what was significant about the habitat that was so favorable and this is discussed below. Because the Cypress Hills constitute an ecological island (similar in this respect to several in Montana east of the Rocky Mountains) and because a considerable proportion of the taxa found appeared J. Bot. Res. Inst. Texas 1(2): 1031 — 1090. 2007 1032 Journal of the Botanical R h Institute of Texas 1(2) to have distributions tied to these ecological islands, it seemed of interest to consider the biogeographic ramifications of the Cypress Hills Crataegus populations particularly in the context of the Great Plains Ho- locene, which is also done. These and related matters form the subject of this paper. PHYSICAL BACKGROUND The Cypress Hills are a regionally significant landform lying between about 111? W and 108? 40' W and 49? 20' N and 49? 45' N, i.e. in the southern part of the western half of the Canadian Prairie Provinces, thus occupying about 165 km E-W x 60 km N-S or about 10,000 sq. km. Their location is shown on Plate 1. The Cypress Hills are presented in three maps the first of which is a colored relief map with forest cover indicated (Plate 2) followed by a detailed sketch map of the hills broken into eastern and western halves (Figs. la, 1b). The Cypress Hills trend east-west and lie between the Canada-United States border, which is to the south, and the Trans-Canada Highway, which is to the north. The topography is gently rolling to almost flat on the summit plateau which has a highest point of 1465 m in the west, nearly due south of Elkwater, Alberta and which descends to levels of only about 1200 m towards the eastern end of the hills in Saskatchewan. The nearly flat-lying prairies at the bottom of the northern scarp lie about 400-600 m below the summit plateau. Along the northern edge of the Cypress Hills is a scarp slope (Fig. 4a), dissected by creeks of vari- ous size, some of which eventually reach the South Saskatchewan River. Both eastern and western limits are also quite sharply defined, with a significant eastern scarp from which arise tributaries of Swift Current. Medicine Lodge Coulee (Fig. 4b), a glacial spillway, slices off the western end of the Cypress Hills in Alberta and is a particularly striking topographical feature. From there, Lodge Creek flows southeastwards into the Milk River in Montana. The southern limits of the hills are, contrastingly, much less sharply defined (Fig. 4a, Plate 3c) and fade into high grassland towards Montana. A further important valley, that of Battle Creek, another glacial spillway, originates near the northern scarp in Alberta, thence flowing southeast through Fort Walsh in Saskatchewan before 11 joining the Milk River east of Havre in Montana while a major river, the east-flowing Frenchman, dissects the hills in their south-eastern quadrant. A considerable variety of flat-lying sedimentary rocks of upper Mesozoic to lower Tertiary (Fig. 3) creates the basis for the generally smooth topography. Some interesting elements include good exposures of the KT boundary clay and upper Cretaceous finds of dinosaurs such as Tyrannosaurus rex. Brown chernozem soils are found in grassland below the tree line and dark brown chernozems on the summit plateau while forest slope soils are typically dark grey luvisols (Henderson et al. 2000). However, there is no obvious relationship between underlying geology, resultant soil types and Crataegus distribution patterns in the Cypress Hills. Rather, Crataegus distribution in the Cypress Hills appears mainly to follow soil moisture patterns. Indeed, moisture has been shown to be the key determinant of regional vegetation distribution in the southern Canadian Prairies and adjacent United States (Looman 1979; Larsen & MacDonald 1995; Hogg 1994). Hogg (1997) calculated a climatic moisture index (CMI) to which the regional vegetation was more closely tied than to any other factor. Hogg's CMI was calculated as mean annual precipitation minus potential evapotranspiration. He showed CMI’s below 30 are for grassland, -30 to -15 (grassland with stunted aspen), -15 to 0 (mixed aspen-grassland), 0 to 15 (aspen dominated woods with conifers) and over 15 (coniferous woodland). The presence of habitats in the Cypress Hills with higher CMIs is the principal reason for the presence of Crataegus there so forest cover is shown on the topographical map (Plate 2) although it is some- what inexactly represented at this scale. The reason for the prevalence of higher CMI’s is connected with the combination of lower temperatures with increasing altitude as well as higher precipitation with increasing altitude in parts of the Cypress Hills, a topic which is elaborated in a discussion of the meteorology. We were not able to draw upon a fine grid of relevant microclimatological data for the Cypress Hills, only one meteorological station being within the hills proper, that at the Cypress Hills Provincial Park HQ, SK. The surrounding prairie stations are, however, quite close by and thus present characteristic regional climatic data which presumably also reflects that for the grassland parts of the Cypress Hills in precipita- tion if not in temperature. The surrounding prairie stations selected are: Medicine Hat, Alta., 45 km NW of Phipps and 0 , Crataegus of Cy] t 1033 > io TT de o lz s m 3 AS => E 2 LÀ M 945 P E] Sy à $ epy ' E oie NY 1 u. 1998109 E t * = 09) 1 $. = ; 6$ iu i 2% 32 : GN d | | o e = AAA] = e M oe ? y Tew "d 3 o to N = on 2 CY 8 k s | p FEDA - = n EN PR : 5 [= 39910 po Sysynd 9 94uey n = Cc c e i E NI i> a a zs Y 39 E HE A w L a Y E = = 5 In E z z E [5] to : E + g E Elkwater; Maple Creek, Sask., at the base of the northern slopes of the hills; Shaunavon, Sask., ca. 25 km E of the eastern scarp of the hills; and Willow Creek, Sask., on the Montana border due south of the centre of the hills. These sites thus collectively and usefully box in the Cypress Hills and show the regional prairie climate to be quite uniform. D £L JUUTTIal OF 1034 es ; Med uJo]seo 223 Lieu 12 GL 7 Gv. 6v oreo’ (ut 082) X99319 ajdeyy b2‘€L MS ~w 0001 o JU o ` NOLL 1035 $p40291 |] "S3UI lr FEMO F} ] W CA 10 ia E 7 TA . Ny EN C hv n == - | | Im E - UJ ere "n 3 ; l1. = £ 1036 Journal of the Botanical R h Institute of Texas 1(2) Although an almost complete lack of me- ERA _PERIOD__EPOCH_AGE* FORMATION MATERIAL teorological stations within the Cypress Hills Neogene |Pieistocene]0.1 1255 II. loess: til : : : Miocene 1 prevents any precise and detailed interpreta- E Oligocene | conglomerate tion of climatic variation, comparison between > ocene E Maple Creek, a drier and warmer site at the e Paleogene sandstone iltst base of the central part of the northern scarp l Paleocene coal. shale at 764 m, and Cypress Hills Provincial Park, nestled in a low valley in the central part of the summit plateau almost due south of Maple rió n ale Creek, and only about 28 km distant, at 1196 m, is instructive. From the mean monthly precipitation and temperatures (Tables 1, 2) for these and other reference stations it may 9n Tu po AS - E M 7 be seen that the area around the Cypress Hills E || Whitemud||[[|) sandstone, clay ee : : : [rp 7 shale Provincial Park meteorological station is both B 7 1 TT Upper Maastrich- : z Uppe cooler (mean monthly difference during the Cretaceous] tian f : o sandstone, shale growing season from April to October of 3.1 B coal deg. C) and substantially wetter (606 vs. 375 mm annual precipitation), thus ecologically more mesic, than Maple Creek. Maple Creek, in spite of being so close to the hills, is neverthe- 68 BF E less typical of the group of regional sites used. | Campanian|>68|= Bears Paw = ||| shale All of the latter thus have similar to slightly *in millions of years warmer temperatures and similar to slightly less Fic. 3. Geoloaical + £f Hlc ¢ All n £ precipitation than Maple Creek, thus illuminat- POE ing the island nature of the mesic parts of the Mie Eque nt Cypress Hills. It is the impact of moisture-laden cold fronts rising up exposed scarps from the north and northwest that accounts for the unequal distribution of precipitation in the Cypress Hills and altitude for the temperature differential. Growing season snow has twice been encountered during our fieldwork over 20 cm deep on parts of the northern scarp face (Plate 5a). The effect of altitude on temperature may also be seen by considering anthesis times, as the following example illustrates. In spring 2003 the temperature contrast between Cypress Hills Provincial Park and the Maple Creek meteorological stations translated into one species of Crataegus (C. cupressocollina) being in full flower on 23 May on the lower slopes of the northern scarp at 892 m, near Maple Creek, as well as in the Frenchman Valley near Eastend at the same date and similar altitude but being still in tight bud (flower buds ca. 2 mm diam., entire inflorescence extension only about 15 mm) near Cypress Hills Provincial Park at 1331m, on 24 May. The same situation was seen a day later, 25'* May 2003, along Battle Creek, Alta., at 1166 m and also as well as at higher elevations near Reesor Lake (1292 m), Alberta. Similar observations may be made each year. CRATAEGUS ECOLOGY The Cypress Hills present predominantly a northern short-grass prairie biome broken by large to small patches of forest and shrubby thickets near creek beds along most of the upper part of the northern scarp as well as in some other draws and valleys more generally (Plates 3, 4, 5b). The overall vegetation was discussed by Breitung (1954) and the forests specifically, in some detail, by Newsome and Dix (1988) as well as by Henderson et al. (2000). Each of these papers is notable for a nearly complete lack of attention to hawthorn habitat, in spite of its extent and significance. Forest is concentrated in the middle and upper parts of the north-facing slopes where it can be continuous for miles (Plate 3a). Smaller patches line the deeper ravines. 1037 FRIPps anu " Y JI SOUTH SASKATCHEWAN VALLEY SUMMIT PLATEAU FRENCHMAN RIVER MONTANA 4— —» AE ap ree TN — 1200 m f " "E 1000m NS An E II ted 3 He Oa aS cite 800 m 700m l l l | 0 5 10 15 20 Horizontal scale (km) a. N-S section of Cypress Hills at about 109°20°W MEDICINE LODGE COULEE SUMMIT PLATEAU 1600 m A 1500 m 1400 m AREA, P din. L 1300 m P o 1200 m LODGE CREEK 1000 m L fi | | l | 0 1 2 3 4 5 Horizontal scale (km) b. SW (left) - NE secti ) dge Coul SUMMIT BONE T PLATEAU CREEK 1100 m py eee 1000 m 900m | | l | 0 1 2 3 4 Horizontal scale (km) c. E-W section across Bone Creek 47*55'N Fic. 4. Topographical ions for three | he Cypress Hills, | I in Plate 2, p. 1045 a) A-B. North-south section running from just east of Maple Creek. b) C-D. Soutt | h Medicine Lodge Coulee. c) E-F, East-west section across Bone Creek Coulee. Aerial surveys indicate that about 5-15% of the Cypress Hills land area is covered by forest or forest-scrub (see map, Plate 2). The trees involved are principally the boreal species white spruce (Picea glauca), found in deeper northern valleys and the steeper scarps, the Rocky Mountain species, lodgepole pine (Pinus con- torta), on drier summit and sub-summit slopes and finally aspen (Populus tremuloides) which forms extensive areas of forest or parkland linking many of the white spruce and lodgepole pine stands, primarily on the upper northern slopes and and the sides of deeper ravines (Fig. 4c). A few other tree species (e.g., willow, boxelder) occur in small quantity in the deeper valleys. The grassland biome outside of the Inter-provincial Park boundaries is farmed and carries a mix of dryland crops such as wheat and alfalfa as well as fallow (in flatter areas), and rangeland (on slopes). Hawthorns are found in all parts of the Cypress Hills (Fig. 6a) but reach their greatest abundance as an almost continuous aspen understorey in the more mesic aspen-dominated sites and in thickets, some- times huge, in north-slope draws and in the deeper coulees. This biome, particularly at higher altitudes and particularly on north and west slopes in the Cypress Hills supports extensive hawthorn thickets along 1038 Journal of the Botanical R h Institute of Texas 1(2) Taste 1. Climatic data for Cypress Hills and nearby localities. Mean monthly precipitation (mm) for high elevation northern scarp compared with surrounding prairies. J F M A M J J A S 0 N D Total Cypress Hills (1196 m) 35.31 281 41.9 405 768 638 696 475 582 367 35.1 407 6068 Maple Creek (764m) 258 171 189 305 45.7 578 456 354 380 182 179 242 3753 Medicine Hat (717m) 17.8 102 164 252 425 575 408 29.7 368 15.1 15.0 20.0 304.0 Shaunavon 2 (914m) 188 128 233 249 572 685 524 364 31.1 184 169 260 3846 Willow Creek (861m) 124 78 160 156 471 606 413 309 323 163 170 166 3138 NA | Taste 2. Climatic data for Cypress Hills and nearby localities. Mean monthly temperature (9C) for high elevation northern scarp compared with surrounding prairies. J F M A M J J A S 0 N D mean Cypress Hills (1196 m) -10.7 -7.8 -37 28 9.0 Sl 16:2 160.0 o 43 “38 88. 30 Maple Creek (764m) -10.8 -70 -15 58 Miss os. d97- 190° 128. 74 AD "815 25.6 Medicine Hat (717m) -108 -69 -12 63 124 170 198 191 129 74 Sco o> Shaunavon 2 (914m) -110 -6.8 -19 56 11215 5: 118:2: å Sos 39 -93 44 Willow Creek (861m) -117 -80 -15 58 116 162- 192. 188. 123 56 38 -97 46 creeks and in draws (Plate 4a). Such creek thickets represent the bottom part of a catena Morison et al. (1948) where groundwater regimes override topland CMI’s and create an interdigitating habitat mosaic this part of which is particularly suitable for hawthorn. Hawthorn numbers are thus concentrated in all those sufficiently mesic sites not dominated by evergreen conifers and CMIs of 10-15 for topland sites may be inferred. The abundance of Crataegus under aspen (Plate 4b) in the Cypress Hills represents a new habitat for Crataegus so far as the literature is concerned. This kind of situation, relatively mesic and under dappled or light, but not heavy, shade presents an analogue to the rather sparser Crataegus understorey below oak and pine that is commonly encountered in the southeastern United States and is to some extent at odds with the relatively often seen observation that Crataegus is shade-intolerant. The high negative correlation with fruticose Salix species in the Cypress Hills, the latter abundant in some marshy bottomlands may be due to hawthorns’ poorer tolerance of periods of waterlogging even though many of these willow habitats apparently dry out extensively during summer. In the Cypress Hills hawthorns are virtually never found in the open prairie away from draws and similarly mesic sites and their distribution there is in line with the general interpretation of most North American hawthorn species being mesophytes rather than being particularly xeromorphic. Possibly also a significant factor in the regionally extraordinary abundance of hawthorn in the Cypress Hills is the absence of Juniperus, alternate host of the damaging Gymosporangium rust. Injury by this rust is infrequent in the Cypress Hills and to cause infection Gymosporangium would need to migrate into the area annually. ORIGINS OF THE CYPRESS HILLS WOODY FLORA Tas] 3 The origins of woodland in the Great Plains generally and the Cypress Hills particularly are topics of direct relevance to this paper. Sauchyn and Sauchyn (1991) in an account of the Holocene pollen record of Harris Lake in the Cypress Hills and Porter, Sauchyn and Delorme (1999) in a parallel account of the ostracod record of the same lake inferred vegetation change for the Cypress Hills and set it in a wider context. Harris Lake, elev. 1180 m (Plate 5b), is about 9 ha. in size and lies about 5 km northwest of Adams Lake (mapped, Fig. 1b) just below the steepest part of the northern scarp of the Cypress Hills. Today, Harris Lake has Picea glauca and Pinus contorta forest upslope, some aspen around the lake and grassland downslope towards the north and northeast. It is known to have been in a meltwater channel that existed about 12,000 B.P. during DL: Wat lA f de £ f. ENII AIL 4 Ic I 4 ^L Phipps and 0 , Crataegus of Cy] 1039 the main deglaciation. However, continuous pollen and ostracod records only exist from ca. 9,200 B.P. so the above authors assume that it was in the missing interval ca. 12,000-9,000 B.P. that the main invasion of Picea glauca, a species whose abundance later fluctuated greatly, occurred. Inferences of vegetation from the two papers cited above are slightly different so the more recent paper is used here. Four main paleoenvironments are recognized for the Cypress Hills, I-IV. Zone I, early Holocene, 9240-6400 B.P., had a variable climate supporting aspen parkland; Zone II, Hypsithermal, 6400-4500 B.P., was warm and dry supporting grassland and only small patches of forest; Zone III, moister and cooler, 4500-3600 B.P., had expanding subboreal forest; while Zone IV, 3600 B.P. to the present, represents short grass prairie with islands of trees. An important deduction in the earlier paper is that Picea glauca probably persisted in small patches through the Hypsithermal. This implies continuity for Picea glauca, the most me- sophytic of the three main tree species, at least at low abundances, continuously since its main invasion. It is also notable, however, that when a microscope is applied to the time periods, quite sharp climatic changes have been shown to occur in the region even during the approximately 120 years for which meteorological records were available for this area (Sauchyn & Beaudoin 1998), although the extent to which this impact overall survival of quite long-lived woody species is less clear. Interpetation of the Crataegus record for the Holocene for the Cypress Hills remains inferential, however, as it is insect-pollinated and no pollen evidence has been found, while macrofossils, also generally rare for Crataegus, have not turned up either. What we have independently established, however, is that Cypress Hills Crataegus occur abundantly in mesic aspen woodland (CMIs of ca. 10-15) as well as along numerous creek beds of appropriate, and probably averaging similar, mesicity. It is therefore probably safe to infer that the Cypress Hills, since the time Picea glauca was first present, i.e., ca. 11,000 to 10,000 B.P., have continuously contained at least some habitat sufficiently mesic for Crataegus but that this habitat has probably fluctu- ated greatly in extent in the intervening period. The first suggestion of extensive areas of suitable habitat is therefore coterminous with Zone I, aspen parkland, 9200—6500 B.P. Nevertheless, Zone I was not neces- sarily the first Cypress Hills paleoenvironment to support Crataegus as that would have depended on the presence of similarly suitable source paleoenvironments to the south and southwest, especially in the Bears' Paw Mountains and the Sweet Grass Hills about which parallel information is not available. However, three or four Cypress Hills species seem to have some ability to survive along watercourses in the open prairie (and thus are not restricted to montane areas). These species are Cc. chrysocarpa, sheridana, sheila-phippsiae and perhaps rubribracteolata. Each of the Bears Paw Mountains and the Sweet Grass Hills are much smaller in area than the Cypress Hills, nevertheless the former so far has nine species of hawthorn recorded while the smaller Sweet Grass Hills, only three so far found, in spite of hosting seven conifers compared to two for the Cypress Hills. From such types of consideration, one can deduce that the quantity of suitable receiving habitat, together with its continuity, nearness to source populations, the time available for colonization, as well as later unfavorable climatic bottlenecks are all relevant to the size, persistence and diversity of immigrating populations. Nearness to source populations needs to be further interpreted with respect to dispersal ability of different Crataegus species about which almost nothing is yet known in a precise way. Solutions to the problem of origin will thus be helped by greatly increased knowledge of dispersal ability and detailed habitat preferences among the different species and also by reliable inferences on the age of different Crataegus taxa involved in addition to the other factors mentioned. Therefore, the present nearly comprehensive lack of such hard information prevents for the time being obtaining firm answers about the origin of the Cypress Hills Crataegus flora so, instead one has to resort to inferences derived from general habitat preferences, relative abundance of same and current biogeography. TAXONOMIC PART Introduction Thirteen species of Crataegus occur in the Cypress Hills: C. douglasii Lindl., C. castlegarensis J. B. Phipps and Kennon, C. chrysocarpa Ashe, C. sheila-phippsiae J.B. Phipps & O'Kennon, C. macracantha Lodd. ex Loudon, C. sheridana A. Nels. plus the seven new species described below. Previous authors (Boivin 1967; Moss 1040 Journal of the Botanical R h Institute of Texas 1(2) 1959; Packer 1983; Scoggan 1978) had listed two—cC. douglasii and C. chrysocarpa (sometimes listed as C. rotundifolia Moench) or three (Breitung 1954), adding what he called 'C. columbiana' the only cited specimen of which that we have located proving to be C. aquacervensis, so our paper constitutes a major development in the understanding of this flora. The arrangement of species in this paper is according to small, rather narrowly defined phenetic groups treated as series. However, one of the newly described species (C. rubribracteolata), and the long-overlooked species (C. sheridana) considerably widen the concept of an existing series, in this case Macracanthae. In view of the evolving understanding of correct sequence of bifurcations in the Crataegus evolutionary tree from molecular work, a vague assignation like this seems entirely acceptable for the present. Even so, two new series ‘Cupressocollinae’ and ‘Montaninsulae’ are designated to handle other species that do not fit serial circumscriptions as currently understood. 1 I a J Characters used in the descriptions are the numerous, over 50, morphological standard 1 in careful Crataegus species descriptions. A recent addition to these is the case of bracteoles which have now been systematically recorded by the first author in descriptions of Chinese (originally planned for Flora of China) and North American (to appear in FNA) taxa. More meticulous study (Dvorsky & Phipps 2005a, 2005b) now allow it to be stated that nearly all bracteoles observed in Cypress Hills Crataegus belong to Dvorsky's types la and 1b (revised numbering follows thesis and forthcoming papers) which characterize the majority of Asiatic and North American species. Bracteole type 1 refers to the symmetric type found in all Crataegus species and is usually small, caducous, membranous, narrow or linear and slightly venous. Subtypes la and 1b are characterized by the presence of sessile or short-stipitate glands around the margins, in which the color when young for C. rubribracteolata (Plate 8a) is a solid clear red-brown (hence the name) although this often fades with age. Only a few Crataegus species have intensely colored bracteoles, the most striking being C. phippsii, also western Canadian and adjacent United States, in which they are crimson. Crataegus cupressocollina bracteoles, however, are closer to subtype 1c as they nearly completely lack marginal glands (Plate 6.1c). Nineteen of the more useful of the characters used in the descriptions and the keys are presented in Table 3 to sumarize the manner in which they vary between species. A number of these are continuous characters about which it is difficult to be more than very general in a table. For instance, leaf length in Cypress Hills Crataegus ranges from ‘small’, 3-4 mm long, to ‘large’, ca. 6-8 mm long. However, such characters may be still excellent discriminators, at least among species with dissimilar values. As there is nearly always fairly significant variation in leaf size on a given specimen the terms are limited to a central tendency, rather than including the extremes. Such central tendencies become more useful discriminators. Numerical equivalents are provided for the central tendencies in the caption to Table 3 while subjective terms are used for shorthand purposes in the table. An exception involves density of indumentum which is too complicated to assess numerically with any ease and, even if it were done, would provide significant difficulty for the user. Eight of the continuous variables appearing in Table 3 were those of particular value in separating certain species. These were sampled over a substantial range of individuals if such was available in order to calculate their ranges, means and standard deviations. These results are presented as box plots in Figures 5a to 5c and are discussed at relevant points in the text. We also try to give some measure of precision to the degree of leaf incision, the ‘leaf incision’ index or ‘LIT CIFT in Latin), which represents the percentage of width of the lamina diminished by a sinus as measured by the line from the midrib to that sinus parallel to the veins to the adjacent lobes and compared to the same line projected through the sinus to the intersect of the adjacent lobes. This may vary from 0% (no lobing) to 100% (blade cut to midvein). Identification In identifying Cypress Hills Crataegus, whether in the field or herbarium, most of the characters used in the descriptions may be valuable but additional characters, hard to specify precisely, often of stature, branch- ing pattern, foliage color and reflectivity, planeness and size of leaves at anthesis, precise anthesis order, 1041 IAW DL ‘(asuap ÁA) s 01 (snouqe|D) 9 :e»ue»seqnd Buo| 21 juenbay = 4 yep =P HOyYS = us “+p =] '€-Z — 9S '|-0 =O ‘Ueda 'ouenbay puejb əjonəd 9|091381Q = Peq jubuq = q 1>e/q = Iq adund = ind '96 UI |[1 Xew :u3dep 8qo| 'eeurunoe-s ‘JUN|G=| :ssaudueys aqo| sok =A SUM 2 M KIA — ^ K||ensn = sn unipeul = W 'UJUJG/-SG =] ‘WW 09-04 =W ‘WW Gy-SE =S :9BUE) [e Juan ‘aINjJewW ‘yua Jea] [eanas = ‘AS pus =S D9AJn29J = Dad pa=1 Apunbing = bing ‘WU +0 = 7] ‘WW Op-Sz =W ‘WLU OE-S | =S :y1Bua] uoy} yuid = xd juajed =ed obuejJo = JO ¡quoiseddo=0 pessaudde = dde :san¡en jo uoneue|dx3 ‘SUORE SIW A208 JO ISƏM SIUM AjjEWOU y "SIN ÁADOY JO sam IBUEIO + z : : : A Á A - - - A Á Á A UOISOJ3 [21318] 3 v-t Vane P vy-t v-t Ex GE a Ge (Sire Vine E S-t ‘OU Jonu ed jed ‘Yor udde-yed yaljed ys-1dde Jddex dal-jed ‘W jed ^N 291 ^N ys ys ys 7 JIN 'sego|-xA|e» t (S)osn 0 v-0 Sp Gat S-F 0 0 S-E Em red 0 0 e»ueoseqnd NIJ pal deep pal pal pal pau uq pal uq pal pal Jnd | jnd-ng ind Iq-und Ig IQF 372| OJO) YN pal +paJ paJ-JO pa1-10 pal pal +P9J pa Hing pel yp pel uunyd und Ig Hing Alea JO|OD 3n M M M M M M Ad yd yd yd yd yd yd M JO|O) Jayjue OL 0l 0c OL OL 0L OL OL OL OL Ol OL OL 0l ‘OU uauJeis 81-91 9|-vl 8l GOL Sl CIEE LC-41 9l gizel [C EE Sis Slc GEE gl=sSl (Ww) "UJelp ‘y el el al el e| e| el el el e| el e| 21/21 edÁA “Peq a v-€ € Po € S e Cl € 3 € € € S “bad 12eJg S 3 01 S-y S S S v-z E-Z S- v-z E-7 0 €-~ aduadsaqnd ped L e 9 8 OL 8 SL-OL OL OL Ll $8 OCc-8 | SC-0l OL (ueipeuJ) ‘yusi Jq-i 1q ‘plu JqTp J97J JP 1q "plu APA APA APA APA APA Aq p-4g Jq poom pjo JÁL WN W-S W-S HN W-S TW W W-S S S WN S $ WN u1bue| WOU} ‘AOS 'A9S "NOS W-4 WAS W- O 'A9S-O 1-0 +0 ‘AS © l l spuej6 Jejoned Vee S- S- € S-€ é €-Z ESC € "ed iv ESSE S-y sseudieus eqo| v ESE ¡E S-r v 9-5 S-€ v-E ESC ví be (b-JE-0 x0 v-t 3pis/ou 3qo| SC-SI O£-S1 SE-SC OCO ££-SC SC-SI SC-SI Oc-S Sese Te 22S 1501 OCSI e 'sn Xew Xew "xeuJ Xew HD Xew Xew SM Xew 'sn 'sn Xew (IIT) azis eqo| WN S S TN WS W-S W W-S S WN WN S S WN 9ZIS jee| 3 S a 5 ES ES = R 2 = S S g S & e S gy R. S S S & S 2 2 S E $ E: E SE a. j E] = = = 3 S S S 38 = S 2. S T mp pa alu Y "SUJOYIMEY S|IIH SSe1d A5 BUIAJNUIPI ur INEA YO sonsue1oeaeup ¡EDIBOJOYANON “E 318v]. 1042 Journal of the Botanical R h Institute of Texas 1(2) Thorn Length (mm) Thorn Curvature (deg.) K K x = K 607 m 15.05 Y : K Y K K 8 a 507 | 10.0% i K Ale 404 | 8 po E 30-2 | + | 5.04 F T T [| LI 204 | T K Y x 104 Dg y L| [| L| [| [| [| L| T AQUA CSTL CUPR DGL PURP RVA RVP SHPH SHSK T T T T T CSTL DGL PURP RVA RVP Species Species Fa. 5a. Box pl AES howing TT 4 NU ; pairt REE eee ee TUAM AW : 1501 pee J ; pair! MA MADE i : nde ).(left f vios TE " ANITA C. aquacervensis, CUPR = C. apres, CHRYS = € chrysocarpa, CSTL =C. castlegarensis, DGL = C. douglasii, PURP — C purpurella, RBR= = C sheridana, SHPH = C. sheila-phippsiae var. sheila-phippsiae, SASK = Leaf Length in Flowering Specimens (mm) [] T i T i [] [ U CHRYS PURP RBR RVP SHER SHPH SHSK URSO Species C rubribracteolata, RVA = C. rivuloadamensis, RVP = C. rivulopugensis, SHER = C sheila-phippsiae var. saskatchewanensis, nu = C ursopedensis. Leaf Length in Fruiting Specimens (mm) 907 Habitat a - Thickets b - Aspen Forest € - All 4 mi LSL T T T T T T T T CHRYS PURP RBR RVP SHER SHPH SHSK URSO Species Fic. 5b (box plots continued). (left) Leaf length, flowering specimens (mm). In the case of the two varieties of C. sheila-phippsiae measurements n RS Rn E PE c APA eden eta, : 1 e bud) t ee Fig. 5a for y y Jl Y acronyms of taxa. Length:Width Ratios, Leaves Fruiting Specimens 1.804 1.604 ] au 1.404 +: im | 1.20” T 1.00” I I I I I I I CHRYS PURP RBR RVP SHER SHSK URSO Species tcl 1043 Leaves, No. of Lateral Veins 21/5 Length of Longest 24 We 671 5-2 K 4= ae 3s y Į PURP RVP Species Efi Tat, PR y laft) | £ : I b. G IN J J V r acronyms of taxa. Length-Breadth Ratios, Fruit 1.505 1.25™ 1.005 I I CSTL DGL Species Fic CA (b pl + "A d) ia * * I g Lf ? el See Fig Sa for acronyms of taxa. idth. (right) Leaves, number of lateral veins more than 10% length longest. See Fig. 5a for etc. frequently permit recognition from a distance before key characters can be individually observed. This is often referred to as the ‘facies’ of a plant and its recognition comes with practice. Nevertheless, the facies has the potential to reinforce a species concept based on technical characters. Leaf shape and size are somewhat variable within species but are nonetheless important field characters. However, while extension shoot leaves are quite often in some species close to typical shape and size their considerable potential for plasticity suggests great caution in their use. Nearly all short shoot leaves found in the Cypress Hills belong to the principal leaf-shape group in North American Cra- taegus (more or less shallowly lobed, no veins to sinuses) but even so there may be significant specificity of detail. By far the most abundant form of Crataegus chrysocarpa for instance, should be quite easily recognizable from sterile material. And in the case of C. ursopedensis, the stimulus for the original search for came from a single sterile specimen with huge leaves noticed in her- barium of the University of Montana at Havre at a time when the remainder of its characteristics were quite unknown. It is, nevertheless, quite difficult to describe the exact form of leaves with the precision required for discrimination, especially taking into account some variation, and in this context the illustrations are intended to help. Also note that at anthesis leaves are still in a state of expansion and may be much smaller than at maturity. Anther color is important but anthers may be blackened by frost during morphogenesis and in any case usually become brownish by late flower. Nearly all Cypress Hills Crataegus taxa possess a mode of 10 stamens and only one, C. sheila-phippsiae, 20. ‘Ten’ and ‘twenty’, seemingly precise values, may 1044 Journal of the Botanical R h Institute of Texas 1(2) a WE Lake Riverhurst Hussars? * Beechy E i ‘ tra! Cu ILION IR LS THE DINOSAUR PROVINCIAL PAR Little Bo vers » servoir Lake " Vauxhall « 7 trons “Turin Milk Ridg Reser mer wS MILK Rivep £ RIDGE Mil E cs s Hiysdale Q. mm FORT BELKNAP INDIAN RESERVA TIP PINEY BUTTES Ne Ka E PLATE 1 T, kl £ tnt +kosanteal Mantana +) All Camel L Ļ : I : Lahn f^ Hills Tonoaranhical T reser I kinhiinh J EA ^n 1nnn la + IANN »nnn layl £L r P 1^^n Dalinf r YICCH— IVY I LATATA "n cnlorina: area CULO. Greer by shading. Scale: 100 km — 31 mm. n be taken as approximate, morphogenetic control being more or less imprecise in this matter. For instance, both stamens and style numbers may be radically reduced by low temperature damage during early flower development. Fruit color at full ripeness is basically either in the purple to black range or red. However, earlier, the purple to black fruit type may be plum-red or similar, while over mature red-type fruit may be a similar deep red. Also, some red-type fruit may be orange-red young. Therefore, with changing fruit color, comparison must be made at comparable points in the season. Pubescence of the inflorescence branches may be partially lost through the summer, generally leading to more variable indumentum in infructes- cence than inflorescence branches. The lateral face of the nutlet is another important characteristic to use in identifying Crataegus but it is easy to go wrong with this if the sides are not properly cleaned. Scraping the flesh carefully off the sides of typical (but not very narrow nor deformed) nutlets does not automatically reveal the pitting as this can remain full of flesh. So a fine probe must be used to see if excavation is pos- sible. Species such as C. chrysocarpa and C. sheila-phippsiae have plane to slightly convex (plump) or very shallowly concave sides, while others such as C. macracantha and C. douglasii have pitted or 'eroded' sides, the erosion of which may be quite irregular. The typical states for these characters are shown for each of 1045 INV DL F IPPS ans Y "(p 514 aos) suonpas jo suonisod 918 5-4 pue q-) 'g-v seur] *SINOJUO) W OSZL pue w OOOL 9u Huey | 8 "JS “DH Wed jepuroag SIIH ssaJdf) = ¢ “ys "pue se3 = Y :L£Z/LC XS V f ‘yao ejdew = € ‘gy oye] JOsaay ‘dy uayuu = 7 ‘Lp gy uo Á 19183 = L save sdew oue yim Hued y syuiod I 1 'pue|poow y 119618130 uonnquisip Buneoipur (uəə16 ur) Keano uim dew ¡e>1yderbodo) siji ssaadÁ) *z 31114 N M.601 itute of Texas 1(2) £L Journal of 1046 Pate? 6 Wille hahitate 1a) Aal Ranet Adame Fraak CK uith vallan fI T E Id ah Di fd arl yl Pr y y g I g (dark) i Tee a M PEE E a e Gained naar Flikwatar ADIRE (pale), grass | g g I i g " i ; £ era eT yeaa Hand ; laval land all vicihla: r, SE, I I g f pi g g 3. £, J MD e I J sel FRAAIE | I 5f £4 üt © xb M £, L | I£, J 4 g | g g pr pes (foreg 1275 m), SK. -—Y Y pan ATEN 1047 J OL: J f^! WM y + £f. Lil AIL FIHpp» anu , Y JI r riick DEPT. PLATE 4. Cyr qe : ann J Creek grassland, and aspen woodland (background). 1(2) 1@Xd5 "U gcc LAO “£007 das cL ‘YS 991) 1ƏJLYS je 493uno»ue £f T, FE 101) pue (1,08) 10ny (2 'z “bly uo UMOYs aye] swepy Jo MN WY OL “2? sr axe] SHIEH “y007 das “Aaja W 9g LL 3e MS Ájoreuut IAs nez9b3A auar0jOH SIIH 1} ‘YS eye] sued (q ls snba0301) Auew jo sisayue [nj Key ut s10y p Aq pe»ueued ] osje seu Kneay Ápejius Duio1b ay} Jo | ipe1dun əy} sayiid Dold siy, “W DOLL ‘Aaa ‘LZ WS JO} pne | | qugna snbaD301) *£007 LL das sawwNs ui MOUS (e :SNOLIBA *c 21d £L JUUITIdI OF 1048 ET X ^ PEA e i vere $ + ki m A x v 3 f RU. O us itd 1049 INV T mp pa alu Y DI 6 ! i Bud luu NAAA 4 rd LL LELEEEELEEEEEEELEEE EL L I udis inf TITIT T impri | 6 J J J > V TER £ > = x M = uw = a n = D E D S < “oO y ` c =< x es c n c L € = = ee n. h- 4 O — eR: c E Sz Sc S c O & 4 Sc S pe = n c c E uod a 2 — E e Epa] = € m vu mc = E © ` e c = ‘ I > c LE a YU AE oer 4 onm. w — A + os o = y nes 5 = 2 SES eom ES 1050 Journal of the Botanical R h Institute of Texas 1(2) EPA tE, £4 : £ J H £f. Use f. Wal Wt /IDD 0 y orno & 0'K 8508), near Eastend, SK. b) C. Prate 6.2. P purpurella (JBP & 0'K 8984), near Loch Lomond, CHPP, SK. Phipps and O'Kennon, Crataegus of Cypress Hills, Alberta and Saskatchewan Puate 7.1. Portraits of fruiting specimens of Cypress Hills ae a C cupressocollina (JBP 9156) at the rarely observed reddish stage; SK 271 above Adams Creek. b) C. cupressocollina (uncollected specimen at mid- ripe stage, later becomes glossy purple-black); location close to last. c) C. castlegarensis (JBP 2 West Butte, Sweet Grass Hills, MT. d) C. pupa P 0'K 8792), Elkwater, A te of Texas 1(2) P" £L JUUITIdI OF 1052 £n 'gy 'Ilo9QgeAun EHIE 113 AO0V L0 Gdl ) ~ " A A fh 2 (p “AS 99.) swepy a I-FII*'V/CZO AU 0 ddl] ^ £ c daan Á lar m F " A 311110 MY 0 0401) TI indand *) (q EE IU . F H Ves PsN d (Liv Phipps and 0K ] TAS £f MAT ; AIL 4 IC | aera | 1053 D EE SES n £r. TT ir - HDD O VF 0701. EM " h: E 1 Puate 7.3 gsi yp g ) (J J; , AB, b) macracantha {IDD O ni orno, Li: AL JAA AAT. O 124,11 J I a: £, 12? f. MAI) I O C L sy SAI, = K vor J at Y T u J J "T UWO); Cy Hills pl g Ily have smaller leaves but of tl g | shape. d) C. rubrit lata (JBP 8819); bundant, tightly MIA TWIST Ien " A , n TET o m SK , J rr at the Cypress Hills taxa in Table 3. Thus, careful users have an excellent chance of correct identification from good quality median-type material. Nevertheless, due care must be used! Following is the formal taxonomic treatment with keys to the species in fruit and in flower, detailed protologues for the new taxa, variably detailed treatments for the previously known species and line draw- ings and range maps (limited to the Cypress Hills) for all taxa treated. The maps on Plate 2 and Figures 1 and 2 provide many defined locations which may be helpful in the inspection of the range maps. Conspectus of Crataegus taxa recognized in the Cypress Hills ser. Cupressocollinae J.B. Phipps @ O'Kennon 1054 Journal of the Botanical R h Institute of Texas 1(2) £f. Ulle f. xr ob [IDDO n'y 0000 kaikes | £ feb Y J 7 Rv 21 UD Udal Ranch, Adams Creek, SK. b) C. sheila-phippsi katch IBP & 0'K 8848) ] th of Hay Cr., SK. ¢) C chrysocarpa (JBP rere V 21 F 4 Se g ILE h nf EIL + B.d)C ursopedensis (UBP & 0'K 8902); Oakes Ranch, south of SK 271. 1. C. cupressocollina J.B. Phipps & O'Kennon ser. Douglasianae (Rehder ex C.K. Schneid.) Rehder 2. C. douglasii Lindl. 3. C. castlegarensis J.B. Phipps & O'Kennon ser. Purpureofructi J.B. Phipps & O'Kennon 4. C. aquacervensis J.B. Phipps & O'Kennon ser. Montaninsulae J.B. Phipps & O'Kennon 5. C. rivuloadamensis J.B. Phipps & O'Kennon 6. C. purpurella J.B. Phipps & O'Kennon 7. C. rivulopugnensis J.B. Phipps & O'Kennon ser. Macracanthae (Loudon) Rehder, and others with red-fruited, eroded nutlets 8. C. macracantha Lodd. ex Loudon 9. C. rubribracteolata J.B. Phipps & O'Kennon 10. C. sheridana A. Nels. ser. Rotundifoliae Eggl., extended 11. C. chrysocarpa Ashe 1055 12. C. sheila-phippsiae J.B. Phipps & O'Kennon var. saskatchewanensis J.B. Phipps & O'Kennon 13. C. ursopedensis J.B. Phipps & O'Kennon KEY TO CYPRESS HILLS HAWTHORNS IN FRUIT (NOTE: FULLY RIPE FRUIT COLORS GENERALLY OCCUR AFTER ABOUT SEP 10) pu fruit burgundy to black (may be reddish-plum or similar color earlier). Sides of nutlets clearly eroded. 3. Thorns 3-5 cm long; fruit ellipsoid, glossy, pendent, frequently in large clusters; calyx-lobes long, prom 158: nent, narrow, + spreading aaco 3. Thorns 1.5-5.5 cm long; fruit ellipsoid to suborbicular or turbinate, not especially glossy, m both broad. hl | | pendent and in large clusters, size variable; calyx-lobes short to longer, relatively br 4. Thorns 1.5-3 cm long; calyx-lobes very short, + appressed; fruit in early September purple to blac Dy dead black by 1% Septembe lly + elli 1; infructescence branches glabrous; plant habit tall and thin; double and ane thorns not observed 2. C. douglasii 5. Fruit purplish or vinous in early September, very dark purple to almost black later, subglobose; infructescence branches variably pilose (difficult to see in some specimens); plant habit gener- ally dense and twiggy; double and triple thorns often present in small numbers, occasionally undant C. castlegarensis 3. 4, ud: 3.0-5.5 cm long; calyx-lobes a little larger, recurved rather than appressed; fruit in early September burgundy 6. C. aquacervensis 2. Sides of nutlets + mi to very shallowly concave Sur usually 1.5-3(-4) cm long, usually + stout, often + M and conical; pubescence of infru- +d le reddish plum to bur 4. C. rivuloadamensis 6. Thorns 2. ic 5.0 cm long, slender, ly always recurved; TE pubescence nil to moderate: fruit reddish or deep purple, glabrous to somewhat pubesce 7. Thorns mainly 2-3.0 cm long; leaves (3-)4-6- Pin per A often only 1.0-1.25 times as long as wide; fruit deep purple fully ripe, burgundy e 6. C. purpurella Te Pod. 2.5-3.5(-4) cm long; main lateral veins due usually 5-8 per side; leaves generally at least 7. C. rivulopugnensis as long as wide; fruit BEC red a ripe, red earlier ie ie fruit vue to dull red, occasio! rally somewhat plum re Nutlets distinctly eroded latera 9. ld ebr ovate t | insim| maturity; M lobes fairly deeply UM my d eglandular 8. C. macracantha 9. Leaves shallowl uit | dular but barely serrate; petioles glandular. NOME t | | sharply acute; thorns stout, usually 3.5-5.5(-7) 9.C cm long; calyx-lobes normaly PA eN to fruit. Ded veins not sig! ficat tly in pressed at maturity; calyx- lobes glan- C. rubribracteolata 10. Leaves relatively broa e slender, usually 4 E cm long; calyx-lobes in fruit usually + spreading their tips at most acute; ua ns 0. C. sheridana mo 8. ee eee ally. 11. Leaf-blades 3-5 cm long; thorns slender to moderately stout, usually 3-4 cm long. (Lak horns as above, or stouter and longer, 3-5 cm long (anthers ivory). 13. Infructescer ariably pubescent; ca. 10 stamen bases C. chrysocarpa 13: ee Hanon ca. 20 stamen bases T 12. Thorns slender, recurved, 2-4 cm long (anthers pink) El: 12. C. sheila-phippsiae var. saskatchewanensis 7.C. rivulopugnensis 11. Leaf-blades to 7 cm long; thorns stout, usually + recurved, 4-7 cm long. 14. Blades rhombovate, usually flat or concave C. ursopedensis 14. Blades ovate to broad elliptic, often convex KEY TO CYPRESS HILLS HAWTHORNS IN FLOWER OS Bs 2, 13a. (see note after sp. 13) 8. C. macracantha lul tinctly glandular-serrate 2 potios s andul D may oe ate small). 3 1.5-3 cm a Vd straight. "s lai 2. C. douglasii 4. c mne to densely pubescent. 1056 Journal of the Botanical R h Institute of Texas 1(2) 5. Flowers 12-15 mm diam; leaves lly not sharply lobed 3. C. castlegarensis 5. Flowers 17-21 mm diam; leaves Pay lobed 5. C. rivuloadamensis 3. Thorns 3- Y cm long, usually recurved. 6. Flowers 17-19 mm diam; inflorescences moderately hairy; thorns stout, 3.0-5.5 cm long aquacervensis 6. Flowers 11-17 mm diam.; inflorescences subglabrous to somewhat hairy; thorns slender, 3-3.5 cm long. 7. Leaves (3-)4-6 veined per side; blades mostly 2.5-4.0 cm long (fruit purple) 6. C. purpurella . Leaves 5-8 ^us per side; blades mostly 3-5 cm long (fruit red) 7. C. rivulopugnensis . Anthers white to cre 8. Twoto five year a m often orange-brown; bracteoles numerous, of varying size and form, + eglandular 1. C. cupressocollina 8. Two to five year old twigs gray or dull brown; bracteoles sparse to numerous, of not greatly varying form, margins glandular 9. Bracteoles in young inflorescence strongly rust-red; expanding bud-scalesred — |] ^ — 9.C.rubribracteolata 9. Bracteoles in young inflorescence much paler though may be tinted pinkish or reddish, particularly towards the edges; expanding bud scales colored or not, but never richly reddish throughout 10. Flowers 20 mm diam,; thorns stout, 4-7 cm long usually recurved 1. Many leaves at early anthesis very small («3 cm long), flabellate, with very prominent veins; blades + flat on expansion, or slightly keeled 13. C. ursopedensis . Leaves not unusually small at early anthesis, » 4 cm long, not flabellate, venation not especially prominent; many blades oft onv 13a. (see note after sp. 13) 10. Flowers 15-18 mm diam; o slender to stouter, mainly 3-4(-5) cm long, recurved or + straight. 12. Pedicels subglabrous; stamens 20 12. C. sheila-phippsiae 12. Pedicels pubescent; stamens ca. 10. 13. Blades often only 0.8-1. le as] with widest point near hyt der isely pubescent 10. C. sheridana 13. Blades 0.6-0.8 x as wide as | broadest near | hypanthi nly densely pubescent in a few aberrant individuals 1. C. chrysocarpa series Cupressocollinae J.B. Phipps & O'Kennon, ser. nov. Ter species: Crataegus cupressocollina J.B. Phipps € Similis ser. Purpureofructuum his proprietatibus exceptis: Planta gracilis; spinae longiores (ad 7 cm.), curvatae, tenues. Bracteolae paucae-glandulares vel fere eglandulares, et interdum ut videtur se versantes gradatim in squamas gemmarum. Stamina 10, antheris albis. Fructus in fasciculis pendulis, nitentes, atropurpurei, nigri in maturitate plena, purpurei in juventate; lobi calycis + patentes et acuminati ubi non erosi. Like ser. Purpureofructi except: Plant slender; thorns longer (to 7 cm), curved, slender. Bracteoles few-glandular, or nearly n and in some cases apparently grading into bud-scales. Stamens 10; anthers white. Fruit in pendulous clu , glossy, deep purple, nearly black when fully ripe, purple younger (initially plum-red); NAE + patent and pointed when not eroded. A sole species, below. 1. Crataegus cupressocollina J.B. Phipps & O'Kennon, sp. nov. (Plates 6.2a, 7a,b; Fig. 6). tv: CANADA. SASKATCHEWAN: Cypress Hills, Maple Creek Rur. Mun., thickets near Hay Creek, alt. 889 m 23 May 2003, J.B. Phipps & R. O'Kennon 8501 (HOLOTYPE: UWO; IsoTYPES: CAN, DAO, MO, SASK, TRT). Frutex, + erectus, 2.5-6 m altus; n 3-5 p cm o dE + numerosae vel paucae, ad duos annos nitenter atro-brunneae, + rectae i, virido-rubri, annotini nitenter rubro-brunnei vel atro- vel « valde curvatae 1 p i T i 1 i i is. Folia decid tioli 1-1.5 em longi in anthese, leviter pubescentes brunnei; in juventute, pauco- VPN. NUN 3-6(-8) cm longae in anthese, majores in maturitate, + ovatae vel late- dE in forma generali; apex acutus, basis late cuneata; latera 3-4 M acutis vel DAY d: iUd max. IFI 1596; I 3-4 2113 1 quad nervis per latus in foliis parvoribus, 5-7 in fi numerosis adpresso-pilosae d ERE in iuventute, + = glabrescentes fere glabrae Md sed aliquantulum pilosae O secundum nervum caducas, membranaceas ntralem: + teni lineares, pallide brunnescentes, usiter fere TEES ipd ed 15- 18 n mm diam.; po extrinsecum unum lobi calycis pilosi adaxialiter, glab ialiter, triangulares gini iter glandulo DL: Wat lA f de £ f. ENII AIL 4 e 4 Phipps and 0 kate! 1057 Fic. 6. Line drawing of C cupressocollina SI i ]: Ph ipp 2 0'K 8488, 8489, 8505, 8957 (fl ina): Phi 2 0'K 8618 (fruiti € d * J FF y e " Peery pee £ z " lnc M SCR NE i m J F y F FI HHTUICOCCIICC of JBP & 0'K 8469. Otl [ hole leaf abaxial surf. | [ I y gl fad f calvx-lobe, fruit and nutlets. S. Laurie- ) Bourque del.; scale bars equal 1 cm. 1058 Journal of the Botanical R h Institute of Texas 1(2) | D 1 A | it | ( MS y nung Qu q « i Er OS é Fic. 7. Map of C | Hing distribution in tl Cyt Une D | for Fin 2 (vide disputationem), antheris albis; styli 3-5. Infructescentia cum ramis pendulis; fructus 7-10 mm diam., usiter perfecte ellipsoidei, vinosi in Augusto exeunte-Septembro ineunte, atropurpurei in Septembro exeunte, glabri; lobi calyci angusti, + acuminati, patentes; pyrenae 3—5, dorsaliter vadositer sulcatae, lateribus cum foveis irregularibus vel cicatricibus diagonalibus. Bush, + more or less erect, 2.5-6 m tall; thorns 3—5(-7) cm long, fairly numerous to few, at 2 yrs shiny blackish-brown, + straight to quite strongly recurved, medium thickness; extending twigs at first sparsely pilose, greenish red, at 1 yr shiny, deep to reddish-brown, older orange-brown overlaid with gray. Leaves deciduous; petioles thinly pubescent young, 1-1.5 mm long at anthesis, with a few glands; laminas 3-6(-8) cm long at anthesis, larger at maturity, + ovate to broad-elliptic in general shape, acute at the tip, broadly cuneate at the base, sides with 3-4 pointed to somewhat rounded lobes, max LII 1596; venation craspedo- dromous, 3-4 veins/side in smaller leaves, 5-7 in larger; margins with numerous small teeth, gland-tipped at least at first, appressed hairy adaxially young, nearly glabrous below except for some axillary pilosity along the midvein; thinish. Inflorescences 5-15 flowered, the branches thinly to moderately densely pilose, bearing numerous, caducous, membranous, linear, pale brownish, generally nearly eglandular bracteoles. Flowers 15-18 mm diam.; hypanthium externally glabrous; calyx-lobes adaxially pilose, abaxially glabrous, triangular, margins finely glandular-serrate, + richly anthocyanic; stamens 0-10 (see discussion), anthers white; styles 3-5. Infructescence with pendulous branches; fruit 7-10 mm diam., generally nearly perfectly ellipsoid, burgundy in late August to early September, glabrous; calyx-lobes narrow, + acuminate, spreading; nutlets 3-5 dorsally shallowly grooved, sides + irregularly pitted or scarred. Common name.—Cypress Hills Hawthorn. Distribution (Fig. 7).—Found throughout the Cypress Hills of Saskatchewan in suitable habitats together with other species of Crataegus. In Alberta it is only known from a few specimens in the Battle Creek valley and one from Spruce Coulee Road. Outside the Cypress Hills we have recorded it from a couple of speci- mens in the Bear's Paw Mountains, Montana. The first known collection is by Breitung (1947), who called it C. columbiana, in the Cypress Hills Provincial Park, Saskatchewan. Eleven other collections were made pre-dating any of ours. These had been generally ascribed to C. douglasii but very substantial differences exist, see below, keys and box plots. In fact, it was the first author's observation of this interesting form of ‘C. douglasii’ in1998 at SASK that provoked our first visit to the Cypress Hills. Phipps and 0K ] TAS g £f LU: ; AIL 4 IC Ares | 1059 Fic. 8. Line drawing of leaf sp f C douglasii (upper), and C. chry pa (1 ). Speci 1: C. douglasii—left, unlobed common form in Cypress Hills DL i | ON‘, 0939 IRAIA) igl L | ly lal I£, DL i | O n'/y, Yodo do F ad (UWO); FÉ chrysocarpa—left, very lanate form, rare ; Cypress Hills, Phipps & 0'K 8237 (UWO); right, less put form, in the Cypress Hills, Phipps & 0'K 88991 as Bourque del.; scale bars — 1 cm. Foranew species ina fairly well collected region (Es lli is surprisingly listi i ] IMOPPCOVOU very common in many parts of the Cypress Hills. It is a tall, willowy, upright species, distinct in leaf form, with characteristic thorns and two to five year old orange-brown bark. The flower closely resembles that of species in ser. Douglasianae ptin anther colour and the plant is dark-fruited with laterally eroded nutlets like all members of that series but has much longer thorns (Fig. 5a.1). The ripe fruit of C. cupressocollina is quite glabrous and is plum-red to purple-brown in late August after which it ripens to a glossy purple-black. The large, pendulous clusters of fruit at once draw attention, as do, in flower, the reddish tips of the calyx lobes and the soft red-brown of the often nearly eglandular bracteoles approaching Dvorsky type 1c (Plate 6.1b). The bracteoles in this species appear to have some transition to the bud scales. The latter, when expanding, are also rather strikingly colored, although less so than in C. macracantha. Crataegus cupressocollina fits no existing series as currently conceived, but it might be placed in an expanded ser. Purpureofructi. Therefore, we are placing it in its own series, Cupressocollinae. The specific epithet refers to the Cypress Hills where it is far more abundant than anywhere else. Crataegus cupressocollina is, by a small margin, the first to flower of the local species. In spring 2003 extensive damage was observed to reproductive parts: black and shrivelled styles, blackened anthers and reduced stamen count (instead of being usually + 5 or + 10 as expected), which was probably a consequence of an extended period of low temperatures earlier in May which included values of -15? C at Cypress Hills Provincial Park meteorological station. series Douglasianae (Rehder ex C.K. Schneid.) Rehder. Wee species: Crataegus douglasii Lindl. 1060 J lof the Botanical R h Institute of Texas 1(2) Tow A n" «m NE) li i Y Fic. 9. Map of C. d Inet? Hictributian inthe f Mille R for Eia 2 J at T J Bushes or small trees; thorns very short (1 cm) to short (3 cm). Flowers small-medium (12-18 mm diam.), pink-anthered. Fruit smallish (6-10 mm diam.), when ripe very dark purple or black, paler, but not red when younger, calyx lobes short, blunt, + appressed, nutlets with pitted sides. Five species, mainly western montane North America, one disjunct to Great Lakes. The two species here are readily distinguished in the Cypress Hills by inflorescence pubescence and fruit shape (Fig. 5c) and to some extent by thorn characters and facies. 2. Crataegus douglasii Lindl., Edwards Bot. Reg. 8:1810. 1836. (Plate 7.2a; Fig. 8, upper). An often somewhat upright bush in the Cypress Hills, to 6-7 m; thorns short; inflorescence glabrous; an- thers pink; ripe fruit black. Other characteristics as key. A detailed description may be found in Phipps and O’Kennon (2002) and a further color illustration in Phipps et al. (2003). Common names.—Black Hawthorn, Douglas Hawthorn. Distribution (Fig. 9).—Widespread west of the Rocky Mountains, discontinuous in higher land through the southern prairies and northern Great Plains and in lowlands bordering the upper Great Lakes. Crataegus douglasii is locally common in the Cypress Hills in mesic habitats, particularly below aspen. It is often the only hawthorn in mesic north-facing high altitude draws over 1200 m and is rare to absent in drier hawthorn habitats or lower altitude thickets. In the Cypress Hills, C. douglasii is a more willowy and lanky plant than C. castlegarensis. Two leaf forms exist in the Cypress Hills, one with nearly unlobed blades, the other and rarer form with sharp and deep lobes. 3. Crataegus castlegarensis J.B. Phipps & O’Kennon, Sida 20:121. 2002. (Plate 7.1c, d). Distinguishing characters per key. A detailed description and illustration may be found in Phipps and O’Kennon (2002). Common name.—Castlegar Hawthorn. Distribution (Fig. 10).— Crataegus castlegarensis has a similar distribution to C. douglasii except that it 10'K TAS £f. LU: J AIL " IC batal 1061 Fic. 10 Map of C leg DOE SNNT Mille R for Eia 2 is not found in the Great Lakes region. The distribution in the Cypress Hills is concentrated above 1100 m and the species is perhaps more heliophilous than C. douglasii. This is the species that most likely has double, triple or even quadruple thorns. Sometimes the majority of thorns on the twigs are of this type but more likely the character will just be seen on a number of twigs or even not observed. In the Cypress Hills this species is usually a much denser bush than C. douglasii and the thorns average slightly longer than in that species (Fig. 5a.1) although there is some overlap. series Purpureofructi J.B. Phipps @ O'Kennon. Tre species: Crataegus okanaganensis J.B. Phipps & O'Kennon. Medium-large bushes to small trees; thorns usually moderately short (2.5-5 cm), somewhat stout. Flowers 12-16 mm diam., stamens usually 10, anthers DIS. mul often brilliant red when subripe, alternatively + vinaceous, deep FA to purple-black ripe; fruiting patent, somewhat accrescent; nutlets with + pitted sides. Seven species, western montane North America. 4. Crataegus aquacervensis J.B. Phipps & O'Kennon, sp. nov. (Plate 7.2c; Fig. 11). Tv: CANADA. ALBERTA: Cypress Hills Provincial Park, near entrance gate to Horsehoe Canyon, Elkwater, alt. 1302 m, 7 Jun 2005, J.B.Phipps & R.O'Kennon 8989 (HOLOTYPE: UWO; isotypes DAO, ALTA, UAC). Frutices, dense ramosi, 2—4 alti li pluri tl ino, f labri ini liocriter-vel atro-b i; V A it t cir 6.5) cm longae, + PEE peratro-castaneae. Folia decidua; petioli 30—4096 eeu laminae, EM m in sulco adaxiali, ferentes aliquot sessiles glandes; laminae 3-3.5 cm longae in anthese, 4—7 cm longae in maturitate, ovatae vel trullato-ovatae vel late np in e penal bases cuneatae vel late-cuneatae; acuti; acuti aa 4 per latus, max. IFI 15-259 g p 1 a 1 Pss. » | Ew] korme acutis vel subacutis; I p pra, venis (3-)45 p I SEI en 1 e | T Tt E d ] = & 1 fa = scabro-{ te I jue g I Inflorescentiae 5-12 fl i ali i 1 ] ] 1 f d membranaceas, pallide | + lineares glandulo-marginatas etus sores 15-18 mm diam.; E subglabrum vel Du vel Ede eae lobi pap lahri enm +tinfr nallidi 1h es P| glandulo- gend vel-serrati, 2-5 mm longi, c EU: petala alba, + 10 1 ; 13 a Pedicelli tuti EE n x pubescentes. Fructus 9-12 mm alti, ellipsoidei a late ellipsoidei vel turbinati, rubro-prunini primo, pr in pleno matu- 1062 Journal of the Botanical R h Institute of Texas 1(2) Fic. 11. Line drawing of C aquacervensis. Speci 1: Phipps & O'K 8753, 8769 (fl ina): Phipps & 0'Ke 8529 8549, 8803 (fruiting) AA L L I I £-L ea | £. L LASA o d 1 £ fl : (pee eee PE | fruitand nutlets. S. Laurie-Bourque F F J J J del.; scale bars equal 1 cm. n 10'K TAS £f. MAN J AIL " IC batal 1063 X E 4 £ £ Y x E Ng Fic. 12. Map of C. icdictrihntinnintha ( uic R for Fia 2 1 YN F J ritate, subglabri vel moderate pubescentes; lobi calycis typi to-recurvati, 2.5-4 mm longi; pyrenae 3—4(—5), faciebus lateralibus varie sulcatis, quamquam saepe vadositer. Densely branched shrubs, 2-4 m tall; extending shoots anthocyanin-rich, nearly glabrous; at one year medium to darker brown; older gray to dark gray; thorns generally abundant, 3—5.5(-6.5) cm long, + stout, usually recurved, at two years shiny, very dark red brown. Leaves deciduous; petioles 30-40% length of blade, pubescent only in the adaxial sulcus, bearing a variable number of sessile glands; blades 3-3.5 cm long at anthesis, 4-7 cm at maturity, ovate or trullate-ovate to broad-elliptic in general shape, base cuneate to broad cuneate, apex acute; lobes (2)-3-4, sharp, per side, max LII 15-2596; margins with many small acute to subacute as venation asas ileus cy impressed, (3—)4—5 per side; adaxial surface + densely apy ; mostly glal lly except for occasional dense fine pubescence along the sides of ES veins. Inflorescences 5-12 flowered; bene somewhat thinly to moderately densely pubescent, bearing , light brown, + linear, gland-margined bracteoles. Flowers 15-18 mm diam.; hypanthium subglal thin] y or moderately pubescent; calyx-lobes denticulate to glandular serrate, 2.5 mm long, VN glabrous on both sides, pale with some anthrocyanins or light brown; pet- als white, + circular; stamens ca. 10, anthers pale pink or pink, styles 3-4. Fruiting pedicels subglabrous to moderately densely pubescent. Fruit 9-12 mm high, ellipsoid to broadly ellipsoid or turbinate, reddish- plum early, dull purple when fully ripe, subglabrous to moderately densely pubescent; calyx-lobes typically erecto-recurved, 2.5-4 mm long; nutlets 3—4(—5), sides variably grooved, though often shallowly. Common name.—Elkwater Hawthorn. Distribution (Fig. 12).—Crataegt vensis is widely distributed in the northern part of the Cypress Hills from the Adams Creek area and Fort Walsh, north and west. The first record seems to be by McCalla (see cited specimens) from near Elkwater, Alberta, in 1930. Crataegus aquacervensis is most superficially similar to C. rivuloadamensis, C. rivulopugensis and C. purpurella, although it differs from all three in the + eroded lateral faces of the nutlets and differs from the last two in the longer and stouter thorns, greater hairiness of inflorescence and flower, and from C. rivuloadamensis in the larger, curved thorns. All four have different fruit color. Crataegus purpurella ripens 1064 Journal of the Botanical R h Institute of Texas 1(2) ds 13. Line drawing of C. rivuloadamensis. Speci 1: Phipps & O’K 8739, 8963 (fl ina): Phipps & O’K 8224, 8543, 8907 (fruiting). F FF J J FT Ee ST RENT < (RA? Seal PERR AE REIR d fi : CREDAT eee ECT , fruitand nutlets. S Laurie-Bourque tt F 7 J , , del; scale bars equal 1 cm. nL NV f de £ f. ENII AIL de Je I 4 ^L Phipps and 0 , Crataegus of Cy] 1065 from burgundy to deep purple, C. rivulopugnensis is essentially red, C. rivuloadamensis is deep red to rich burgundy and C. aquacervensis ripens from deep red to deep purple. Crataegus aquacervensis is distinctive also on account of possessing clearly eroded sides to its nutlets and for this reason might be best placed in series Purpureofructi. Crataegus aquacervensis is, however, not a particularly typical member of this series. The species is named for the settlement of Elkwater in the north-west of the Cypress Hills near where C. aqudcet vensis is particularly abundant. Series Montaninsulae J.B. Phipps & O'Kennon, ser. nov. Wer species: Crataegus purpurella J.B. Phipps & O'Kennon. Frutices vel interdum arbores D un uon tenues, longitudinis onus n 4 En iu adaxialiter NU GP. dissimilia a lO, antherisroseis. F pyrenarum o E E MA UM levibus vel Ente eU Shrubs or occasionally small trees; thorns typically slender, of medium length (2-4 cm). Leaves subglabrous above, unlike most Crataegus species in the region. Stamens 10, anthers pink. Fruit of medium size, purple or red; sides of nutlets smooth or slightly concave. The description covers two species, C. purpurella and C. rivulopugnensis; C. rivulodamensis, much hairier, with large fruit and frequently, short, stout, + conic thorns, is also treated here but may not be very closely related. AII three are, on present knowledge, Cypress Hills endemics. 5. Crataegus rivuloadamensis J.B. Phipps € O'Kennon, sp. nov. (Plate 7.3a; Fig. 13). Tv: CANADA. Algerta: Cypress Hills, Cypress Hills Provincial Park, eastern slopes Medicine Lodge Coulee, Murray Hill Road, 1182 m, 15 Sep 2004, J.B. Phipps & R. O'Kennon 8800 (HoLoTyPE: UWO; isorvrEs: CAN, DAO, ALTA, SASK, TRT) Frutex, late-ramificatus, vel arbor parva, 2.5—4(—5) m alta; spi 1.5-2.5(-3) cm longae tae (brevi j t ) politae per atrorubro-brunneae post secundum annum, veteriores atrocinereae; ramuli prorecti pubescentes, rubentes, atrovirides; anno- tini nitenter atrorubro-brunnei; ramuli post secundum annum atrorubri et cinerei; veteriores cinerei. Folia decidua; petioli 25-35% uus laminae, tenuissimi, dense mos in i M M UE vel cum paucis glandibus sessilibus, vel cum d basem; maturitate 4—8 cm longae, + ellipticae vel late-ellipticae vel on. ue in forma generali; bases cuneatae vel bp cuneatae; gee cuneatae; lobi 2-4 per latus, max. IFI (0-)5-20(-25%), apicibus subacutis vel late cuspidatis; margines cum d serratis vel serratis; venati p P venarum E principalium lateralium 4-6, vix i l ia adaxialis « d | l so-scabra pu venas, superficiebus ie = cum dispersis pilis m RM 4—10 ce rami dense CAES membranaceas, + rubras, angustas, dense glandulo-marginatas bracteolas. Flores 17-21 mm diam.; hypanthium dense pubescens sed interdum in apice; lobi calycis late triangulares, + 3 mm longi, glabri adaxialiter, pubescentes abaxialiter, marginibus glandulo-denticulatis, x viridibus; petala suborbiculata, pallidissime cremea; stamina 10, antheris roseis; styli 3—5. Infructescentiae cu 5mm longi, anguste-triangulares, + patentes, recurvati; pyrenae 3-5, l l l vadositer E in (1- ia a -fructatae. Fructus 10-12 mm alti, subglobosi vel late turbinati, subviolacei, serius atrorubri, rubroprunini et vinacei, al E concavis, fort Shrub, wide-branching, or small tree 2.5—4(—5) m tall; thorns 1.5-2.5(-3) cm long, + straight, the shorter conical and very stout, glossy very dark red-brown in second year; older dark gray; extending shoots pu- bescent, dark reddish-green; one year old shiny dark red-brown; two year old dark reddish and gray; older gray. Leaves deciduous; petioles 25-35% length of blade, quite slender, densely pubescent in sulcus abaxi- ally, none to few sessile glands, or stipitate glands at leaf base; laminas half-grown at anthesis, at maturity 4—8 cm long, + elliptic or broad-elliptic to rhomb-elliptic in general shape, base cuneate to narrow cuneate, apex acute; lobes 2-4 / side, max. IFI (0-)5-20(-25)%, tips subacute to broadly cuspidate; margins with small crenate-serrate or serrate teeth; venation craspedodromous, 4—6 pairs of hardly impressed main lateral veins; pubescence adaxially + dense appressed scabrous-pubescent only, persisting, abaxially appressed scabrous-pubescent on the veins, the surfaces glabrous or with scattered hairs young. Inflorescences 4-10 1 flowered; branches densely pubescent bearing caducous, , somewhat reddish, narrow, densely gland-margined bracteoles. Flowers 17-21 mm diam.; hypanthium densely pubescent except sometimes at the top; calyx-lobes broad triangular, + 3 mm long, abaxially glabrous, adaxially pubescent; margins glandular-denticulate, greenish; petals suborbiculate, palest cream; stamens 10, anthers pink; styles 3-5. Infructescences few (1—5(-6))-fruited. Fruit 10-12 mm tall, subglobose to broad-turbinate, pale lake, turning 1066 J | of the Botanical R h Institute of Texas 1(2) MW A AW Fic. 14. Map of C. rivuload fc lictriburtian in tha (^ TERES for Eia 2 through deep red to reddish plum and burgundy, generally rather hairy; calyx-lobes 4—5 mm long, narrow- triangular, + patent to recurved; nutlets 3-5, sides plane to very shallow concave, back strongly ridged. Common name.—Adams Creek Hawthorn. Distribution (Fig. 14).—Widespread in the upper (mainly above 1150 m) parts of the Cypress Hills with the great majority of the records found west of SK271 and SK615. Occurs in thickets, in open places and under Populus species. Not recorded from elsewhere. The extremely dark young wood and short, often + conical thorns show a superficial resemblance to species of the Eurasian section Sanguineae. However, the unpitted nutlets are a marked difference. The flow- ers appear to be palest cream rather than pure white. Crataegus rivuloadamensis is named for Adams Creek along whose upper reaches it is quite plentiful and where we first recognized it as distinct. Early collections of C. rivuloadamensis are by Breitung in 1947 and Moss in 1952 (see cited specimens for both). They both come from unspecified locations in the west of the Cypress Hills in Alberta. Breitung called his specimen C. columbiana and Moss called his C. chrysocarpa. 6. Crataegus purpurella J.B. Phipps & O'Kennon, sp. nov. (Plates 6.2b, 7.2b; Fig. 15). Tv CANADA. Saskarcu- EWAN: Cypress Hills, Centre Block, spillway of Loch Lomond, alt. 3930 ft, 15 Sep 2003, J.B. Phipps & R. O'Kennon 8557 (HOLOTYPE: UWO; isotypes: CAN, DAO, SASK, TRT). Frutex dense ramunculiformis vel arbor parva, 2-3.4 m alta; ramuli prorecti + glabri, potius anthocyananei; annotini mediocriter vel atrobrunnei, aliquantum nitentes, cinerei post secundum annum; spinae frequentes, 2-3.5 cm iun c rectae de minus curvatae, T5 atrobrunneae annotinae. Folia decidua; petioli 30—4096 longitudinis laminae, tenues i, ferentes 1-paucos, parvos, sessiles glandes; laminae 2.5-4.0 cm longae in anthese, 3.5—4.5(—5) cm in maturitate, + late rhombeae vel late ellipticae in forma generali, 0.7-1.0 latiores quam longa in maturitate; bases cuneatae, apices acutae vel subacutae; lobi 2-3 per latus, max. IFI 15-2596; Mcd cum pu acutis vel us dentibus; venatio POL PARE venis (3 mis : x latus; d eoo sca- br ialiter + glabrae subglabri vel sparsim pilosi, mus caducas, membranaceas, pallide brunneas, angustissimas, uri ndulo-marginatas bracteolas. Flores 13-16 mm diam.; diced RTT obi calyci 3 mm s mini Fora a abaxialiter glabri, marginibus 10 e ulatis . Infructescentiae 1-4 fructatae; pedicelli subglobosi vel sparsim pilosi. s (9211-12 mm alti, ellipsoidei vel subglobosi, E vel rubro-vinosi in juventute, 1067 /f JN m 0012 [fl ONY, 8614, 8871 (fruiting). Mags show whole Fic. 15. Line drawing of C purpurella [ equal 1 cm. , fruit and nutlets. S. Laurie-Bourque del.; scale bars 1068 J lof the Botanical R h Institute of Texas 1(2) N Ng "SOME ds Å Fic. 16 Map of C purp Ila A HI : : r Or Wille D [ for Fia. 2 atropurpureae in maturitate; lobi calycis 4-4.5 mm longi, acuti, erosi vel patentes; pyrenae 3-4, lateribus sulcatis longitudinaliter vel vadositer concavis, dorsaliter vadositer x profunde sulcatae. Dense twiggy bush or small tree 2-3.5 m tall; extending twigs + glabrous, rather anthocyanic; at 1 year medium to deep, somewhat glossy brown; two year old twigs gray; thorns 23.5 cm long, rather frequent, straight to slightly curved, slender to somewhat stouter, deep brown at one year, nearly black at two years. Leaves deciduous; petioles 30-40% of length of blade, slender, pubescent in adaxial sulcus, bearing 1-few small sessile glands; blades 2.5-4.0 cm long at anthesis, 3.5—4.5(—-5) cm at maturity, x broad rhombic to broad elliptic in general shape, 0.7-1.0 x as broad as long at maturity; base cuneate, apex acute to subacute; lobes 2—3/side, max. LII 15-2596; margins with very small subacute to acute teeth; venation craspedo- dromous, (324-6 per side; appressed scabrous-pubescent adaxially young, often +glabrescent later, + glabrous abaxially. Inflorescences 6-15 flowered; branches subglabrous to thinly pilose, bearing caducous, membranous, pale brown, very narrow, gland-margined bracteoles. Flowers 13-16 mm diam.; hypanthium glabrous to sparsely pilose; calyx-lobes 3 mm long, triangular, adaxial surface pubescent, abaxial surface glabrous, margins glandular-denticulate, generally pale green; petals white, circular; stamens 10, anthers pink; styles 3-4. Infructescence with 1-4 fruit, pedicels subglabrous to thinly pilose. Fruit (9211-12 mm tall, ellipsoid to subspherical, deep red to red-burgundy younger, deep purple fully ripe; calyx-lobes 4—4.5 mm long, pointed, spreading or eroded; nutlets 3-4, longitudinally grooved or very shallowly concave, shallowly or more deeply furrowed dorsally. Common name.—Loch Lomond Hawthorn. Distribution (Fig. 16).—C. purpurella is a species widespread in the eastern half of the Cypress Hills with an eastern limit near the provincial park headquarters in Saskatchewan and is not known elsewhere. It is common in thickets along north slope draws. The most similar species is C. rivulopugnensis, with which C. purpurella is allopatric. Both are slender, delicate bushes in the shade of aspen but much denser in the open and both have shortish slender thorns. Crataegus purpurella differs from C. rivulopugnensis in generally much smaller, fewer-veined leaves and greater thorn curvature (Figs. 5b.4, 5a.2 ) as well as in the deep purple, rather than red, fruit at maturity. The name 1069 c Y. DE 0 fv, Q7OC [fruit Y M | hal Fic. 17. Line drawing of C. rivulopugnensis. g pug EVER | ruit and nutlets. S. Laurie-Bourque del.; scale bars sF F 7 equal 1 cm. 1070 J lof the Botanical R h Institute of Texas 1(2) 10W] A ME) f j wi Y Fic. 18. Map of C. rivulop ds RT "THIS for Ein 2 refers to the deep purple fruit fully ripe, the small leaves and the delicacy of the species in the usual form in which it is seen. 7. Crataegus rivulopugnensis J.B. Phipps & O'Kennon, sp. nov. (Plates 5c, 7.2d; Fig. 17). tee: CANADA. SASKATCHEWAN: Cypress Hills, Reno Rur. Mun., high draws above Shafer Creek, alt. 1258 m, 13 Sep 2003, J.B. Phipps & R. O'Kennon 8533 (HOLOTYPE: UWO; isotypes DAO, CAN, SASK, ALTA,TRT Frutex vel arbor parva, 2-5 m alta, gracilis nisi in loco aprico; ramuli prorecti primum sparsim pubescentes, glabrescentes; annotini atrorubento-vel atropurpureo-brunnei; mediocriter brunnei post secundum annum, veteriores atrocinerei; spinae frequentiores vel frequentissimae, 2.5-4 cm longae, rectae vel recurvatae, tenues; nitentdissimae atrorubro-brunneae post secundum annum. Folia decidua; petioli 25—3096 longi quam laminae, pubescentes solo in sulco adaxiali, cum paucis vel pluribus glandis sessilibus; laminae 4—6 cm longae in maturitate, aliquantalum minus in anthese POE late eue vel ovatae in forma ap basis cuneata ie late neata vel curvata; apex acutus; lobi 3-4 per latus, max. IFI 15 venatio craspedodroma, venis lateralibus us principalibus 5-8 per latus, le mu EUR adaxialiter ur vel sparsim pubescentes in maturitate; abaxialiter perf Inflorescentiae 5-15 Dora es MR s cs piles ferentes a etiam in n jene md du uem pallide brunneas, glandulo p ) 16mm n hypanthium glabrum vel + pilosum ad basem; lobi calycisca ca. 4mm ur triangulares, irid ginibus pallidi g glandulo-denticulatis, adaxialiter sparsim pilosi, al ularia, alba; stamina ca. 10 heris pallid is vel i li 4 in floribus visi. Infructescentiae 1-4 fructatae; e. abo vel moderate pubescentes; fructus 8-10(— 12) mm alti, psoe a suborbiculares, rubri primo, claro rubri vel atrorubri in maturitate TR pon Ha w a ds mn tr ads pendio serrati, e oe NG vel recurvati; pyrenae 3-5, d in pyrenis angustissimis. Bush or small tree 2-5 m tall, slender unless in full sun, extending twigs thin-pubescent at first, glabrescent; one-year old twigs deep reddish- or deep purple-brown; two-year old twigs mid-brown, older deep gray; thorns moderately to very frequent, 2.5-4 cm long, straight to recurved, slender, bright shining dark red- brown at two years. Leaves deciduous; petioles 25-30% of length of blade, pubescent only in the adaxial sulcus, with few to several small sessile glands; blades 4-6 cm long at maturity, a little less at full anthesis, broad elliptic to ovate in general shape; base cuneate to broad cuneate or rounded; apex acute; lobes 3-4/side, ; venation craspedodromous, main T 1 max. LII 15-2596; margins with 1 INV. ,C " £f. MAN J AIL " IC batal 1071 X E 4 - £ € Y x Fic. 19. Map of C ha distribution in the Cyj Hillc R | for Fia 2 1 J un by | El lateral veins 5-8 pairs, slightly impressed; p thinly to moderately densely scabrous-hairy young, subglabrous to thinly scabrous pubescent at maturity, abaxially completely glabrous young except for sometimes small axillary tufts of hair. Inflorescence 5-15 flowered; branches very thinly to moderately pilose, bearing sparse, even when young, caducous, very narrow, membranous, light brown, gland-mar- gined bracteoles. Flowers (only few plants seen at full anthesis) ca. 16 mm diam.; hypanthium glabrous to somewhat pilose towards the base; calyx-lobes ca. 4 mm long, triangular, margins glandular-denticulate, adaxially sparsely pilose, abaxially glabrous, green with pale edges; petals + circular, white; stamens ca. 10, anthers pale pink to pink; styles 4 in flowers seen. Infructescence 1-4 fruited; branches glabrous to moderately pubescent; fruit 8-10(-12) mm tall, ellipsoid to suborbicular, red early, bright to deep red fully ripe, glabrous to thin pubescent; calyx-lobes triangular, glandular-serrate, spreading—reflexed to recurved; nutlets 3-5, dorsally furrowed, sides + plane to shallowly and smoothly concave, the concavity deepest on narrowest nutlets. Common name.—Battle Creek Hawthorn. Distribution (Fig. 18).—Crataegus rivulopugnensis has a wide distribution in the western part of the Cypress Hills from Adams Creek westwards. It is not known elsewhere. There appear to be no earlier col- lections than our ow Although the ot of the ripe fruit is quite different (red vs. deep dion e E is in other 11 15 respects similar to C. purpurella, differing most obviously in its almost exactly distribution, gener- ally greater leaf size and number of lateral veins per side (5-8 vs. 4-6). Both these species have less hairy adaxial leaf surfaces than most other Cypress Hills hawthorns, indeed they may be adaxially subglabrous at maturity. series Macracanthae (Loudon) Rehder. Tre species: Crataegus macracantha Lodd. ex Loudon. Shrubs, sometimes large, seldom tree-like; thorns stout, of medium (3-5 cm) to great (occ. 11.5 cm) length. Petioles eglandular. Generally very hairy in the inflorescence; anthers pink or white; calyx-lobes with gland- tipped very short to long teeth. Fruit orange-red or red when ripe, nutlets deeply pitted laterally. 1072 Journal of the Botanical R h Institute of Texas 1(2) b FF J FF Fic. 20. Line drawing of C rubribracteolata. Speci 1: Phi & 0'K 8650, 8656 (fl ina): Phi & 0'K 8819, 8823 (fruiting) Mags £l EET 1L I Klan sai £. F F 4 J Ink show emerging leaves with enlarging bud-scales, young i y I leaf adaxial surface, fl i ] caly , fruit and nutlets. S. Laurie- Bourque del.; scale bars equal 1 cm. Phipps and 0 , Crataegus of Cypress Hills, t 1073 Three species as normally understood, widespread in the United States and southern Canada to which C. rubribracteolata and C. sheridana are provisionally added. Both the latter have red fruit and pitted nutlets but the former has highly glandular petioles and the latter also has glandular petioles but not very stout thorns. Neither of these characters fits the traditional concept of the series. Both of these additional species also have quite strong superficial resemblances to C. chrysocarpa. 8. Crataegus macracantha Lodd. ex Loudon, Arbor. Frutic. Brit. 2:819. 1838. (Plate 7.3b). Crataegus macra- cantha Lodd., nomen nudum, in Cat. Pl. 41, 1830, per macrocantha. Crataegus succulenta Schrad. ex Link, Handb. Erkennt. Gewáchse 2:78. 1831 (in part). Crataegus occidentalis Britton, Bull. New York Bot. Gard. 1:448. 1900 A recent line drawing of this species may be found in Phipps (1997). Differentiating characters per key and it is one of the most easily identified species of the Cypress Hills with its eroded nutlets and eglandular petioles. Crataegus macracantha is the 10-stamen counterpart of C. succulenta which species is entirely eastern. In western North America west of the Rocky Mountains C. macracantha is almost always white-anthered but in the Cypress Hills it is always pink-anthered. Common name.—Succulent Hawthorn, in part. Distribution (Fig. 19).—Crataegus macracantha is a fairly common hawthorn in the Saskatchewan part of the Cypress Hills although mainly restricted to the lower altitudes and more xeric habitats, like C. chrysocarpa. It had been reported for that province, but not specifically from the Cypress Hills, by Harms (2003) under C. succulenta, but it is not in Breitung's (1954) list nor seen among his collections. Crataegus macracantha may flower later, or much later, than sympatric C. chrysocarpa and is also char- acterized by bright coral-red expanding bud-scales (Plate 6.13) when C. chrysocarpa is already leafed out as well as by eglandular petioles and laterally pitted nutlets. This species generally has very stout, recurved thorns and extremely dark one year old twigs and thorns. 9. n am rubribracteolata J.B. Phipps & O'Kennon, sp. nov. (Plate 6.1c, 7.3c; Fig. 20). T»: CANADA. ASKATCHEWAN: Cypress Hills, Maple Creek Rur. Mun., roadside off SK 724 near Downie Lake, alt. 860 m, 17 Sep 2004, J.B. Phipps & R. O’Kennon 8819 (HOLOTYPE: UWO; isotypes: CAN, DAO, SASK). Frutex erectus, multiramosus in aperto us ramosus in umbra, 2-3 m alt | g I ] forul li prorecti appresso-] b t i br unnei; post sec undum annum rubro-brunn i teri i i vel atrocinerei; spinae plerumque numerosae, 3.5—5.5 cm longae, generaliter ro rectae vel leviter curvatae; + validae; 1 ] j brunneae. Folia decidua; APN plerumque 30-35% longitudinis la 1 i lco adaxiali, ferent lt ; laminae 2.5-3.5 c d in anthese, in maturitate 4—5 cm longae in h 5-7 cmin n * ovatae a ovato- as basis cuneata vel late cuneata; apex acutus vel acuminatus; lobi 5-6 per latus, acuti vel acuminati, max. IFI 25-35%; margines cum De ae acutis dentibus, cae pe in maturitate; venatio raspedodroma, venis eae (526-7 perl br L praeter o ilos secundum mediam venam proximal Inflorescentiae (628-12 e ramuncul + tomentosi, ferentes 1 E ] z I g g Flores 12-15 mm diam.; hy paru im +t t ld 1 lobi calycis 3—5 ee triangulares inil landulo-denticulati 1 1 >J: ] TE y Real S E deed 1 1h abundantes, caducas, angustas I alba; stamina ca. 10, antheris oues vel cremeis; eae 4. Infructescentiae e plerumque 3 3-10 f licelli den ld 1 | E y ARCO 1 1 Fructus 9-13 mm alti, 1 I g j 1 E nisi aurantiaco-rubri in is sparsim vel KE ] | lobi calycis pl i, 3-5 mm longi; pyrenae 3-4(5)cm. altae, + profunde concavae lateraliter, dorsaliter ST Erect multistemmed bush in open, wide branching in shade, 23 m tall; expanding bud-scales bright ru- fous-red; extending twigs appressed pubescent; at one year mid-brown; at 2 yrs dark red-brown, older gray or dark gray; thorns generally plentiful, usually 3.5—5.5 cm long, straight to slightly curved, + stout, at two years dark blackish-brown. Leaves deciduous; petiole usually 30—3596 length of blade, densely pubescent in the adaxial sulcus, bearing sessile glands in the open, fewer in shade; blades + ovate to ovate-trul- late, 2.5-3.5 cm long at anthesis, at maturity 4—5 cm long in open, 5-7 cm in shade; base cuneate to broad cuneate; tip acute to acuminate; lobes (4—)5—6 per side, acute to acuminate, max. LII 25-33%; margins with 1074 Journal of the Botanical R h Institute of Texas 1(2) 10w] A ae In A | T eL. | i 1 1 N "H 1 2 S 1 | í A | | ABA | ES A QU ` mm Xs nt 0 i ' a © ' | QS E g. j d e PT M (1 : Ni EE ' Fic. 21 Map of C. bril I listribution in tl Or Hille R i for Ein. 2 small, regular, acute teeth often subacute at maturity; venation craspedodromous, lateral veins (5-)6—7 per side; adaxial pubescence + dense scabrous-appressed at anthesis, variably persistent to maturity, abaxially glabrous except for sparse hairs along the mid-vein proximally. Inflorescences mainly (68-12 flowered; branches + tomentose, bearing abundant, caducous, narrow, membranous, deep orange-red or fading, densely gland-margined bracteoles. Flowers 12-15 mm diam.; hypanthium + tomentose or densely pubescent; ca- lyx-lobes 3-5 mm long, triangular, margins glandular-denticulate, pubescent adaxially, glabrous abaxially, greenish or pale, the tip somewhat reddish or paler; petals circular, white; stamens ca. 10, anthers ivory or cream; styles 3—4. Infructescences usually 3-10 fruited; branches densely or very densely pubescent. Fruit 9-13 mm tall, generally in + compact clusters, + ellipsoid to subglobose, red early, bright red fully ripe or more orange-scarlet in shade, sparsely, or more usually fairly densely pubescent; calyx-lobes usually ap- pressed, 3-5 mm long; nutlets 3-45), + deeply concave laterally, dorsally furrowed. Common name.—Red Bracteole Hawthorn. T Distribution (Fig. 21).— Crataegus rubribracteolata is common to abundant throughout the Cypress Hills in ue habitats. We have one record from Montana, Hill Co., Bear’s Paw Mountains. Crataegus 1 rubribracteolata was first collected by S.E. Clarke in 1937 at Piapot, M O We have so far noted no other collections of this species besides our own. The characteristic upright habit of this species when grown in full light helps recognition. It is very similar in leaf shape, inflorescence and indumentum to C. ursopedensis, though the mature leaves average smaller except in shade forms and the inflorescence indumentum is denser. The much shorter leaves of high-light thicket situations (Fig. 5b.2) readily contrast with those of C. ursopedensis but in more shaded habitats the larger leaf dimensions of C. rubribracteolata are similar to C. ursopedensis. It is thus often quite easy to confuse with that species unless technical characteristics are carefully considered. The flowers are smaller and in more compact inflorescences and the bracteoles and bud scales strikingly orange-red in C. y een | rubribracteolata (particularly striking in the young inflorescences, Fig. 6.1c), whereas the bracteoles of C. ursopedensis are pallid or only lightly tinged with green or pale red, especially towards the tips and much more heavily gland-margined (Fig 6.1d). Intense reddish tints may also be seen as the tips of calyx-lobes in Phipps and 0K ] TAS £f LU: ; AIL 4 IC Ares | 1075 D yoja Fic. 22. Line drawing of C. h » c : I. Dh 0 MY, 2002214 : Y. Dh: 0 NY, 000NM latinn) AA L AAA ae | nl Ļ PEE D e| £. fl . per lal surface F F 7 J , , 1076 I Lat +l Dat eal D hi PC f Texas 1(2) Tow A Fic. 23 Map of C. h id Ji il H H 1) O Wille D [ for Fia. 2 young flowers of C. rubribracteolata. In fruit the shortish calyx-lobes are practically always tightly appressed and the nutlets have a well defined lateral concavity. The species is named for the reddish bracteoles. This species and C. cupressocollina are the two earliest Cypress Hills hawthorns to flower. 10. Boon sheridana A. Nels., Contr. Rocky Mtn. Herb. 4:370. e Tar 7. 3d; Fig. meee Tyee: U.S.A. Wro NG: Sheridan, 25 Jul 1901, A. os 8673 (LecTOTYPE designated here: RM!). US for a p of 24 July. Bush 23.5 cm tall; extending shoots pubescent, at one year light brown to olive-tan or mid-brown, at two years dark gray or reddish brown overlaid dark gray, older gray; thorns + frequent (2.5—)3—5(-7) cm long, usually recurved, slender to moderately thick, at two years very dark shiny red-brown. Leaves deciduous; petioles 40-50 96 length of blade, + pubescent, especially in the adaxial sulcus, bearing several to many small sessile glands; blades 3.5—5(-6) cm long at maturity, nearly full grown at anthesis, at least some usu- ally broad ovate or suborbicular in general shape, remainder usually + ovate; base cuneate, broad-cuneate or + truncate to subcordate on wider leaves; tip acute; lobes (324(—5) per side, acute and shallow, max. LII 10—20(-25) 96; margins with small, subacute teeth; adaxial surface densely appressed scabrous-pubescent young, variably persistent; abaxially with glabrous surface, veins glabrous to + pubescent, this barely persist- ing. Inflorescences 5-15 flowered; branches + densely pubescent, bearing caducous, membranous, pallid, narrow, gland-bordered bracteoles. Flowers 14-16 mm diam.; hypanthium + tomentose below, dense short- pubescent higher; calyx-lobes triangular, + pubescent adaxially, glabrous abaxially, greenish, the margins glandular-serrate; petals + circular, white; stamens ca. 10, anthers ivory; styles usually 4—5. Infructescences 1-6 fruited, branches usually densely pubescent. Fruit 9-11 mm tall, usually broadly ellipsoid to suborbicular, bright red, + pubescent (? very rarely subglabrous), calyx-lobes + spreading to reflexed; nutlets usually 3 or 4, sides + roughened with shallow irregular pitting or diagonal scarring, dorsally + deeply furrowed. Common name.—Great Plains Hawthorn. Distribution (Fig. 23).—Crataegus sheridana is a quite abundant hawthorn in the Cypress Hills and nearly all our records are found in the high light thicket situations, rather than aspen. UWO also has definite records 1 INV. ,C " £f. MAT J AIL " IC batal 1077 X E 4 - £ € pa x E Xu -— EX v BN Fic. 24 Map of C hr) 1 Jistribution in tl Or Hille R | for Eia. 2 from Saskatchwan, Montana, North and South Dakota, Wyoming and west-central Minnesota. There are also other records from Saskatchwan in prairie regions away from the Cypress Hills, such as Swift Current. Until confusion with the C. chrysocarpa complex of the Great Plains and Prairies is fully cleared up we will only have an incomplete knowledge of its distribution, but it does seem to be widespread in this region. Crataegus sheridana has strong superficial resemblances to the chrysocarpa group but is distinguished by its laterally pitted nutlets. It has tends to have shorter thorns than other hawthorns in the red-fruited, eroded-nutlet group and is also distinguished from most specimens in the Cypress Hills chrysocarpa group fi by its particularly densely pubescent to tomentose i d hypanthium. In addition, once known, the characteristic leaf shape makes it very recognizable. Nelson caused some confusion in citing as a cotype his specimen 1204 (RM), which is a perfectly typical specimen of C. macracantha. Unfortunately, the holotype, which was obviously quite different from the cotype, and whose fruit I did not dissect for Phipps (1998), does possess the superficial resemblance to C. chrysocarpa mentioned above and I therefore wrongly synonymized it with C. chrysocarpa in that paper as E.P. Kruschke and W.W. Eggleston both did on their annotation labels. series Rotundifoliae (Eggl.) Rehder, Sect. Rotundifoliae Egg]. Tve species: Crataegus rotundifolia Moench. Here treated as restricted to C. chrysocarpa and immediate relatives. Shrubs, seldom large; petioles glandular; leaf-blades somewhat small; flowers of medium size; anthers pink or white; ripe fruit red; nutlets smooth laterally. Two species, in the restricted sense of the series, both found in the Cypress Hills. Western Canadian taxa were reviewed in Phipps and O'Kennon (2004. Crataegus ursopedensis may not belong but is provisionally placed here. It has much larger leaves (5.5-9.5 cm long), generally longer thorns (4-7 cm) and the lateral faces of nutlets may be somewhat concave. 11. Crataegus chrysocarpa Ashe, Bull. North Carolina Exp. Sta. 175:110. 1900. (Plate 7.4a,b; Fig. 8, Crat lifolia M h, in part, sens. auctt. Amer 1078 Journal of the Botanical R h Institute of Texas 1(2) F +f] Fic. 25. Leaf spectrum of C. ‘Brett Gaff’ (Phipps & 0’K 8537, 8817), top, and compari f C. sheila-phippsi ieties, bottom. Specimens used LE DIN yy te 70154, 8338 eee ace RA AU AHEAD. Q657 90 PP (var choila-nhinncino. FTI * fl (var ). S. Laurie-Bourque del.; scale bars equal 1 cm. J Bush, often thicket-forming, 1.5-2.5 m in open, 2-4 m under aspen; extending twigs thinly pubescent, often reddish; at one year tan to mid-brown; two-year old brown, grayed, older gray to gray-brown; thorns moderately frequent, mainly 3-5 cm long, straight or more often curved, usually + slender to moderately stout, dark shiny red-brown at two years. Leaves deciduous; petioles 30-45% length of blade, usually glabrous (but occ. dense-pubescent), except for adaxial sulcus, bearing small sessile glands; blades 3-4.5 cm long at maturity, 2-3.5 cm at anthesis, rhombic, rhombovate or trullate-ovate in general shape, base cuneate, apex acute; lobes 2-3(—4)/side, max. LII mainly 15-25 96; margins with very small acute to subacute teeth; vena- tion craspedodromous, usually 3-4 main veins per side; adaxially + densely appressed scabrous-pubescent young, variably persistent till maturity; abaxially glabrous on the surface, main veins glabrous to thinly pubescent. Inflorescences 4-10 flowered; branches usually moderately to very densely (less commonly) pubescent, bearing caducous, small, pallid to light brown, narrow-oblong, gland-margined bracteoles. Flowers 14-16-18) mm diam.; hypanthium usually glabrous to thin-pubescent, especially below, rarely densely pubescent; calyx-lobes triangular, 3-4 mm long, adaxially thinly pubescent, abaxially glabrous, margins glandular but barely serrate, pallid to light green; petals + circular, white; stamens ca. 10, anthers ivory; styles 3-4. Infructescences 1-5 fruited; branches subglabrous to moderately pubescent. Fruit 9-11 mm high, broad-elliptic to subglobose, bright orange-red, finally becoming bright or deep red; calyx-lobes spreading; nutlets 3-4, laterally + plane or sometimes shallowly concave, dorsally grooved. Common name.—Fireberry Hawthorn. Distribution (Fig. 24).— Crataegus chrysocarpa is one of the two most wide-ranging hawthorns in North America, reaching from the inland valleys of Oregon, Washington and British Columbia to the Atlantic coast in a broad band on both sides of the Canada-United States border and as such it is rather variable overall n 10'K TAS £f. MAN J AIL " IC batal 1079 X E 4 £ £ Y x Fic. 26. Map of C. hoil, phi ; Inia a TA ED for Fia 9 though not particularly so in the Cypress Hills. In the Canadian Prairies as a whole C. chrysocarpa is by far the most widespread and common hawthorn while in the Cypress Hills it is most abundant at the lower altitudes, most usually in thickets in draws lacking shade of aspen, but is also found in light shade. As one reaches the driest sites in which hawthorns occur in the Cypress Hills C. chrysocarpa will be one of the few hawthorn species present. The Cypress Hills forms mostly represent a regionally common form for the species with broad-ovate to rhomb-ovate leaves, + hairy inflorescence branches, hypanthium glabrous or at most slightly hairy below, shortish thorns and 10 stamens with white anthers. Occasional variation in pubescence is found with a lo- cally rare, particularly hairy form densely pilose on the inflorescence branches and flowering hypanthium, usually quite persistent into fruit. Ripe fruit is also occasionally golden, instead of bright or orange red. Old fruit may be dark red. The type has rather hairy, + globose fruit and four plane-sided nutlets, representing a common leaf-type in the species. Crataegus chrysocarpa is of lower stature at maturity than most of the other species treated here. It has been badly confused in the past with C. rubribracteolata and C. sheridana, which see for comments. 12. eei (rete ie B. Phipps & O'Kennon var. saskatchewanensis J.B. Phipps & O'Kennon, v. (Plate 7 25). Tre: CANADA. SASKATCHEWAN: Cypress Hills, lower northern slopes, Maple Creek Rural. Ne pé dns M Me. Hay Creek, alt. 856 m, 24 May 2004, J.B. Phipps & R. O'Kennon 8654 (HoLoTYPE: UWO; ISOTYPES: CAN, SASK, DAO) Differt constanter a var. sheila-phippsi lo mi boriformi; spinis pl lto longioribus (3-6 cm vs. 2.5-4 cm) et multo lidioribus; folii ] inoril 2 5- 15 cm vs. 4-5 = ramulis inflorescentiarum sparsim a P : ; 1; 1 : aes ME “1 1 ats vel Scaflatinis. Thicket-forming shrub 1.5-3 m tall; extending shoots green, glabrous; one year old twigs smooth, dark brown, older gray or dark gray; thorns usually frequent, 3-6 cm long, straight to somewhat recurved, moderately stout, shiny dark red-brown at two years, darker with age. Leaves deciduous; petioles 25-40% length of blade, + glabrous except pubescent in adaxial sulcus, bearing several small, sessile glands; blades 2—4(—5) cm long, about half that size at early anthesis, rhombic, rhomb-ovate or rhomb-elliptic in general 1080 I Lnfthn Dat ee >) JOUTTIal OF f Texas 1(2) »— NS SC. Fic. 27. Line drawing of C ursopedensis S Ph ipp 2 0'K 8741a, 8949, 8975 (fl g Ph ipi 2 0'K 8547, 8587 (fruiting). V. gi £1 L "A gl L I Mags ow hala I £ nL CENT] £. h I £ J LEN i surface, fl lyx-lobe, fruit and nutlets. S. Laurie-Bourque del.; scale bars equal 1 cm. Phipps and 0 , Crataegus of Cypress Hills, t 1081 shape, base cuneate to broad cuneate, apex acute to acuminate; lobes 2-3 per side, these acute to acuminate, max LII 15-30%; margins with numerous, small, acuminate to acute teeth; venation craspedodromous, 4-5 main lateral veins per side; adaxial surface + densely appressed-pubescent young, this usually persistent to maturity, abaxially xd except sometimes for a few thin hairs on the major veins. Inflorescences 3-10 flowered; ł I ery sparsely pilose proximally, bearing caducous, narrow, membranous, pallid, densely gland- oad oo Flowers 16-20 mm diam.; hypanthium d E ia ca. 3mm long, triangular, glabrous on both surfaces, margins glandular-denticulate, greenish; petals + circular, white; stamens about 20, anthers white or cream; styles usually 3-4. Infructescences with 1—7 fruit; pedicels glabrous. Fruit 10-12 mm tall, turbinate to subglobose, orange red to bright red, sometimes blotched green at maturity; calyx-lobes erose or reflexed; nutlets 3-4, plane laterally, grooved dorsally. Common name.—Saskatchewan Hawthorn. Distribution (Fig. 26).—This is a generally common species at the lower altitudes («1100 m) in the eastern Cypress Hills, appearing more abundant in thickets than in aspen. It is also known from prairie areas in Saskatchewan well to the north of the Cypress Hills but its overall distribution remains unclear due to its earlier confusion with C. chrysocarpa. Ledingham made the first Cypress Hills collection (# 8922) in the Saskatchewan Centre Block, but named his specimen C. douglasii. Crataegus sheila-phippsiae var. saskatchewanensis is a typical member of the C. chrysocarpa group with its + rhombic, small leaves, relatively low shrubby stature and plane-sided nutlets. As a species it is distinguished by 20 stamens and + glabrous pedicels. The new variety here described differs from the type variety, which is much localized and restricted to the northern British Columbia Okanagan, by its less tree-like habit, generally longer thorns, smaller leaves which at early anthesis are particularly small and white rather than pink anthers (Figs. 5a, 5.b1) In Cypress Hills populations it is surprising how easy this species is to identify in flower or in fruit, once known, without having to approach close enough to note its lack of inflorescence pubescence and 20 stamens. 13. Crataegus ursopedensis J.B.Phipps & O'Kennon, sp. nov. (Plates 6.1d, 7.4d; Fig. 27). Te: U.S.A. Mon- TANA. Hill Co.: Beaver Creek Park at 1220 m., below aspen, 19 Sep 2003, J.B. Phipps & R. O’Kennon 8589 (HoLoTYPE: UWO; ISOTYPES: C.AN, DAO, MO, SASK, TRT, US). Frutices 2-34) m alti; squamae expansae gemmarum aurantiaco-cervicolores vel pnm ramuli Mich DM annotini nitenter Hr D post secundum annum rubro-brunnei cum maculis blancis t 6 D (324-7 cm longae, post secundum annum nitenter peratrorubro-brunneae, + validae, + rectae vel recurvatae. Folia COEM petioli 30-45% Un laminae, sulco adaxiali RNC in maturitate, > gabro cum paucis vel + numerosis MUN E 1 ovatae vel ovat laminae 5.5—9.5 cm longae (saepe solo cuneatae vel angusto-cuneatae; apices acuminati; lobi 4—5 per nee acuti vel acuminati, max. IFI 25-35%; margines cum numerosis parvis, « acutis dentibus; venatio craspedodroma, plerumque 5 venis principalibus per latus; iv DL adaxialis dense appresso- scabro-pubescens in juventute, in maturitate PAS pubescentia abaxialis glabra nisi interdum sparsim vel moderate dense pu- bescens secundum venas MN E in pus Inflorescentiae (1-)5-12 floratae, lato-patentes in OR plena; ramuli moderate dense pubescentes g llipticas, cremea d rirides, saepe suffusas pallide rubras distaliter praesertim, perglandulosas I 1 l l Se V i. Flores 15-18 mm diam.; hypanthium varie pubescens infra, + glabrum supra; lobi c adds s4—5 mm De NP DR oe m vel scariosi, adaxialiter leviter pubescentes, abaxialiter glabri, marginibus glanduliferis vel glandulo-denticulatibus; petala + circularia, alba; stamina ca. 10, ds eburneis; L I styli 3-4. Infructescentiae 1-8 fructatae; ramuli pubescentes, interdum i tenuiter. Fructus 8-11 mm alti, p idei vel ub oblongi, varie pubescentes, aurantiacc ri prim I l + appressi, 3-4 mm longi; pyrenae 3-4, lateribus + al vel e concavis, sulcatae nm dis Bushes 2—3(—4) m tall; expanding bud-scales orange-tan or paler; extending twigs pubescent; one year old shiny red-brown; two year old red-brown with white flakes, older gray; thorns usually plentiful, 84-7 cm long, at two years shiny very dark red-brown, fairly stout, + straight to recurved. Leaves deciduous; petioles 30—4596 of length of blade, adaxial sulcus pubescent at maturity, otherwise glabrous, with few to + numerous sessile glands; blades 5.5-9.5 cm long (often only 2-3 cm at early anthesis when also very flabellate), ovate to ovate-trullate in general shape; base cuneate to narrow cuneate, apex acuminate; lobes 1082 J | of the Botanical R h Institute of Texas 1(2) Fic. 28. Map of C lensis distribution in the C Hille R for Fin 2 d Ls at F g 4—5 per side, acute to acuminate, max. LII 25-35%; margins with numerous small, + acute teeth; venation craspedodromous, usually 5 main veins per side; adaxially densely appressed scabrous-pubescent young, at maturity subglabrous, abaxially glabrous except for sometimes thin to moderately dense pubescence along the main veins young. Inflorescences (1-)5-12 flowered, widely spread at full anthesis; branches moderately densely pubescent, bearing caducous, narrow elliptic, cream to pale green, often suffused light red especially distally, very glandular, membranous bracteoles that appear to grade into bud scales. Flow- ers 15-18 mm diam.; hypanthium variably pubescent below, + glabrous above; calyx-lobes 4-5 mm long, triangular, pale green to scarious, adaxially slightly pubescent, abaxially glabrous, margins glandular to glandular denticulate; petals + circular, white; stamens ca. 10, anthers ivory; styles 3-4. Infructescence 1-8 fruited; branches pubescent, sometimes only thinly so. Fruit 8-11 tall, mainly ellipsoid or oblong, variably pubescent, orange-red early, later orange-red to bright red; calyx lobes spreading to + appressed, 3-4 mm long; nutlets 3-4, sides + plane to slightly concave, dorsally furrowed. Common name.—Bear's Paw Hawthorn. Distribution (Fig. 28).— Crataegus ursopedensis is found throughout the Cypress Hills in suitable habitats though it seems most common in Saskatchewan. There are fine stands of it also in the Bear's Paw Mountains, Montana, along Beaver Creek at around 1200 m where the first record was discovered by us after seeing a sterile specimen at the herbarium at Montana State University at Havre. Other than our own collections this is the only known specimen. The species is named for the Bear's Paw Mountains although it is far more abundant in the Cypress Hills. It is found in the usual habitats for Crataegus in the Cypress Hills. Bear's Paw Hawthorn can have strikingly large, deeply and sharply incised leaves to 9.5 cm long under favorable conditions although they are often rather smaller and then difficult to tell, particularly at maturity, from the Red Bracteole Hawthorn. The leaves tend to be more glossy at maturity than in that species, the inflorescence is more open, the petioles are relatively longer, but materially the bracteoles much paler (Plate 6.1d) and,the nutlets sides more nearly plane than in C. rubribracteolata. This ensemble of characteristics appears to relate C. ursopedensis to C. chrysocarpa and other species in that part of ser. Rotundifoliae. Also see discussion under C. rubribracteolata. DL: J AIV f de £ f. ENII AIL 4 Ic I 4 ^L Phipps and 0 , Crataegus of Cy] ; 1083 13a. unnamed variants Two variant forms have been seen that will key out under C. ursopedensis. Both have broadly elliptic leaves. These are: G) a very hairy form with dark green, rather glossy, convex leaves rather smaller than and gener- ally differently lobed from C. ursopedensis, and with bright red fruit, labeled ‘Brett Gaff in Fig. 25 (upper), only with certainty at Shafer Creek, SK but possibly more widespread as in the unnamed specimen in Fig. 4b; and (ii) a form with often fairly deeply impressed veins, leaves often smaller and rather more sharply lobed than in C. ursopedensis, from localities in the area of the headwaters of Dimmock Creek, Bridge Creek and Skull Creek as well as from Bone Creek Coulee, all in the northeast of the Cypress Hills. More material and further study will be required to clarify their taxonomic status. BIOGEOGRAPHY Thirteen species of Crataegus are now reported for the Cypress Hills. These species vary greatly in their overall distribution from continent-wide species to, as far as is presently understood, endemic species. With such widely different distributions it may be anticipated that there is some degree of niche differentiation with respect to their habitats but so far there exist no studies on any species of Crataegus that address such matters in any detail so that only a few broad generalizations are possible. These derive mainly from observ- ing situations where species are sympatric. Even so, in the Cypress Hills, as in many parts of the range of Crataegus, different species, often of similar stature, may be found flourishing in the same thicket and other such thickets may also hold a similar mix of species so that it is not merely some random association that is observed but that their ecological requirements are broadly similar. In spite of this some differences have been observed which will be alluded to later. The main biogeographic patterns observed are as follows: a). Continent-wide taxa The only species in this group are C. chrysocarpa and C. macracantha both of which extend from intermontane regions quite close to the Pacific Ocean across to the Atlantic. Crataegus chrysocarpa is the more northerly of the two and throughout its range it is the most northerly species of Crataegus on the continent. It does not extend very far south into the United States except at altitude. On the other hand, C. macracantha does not extend very far into Canada and its southern limits are consistently hundreds of kilometers to the south of C. chrysocarpa in the United States. Crataegus chrysocarpa occurs somewhat locally in more mesic sites in the dry prairies whereas C. macracantha is less likely to occur there. b). Cordilleran taxa Two species are primarily Cordilleran, the rather variable C. douglasii and the much more uniform C. castlegarensis. Both have range extensions to disjunct populations in mesic areas to the east of the Rocky Mountains. c). Taxa restricted to mesic montane islands in the Great Plains Six or perhaps seven species are in this group and each is of quite restricted range. All are newly described in this paper. They are C. cupressocollina, C. rivuloadamensis, C. rivulopugnesis, C. purpurella, C. aquacerven- sis, C. ursopedensis and perhaps C. rubribracteolata (this last may get referred to the next group eventually). Crataegus rivuloadamensis, C. rivulopugnesis, C. purpurella and C. aquacervensis are at present knowledge restricted to the Cypress Hills while the other two have also been found in the Bears Paw Mountains. It may be noted that, in spite of their restricted ranges, all are locally abundant species in the Cypress Hills. Indeed, C. cupressocollina is one of the commoner and more distinct species found there where it is much more abundant than C. douglasii. d). Taxa primarily of the Great Plains but not confined to mesic montane islands The most striking example in this category is the resurrected species C. sheridana which has a wide but cur- rently incompletely understood range from Wyoming to Saskatchewan. In addition, there are the inadequately documented C. sheila-phippsiae var. saskatchewanensis, new in this paper and only from Saskatchewan so far Maec (var. sheila-phippsiae is from British Columbia) and C. rubribracteolata also new to this paper and which is 1084 Journal of the Botanical R h Institute of Texas 1(2) known from the Cypress Hills and Bears Paws with suggestions of occurrence in the surrounding prairie areas. Comparisons with Crataegus floras in montane islands in Montana have to be treated with caution as the Montana Crataegus floras are not yet been studied in the same detail. The Bears Paw Mountains have the same suite of species as the Cypress Hills except, naturally, for the latter's endemics and appear to have no species not found in the Cypress Hills. In the Bears Paws hawthorns are not nearly as abundant as in the Cypress Hills and are mostly restricted to the Beaver Creek valley which flows north towards Havre. In the still smaller area of the Sweet Grass Hills, only three species of hawthorn have been found, all widespread species, while in the Highwood Mountains five more or less widespread hawthorn species (C.douglasii, C. castlegarensis, C. macracantha, C. sheridana and C. chrysocarpa) are now known. However, at this juncture, only the Crataegus flora of the Bears Paw Mountains, among the Montana montane islands, is reasonably well known to us. In summary Crataegus in the Cypress Hills is much more speciose and seemingly far more numerous in individuals than any other discrete area between the Rocky Mountains and Minnesota at which point mesicity in topland soils begins to get back to optima for Crataegus. There is no single predominant bio- geographic relationship of this Crataegus flora although it should be noted that are there no species present with primarily eastern distributions. The Crataegus species of the Cypress Hills appear to range from fairly drought tolerant, e.g., C. sheridana, C. chrysocarpa, to more mesophytic types characterised by the Cypress Hills endemics, C. cupressocollina and the race of the variable C. douglasii that occurs there. ORIGINS OF THE CYPRESS HILLS CRATAEGUS FLORA As discussed in the section on origins of woodland in the Cypress Hills (pp. 1038-39), in the absence of firm data from the fossil record, etc., or convincing arguments from molecular evidence, one can only guess at times of origin of different species based to some extent on phenetic similarities and distributions. There is no direct evidence bearing on the age of any of the Crataegus species restricted to montane mesic islands such as the Cypress Hills and the latter, although not completely glaciated (parts of summit plateau remained unglaciated), have only presented conditions suitable for hawthorns over the last 10-12,000 years at most. Indeed the Holocene Zone I, from about 9000 to 6400 B.P., is a more likely starting point for Crataegus, taking into account general Crataegus temperature and mesicity preferences, and it is also likely that haw- thorn beginnings in the Cypress Hills were several events, not one. This position is supported by the widely different biogeographic patterns of the various species and their ecological differences so far as understood. 1 1 However, it is possible that the remarkable restricted distributions of C. cupresso-collina, C. rubribracteolata, C. ursopedensis, C. aquacervensis, C. rivuloadamensis and C. purpurella, all at least locally common in the Cypress Hills, may reflect relatively recent evolution, even if not entirely Holocene. An interesting feature for which it is not possible to determine the explanation at present is the differing distribution patterns among the Cypress Hills hawtl within the hills themselves. Broadly speaking, most taxa occur more or less throughout the hills from west to east in suitably mesic hawthorn habitats, although there is preference for a higher altitude distribution pattern in some species and a lower altitude pattern in others. Examples of this are noted with particular species in the text. However, four of our taxa, C. sheridana, C. sheila-phippsiae, C. cupressocollina and C. purpurella, are found predominantly in the Saskatchewan part of the Hills, see distribution maps (Figs. 7, 16, 23, 26), while three, C. aquacervensis, C. rivuloadamensis and C. rivulopugensis, are found mainly west of a line through Adams Creek (Figs. 12, 14, 18). These patterns might reflect relatively recent starting points within the Cypress Hills as there is no discernible significant east-west ecological difference between the eastern and western halves. A second possibility is that the endemics might be nothing more than relict species, unlikely though this might seem. Such a view would be more conformable with the observation that no other plant endemics are known from the Cypress Hills. No other plant genera, so far as we know, have species that make this distributional distinction. At pres- ent it is perhaps plausible, at most, that the Cypress Hills endemics originated during the Holocene but not necessarily within the Cypress Hills. In the cases of C. sheridana and C. sheila-phippsiae, however, the Phi n J OK TAS £f LU: ; AN 4 IC EL 1085 reduced distributions observed to date are perhaps only an artefact of undercollecting along a number of rather inaccessible but otherwise seemingly appropriate north-draining draws in Alberta. The existence of pairs of rather similar Crataegus species in the Cypress Hills flora might also be taken to suggest relatively recent divergence or alternatively, if any are of are of hybrid origin, similar but different origins in different locations. These pairs are: [C. purpurella* and C. rivulopugnensis*] [C. aquacervensis* and C. rivuloadamensis*] [C. ursopedensis* and C. rubribracteolata*] == [C. chrysocarpa and C. sheila-phippsiae var. saskatchewanensis of which the asterisked taxa are restricted to the Cypress Hills and their immediate surrounds or the Bears Paws as well. The existence of such rather similar pairs might also lend support to the idea of recency of origin although it must be noted that in all these our cases the individual species can be straightforwardly differentiated with adequate material. This, together with their relative abundance therefore suggests that they are (already) behaving as good species. The biogeographical patterns noticed here for Crataegus in the Cypress Hills have not received previous mention either for Crataegus or other plant genera. CONCLUSIONS The Cypress Hills flora contains 13 distinct Crataegus species, 10 or 11 more than in the literature. Of these, seven species and one variety are new to science. Although some specimens of all but two (C. castlegarensis and C. macracantha) of the Cypress Hills hawthorns had been collected by earlier workers (see cited specimens) only very few species of these had been recognized by them as being present. Crataegus cupressocollina, C. aquacervensis, C. rivuloadamensis, C. rivulopugnensis, C. purpurella, C. rubribracteolata, C. ursopedensis and C. sheila-phippsiae var. saskatchewanensis are new to science while C. sheridana is resurrected from its type description where it s duc for over 100 years except for occasional appearances as a synonym and this species is ther ly also new to general knowledge. All the new taxa can be straightforwardly distinguished with m material even if they have superficial similarity. The substantial numbers of individuals of each suggest that even in the case of the newly described taxa, they are behaving as distinct species. Boxplots of 8 selected continuous variables add to this conviction. Five of the previously known species are wide-ranging taxa that cross both major vegetation and floristic boundaries of Barbour & Christenson (1993) and Thorne (1993) and two of these, C. castlegarensis and C. sheila-phippsiae, represent significant extensions of previously known species. Two of the wide-ranging O species (C. douglasii and C. castlegarensis) are western cordilleran to mid-continent taxa and the other two, C. chrysocarpa and C. macracantha, constitute the two most wide-ranging North American Crataegus species, occurring from the western cordilleras to the Atlantic. One of the species newly recognized as present, C. macracantha, proves to be quite common in the Cypress Hills particularly at lower altitudes on the northern slopes, but does not appear to have been previously collected there, while C. sheridana, on present evidence, is also widespread in the Great Plains. Of the seven species new to science, four are, at present knowledge, Cypress Hills endemics, three are almost endemic species (a few plants are also known from Bears Paw Mountains, MT). Crataegus cupressocollina, new to science and arguably the most distinct of all the Cypress Hills species, may be the most abundant hawthorn in the Cypress Hills. The vegetation of the Cypress Hills is discussed for the purpose of better understanding hawthorn habitat. This is done in the context of 'CMT, a moisture index which apparently is the best single predictor of distribution patterns of plant species for the Great Plains. Relevant CMIs range from negative (prairie grassland) to 15+ (coniferous forest). Aspen occurs mainly between 1 and 15. Hawthorns thrive in the more mesic aspen woodland with a presumed CMI of about 10-15 and in creek thickets in prairie grassland. It is clear that in the Great Plains, at least, Crataegus responds more to a particular position on a moisture gradient (including accepting supplemental water as substitute along creek beds) and to a particular light regime than to precise soil type except for rejecting sandy soils. Thus the mesic island of the Cypress Hills becomes a suitable home to an estimated millions of hawthorns, far in excess of any other bushy plant. AII 1086 Journal of the Botanical R h Institute of Texas 1(2) the species discussed are mesomorphs with the most xerophilic tendencies being found in C. chrysocarpa, C. rubribracteolata and C. sheridana. The origins of the woody vegetation of the Cypress Hills are discussed and are entirely post-glacial. The successive zones of Holocene vegetation are summarized from work by David Sauchyn and others and extrapolation used to consider possible times of arrival of Crataegus species in the context of their varied biogeography and other factors. Because Crataegus fossilizes poorly (no relevant fossils have yet been recorded) speculation about arrival time has to be made on such matters as the pollen record for other woody species. From this line of thinking the level of endemism unearthed suggests the possibility that the Cypress Hills may represent an active area of recent hawthorn evolution. If so, it would be helped by the favorable clim and habitat that has allowed huge populations to develop, a situation not unlike the northern Okanagan of British Columbia, another location of favorable climate and large Crataegus populations at the same latitude, the Crataegus flora of which has been extensively investigated by the authors. APPENDIX FURTHER CITED SPECIMENS Notes.—Arrangement of taxa is alphabetical; non-JBP numbers with collection information in full; JBP numbers data abbreviated to season and province; numbers with asterisk = Alberta collections; without asterisk = Saskatchewan collections; subtotals given for last two categories. Most JBP numbers are Phipps and O'Kennon collections. 1. Crataegus aquacervensis a) non-JBP specimens (12 Alberta: Cypress Hills, no locality, 14 Aug 1947, Breitung, AJ. 5597 (DAO). Cypress Hills, hillside above Elkwater Lake, 16 Jun 1930, McCalla, W.C. E3728 (ALTA). Saskatchewan: Cypress Hills Park, no locality, 27 May 1949, Budd, A.C. 1305 (xerox) (SASK). Cypress Hills Park, no locality, 20 Aug 1935, Bolton, J.L. in SASK 111546 (xerox). Cypress Hills, no locality, 15 Aug 1926, Clarke, S.E. in SASK 111552 (xerox). Cypress Hills, no locality, 15 Aug 1936, Bolton, J.L. 186 (xerox) (SASK). Cypress Hills Park, no locality, 14 Jul 1947, Breitung, AJ. 4776 (xerox) (SASK). Cypress Hills, no locality, 07 Sep 1950, Budd, A.C. 1927 (xerox) (SASK). Cypress Hills Prov. Park, Centre Block, along beaver-ponded stream(s) of Loch Lomond, 10 Jul 1986, Harms, V.L. & R.M. 36371 (xerox) (SASK). Cypress Hills Prov. Park, Centre Block, along beaver-ponded stream(s) of Loch sd 10 Jul 1986, Harms, V.L & R.M. 36370 (xerox) (SASK). Piapot, Bear Creek Forest Reserve, 02 Jun 1938, Budd, A.C. 548 in SASK (xerox). b) n. Al (260779): Aug-Sep 2001:8227*. Aug-Sep 2003:8515*,8521*, Fan ia E 8528*, 8530*, 8532*. May-Jun 2004: 8723*,8727*, 8738a, 8754, 8755, 8772*, 8774*, 8775*, 8778*. Aug-Sep 4: 8791*, 8794*, 8796*, 8804*, 8810*, 8831*, 8906, 8912. May-Jun 2005: 8942, 8967, 8969, 8971, 8989*, 8991 o 9002*. RC 2. Crataegus castlegarensis a) non-JBP specimens b) JBP numbers (10*, 20): Aug-Sep 2001: 8221*, 8226*, 8228*, 8229*, 8230*, 8231*, 8232, 8236, 8244. Aug-Sep 2003: 8523*, 8531*, 8536, 8546, 8571,8613, 8626, 8792*. May-Jun 2004: 8681, 8683, 8738, 8746, 8750, 8761, 8773*. Aug-Sep 2004: 8844, 8881, 8894. May-Jun 2005: 8932, 8982, 9003. 3. Crataegus chrysocarpa a) non-JBP specimen Alberta: Cypress Hills Prot: Park, S of Girl Guid p, 09 Jun 1964, de Vries, B. 1896 (DAO). Cypress Hills Prov. Park, along small trail E of main campsite, 06 Jun 1964, de Vries, B. 1918 (DAO). Cypress Hills, on Brown Ranch along Ventre Creek Valley, N of Cypress Hills, 29 May 1978, Bradley, C. 84434 (ALTA). d Creek Ranch a 10 Jun 1975, Klar, A. 1078 (ALTA). Cypress Hills Prov. Park, Braun Coulee, Nott's Coulee, 13 Jun 1945, Cormack, R.G.H. 62 (ALTA). near Manyberries, 22 Jun 1946, Moss, E.H. 7016 (ALTA). Saskatchewan: N side of due Hills ca. 20 km E of Elkwater, 15 Sep 1995, Macdonald, 1.D. in SASK 137785. Cypress Hills, Fort Walsh, 20 Aug 1947, Breitung, AJ. 5692 (DAO). b) JBP numbers (16*, 23) Aug-Sep 2001: 8217*, 8234, 8235, 8239, 8583*. Aug-Sep 2003: 8525*, 8608, 8622, 8846. May-Jun 2004: 8644, 8662, 8684, 8694, 8702, 8704, 8710, 8712, 8714, 8716, 8725*, 8741, 8744, 8758, 8770*, 8776*. Aug-Sep 2004: 8797*, 8799*, 8814*, 8815*, 8846*, 8824*, 8826*, 8838*, 8885. May-Jun 2005: 8933, 8953, 8961*, 8980, 9006* DL: J f/f f de £ f. ENII AIL 4 Ic I 4 ^L Phipps and 0 , Crataegus of Cy] 1087 4. Crataegus cupressocollina a) non-JBP specimens (12 Saskatchewan: Cypress Hills Prov. Park, W Block, 21 Sep 1984, Ledingham, G.F. 8932 (USAS). Cypress Hills Park, Centre Block, 28 Aug 1990, de Vries, B. in USAS 278390. Cypress Hills, Battle Creek Ranger Station, 17 Aug 1947, Breitung, AJ. 5672 (ALTA). Cypress Hills, W block 14-112 km SE of AB border, 23 Jun 2003, Harms, V.L. 44175 (SASK). Cypress Hills Park, Battle Creek Ranger Station, 17 Aug 1947, Breitung, AJ. 5672 (DAO). Cypress Hills Park, no locality, 25 Jul 1949, Breitung, AJ. 8065 (DAO). Cypress Hills Park, valleys, 14 Jul 1947, Breitung, AJ. 4776 (DAO, SASK). Cypress Hills Park, shore of Loch Leven, 25 Jul 1949, Breitung, AJ. 8065 (CAN, SASK). Cypress Hills Park, W Block, 16-17 Jun 1951, Porsild, A.E. 17920 (CAN). Cypress Hills Prov. Park, along Lake Leven near Park Hdats., 27 Sep 1969, Boyce, R. in SASK 40840. Cypress Hills, no locality, 20-25 Jul 1949, Carmichael, L.T. 336 (DAO). Cypress Hills Prov. Park, Centre Block, vicinity of Loch Lomond, Jun 2003, Nagel-Hisey, M. 1 (UWO). b) JBP numbers (4* Aug- F 2001: 8241, 8243, 8246, 8247, 8249, 8250. May-Jun 2003: 8485, (uus 8491, 8493, 8496, 8501, 8503, 8504, 8506, 8508, 8509*, 8510*. Aug-Sep 2003: 8519*,8538, 8539, 8545, 8550, 8553, 8554, 8561, 8562, 8566, 8567, 8568, 8569, 8570, 8572, 8576, 8578, 8580, 8611,8615, 8616, 8617, 8619 (2), 8630. May-Jun 2004: 8648, 8660, 8666, 8671 1,867, 8679, 8689, 8708, 8715, 8730, 8767. Aug-Sep 2004: 8845, 8852. May-Jun 2005: 8923, 8931, 8977, 8981, 8993* 5. Crataegus douglasii a) non-JBP specimens (1 Alberta: Cypress Hills prow Park, Nicholls Spring, 27 Jul 1945, Cormack, R.G.H. 304a (ALTA), S side of Cypress Hills, ca. 6 km S of Elkwater, plateau, south slope, 16 Aug 1995, Macdonald, I.D. 950816a1 (ALTA). Saskatchewan: Swift Current District 20 mi S &5 W of Swift Current, 06 Aug 1952, Boivin, B. & Alex, J.F. 10067 (USAS). 5 mi NE of Fort Walsh, W block of Cypress Hills, near road to Fort Walsh Historic Site, 06 Jul 1981, Ledingham, G.F. & Naturalist Class 7477 (USAS). Cypress Hills Park, no locality, 25 Jul 1948, Ledingham, G.F. 48-687 (USAS). Cypress Hills Park, centre block, 30 Aug 1990, de Vries, B. in USAS 78090. Cypress Hills Prov. Park, Centre Block, S of Loch Lomond, E of Ben Vannock Drive, 20 Sep 1984, Ledingham, G.F. (USAS). Cypress Hills Prov. Park, Centre Block, along beaver-ponded stream(s) of Loch Lomond, 10 Jul 1986, Harms, V.L& R.M. 36343 (CAN). Cypress Hills Park, no locality, 14 Jul 1947, Breitung, AJ. 4776 (ALTA). Cypress Hills Park, no locality, 13 Jul 1947, Breitung, AJ. 4733 (ALTA). Cypress Hills, W block, 32 mi SW of Maple Creek, Univ. Regina Field Station, 12 Jun 1973, 1 edingham G.F. 5388 (USAS). b) JBP numbers (3*, 13): Aug- F 2001: 8240, 8242, 8245. May-Jun 2003: 8500. Aug-Sep 2003: 8552, 8612. May-Jun 2004: 8670, 8698, 8699, 8705, 743, 8764*. Aug-Sep 2004: 8853. May-Jun 2005: 8962*, 8964*, 8983 == — 6. Crataegus macracantha a) non-JBP specimens (0). b) JBP numbers (4*, 16): May-Jun 2003: 8494, 8502. Aug-Sep 2003: 8541, 8551, 8575, 8609. May-Jun 2004: 8649, a 8667 (2), 8675, 8703. Aug-Sep 2004: 8830*, 8862, 8874, 8888. May-Jun 2005: 8919, 8954, 8965*, 8997*, 9017 7. Crataegus purpurella a) non-JBP specimens (5): Saskatchewan: Cypress Hills Prov. Park, Center Block, along beaver-ponded stream(s) of Loch Lomond, 10 Jul 1986, Harms, VL &R.M 36370 (DAO, SASK); same date and locality, Harms, V.L & R.M 36371 (SASK). Cypress Hills, no locality, 16 Jun 1964, Newsome, R.D. 386-64 (DAO). Cypress Hills Prov. Park, Loch Lomond, E side of dam spillway, 09 Jun 2004, Nagel-Hisey, M & Goett, R. 3 (UWO). Cypress Hills Prov. Park, Centre Block, vicinity of Loch Lomond, 09 Jun 2003, Nagel-Hisey, M. 2 (UWO). Cypress Hills Prov. Park, Loch L E side of dam MA 09 Jun 2004, n Hisey, M. & Goett, R. 2 (UWO). b) JBP numbers (0*, 41): Aug- me 2001: 8248, 8251, 8252. May-Jun 2003: 8490, 8495, 8499. Aug-Sep 2003: 8555, 8557, 8558, 8559, 8610, 8610a, ay-Jun 2004: 8653, 8674, 8686, 8690, 8697, 8707, 8742, 8747. Aug-Sep 2004: 8700, 8856, 8858, 8860, 8867, BUR 8870, 8872, 8873, 8878, 8883, 8892. May-Jun 2005: 8927, 8928, 8929, 8938, 8955, 8984, 8985, 8988. 8. Crataegus rivuloadamensis a) non-JBP specimens Alberta: Cypress Hills, W slope, in coulees, 13 Aug 1947, Breitung, A.J. 5502 (DAO). W side of Cypress Hills, side of coulee, 27 52, Moss, E.H 10089 (ALTA, CAN). Saskatchewan: Cypress Hills Prov. Park, Fort Walsh, 20 Aug 1947, Breitung, AJ. 5704 (DAO). Cypress Hills Prov. Park, Loch Lomond, E of dam spillway, 09 Jun 2004, Nagel-Hisey M. & Goett R. 1b (UWO). Cypress Hills ie Park, Loch Lomond, E of dam, 09 Jun 2004, Nagel-Hisey M. & Goett R. 1a (UWO). b) JBP numbers (21*, Aug-Sep 2001: oe pom May-Jun 2003: 8512*. Aug-Sep 2003: 8516*,8518*, 8544, 8548, 8556, 8627,8631. May-Jun 1088 Journal of the Botanical R h Institute of Texas 1(2) 2004: 8722*, 8734, 8737, 8749, 8752, 8762*, 8763*, 8765*, 8766*, 8768*, 8777*, 8782*, 8805*. Aug-Sep 2004: 8805*, 8807*, 8808*, 8811*, 8832*, 8837*, 8843*, 8868, 8900. 9. Crataegus rivulopugnensis a) non-JBP specimens (1): Saskatchewan: opie Hills, W block, %4-1% km SE of AB border, 23 Jun 2003, Harms, V.L. 44174 (SASK). b) JBP numbers (23*, 12): Aug- T 2003: 851 7%, 8520*, 8533, 8542, 8628, 8629, 8632*. May-Jun 2004: 8728*, 8748, 8760, 8771*. Aug-Sep 2004: * 8800*, 8802*, 8806*, 8809*, 8827*, 8835*, 8836*, 8839*, 8840*, 8842*, 8877, 8895, 8908. May-Jun 2005: 8973, oe 8992*, 8994*, 8996*, 8999* (2), 9004, 9005*, 9007*, 9008*. 10. Crataegus rubribracteolata a) non-JBP specim Alberta: SE Rees ea tel 12 Jul 1935, Moss, E.H. 3270 (ALTA). MA Hills bd. up Suicide ue Suicide Creek, E of Elkwater PO, 1945, Cormack, R.G.H. 46512 (ALTA). Saskatchewan: ca. 10.1 mi S of Maple Creek, near bridge crossing Nn due ee #21, 10 Jul 1969, Zubrowski, S.J O SE of Tompkins, de May 1969, Looman, J. 12180 (SASK). b) JBP numbers (13%, 33): Aug- a 2001: 8217a*. dr 2003: 8497. Aug- 3 2003: 8540, 8560, 8564, 8565, 8573, 8623, 8624, 8625. May-Jun 4: 8636*, 8645, 8647, 8658, 8659, 8663, 86 673, 8676, 8678, 8680, 8688, 8709, 8713, 8719*, 8720*, 8721*, 8728a*, Pus 8780*, 8781*. Aug- x. 2004: 8798*, dn 8820*, 8821*, 8834*, 8849, 8850, 8851, 8852, 8854, 8864, 8896, 8903. May-Jun 2005: 8950, 8986 11. Crataegus sheila-phippsiae var. saskatchewanensis a) non-JBP specimens (1): Saskatchewan: Cypress Hills Prov. Park, along trail into Lone Pine Campground, E of Ben Vannock Dr., Centre Block, S of Loch omond, 20 Sep 1984, Ledingham, G.F. 8922 (USAS). b) JBP numbers (0*, 22): Sa 2003: 8486. Aug-Sep 2003: 8577. May-Jun 2004: 8646, 8651, 8654, 8655, 8664, 8682, 8696, 8740. Aug-Sep 2004: 8859, 8861, 8864, 8865, 8879, 8889, 8891, 8893 (2). May-Jun 2005. 8924, 8940, 8956. 12. Crataegus sheridan a) non-JBP specimens AS all specimens at UWO from outside Cypress Hills (18): [Note that a large number ofthe cited records are from cultivated plans in North Dakota. As these have very reliable provenances they have been arranged according to "m source states and counties.] dalla Big Stone Co.: 3.5 mi SE of Ortonville, Big Stone NWR, 6 Jun 1997, W.R. Smith 26583 (UWO); same loc., 22 Sep 7, WAR. m 26877 (UWO). "m ec Seed from North Dakota, Sheridan Co., cultivated North Dakota, Burleigh Co., E of Bismarck, 01 Oct 1993, Knudson, M. 9005841 (UWO). Seed from North Dakota, Sheridan Co., cultivated North Dakota, Burleigh Co., E of Bismarck, 01 Jun 1993, Knudson, M. 9005841 (UWO). Saskatchewan: Maple Creek, 14 Jul 1947, Breitung, AJ. 4875 (DAO). Cypress Hills, Birch Creek Ranger Station, 15 Jul 1947, Breitung, A.J. 4857 (ALTA, DAO). Ravenscrag, fossil site, 10 Oct 1987, Lepage, 144 (SASK). Swift Current, 26 May 1936, Bolton, J.L. in SASK 111550. South Dakota: Seed from South Dakota, Butte Co, 1 mi S of Castle Rock Butte, cultivated North Dakota, Burleigh Co., E of Bismarck, 22 May 1991, Knudson, M. 9005839 (UWO). Seed from South Dakota, Day Co, cultivated North Dakota, Burleigh Co., E of Bismarck, ?? Oct 1989, Knudson, M. 9034984 (UWO). Seed from South Dakota, Day Co, cultivated North Dakota, Burleigh Co., E of Bismarck, 01 Oct 1993, Knudson, M. 9005797 (UWO). Seed from South Dakota, Day Co, cultivated North Dakota, Burleigh Co., E of Bismarck, 01 Jun 1993, Knudson, M. 9005797 (UWO). Seed from South Dakota, Dueul Co., cultivated North Dakota, Burleigh Co, E of Bismarck, 21 May 1993, Knudson, M. 9005778 (UWO). Seed from South Dakota, Dueul Co., cultivated North Dakota, Burleigh Co, E of Bismarck, 01 Oct 1993, Knudson, M. 9005778 (UWO). Seed zu South Dakota, Marshall Co., cultivated North Dakota, Burleigh Co., E of Bismarck, 01 Oct 1993, Knudson, M. 9005850 (UWO). Seed from South Dakota, Hamlin Co, E side of L. Poinsett, cultivated North Dakota, Burleigh Co., E of Bismarck, 22 May 1991, Knudson, M. 9005798 (UWO). Seed from South Dakota, Marshall Co., cultivated North Dakota, Burleigh Co., E of Bismarck, 01 Jun 1993, Knudson, M 9005850 (UWO). Seed from South Dakota, Marshall Co., cultivated North Dakota, Burleigh Co., E of Bismarck, 01 Jun 1993, Knudson, M. 9005849 (UWO). Seed from South Dakota, Marshall Co., cultivated North Dakota, Burleigh Co., E of Bismarck, 01 Oct 1993, Knudson, M. 9005849 (UWO). Wyoming: Seed from Wyoming, Crook Co, cultivated North Dakota, Burleigh Co, E of Bismarck, 01 Oct 1993, Knudson, M. 9034980 (UWO). Seed from Wyoming, Crook Co., cultivated North Dakota, Burleigh Co, E of Bismarck, 20 May 1993, Knudson, M. 9034980 (UWO). Crook Co, Black Hills, along Inyan Kara Cr. ca. 5 air mi E of Keyhole Reservoir, ca. 15 air mi W of Sundance, 18 Sep 1984, Nelson, B.E. 12108 (UWO). b) JBP numbers (1*, 22): Aug-Sep 2003: 8492, 8574. May-Jun 2004: 8669, 8672, 8687, 8706, 8756. Aug-Sep 2004: 8718*, 8736, 8745, 8866, 8875, 8886, 8902, 8909. May-Jun 2005: 8918, 8920, 8921, 8941, 8952, 8958, 8970, 8974 ~ Phi ipps an id O'K C 4 g £f MAT ; AIL 4 IC EL 1089 13. Crataegus ursopedensis a) non-JBP specimens (1): Saskatchewan: Skull Creek, 12 Jun 1969, Looman, J. 12520 (SASK). Cypress Hills, Piapot, 08 Sep 1937, Bolton, J.L. in SASK 111549, b) JBP numbers (8*, 25): Aug-Sep 2003: 8534, 8563 (4), 8621. May-Jun 2004: 8717*, 8729*, 8731, 8735, 8757, 8779*. Aug-Sep 2004: 8818, 8828*, Bod 8847, 8857, 8876, 8890, 8904, 8905, 8910, 8911, 8913. May-Jun 2005: 8711, 8922 (2), 8930, 8939 (2), 8951 (3), 59, 8968, 8972, 8987, 9000*, 9001*, 9009*. 13a. Crataegus “Brett Gaff” a) non-JBP specimens (0). b) JBP numbers (3%, 18): May-Jun 2003: 8492a. May-Jun 2004: 8637, 8665, 8695, 8701, 8726*, 8733, 8751, 8759. Aug-Sep 2004: 881 3*, 8833*, 8863, 8884, 8887, 8898. May-Jun 2005: 8926 (2), 8934, 8935, 8937, 8939, 8979. ACKNOWLEDGMENTS Thanks are due to the Alberta and Saskatchewan parks authorities for permission to collect in the Alberta and Saskatchewan sections of the Cypress Hills Inter-Provincial Park, to the staff of Cypress Hills Inter- Provincial Park for much help and advice and for several additional collections made when we were unable to be present (collected by Les Weekes, AB, and M. Nagel-Hisey and Rick Goett, SK). Joan Hodgins, when at the T-rex Center, Eastend and Hillary Tarrant, formerly of Elkwater, also made several helpful collections. Anthony Littlewood, Dept of Classical Studies, The University of Western Ontario, is thanked for critically checking the Latin descriptions. Susan Laurie-Bourque of Ottawa again receives our thanks for her beautiful line illustrations. Ian Craig, imaging specialist, Faculty of Science, The University of Western Ontario, greatly assisted creating and organizing the figures. Students Vishaal Gupta and Janak Patel recorded the data and and Scott Hughes ran the program to generate the box plots. The curators of the following herbaria (ALTA, CAN, DAO, SASK, UAC and USAS) are thanked for the loan of specimens. We also want to thank Wayne Litke, proprietor and editor of the Maple Creek Advance Times, who was very helpful when supplies ran short and shared with us his knowledge of Maple Creek, the normal base for our fieldwork. REFERENCES Bangoun, M.G. and M.L. Christensen. 1993. Chapter 5, Vegetation. In: Flora of North America 1. Oxford University Press, New York. Boin, B. 1967. Crataegus. In: Flora of the Prairie Provinces. Provancheria 2(1). BREITUNG, A.J. 1954. A botanical survey of the Cypress Hills. Canad. Field Natural. 68:55-92. Dvorsky, K.A. and J.B. Pmirrs. 2005a. Bracteoles, a new character in Crataegus taxonomy. Canadian Botanical Association Annual Meetings, Edmonton, Alberta. June, 2005. Poster presentation. Dvorsky, K.A. and J.B. Pies. 2005b. Taxonomic studies of bracteoles in Crataegus.16'^ International Botanical Congress, Vienna, Austria. July, 2005. Poster presentation. Harms, VL. 2003. A checklist of the vascular plants of Saskatchewan. University of Saskatchewan Extension Press, Saskatoon, SK. HENDERSON, N., E. Hoc, E. Barrow, and B. Dotter. 2000. Climate change impacts on the island forests of the Great Plains and the implications for Nature Conservancy Policy: The outlook for Sweet Grass Hills (MT), Cypress Hills (AB- SK), Moose Mountain (SK), Spruce Woods (MB) and Turtle Mountain (MB-ND). PARC report. U. Regina, SK. Hoca, E. 1994. Climate and the southern limit of the western Canadian boreal forest. Canad. J. For. Res. 24: 1835-1845. Hoag, E. 1997. Temporal scaling of moisture and the forest-grassland boundaries in western Canada. Agric. For. Meteorol. 84:115-122. LARSEN, C. and G. MacDonaLo. 1995. Relations between tree-ring widths, climate and annual area burned in the boreal forest of Alberta. Canad. J. For. Res. 25:1746-1755. LooMAN, J. 1979. Vegetation of the Canadian Prairie Provinces. 1. An overview. Phytocoenologia. 5:347-366. 1090 Journal of the Botanical R h Institute of Texas 1(2) Morison, C.G.T., A.C. Hoyle, and J.F. Hore-Simpson. 1948. Tropical soil-vegetation catenas and mosaics. J. Ecol. Moss, E.H. 1959. Flora of Alberta. University of Toronto Press. Newsome, R.D. and R.L. Dix. 1968. The forests of the Cypress Hills, Alberta and Saskatchewan, Canada. Amer. Midl. Naturalist 80: 118-185. Packer, J.G. 1983. Flora of Alberta, ed. 2. University of Toronto Press. Phipps, J.B. 1997. Introduction to the red-fruited hawthorns (Crataegus, Rosaceae) of western North America. Canad. J. Bot. 76: 1863-1899. Puipes, J.B. 2005. A review of hybridization in North American Crataegus—another look at the ‘Crataegus problem. Ann. Missouri Bot. Gard. 92:113-120. PuiPPs, J.B. and R.J. O’KENNON. 2002. New taxa of Crataegus (Rosaceae) from the northern Okanagan-southwestern Shuswap diversity center. Sida 20:115-144. Phipps, J.B. and R.J. O'KeNNoN. 2004. Review of Crataegus series Rotundifoliae in western Canada. Sida 21:65-77. Porter, S.C., D.J. SAUCHYN, and L.D. DeLorme. 1998. The ostracode record from Harris Lake, southwestern Saskatch- ewan: 9200 years of local environmental change. J. Palynol. 21:35-44. SAUCHYN, D.J. and A.B. BEAUDOIN. 1998. Recent environmental change in the southwestern Canadian plains. Canad. Geogr. 42:337-353. SaucHyn, M.A. and D.J. SaucHyn. 1991. Paleogeogr., Paleoclimatol. & Paloecol. 88:13-23. ScocGAN, H.J. 1978. Crataegus. In: Flora of Canada vol.3. Queens Printer, Ottawa. THORNE, R.F. 1993. Phytogeography. In: Flora of North America. 1. Oxford University Press, NewYork. THOMAS WALTER TYPIFICATION PROJECT, IV: NEOTYPES AND EPITYPES FOR 43 WALTER NAMES, OF GENERA A THROUGH C Daniel B. Ward dua el ay Gainesville, Fonda 3261 7 USA. ABSTRACT Flora Caroliniana (1788). by Thomas Walter, he fj FA ; 1 pl MEE 2] 1 2 1 classification system of Linnaeus. Many of its species were new to science, d their names remain important in documentation of the southeastern American flora. But Walter dug no pidas Though pae have been chosen for a few of Walter's names, a large proportion of! d here, with selection of appropriate neotypes or E E Tos to represent 43 Thomas Walter names. RESUMEN f. je sue ques 1 1 ; 1 1 Jo 1 hi xaj Flora Caroliniana (1788), por Thomas Walter k = y el sistema de clasificación sexual de Linneo. Mechas de sus especies an ron nuevas para la ciencia, y sus nombres continúan siendo importantes en la documentación de la flora del sureste de América. Pero Walter no mantuvo un herbario. Aunque han sido oe i» p Eve de los nombres i Walter, una proporción grand p | tin L > E E E E L L E 43 D de Thomas Walter. In the 1780s, Thomas Walter owned and operated a rice plantation on the cleared bottomlands of the Santee River, South Carolina, in what is now Berkeley County. Of English origin and a classical education (Rembert 1980), Walter, though far isolated from anyone trained in botany, undertook the compilation of a flora of the plants he saw around him. The resultant book, Flora Caroliniana (1788), is the first treatment of American plants employing the binomial nomenclature and sexual classification system of Linnaeus. The Thomas Walter Typification Project is intended to bring understanding and nomenclatural preci- sion to the plant names published by Thomas Walter in his Flora Caroliniana (1788). Previous numbers of the Project have addressed: the Fraser/Walter folio herbarium, Natural History Museum, London, whose specimens were collected by John Fraser and whose labels often bear Walter's hand (Ward 2006); an inventory of the specimens from this herbarium and elsewhere that have been treated as types of Walter names by other authors (Ward 20072); and a presentation of those specimens from this herbarium worthy of designation as lectotypes or selection as neotypes (Ward 2007b). Documentation that the herbarium is that of John Fraser, rather than of Walter, has been provided separately (Ward 2007c). The present task, of this and the subsequent number, is to select additional specimens from among recent collections that may serve as neotypes or epitypes in support of Walter's names. The International Code of Botanical Nomenclature (McNeill et al. 2006; Art. 9.6, 9.11) provides rules for selection of a neotype where all original materials are missing. The Code also (Art. 9.7, 9.18) permits selection of an epitype where original materials may be extant but are demonstrably ambiguous. Walter left no herbarium, and the great majority of the specimens in the collection often known as the “Walter Herbarium" were largely gathered by John Fraser and probably not used by Walter in his writing. Thus it is to be expected that many Walter names should lack specimens that can be interpreted as their type. But when original materials are missing, later authors have often attempted to make remedy by select- ing specimens from among Fraser's materials. These authors, in good faith, selected Fraser specimens in the belief that they were the basis for Walter's names. Most unfortunately, this belief at times forced these authors to select specimes that were seriously defective, even unidentifiable except for an early notation by Fraser or Walter. However, once selected, a typification cannot be removed except by the laborious and J. Bot. Res. Inst. Texas 1(2): 1091 — 1100. 2007 1092 Journal of the Botanical R h Institute of Texas 1(2) uncertain pathway of conservation of a more suitable specimen, as provided by the Code (Art. 14). Selec- tion of an epitype, though not displacing the defective type, still serves to give stability to the name. Those defective typifications noted previously (Ward 2007a) merit treatment here. For the present task, it was thought wise to obtain all possible materials from a single institutional her- barium, one with adequate Southeastern holdings and an established history of taxonomic and nomenclatural interests. The Harvard University Herbaria (GH) offered their services and the loan of their specimens. By this cooperative action, a large portion of the specimens that will now document the meaning of Walters names will be housed together, available for scholarly use. Itis recognized that a replacement type, either neotype or epitype, should correspond as closely as possible with the materials held by the original author. This obligation is not reduced even by the impos- sibility of certain knowledge of that intent, obscured by time and mortality. Specimens for use in this phase of the Project were chosen with the following guidelines: The curator (GH) was asked to select three (or occasionally more) specimens of the designated species. The specimens were to show diagnostic characters. Insofar as possible, they were to come from South Carolina, ideally from Berkeley County (Walter's home). Specimens that had been annotated were to be preferred, thus adding assurance to the identification. A peculiar feature of Flora Caroliniana, which cannot but detract from the apparent credibility of Walters work, is his repetitious use of “Anonymos” as a generic name. From the standpoint of the value of his new names as seen by botanists two centuries later, Walter made a significant error by his judgment that many species or groups of species which he believed of generic value should await future study, rather than be named by him. Walter employed “Anonymos” for 28 of his genera, containing 44 species. These names, by decision of a much later botanical congress, have been ruled illegitimate, and their epithets are not avail- able for transfer to other genera (Sprague 1939; Ward 1962; Wilbur 1962). Even so, nearly all of Walter's Anonymos names with their accompanying descriptions were used by Gmelin (1791-1792) as the foundation for later legitimate names. The typification of Gmelin's later names, therefore, rests on the materials used by Walter and, where these materials are missing, justify selection of neotypes. Here, 14 of Gmelin's names are typified, each based upon Walter's name and description. In this phase of the Project, a total of 106 specimens are chosen for selection as types, with 101 as neotypes and 5 as epitypes. (For convenience in handling, this number is divided between the 43 names addressed in the present report, and a second report, soon following.) One-third bore annotations by other persons, confirming the identification. The typifications are presented here in the format used previously (Ward 2007a, 2007b), in alphabetical sequence, using the names given them by Thomas Walter. TYPIFICATIONS WALTER'S NAME: Aegilops aromaticum Walter (p. 249) MODERN NAME: Ctenium aromaticum (Walter) Wood Common in eastern SC. Spm. 112-B, labeled “Aegilops” by Walter, was identified as this by Hitchcock (1905: 55), but not designated as type. The specimen is fragmentary. Godfrey & Tryon 42, 23 Jun 1939 - GH (annot. Paul Young 1970), from 12 mi N of Georgetown, Georgetown County, South Carolina, is here selected as NEOTYPE for Aegilops aromaticum Walter, basionym of Ctenium aromaticum (Walter) Wood. Known ISONEOTYPES: BACA DU KRESESMOANY US: WaLTER'S NAME: Aegilops saccharinum Walter (p. 249) MODERN NAME: Dactyloctenium aegyptium (L.) Willd. Frequent in eastern SC. Spm. 112-D, labeled “Aegilops” by Fraser, was identified as Dactyloctenium aegyptium by Hitchcock (1905: 56). The specimen is mediocre. Godfrey & Tryon 1666, 22 Aug 1939 - GH (annot. S.L. Hatch 1993), from 8 mi SE of Kingstree, Williamsburg County, South Carolina, is here selected as NEOTYPE for Aegilops saccharinum Walter. (= Dactyloctenium aegyptium (L.) Willd.). Known isongotyres: BH, CA, CAS, DUKE, MO, NY, PH, US. Ward, Thomas Walter typification IV tyr ] epity 1093 WALTER'S NAME: Aira purpurea Walter (p. 78) MODERN NAME: Triplasis purpurea (Walter) Chapm. Frequent in eastern SC. Hitchcock (1905: 46) accepted Walters diagnosis as representing this species. There is no specimen. Godfrey & Tryon 1570, 17 Aug 1939 - GH (annot. S.L. Hatch 1995), from South Island, Georgetown County, South Carolina, is here selected as NeOTYPE for Aira purpurea Walter, basionym of Triplasis purpurea (Walter) Chapm. Known isongotypes: BH, CA, CAS, DUKE, F, MO, PH, US. WALTER'S NAME: Aletris aurea Walter (p. 121) MODERN NAME: Aletris aurea Walter Common in eastern SC. Spm. 112-E is this species, but consists solely of a flowering scape. The label, “Aletris Aurea," is in Fraser's hand. There is no indication it was seen or used by Walter. Ahles 15423, 25 Jun 1956 - GH, from along SC 63, 5.1 mi W of junction with SC 21, Colleton County, South Carolina, is here selected as NEOTYPE for Aletris aurea Walter. Known ISONEOTYPES: NCU, NY. WALTER'S NAME: Allionia albida Walter (p. 84) MODERN NAME: Mirabilis albida (Walter) Heimerl. [2 Oxybaphus albidus (Walter) Sweet] Rare in SC G counties). Spm. 3-B is this; it was labeled “Allionia xxx” by Walter, and “Albida” by Fraser. The species is only known in fall-line counties, outside of Walter's area; but one is Aiken Co. where Fraser is known to have traveled and collected. This specimen could perhaps be argued to be a lectotype. But since it was not recognized by Walter, one is hard put to believe it was part of material used in his writing. Ahles 55535, 27 Oct 1961 - GH, from North Augusta, Aiken County, South Carolina, is here selected as NEOTYPE for Allionia albida Walter, basionym of Mirabilis albida (Walter) Heimerl. Known tsonrotypes: FLAS, GA, MICH, NCU, NY. WALTER'S NAME: Alopecurus carolinianus Walter (p. 74) MODERN NAME: Alopecurus carolinianus Walter Frequent throughout. Because he considered Walter's diagnosis ambivalent, Hitchcock (1905: 40) concluded WA 7 “this species must remain doubtful.” Even so, it appears impossible to match Walter's words with any other equally likely species. Of the genus Alopecurus, only A. carolinianus occurs in SC with any frequency. Spm. 112-C is fragmentary, but may be this and bears the word “Alopecurus.” Though the writing is by Walter, he would have no need for this specimen in preparing his description. Walter's name is in general use, and Spongberg 67-64, 15 Apr 1967 - GH, from 40-acre Rock, Taxahaw, Lancaster County, South Carolina, is here selected as NEOTYPE for Alopecurus carolinianus Walter WALTER'S NAME: Amorpha herbacea Walter (p. 179) MODERN NAME: Amorpha herbacea Walter Rare in coastal SC, frequent inland. Epithet is not in italics, as was customary for a Walter name; perhaps a Fraser oversight. Though spm. 5-B was labeled “Amorpha fruticosa” by Fraser, it has been annotated by A.M. Vail as A. herbacea. But since its source is unknown and there is no indication it was seen by Walter, Cor- rell 5263, 13 Jun 1936 - GH (annot. Robert L. Wilbur 1959), from near Myrtle Beach, Horry County, South Carolina, is here selected as NEOTYPE for Amorpha herbacea Walter. Known ISONEOTYPES: GA. WALTER'S NAME: Amsonia ciliata Walter (p. 98) MODERN NAME: Ámsonia ciliata Walter Rare on the SC coastal plain, common inland. Spm. 5-A may be this. But the specimen is poor, and since the label (“Amsonia”) is in Fraser’s hand with no evidence it was seen by Walter, Weatherby 6120, 27 Apr 1932 - GH (annot. J. Williams 1992), from Columbia, Lexington County, South Carolina, is here selected as NEOTYPE for Amsonia ciliata Walter. Known 1soneortypes: PH, US. 1094 Journal of the Botanical R h Institute of Texas 1(2) WALTER'S NAME: Anonymos aquatic[a] Walter (p. 109) MODERN NAME: Nymphoides aquatica (Walter ex Gmel) Kuntze Frequent in eastern SC. Walter's name is illegitimate, but his description still serves as the foundation for Gmelin's epithet (Ward 1962). Spm. 59-E, labeled “Hydrocotyle” by Walter, appears to be this, but is poor (a leaf blade without petiole). Godfrey & Tryon 405, 10 Jul 1939 - GH, from 2 mi NE of Lane, Williamsburg County, South Carolina, is here selected as Neotype for Anonymos aquatica Walter and Villarsia aquatica Gmelin (1791: 447), basionym of Nymphoides aquatica (Walter ex Gmel.) Kuntze. Known ISONEOTYPES: CA, CAST US. WALTER'S NAME: Anonymos aquatic[a] Walter (p. 230) MODERN NAME: Planera aquatica Walter ex Gmel. Frequent in eastern SC. Spm. 115-G isa poor scrap, labeled with a single word ("aquatic") perhaps by Walter, a number (“FC 363”) by Fraser, and a Latin phrase (“Monoecia Triandria G. nova”) apparently by Fraser's son. Walters name is illegitimate, but his description still serves as the foundation for Gmelin’s epithet (Ward 1962). Hill 21222, 3 Mar 1990 - GH, from Edisto River, Givhans Ferry State Park, Dorchester County, South Carolina, is here selected as NEOTYPE for Anonymos aquatica Walter and Planera aquatica Walter ex Gmelin (1791: 130). WALTER'S NAME: Anonymos bracteatla] Walter (p. 181) MODERN NAME: Zornia bracteata Walter ex Gmel. Frequent in eastern SC. No specimen has been identified. Walters name is illegitimate, but his descrip- tion still serves as the foundation for Gmelin's epithet (Ward 1962). Wiegand & Manning 1624, 7 Jul 1927 - GH (annot. Robert H. Mohlenbrock, [1958?]), from 3 mi W of Marion, Marion County, South Carolina, is here selected as NEOTYPE for Anonymos bracteata Walter and Zornia bracteata Walter ex Gmelin (1792: 1096). Known ISONEOTYPES: BH. WALTER'S NAME: Anonymos capitat[a] Walter (p. 69) MODERN NAME: Burmannia capitata (Walter ex Gmel.) Mart. Frequent on the SC coastal plain. No specimen has been identified. Walter's name is illegitimate, but the description still serves as the foundation for Gmelin's epithet (Ward 1962). Godfrey & Tryon 1352, 7 Aug 1939 - GH (annot. J. van Benthem 1977; D. Q. Lewis 1994), from 6 mi SW of Moncks Corner, Berkeley County, South Carolina, is here selected as NEOTYPE for Anonymos capitata Walter and Vogelia capitata Gmelin (1791: 107), basionym of Burmannia capitata (Walter ex Gmel.) Mart. Known isouEorvrrs: BH, CAS, GA, PH. WALTER'S NAME: Anonymos caroliniensis Walter (p. 60); nom. illegit. MODERN NAME: Elytraria caroliniensis (Walter ex Gmel.) Pers. Very rare in SC, but one of the two known modern stations is in Berkeley Co. No corresponding speci- men has been identified. Ward (1962) and Wilbur (1962) have verified the modern name. Walters name is illegitimate, but the description still serves as the foundation for Gmelin's epithet (Ward 1962). Godfrey & Tryon 886, 24 Jul 1939 - GH, from Moncks Corner, Berkeley County, South Carolina, is here selected as NEOTYPE for Anonymos caroliniensis Walter and Tubiflora caroliniensis Gmelin (1791: 27), basionym of Elytraria caroliniensis (Walter ex Gmel.) Pers. Known IsoNEOTYPES: CA, CAS, DUKE, MO, NY, US. WALTER'S NAME: Anonymos caroliniensis Walter (p. 91); nom. illegit. MODERN NAME: Lithospermum caroliniense (Walter ex Gmel.) MacM. Infrequent in southern SC. Ward (1962) and Wilbur (1962) have verified the modern name. Walter's name is illegitimate, but it still serves as the foundation for Gmelin's epithet (Ward 1962). No specimen has been identified. Boufford & Wood 23567, 13 May 1985 - GH, from Williston, Barnwell County, South Carolina, is here selected as NEOTYPE for Anonymos caroliniensis Walter and Batschia caroliniensis Gmelin (1791: 315), basionym of Lithospermum caroliniense (Walter ex Gmel.) MacM. Known ISONEOTYPES: CAS, NY. Ward, Thomas Walter typification IV tyr 1 epity 1095 WALTER'S NAME: Anonymos caroliniensis Walter (p. 168) MODERN NAME: Ruellia caroliniensis (Walter ex Gmel.) Steud. Common throughout. Walter's name is illegitimate, but his description still serves as the foundation for Gmelin's epithet (Ward 1962). No specimen has been identified. Ravenel s.n., [1886?] - GH (annot. E. C. Leonard 1928; M.L. Fernald 1944), from the Santee Canal, [Berkeley County], South Carolina, is here se- lected as NEOTYPE for Anonymos caroliniensis Walter and Pattersonia liniensis Gmelin (1792: 925), basionym of Ruellia caroliniensis (Walter ex Gmel.) Steud. Known ISONEOTYPES: DUKE. WALTER'S NAME: Anonymos cassioides Walter (p. 171) MODERN NAME: Seymeria cassioides (Walter ex Gmel.) Blake Frequent throughout. No specimen has been identified. Walter's name is illegitimate, but his description still serves as the foundation for Gmelin's epithet (Ward 1962; Wilbur 1962). Godfrey 8213, 13 Sep 1939 - GH (annot. John E. Williams 1980), from St. Stephens, Berkeley County, South Carolina, is here selected as NEOTYPE for Anonymos cassioides Walter and Afzelia cassioides Gmelin (1792: 927), basionym of Seymeria cassioides (Walter ex Gmel.) Blake. Known IsoNEOTYPES: CA, US. WALTER'S NAME: Anonymos odoratissim|a] Walter (p. 198) MODERN NAME: Carphephorus odoratissimus (Walter ex Gmel.) Hebert [= Trilisa odoratissima (Walter ex Gmel.) Cass.] Common in southeast SC (incl. Berkeley Co.). Although combinations under the name Anonymos are il- legitimate, Gmelin (1792) based his name on that of Walter (Ward 1962). No specimen has been identified. Wiegand & Manning 3168 - GH (annot. Henry Hebert 1966), from 4 mi NW of Early Branch, Hampton County, South Carolina, is here selected as NeEotyrE for Anonymos odoratissima Walter and Chrysocoma odoratissima Gmelin (1792: 1204), basionym of Carphephorus odoratissimus (Walter ex Gmel.) Hebert. WALTER'S NAME: Anonymos petiolat[a] Walter (p. 108) MODERN NAME: Mitreola petiolata (Walter ex Gmel.) Torr. & A. Gray [= Cynoctonum mitreola (L.) Britt.] Common in eastern SC. No specimen has been identified (Leeuwenberg 1974: 9). Walter's name is ille- gitimate, but it still serves as the foundation for Gmelin's epithet (Ward 1962). Godfrey & Tryon 890, 24 Jul 1939 - GH (annot. A.J.M. Leeuwenberg 1974), from 10 mi NE of Moncks Corner, Berkeley County, South Carolina, is here selected as NEOTYPE for Anonymos petiolata Walter and Cynoctonum petiolatum Gmelin (1791: 443), basionym of Mitreola petiolata (Walter ex Gmel.) Torr. & A. Gray. Known IsSoNEOTYPES: CA, CAS, DUKE, F, MICH, NY, US. WALTER'S NAME: Anonymos pinnatla] Walter (p. 103); nom. illegit. MODERN NAME: Petalostemon pinnatum (Walter ex Gmel.) Blake Infrequent in eastern SC. Walter's name is illegitimate, but it still serves as the foundation for Kuhnia pin- nata Gmelin (Ward 1962). Spm. 117-D was labeled *Anonymos 102” in an unknown hand, followed by “FC” perhaps in Fraser's hand; this is a direct reference to Genus 102, Anonymos, of the Flora. Also on the original label is *Kuhniae affinis” in an unknown hand; these words are copied from Walter's subheading for genus Anonymos. Spm. 117-D also bears a modern annotation stating it to be “Type SPECIMEN of Anonymos pinnata Walter." Barneby (1977: 278) cited an unspecified Walter specimen (as “verified by Wemple 1970, p. 26”) as “Holotypus.” In the absence of evidence that Fraser/Walter 117-D [1787] - BM was seen or used by Walter, Barneby's typification is here corrected to NEOTYPE. But spm. 117-Disa slender leafless stem, wholly unsuitable to serve any identification purpose. Godfrey 8001, 7 Sep 1939 - GH, from McBee, Chesterfield County, South Carolina, is here selected as EPITYPE for Kuhnia pinnata Gmelin (1791: 375), basionym of Petalostemon pinnatum (Walter ex Gmel.) Blake, in support of Barneby’s typification (as corrected). Known tsoepitypes: BH, MO, US. 1096 Journal of the Botanical R h Institute of Texas 1(2) WALTER'S NAME: Anonymos quaternatla] Walter (p. 246); nom. illegit. MODERN NAME: Dioscorea quaternata Walter ex Gmel. Common throughout. There is no specimen. Walter's name is illegitimate, but it still serves as the foundation for Gmelin's epithet (Ward 1962). Hill 22286, 15 May 1991 - GH, from Parris Bridge road, N side of Lake Blalock, N of Carlisle, Union County, South Carolina, is here selected as NEOTYPE of Anonymos quaternata Walter and Dioscorea quaternata Walter ex Gmelin (1791: 581). Known IsSoNEOTYPES: NY. WALTER'S NAME: Ánonymos sessifol[ia] Walter (p. 108) MODERN NAME: Mitreola sessilifolia (Walter ex Gmel.) D. Don [= Cynoctonum sessilifolium Walter ex Gmel.] Common in eastern SC. Spm. 117-B is a slender stem of poor diagnostic character, marked with Fraser's number 685. It was labeled *Genus nov. Pentand digyn" by Walter, not recognized by him as his Anonymos “sessifolia.” The specimen has been annotated as “holotype” of Mitreola sessilifolia by A.J.M. Leeuwenberg. Leeuwenberg (1974: 21) then designated no. 685 as “Type.” Walter's name is illegitimate, but his description still serves as the foundation for Gmelin's epithet (Ward 1962). Since Walter would have had living materi- als available, spm. 117-B was surely not used by him in forming his description; it is thus neither holotype nor lectotype. In view of Walter's failure to recognize the plant as his own, Leeuwenberg's typification of Fraser/Walter 117-B (BM) has been corrected (Ward 20072) to neotype for Cynoctonum sessilifolium Gmelin (1791: 443), basionym of Mitreola sessilifolia (Walter ex Gmel.) D. Don. Because the specimen scarcely shows useful diagnostic features, Godfrey & Tryon 929, 26 Jul 1939 - GH, from 3 mi SW of Manning, Clarendon County, South Carolina, is here selected as EPITYPE, in support of Leeuwenberg’s typification (as corrected). Known ISOEPITYPES: CAS, NY, US. WALTER'S NAME: Ánonymos umbroslum] Walter (p. 63); nom. illegit. MODERN NAME: Micranthemum umbrosum (Walter ex Gmel.) Blake Common on SC coastal plain. Pennell (1920: 248) stated Blake (1915: 131) had identified Walter's “type,” an overstatement in that Blake had merely observed *no specimen, but the description is perfectly definitive of this species." Ward (1962) and Wilbur (1962) have verified the modern name. Walter's name is illegitimate, but it still serves as the foundation for Gmelin's epithet (Ward 1962). Bell 3632, 25 Jun 1956 - GH, from Ashton, Colleton County, South Carolina, is here selected as NEOTYPE for Anonymos umbrosum Walter and Globifera umbrosa Gmelin (1791: 32), basionym of Micranthemum umbrosum (Walter ex Gmel.) Blake. Known ISONEOTYPES: NCU, NY. WALTER'S NAME: Arenaria caroliniana Walter (p. 141) MODERN NAME: Arenaria caroliniana Walter Unknown on SC coastal plain, but frequent to west. Probably a Fraser discovery. No specimen has been identified. Cronquist 4932, 3 Apr 1948 - GH, from 3 mi W of Reidsville, Tatnall County, Georgia, is here selected as NEOTYPE for Arenaria caroliniana Walter. Known ISoNEOTYPES: FLAS, MO, NY, PH, US. WALTER'S NAME: Asclepias cinerea Walter (p. 105) MODERN NAME: Ásclepias cinerea Walter Very rare in SC Q counties, but on coastal plain where Walter may have encountered it). No specimen has been identified. Wiegand & Manning 2588 19 Jul 1927 - GH, from Early Branch, Hampton County, South Carolina, is here selected as NEOTYPE for Asclepias cinerea Walter. Known ISONEOTYPES: BH. WALTER'S NAME: Asclepias humistrata Walter (p. 105) MODERN NAME: Ásclepias humistrata Walter Frequent on the SC coastal plain. There is no specimen. Godfrey & Tryon 1069, 2 Aug 1939 - GH (annot. TJ. Rosatti 1987), from Georgetown, Georgetown County, South Carolina, is here selected as NEOTYPE for Asclepias humistrata Walter. Ward, Thomas Walter typification IV tyr 1 epity 1097 WALTER'S NAME: Asclepias lanceolata Walter (p. 105) MODERN NAME: Asclepias lanceolata Walter Frequent on SC coastal plain. Spm. 10-B appears to be this, but was labeled “Asclepias incarnata” by Fraser. It was annotated A. paupercula Michx. [= A. lanceolata var. paupercula (Michx.) Fern.] by A.M. Vail. But since spm. 10-B is a non-typical form of A. lanceolata, it is inappropriate as a type for Walter's new species. Godfrey & Tryon 154, 27 Jun 1939 - GH, from 4 mi SW of Andrews, Georgetown County, South Carolina, is here selected as NEOTYPE for Asclepias lanceolata Walter. Known isowrorvres: BH, CA, CAS, DUKE, MICH, MO, NY, PH, US. WALTER'S NAME: Asclepias pedicellata Walter (p. 106) MODERN NAME: Asclepias pedicellata Walter Very rare in SC (one county: Beaufort); probably a discovery of Fraser's. No specimen has been identified. Mellichamp s.n., 1880 - GH (annot. T.S. Rosatti 1987), from Bluffton, [Beaufort County], South Carolina, is here selected as NEOTYPE of Asclepias pedicellata Walter. Known 1soneotypes: CAS (“1887”), MICH (“1886”), US 1884. “1893”). WALTER'S NAME: Asclepias perennis Walter (p. 107) MODERN NAME: Asclepias perennis Walter Frequent in coastal SC. Walter labeled 10-F as “Asclepias,” and Fraser marked 10-G as “Asclepias an Nevea” (in reference to “white” flowers, or “new”?). By annotation, A.M. Vail has identified spms. 10-F and 10-G as Asclepias perennis. Both are of mediocre quality, and there is no evidence that Walter had occasion to use these specimens of a common species in preparing his descriptions. Godfrey & Tryon 130, 27 Jun 1939 - GH, L from Palmer Bridge, 5 mi ESE of Honey Hill, Berkeley County, South Carolina, is here selected as NEOTYPE of Asclepias perennis Walter. Known isongotypes: CA, NY, US. WALTER'S NAME: Asclepias viridis Walter (p. 107) MODERN NAME: Asclepias viridis Walter It seems unlikely that Walter, even through the agency of Fraser, would have encountered this largely Floridian species. It is presently unknown in the Carolinas and is very rare in GA (Glynn Co.). There is no specimen. The name could be misapplied; but the diagnosis is detailed and distinctive. To preserve usage, Harper 2184, 3 May 1904 - GH, from near Thalmann, Glynn County, Georgia, is here selected as NEOTYPE for Asclepias viridis Walter. Known isoneotypes: MO, NY, US. WALTER'S NAME: Aster carolinianus Walter (p. 208) MODERN NAME: Áster carolinianus Walter Rare in SC (2 counties), NC and GA (one county each). Walter's lengthy and accurate description (“caule fruticoso ramosissimo flexuoso subscandente...” etc.) clearly indicates he had adequate materials. There is no specimen in the herbarium, nor would one of Fraser's fragmentary specimens have been sufficient. Walter may have encountered the plant near Charleston, where it is known to occur. Leonard 2172, 18 Oct 1968 - GH, from along SC 703, NE of Isle of Pines, Charleston County, South Carolina, is here selected as NEOTYPE for Aster carolinianus Walter. Known IsSoNEOTYPES: CAS, FLAS, MICH, NCU, NY, USCH. WALTER'S NAME: Aster squarrosus Walter (p. 209) MODERN NAME: Áster walteri Alexander in Small Common in eastern SC. Spms. 13-C and 15-D appear to be this; both were labeled “Aster” by Walter. Since Walter did not recognize his own species, and better material would have been available to him elsewhere, Boufford 12340 19 Oct 1973 - GH, from 6 km. S of Socastee, Horry County, South Carolina, is here selected as NEOTYPE for Aster squarrosus Walter. Walter's name is a later homonym (non A. squarrosus All., 1785). 1098 Journal of the Botanical R h Institute of Texas 1(2) WALTER'S NAME: Cacalia ovata Walter (p. 196) MODERN NAME: Arnoglossum ovatum (Walter) H. Robins. Frequent in eastern SC. Two varieties of Arnoglossum ovatum have been recognized: var. ovatum, and var. lanceolatum (Nutt.) D. B. Ward. From his description (foliis ovatis"), Walter had the typical variety. No specimen has been identified. Boufford 23094, 11 Sep 1982 - GH, from along SC 162, 3.3 mi SE of 130, SE of Ridgeland, Jasper County, South Carolina, is here selected as NEOTYPE for Cacalia ovata Walter, basionym of Arnoglossum ovatum (Walter) H. Robins. WALTER'S NAME: Catalpa bignonioides Walter (p. 64) MODERN NAME: Catalpa bignonioides Walter Rare on the Carolina coastal plain, frequent and scattered inland. Its few SC stations are surely introductions. Thought by Little (1979) to be probably native in southwest GA, northwest FL, AL, and MS, well beyond the area traversed by either Walter or Fraser. But the tree was in early cultivation, attributed to *Carolina" by Linnaeus (1753: 623) and perhaps known to Walter near the port of Charleston (where now known as an escape). A single broken leaf in the herbarium (spm. 28-D) bears *Catalpa bignonioides" in Fraser's hand. Its poor quality and lack of linkage with Walter justifies selection here of Nelson 18315, 30 May 1997 - GH, from Leesville, Lexington County, South Carolina, as NeotyPe for Catalpa bignonioides Walter. Known iso- NEOTYPEs: USCH. WALTER'S NAME: Chironia decandra Walter (p. 95) MODERN NAME: Sabatia decandra (Walter) Harper [= Sabatia bartramii Wilbur] There is no specimen. Wilbur (1955) replaced the long-familiar Sabatia decandra with a new name, S. bartramii. However, Walter's description of Chironia decandra is closely matched by Wilbur's description of S. bartramii. Though S. decandra (= S. bartramii) is unknown north of central Georgia, it is within the range traveled by Fraser. Further, Walter must have had something, and discarding his name leaves the ambiguity unresolved. Wilbur & Webster 2691, 18 Aug 1950 - GH, from Sparks, Cook County, Georgia, is here selected as NEOTYPE for Chironia decandra Walter, basionym of Sabatia decandra (Walter) Harper. Known isontoryres: US. WALTER'S NAME: Chrysanthemum carolinianum Walter (p. 204) MODERN NAME: Boltonia caroliniana (Walter) Fern. Infrequent in SC (but known in Berkeley Co.). Spm. 31-E was identified as this by Fernald & Schubert (1948: 227), but they did not call it the type. A 3-digit number (“684”) on the label is by Fraser; the name “Chrysanthemum” is in Walter's hand. But since Walter would have had access to this species near his home, there is no reason to believe he used this specimen in preparing his text. Though the specimen is of fair quality, a better one is to be preferred. Godfrey 8155, 11 Sep 1939 - GH (annot. Judy Tate Morgan 1966), from along Santee River, 3 mi NE of Pineville, Berkeley County, South Carolina, is here selected as NEOTYPE for Chrysanthemum carolinianum Walter, basionym of Boltonia caroliniana (Walter) Fern. Known IsONEOTYPES: CAS, US. WALTER'S NAME: Chrysocoma acaulis Walter (p. 196) MODERN NAME: Vernonia acaulis (Walter) Gleason Frequent in SC. Walter's description was identified by Blake (1915: 135). No specimen. Godfrey 8083, 7 Sep 1939 - GH (annot. S. B. Jones 1968), from 1 mi W of McBee, Chesterfield County, South Carolina, is here selected as NEOTYPE for Chrysocoma acaulis Walter, basionym of Vernonia acaulis (Walter) Gleason. Known ISONEOTYPES: INY, US. WALTER'S NAME: Cistus carolinianus Walter (p. 152) MODERN NAME: Helianthemum carolinianum (Walter) Michx. Frequent on SC coastal plain. Spm. 33-B is poor, but may be Helianthemum carolinianum. Daoud & Wilbur (1965: 211) noted they had seen (by photo) a specimen in *Walter's Herb." but did not specify which one Ward, Thomas Walter typification IV tyr ] epity 1099 they intended. Rather than assign spm. 33-B to type status, Weatherby & Griscom 16586, 10 Apr 1932 - GH, from 1 mi S of Murrell's Inlet, Georgetown County, South Carolina, is here selected as Neotype for Cistus carolinianus Walter, basionym of Helianthemum carolinianum (Walter) Michx. Known ISONEOTYPES: US. WALTER'S NAME: Clematis reticulata Walter (p. 156) MODERN NAME: Clematis reticulata Walter Rare in eastern SC. Spm. 34-F is Clematis reticulata. It bears a label, securely attached by the stem inserted through slits, with “Clematis” written by Walter, and “Crispa” added by Fraser. The specimen also bears a second label, less securely attached, with *Clematis Reticulata” by Fraser. Since Walter appeared not to recognize the species, it is unlikely he used this collection in forming his description. Wherry s.n., 7 Jul 1936 - GH, from 4 mi S of Brookland, Lexington County, South Carolina, is here selected as NEOTYPE for Clematis reticulata Walter WALTER'S NAME: Convolvulus aquaticus Walter (p. 94) MODERN NAME: Stylisma aquatica (Walter) Raf. [2 Bonamia aquatica (Walter) A. Gray] Infrequent in eastern SC. Identified “ex char.” by Myint (1966). No specimen. [Spm. 36-E, with its very nar- row leaves, appears to be Stylisma patens (Desr.) Myint.] Radford 24551, 11 Jun 1957 - GH, from Summerton, Clarendon County, South Carolina, is here selected as Neotype for Convolvulus aquaticus Walter, basionym of Stylisma aquatica (Walter) Raf. Known IsoNEOTYPES: FLAS, GA, NCU, NY. WALTER'S NAME: Convolvulus humistratus Walter (p. 94) MODERN NAME: Stylisma humistrata (Walter) Chapm. Frequent in eastern SC. No specimen has been identified. Identified “ex char.” by Myint (1966). Wiegand 6> Manning 2635, 8 Jul 1927 - GH (annot. T. Myint 1961), from Mars Bluff bridge over Pee Dee River, Florence County, South Carolina, is here selected as NEOTYPE for Convolvulus humistratus Walter, basionym of Stylisma humistrata (Walter) Chapm. WALTER'S NAME: Cornucopiae hyemalis Walter (p. 73) MODERN NAME: Agrostis hyemalis (Walter) BSP. = Common in SC. Hitchcock (1905: 38) considered Walter’s diagnosis to be “undoubtedly” Agrostis hyemalis. There is no specimen. Robinson 97, 27 Apr 1912 - GH, from Navy Yard, Charleston, Charleston County, South Carolina, is here selected as NEOTYPE for Cornucopiae hyemalis Walter, basionym of Agrostis hyemalis (Walter) BSP. Known ISONEOTYPES: BH, US. WALTER'S NAME: Cornucopiae perennans Walter (p. 74) MODERN NAME: Agrostis perennans (Walter) Tuckerm. Mostly a piedmont species, but infrequently reaches the SC coastal plain. Walter's diagnosis as this species was accepted by Hitchcock (1905: 38) without question. This is the grass Walter and Fraser hoped to intro- duce into English cultivation, to great profit (Rembert 1980). There is no specimen. Kelly 254, 11 Jul 1995 - GH, from Ft. Jackson Military Reservation, Richland County, South Carolina, is here selected as NEOTYPE for Cornucopiae perennans Walter, basionym of Agrostis perennans (Walter) Tuckerm. WALTER'S NAME: Corylus americana Walter (p. 236) MODERN NAME: Corylus americana Walter Very rare in eastern SC, common in western SC and NC. Probably a Fraser discovery. No specimen. Small s.n., 17 Aug 1895 - GH (annot. John S. Drumke 1964), from Taccoa, Habersham County, South Carolina, is here selected as NEOTYPE for Corylus americana Walter. Known isoNEorYpEs: NY. ACKNOWLEDGMENTS This number of the Project is uniquely dependent upon the collections of a single institution, Harvard University (GH). By the foresight of Merritt Lyndon Fernald, my dear friend Robert K. Godfrey (with Rolla 1100 Journal of the Botanical R h Institute of Texas 1(2) M. Tryon) was sent to the Carolinas for the summer of 1939, charged with the task of documenting the flora that would have once been known to Thomas Walter. The abundant collections that flowed to GH, and thence to other institutions, have long been a staple of monographers and floristic writers. Though no special effort was made to select Godfrey materials, just under one third of the specimens chosen here as neotypes or epitypes have felt his hand. These specimens (and others of similar high quality) passed an initial screening by Walter T. Kittredge (GH), then came to me for final selection. Robert Dirig and Melissa A. Luckow (BH), Andrew Doran (CA), Debra Trock (CAS), Robert L. Wilbur (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) have supported this effort by searching for isoneotypes from within their herbaria. The abstract was efficiently hispanicized by Christine M. Housel (ABT). REFERENCES Barnesy, R.C. 1977. Dalea imagines. Mem. New York Bot. Gard. 27:1-891. BLAKE, S.F. 1915. Some neglected names in Walter's Flora Caroliniana. Rhodora 17:129-137. DAoub, H.S. and R.L. WiLBuR. 1965. A revision of the North American species of Helianthemum (Cistaceae). Rhodora 67:63-82, 201-216, 255-312. FERNALD, M.L. and B.G. ScHugerT. 1948. Studies of American types in British herbaria. Part IV: 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. LEFUWENBERG, A.J.M. 1974. The Loganiaceae of Africa, XII. A revision of Mitreola L. Meded. Rijks Landbouwhoges- chool 74(23):1-28. LINNAEUS, C. 1753. Species plantarum. 2 vols. Stockholm. Lite, E.L. 1979. Checklist of United States trees (native and naturalized). U. S. Dept. Agric. Handb. no. 541. McNett, J., F.R. Barrie, H.M. Bunper, V. DEMOULIN, D.L. HAwkswonrH, K. MARHOLD, D.H. NicoLson, J. PRADO, P.C. SiLva, J.E. Skog, N.J. TURLAND, and J. Wiersema, eds. 2006. The international code of botanical nomenclature (Vienna Code), July 2005. Regnum Veg. 146:1—568. Myint, T. 1966. Revision of the genus Stylisma (Convolvulaceae). Brittonia 18:97-117. PENNELL, FW. 1920 [“1919"]. Scrophulariaceae of the southeastern United States. Proc. Acad. Natl. Sci. Phil. 71:224-291. RemsERT, D.H. 1980. Thomas Walter, Carolina botanist. South Carolina Museum Commission, Bull. No. 5. SPRAGUE, T.A. 1939. Analysis of the binary combinations published under Anonymos by Walter, Fl. Carol. (1788). Bull. Misc. Inform. Kew 1939:331-334. WaLtER, T. 1788. Flora Caroliniana. London. War, D.B. 1962. The genus Anonymos and its nomenclatural survivors. Rhodora 64:87-92. Warb, D.B. 2006. The Thomas Walter Typification Project, |. Observations on the John Fraser folio. Sida 22:1111- 1118. Ward, D.B. 2007a. The Thomas Walter Typification Project, Il. The known Walter types. J. Bot. Res. Inst. Texas 1:407-423. War, D.B. 2007b. The Thomas Walter Typification Project, Ill. Lectotypes and neotypes for 20 Walter names, as recognized in the Fraser/Walter herbarium. J. Bot. Res. Inst. Texas 1:425-430, Warb, D.B. 2007c. The Thomas Walter Herbarium is not the herbarium of Thomas Walter. Taxon 56:971-926. WILBUR, R.L. 1955. A revision of the North American genus Sabatia (Gentianaceae). Rhodora 57:1-33, 43-71, 78-104, WiLgur, R.L. 1962. The identity of Walter's species of Anonymos. J. Elisha Mitchell Sci. Soc. 78:125-132. ES COTYLEDON AREOLES IN THE FABACEAE SUB TRIBE CETORIUNAE James A. Lackey xu E d NMNH MRC- 166, PO r 370 12 Washington, DC 20013-7012, U.S.A. lackeyj@si.edu ABSTRACT Of the five genera of Phaseoleae subtribe Clitoriinae, four genera; Centrosema, Periandra, Clitoria, and Barbieria; were sampled for cotyledon areole presence or absence, and for bd sus anatomical characters in mature ans seeds. CHT: seeds were unavailable. All o b Md d pt in Clitoria subgenus B f Clitoria subgenus M 1] Neurocarpum circular to ovate, and approximately medial, and their presence or absence, position, and size are beeen ae to RA PeT occurrence. The limited sampling indicates that the subtribe has diverse seed anatomy, eviden by several di Centrosema and Periandra have only spongy cotyledon mesophyll, a Mein fr h id a a lt heid bar r extension. Barbieria has two hilar tongues, palisade mesophyll, and a viscid, clear, colorless, and thermoplastic epitesta. Of the three subgenera of Clitoria, subgenus Clitoria has palisade pM subgenus Bractearia has only spongy mesophyll and two hilar tongues; and subgenus Neurocarpum has only spongy mesophyll, two hilar tongues, and the same epitesta as Barbieria Key Wonps: Cotyledon areoles, endosperm, galactomannan RESUMEN De los cinco géneros de Phaseoleae subtribu Clitoriinae, cuatro; Centrosema, Periandra, d P Barbieria; se muestrearon para la presencia o ausencia de areola en el cotiledón, y otros caracteres anatómicos en las semillas durmi ntes. Las semillas de Clitoriopsis no estuvieron disponibles. Todos los especimenes examinados tienen areolas en el cotiledón excepto Clitoria subgénero Bractearia y dos especies de Clitoria subgénero Neurocarpum. La mayoría de las areolas del cotiledón son pequeñas, de circulares a ovadas, y aproxima- amente mediales, y su presencia o ausencia, posición, y tamaño están directamente relacionados con la presencia de endospermo. El muestreo limitado indica ues la subtribu tiene una anatomía seminal diversa, evidenciada por varios caracteres distintivos en varias gue dV Centrosema ut til a RD ME una E estructura lenticular de p y una t id n 59 ati ] 1 p clara, sin color, y termoplástica. De los tres oe de Clitoria, el cado o Clitoria ti 5fil palizad; el subgé Bractearia tiene t Sfilo esponj y dos lenguas hilares; y el subgénero Neurocarpum tiene úni t ófilo esponjoso, dos lenguas hilares, y la misma epitesta rt INTRODUCTION The cotyledon areole, named by Endo and Ohashi (1997, 1998a, 1998b, 1999a, 1999b), is a small spot of projecting epidermal cells on the abaxial cotyledon surface of dormant mature seeds of many Leguminosae subfamily Papilionoideae, but absent in subfamilies Caesalpinioideae and Mimosoideae. Cotyledon areole cells, and subtending cells, have different size, shape, and several peculiar anatomical and chemical char- acteristics compared with surrounding cells. The presence or absence, shape, and position seem to mark certain genera, tribes, and groups of tribes. In the Clitoriinae, Endo and Ohashi (19992) found medial coty- ledon areoles in one unidentified species each of Clitoria and Centrosema. They established that the soybean structure called a “pit” (Dzikowski1936, 1937; Miksche 1961; Yaklich et al.1984, 1986, 1987, 1989, 1992, 1995, 1996, 1998; Baker & Minor 1987; Ma et al. 2004) is the same as a cotyledon areole. Beck's (1878) "Aleuronfleck," described and illustrated in detail, also matches the cotyledon areole. Cotyledon areole function is unknown (Ma et al. 2004). Despite incidental references to Clitoriinae seed anatomy as part of larger projects, studies of specific features, or reviews at a generic level (Pammel 1899; Martin 1946; Lersten 1981, 1982; Kirkbride et al. 2003), there is no basic anatomical study of these seeds. J. Bot. Res. Inst. Texas 1(2): 1101 — 1118. 2007 1102 Journal of the Botanical R h Institute of Texas 1(2) Because cotyledon areole presence or absence, shape, and position seemed to have taxonomic signifi- cance, because of the lack of basic comparative seed anatomy in Clitoriinae, and because cotyledon areoles have not been studied in relation to other features of these seeds, the current study has the following aims: 1) to survey for cotyledon areole presence in the subtribe; 2) to study basic seed anatomy to determine if cotyledon areoles are correlated with other seed features; and 3) to evaluate if any of these characters could yield data of systematic significance. MATERIALS AND METHODS Studied seeds were deposited at BARC or US (Table 1). All specimens were assigned a unique number, pre- ceded by the letters JAL. Plant names and authors follow treatments given by Fantz cited in the Discussion. In Clitoria species, the subgenus name is indicated in parentheses. Seeds were examined dry, both externally and internally, with a Wild M5 stereomicroscope. Seeds were examined whole or cracked longitudinally or transversely with a razor blade and a custom-made cutting block or with a miniature angled end cutter (Tronex 5083, reground to zero microbevel). For enhanced vis- ibility, a drop of toluidine blue O solution, 0.05% in 1M phosphate buffer pH 6.76, was sometimes applied to the abaxial surface of dry cotyledons to color most epidermal cells more strongly than the more lightly staining cotyledon areole cells. Sketches of seeds were made with a camera lucida microscope attachment. Line work was done in Adobe Illustrator. Conventions for distinguishing features by stippling and other patterns were adapted from those of Schleiden and Vogel (1839), Corner (1951), and Smith (1981, 1983). Views within each drawn sample may have come from different seeds, and thus may differ in size and shape. Photographs were taken with a digital camera on a dissecting or compound microscope, and sometimes processed with extended depth of field software. Orientation terms follow the definitions of Sterling (1954). Median plane is the plane passing through the hilar groove, micropyle, and chalaza and perpendicular to the hilum surface. Transverse plane is any plane perpendicular both to the median plane and to the hilum surface. Frontal plane is any plane parallel to the hilum surface, and therefore perpendicular both to the median and transverse planes. Anterior is towards the micropyle end of the seed. Posterior is towards the lens end of the seed. Dorsal is the side of the seed farthest from the hilum. Ventral is at the side of the seed nearest the hilum. Lateral is at the surface of the seed on either side of the median plane. Right and left are from the point of view of an observer oriented as the seed is oriented, with top to the anterior, bottom to the posterior, dorsal to the back, and ventral to the front. Pod orientation conventions for ventral, dorsal, anterior, posterior, right, and left, are the same as for the seed, with anterior towards the stigma, posterior to the base, ventral towards the suture bearing the ovules, and dorsal towards the opposite suture. Seed terminology and abbreviations were adapted from Schleiden and Vogel (1839), Corner (1951), Gunn (1981), and Kirkbride et al. (2003): al = aleurone layer (outer endosperm layer), ar = aril (strophiole and caruncle of some authors), arp = antiraphe, c a = cotyledon areole, c l= crushed endosperm layer (inner endosperm layer, third layer), e pal = counter palisade, ch = chalaza, cot = cotyledon, cu = cuticle, em = embryo, en = endosperm, eph = epihilum, ept = epitesta, f = funiculus, h = hilum, h gl = hourglass cells, h gr = hilar groove, h t = hilar tongue, I = lens (strophiole of some authors), 11 = light line, m = micropyle, m l = middle endosperm layer (swelling layer), pal = palisade (Malpighian) cells, p = plumule, p m = palisade mesophyll, r = radicle, r a = rim-aril, rp = raphe, r v b = recurrent vascular bundle, t b = tracheid bar, and v b - vascular bundle. Format for seed descriptions generally follow Kirkbride et al. (2003). Descriptive terms for cotyledon areole position follow Endo and Ohashi (19993); the cotyledon is divided into five equal parts from the base to apex: basal, basal-medial, medial, medial-apical, apical. Presence of starch was tested by depositing one drop of Lugol's iodine solution (iodine 0.2 g, potassium iodide 0.3 g, water 30 ml) on the cotyledon surface cut in cross section. Starch grains produced a purple to black color within seconds, when observed under the dissecting microscope. Hydration of seeds was aided with Pohlstoff (Aerosol OT 1 ml, methanol 25 ml, water 74 ml). Uncoated seed material was observed with a Philips XL30 ESEM. Lackey, Cotyledon areoles of Clitoriinae TasLe 1. Specimens studied. 1103 JAL Species Specimen 1261 Barbieria pinnata (Persoon) Baill. Misc. US (BARC) 1271 Barbieria pinnata (Persoon) Baill. Sinteris 5942 (BARC) Puerto Rico 1262 Centrosema arenarium Bent IRI 1355 (BARC) Brazil 1267 Centrosema brasilianum (L.) Benth. Rodrigues 643 (US) Brazil 1268 Centrosema paseuouium Mart Pittier 13474 (US) Venezuela 200 Pi ind (Turp.) Benth. PI 247478 (BARC) Belgian Congo 202 ntrosema plumeri (Turp.) Benth. P| 322329 (BARC) Brazil 1266 o plumeri (Turp.) Benth. Nee 9492 (BARC) Panama 208 Centrosema pubescens Benth. PI 286289 (BARC) Ivory Coast 209 Centrosema pubescens Benth PI 279594 (BARC) 210 Centrosema pubescens Benth PI 219833 (BARC) Ceylon 211 Centrosema pubescens Benth PI 212980 (BARC) India 1015 Centrosema pubescens Benth Pollard et Palmer 328 (BARC) Cuba 1264 Centrosema pubescens Benth PI 308555 (BARC) 1265 Sia pubescens Benth PI 337079 (BARC) 282 ntrosema sagittatum (Willd.) Malme Pederson 2796 (BARC) Argentina-MO 500 o sagittatum (Willd.) Malme Pederson 2796 (BARC) Argentina-K 1263 Centrosema sagittatum (Willd.) Malme Shafer 481 (US) Cuba 204 Centrosema PI 200731 (BARC) 203 mds virginianum (L.) Benth. PI 322356 (BARC) Brazil 205 Centrosema virginianum (L) Benth. PI 322351 (BARC) Brazil 224 dietis virginianum (L.) Benth. 55-17 Urbana (BARC) 1190 rosema virginianum (L.) Benth. Wolff 3138 (US) Texas 1594 zs amazonum Mart. Ex Benth. Martinelli 7263 (US) Brazil 1595 Clitoria P eeu Harms Hahn 5612 (US) Guyana 296 Clitoria laurifolia Poir. CPI 34890 CSIRO (BARC) 1599 Clitoria na. Bain s.n. (US) Tennessee 295 Clitoria decae Pers, CPI 35685 CSIRO (BARC) 197 Clitoria ternate P| 209315 (BARC) 1270 Periandra Heine Benth. Pires 58051 (US) Brazil 252 Periandra heterophylla Benth. P| 322570 (BARC) Periandra heterophylla Benth. 1269 Periandra heal Benth. 1592 Periandra heterophylla Benth. 1593 Periandra ae (Vell) Taub. 1596 Periandra mediterranea (Vell.) Taub. Philcox et Fereira 3874 (BARC) Brazil IRI 1779 (BARC) Braz Irwin et al. 15211 (US) Brazil Irwin et al. 15430 (US) Brazil Irwin et al. 7969 (US) Brazil RESULTS Because seed morphology in the Clitoriinae is diverse, four detailed descriptions are given for Centrosema virginianum, Clitoria (Clitoria) ternatea, Clitoria (Neurocarpum) rubiginosa, and Barbieria pinnata. Other species generally correspond to one of these four detailed descriptions, and will thus be described in notes and dif- ferential diagnoses. Results are summarized (Table 2). Size variation from specimen to specimen and seed to seed is common. Measurements should be regarded as a general guide and not a comprehensive indication of variation for the species. Specimens of several species were inadequate in quantity and quality. Centrosema virginianum. Seed reniform-oval to cylindrical; ca 4.3x1.5x2.5 mm (lengthxwidthxheight) (Fig. la); brown with brown hilum; sink in water; micropyle oriented towards stigma in pod. Seed externally with prominent hilum, anterior micropyle and posterior lens (Fig. 2e). Hilum oval ca 1.3x0.7 mm; surrounded by a raised rim. Rim-aril brown, mostly of ca 0.05 mm raised bead; with hilar tongue ca 0.7 mm wide by 0.1 mm long positioned from mid-hilum extending toward anterior end. Hilar tongue on left in about half the seeds; on right in about half the seeds. Counter-palisade surface naked or Texas 1(2) £f T, P" D £L JUUITIdI OF 1104 ‘sapeds Jo Ids! Iq 'SANISOAO = UE) :oAJquua ui SUIULARUE) "ue? ‘BAIUSOd = up1eis :eAneDau 10 an sod 1591 Yes UOPa|Ajod 'upieis 3uasjedde Jake] e|ppiuu = TW suaredde Jakej usdsopus e|ppiuu “wsdsopuz 1196 = 5 eabida = 3 'JeebodÁu = H :uon -euiuueb “Wan “A ¡eixeqe pue A|jerxepe anssn apesijed = apes!e4 'uone1uasu 1y} ÁBuods = K6uods : | kydosaw UOpa¡A10) os :e1591ido PIDSIA = pIDSIA ‘JUasa yl 3uesqe JO JUasaId Bot DE Jequunu anBuoi sey anuo) JejIH uasaid "M ¡esop = BUIM eq playden 'g| àussaud spiceuben penid = SpIO9UpeJ| :S||32 SUS] ‘SUdT ]eipauJ-|eseq = Wg "]eipeuJ = W :3/031¥ UOPajA10) ^v) 'DUDIDUJ DUO)I[) JO} 31815 = () *e3e1s PPIP WJU, = q-e ‘SIIS aJd NU = qe 'ureyie»un si UONEAJasqo = į '1uesqe = -‘Juasald = + ‘ajqeyiene eyep ou 93e2Ipul 31815 J9120Jeup JOY Pu S00€ 914U2S ‘386 1 ‘C66 | z1ue4 LOL ale sjunoo sopads pue uonnquisig 4/61 Kaye" u10JJ S| a2uesaud BUIURACULD (1861) eng WOM SWIS} YUM ‘786 | 1819 Y9A 668 | [BWW 1681 WIOH ‘886 | 'e6/6| 204 p/6| 1epneg WoL IĮ Snol pue XJOM 1U9JJD2 WO. SINSIY OOIX9lA l 'e2u9uly 59 S s TW 9pesi[eg DIOSIA OML 2 ; Wd e1191q189 eISv undje»ounaN vc 'iseoueuv - (Queis QW) - (3) H Ábuods pDSIA — (BUD-)OM| = - (W8)- 'Bqns eno» ODIXIN ees} eg 67 'eoueuly 5$ S - U2Je1S - j Kbuods - OM| - - - 'bqns emoy e110311) 6 ¿PLOM PIO E TW j 9pesi|eg : O) W 5qns emo) 9 elAljog ‘zeig - . YDJe1s/- IN 5/3 A®uods - 3UuO Bul Splo3yel] WN eipuenəd 9€ seouauly ue) - JW 3/H A®uods - 3UuO OuMW Splo3yel] W ewəso1zuə) ənuo} dds uonnquisiq ue) y2183S uuadsopuy w139 uopəjÁ}o) exsajid3 Je|H aL sua] Ww uoxeL 'SISIDRIEYI paas Jo Kewuns 'z Hav], Lackey, Cotyledon areoles of Clitoriinae 1105 Fic. 1.C lucida drawings of Clitorii Is. Each figure i posed of the following pper left : Is top of page, fseed showing hilum. In up; few avian ieee RET g right side of page. In upper rightis longitudinal section through median plane of seed, anterior to right side of page, showing interior of testa with adl lost lining tl bry ity, embryo removed. Tracheid bar and any ids of | tral side indicated by g. Im f cotyled | p "m T " "OST End hy | linal | | In| ight is left side of emt Cotyledon Y F T L4 » L4 Ii 2 L4 d I J IL 1 LL LEI I I nfs L 1554 J [ fi g +L gl * AUI ERII Tracheid à am LEE | by hatching, Endos; F | by stippling, Pal : | by parallel lines. Spongy phyll Each figure is based on tl ple, but individual vi y be from different seeds. Scale bars are all 1 mm. Cross sections are generally larger DER RENE T e M Us 4 IAI 1190 iandra heterophylla JAL 252. c. Citori fat IAI 203. € Barhi 197. d. Clitoria amazonum JAL 1594. e. Clitoria rubigii 1106 Journal of the Botanical R h Institute of Texas 1(2) Fic. 2. as under dissecting (a-g, i-o) and 1 (h) mi pe of Clitorii Is. a. C ium JAL 1262. b. Centrosema brasilianum JAL 1267. c. Centrosema pascuorum m JAL 1268, d. Centrosema sp. JAL 204 . f-g. a JAL 252. h. Periandra AROPE RE i-j. UM amazonum pon k. Mod laurifolia os 296. 1-0. ele tad JAL 1 are 1 mm (a-g, i-0) or 100 um (h). a Tracheid bar with extension spans entire Lackey, Cotyledon areoles of Clitoriinae 1107 covered with infrequent minute white cellular funicular remnants. Vascular bundle scar between posterior end of hilar groove and rim-aril; white; raised; prominent. Hilar groove visible; when open, allows view of white tracheid bar below; opens and closes in seconds in response to humidity; air bubble often forms on hilum upon immersion in water. Micropyle outside rim-aril, ca 0.05 mm long; Y-shaped when dry, with arms of Y towards hilum. Lens slightly perceptible as ca 0.3 mm cleft; 0.4 mm posterior to hilum. Lateral endosperm deposits sometimes perceptible externally as minute testa swellings. Epitesta deposits wanting. Seed in transverse section through the middle of the hilum with testa, endosperm, and two cotyledons of the embryo. Testa 0.05 mm thick dorsally to 0.3 mm thick at the hilum. Palisade (Malpighian) layer of one cell-layer thick; thinner dorsally; light line immediately under the outer surface except at the hilum where it shifts inward. Hourglass layer immediately interior; of a single cell layer; imperceptible dorsally, thin laterally, thicker ventrally, disappearing under the hilum. Parenchyma layer imperceptible dorsally and ventrally with 50x optics. Hilum testa much thicker and more differentiated than dorsal and lateral testa. Rim-aril and extension into hilar tongue top counter-palisade at outside edge. Funicular cell remnants few; ventral to counter-palisade. Hilum palisade and counter-palisade curved upwards towards hilar groove. Light line in hilum palisade about one-tenth down from outer surface; touching each other at the lips of the hilar groove when groove closed. Tracheid bar white; ovate; immediately below hilar groove; sometimes minutely cleft ventrally; attenuated dorsally into a contiguous or almost separate wing which touches or nearly reaches the endosperm. Parenchyma below hilum of columns of loosely joined dark cells with large air spaces. Endosperm dorsally and ventrally thin; laterally forming two thick pockets 0.05x0.5 mm, about equal to the thickness of the testa, aleurone layer imperceptible, middle and inner layers tan. Cotyledons off-white; concave on part of abaxial face to match expanded endosperm; mesophyll of spongy parenchyma, some palisade orientation of adaxial cells; starch test negative; vascular bundles 6 or 7 per cotyledon, middle one larger. Seed in longitudinal section reveals the same basic testa, endosperm, and embryo structure and de- scription will not be repeated from the transverse section unless there is additional information or greater clarity in this view. Testa palisade tissue slightly thicker above radicle than elsewhere. Vascular bundle from funiculus joins posterior end of the tracheid bar and extends into the raphe vascular bundle. Lens tissue between raphe vascular bundle and Malpighian layer of periclinal and scalariform-thickened white tracheoids similar to anticlinal cells of tracheid bar. Endosperm adnate to and lines the entire inner surface of the testa; transparent and almost colorless or slightly tan when uncut, allowing view of inner surface of testa through endosperm, white where cut or broken; when dry forms an impression of every feature of the embryo, down to the cellular level; thicker areas especially between radicle and cotyledon, and in a lateral thickening both left and right; cotyledon areole impression from embryo on each lateral endosperm thickening; swells on addition of water, especially hot water, becomes gelatinous, and embryo impression structure lost. l hich R | I ll ] | | | I 11 i i y I hal I | gl i tE La surface. ay eee Pape ye wen g bul fendost y Ker icin enn gimp fcotyled 1 I DOR , DM T ligt ly liaj apical position dl g Jinal i f | i f ] showing white tracheoid lens tissue. e, Ventral exterior iew of hil howi I hil hilar tongue, rim-aril, and funicular vascular aundie scar us posterior Wy the hilar (NUOVE. bind is ae open, allowing view of the aie tracheid par DN f. Transverse section of ventral g i wit! g g g right c cotyledo on P oved. Embry is ex Í. Posterior porti EE m cibla h Hane T eee ee P toluidine blue. A fl f pid | ti itl f ll | ighti f 0 fad l phyll cell [ palisade-li {indicated by € Jinterior of testa, s gk P lar bundles.j. T + of ventral testa, showing tl | lition, in which tl heid bar sj ly a porti ftl l hid k. Vi l ior view of hilum, showing two hilar tongues Epi Id i 1 hil |. Lateral ior vi f , Showing clear, colorless, viscid epi I deposi ait aa foreign meaner m. venal exterior view o nium, Dr two hiar tongues E Epitestal deposi ] hilum. n. Cotyledon areole eral light. o. lus al ter-palisade, seed held in position near funiculus by two hilar tanques Vascular bundle from funicul ] fhil ly breaks off at hil f Journal of the Botanical Research Institute of Texas 1(2) Fic. 3. ESEM of Clitoriinae seeds. a-b. Centrosema arenarium JAL 1262. c-d. Centrosema plumieri JAL 1266. e—f. Centrosema virginianum JAL 203. g-i. Periandra heterophylla JAL 252. j-k. Clitoria amazonum JAL 1594. |. Clitoria laurifolia JAL 1594. m. Clitoria rubiginosa JAL 295. n. Clitoria termatea JAL 197. o. Barbieria pinnata JAL 1271. Scale bars are 10 um in h-i, 100 um all others. a. Inner testa surface, showing cotyledon areole impression into endosperm and enlarged middle endosperm layer. b. Longitudinal section of lens cells. Tracheoids external to raphe vascular bundle. c. Abaxial Lackey, Cotyledon areoles of Clitoriinae 1109 Embryo consists of two cotyledons, radicle, and large plumule between cotyledons. Radicle oriented towards micropyle. Cotyledon areole ovate, medial, ca 0.3x0.3 (Fig. 3e), and immediately internal to lateral endosperm thickening (Fig. 36), stains about equally as surrounding cells with toluidine blue; imbibition damage of surrounding cells severe. Cells arranged more regularly in cotyledon areole than surrounding epidermal cells. Centrosema arenarium. Seed asin Centrosema virginianum. Seed spherical to cylindrical; ca 5.5x4.5x5.5 mm; brown with brown hilum. Hilar tongue minute or wanting. Micropyle oval. Counter palisade covered by material that appears to be fungal hyphae. Lens tracheoid cells (Fig. 3b) contiguous with tracheid bar by attenuated white tissue. Cotyledon areole (Fig. 3a) ovate, medial to medial-apical, ca 0.5x0.8. Centrosema brasilianum. Seed as in Centrosema virginianum. Seed reniform-cylindrical; ca 4.0x2.2x2.8 mm; brown to brown-tan mottled with brown hilum. Micropyle Y-shaped to broadly oval. Lateral testa exterior usually conspicuously swollen at site of endosperm deposits (Fig. 2b). Cotyledon areole circular, medial, ca 0.2x0.2 Centrosema molle. Seed as in Centrosema virginianum. Seed reniform-cylindrical; ca 5.0x2.7x3.2 mm; brown or brown-tan mottled with brown hilum. Micropyle Y-shaped to broadly oval. Cotyledon areole broadly ovate, medial, ca 0.2x0.2. Centrosema pascuorum. Seed asin Centrosema virginianum. Seed reniform-cylindrical; ca 4.0x1.5x2.5 mm; brown with brown hilum. Micropyle Y-shaped to broadly oval. Lateral testa exterior sometimes slightly swollen at site of endosperm deposits; interior cotyledon areole impression prominent (Fig. 2c). Cotyledon areole broadly ovate, medial, ca 0.2x0.2. Centrosema plumieri. Seed as in Centrosema virginianum. Seed spherical compressed; ca 7.0x5.0x6.0 mm; brown with brown hilum. Hilar tongue one-fourth length of hilum. Micropyle M c to end oval. Cotyledon mesophyll mostly spongy, one palisade-like layer on adaxial face. Cotyledon ar medial-apical, ca 0.5x0.5 (Fig. 3c,d). Centrosema sagittatum. Seed asin Centrosema virginianum. Seed reniform-cylindrical; ca 6.5x3.0x3.7 mm; brown with brown hilum. Micropyle broadly oval. Cotyledon areole broadly ovate, medial, ca 0.7x0.5. > Periandra densiflora. Seed as in Centrosema virginianum. Seed reniform-compressed; ca 3.5x?x3.5 mm (seeds immature, width uncertain); brown with brown hilum. Seeds not dissected, anatomy unknown. Periandra heterophylla. Seed as in Centrosema virginianum. Seed reniform compressed, ca 4.8x1.9x2.9 mm, brown or brown-tan mottled (Fig. 1b). Funicular remnants on counter-palisade sparse to common, especially visible immediately inside the rim-aril. Lens cells (Fig. 3h) with scalariform thickenings, remi- niscent of tracheid bar (Fig. 2f, 31), which has reticulate thickenings or pits. Cotyledon mesophyll mostly spongy, 1-2 palisade like layers on adaxial face, several irregular palisade-like layers on abaxial face (Fig. 2h). Starch test negative or faint. Cotyledon areole circular, basal-medial, ca 0.2x0.2 (Fig. 3g); almost im- perceptible under 50x. Plumule small to medium. Periandra mediterranea. Seed as in Centrosema virginianum. Seed reniform compressed, ca 5.8x1.7x4.3 mm, brown with tan hilum. Funicular remnants on counter-palisade sparse. Cotyledon mesophyll spongy. Starch test negative. Lens tracheoid tissue poorly developed, 0.05 mm thick. Cotyledon areole ovate, medial, ca 0.4x0.2; almost imperceptible under 50x. Plumule small. Clitoria (Clitoria) ternatea. Seed reniform to cylindrical; ca 5.6x2.9x4.2 mm; brown or brown-black mottled with brown hilum; sink in water (Fig. 1c). T POE "m ae O T See eee i --— cords areole. e. Ab ial tvled f ith cotvled | d pe Ils. f. | "EP face fi n in (e) with rn fe impression yl 9 i E i igh ; E ; ue i. Longitudinal section of tracheid bar. j. Abaxial cotyled face with impression of testa veins, k Testa, showing that the surface is not smooth. l. - ^ L Ļ A ‘tact Enitacts h fı + ai J b T b b Akawv:31 +tulard eb "XU £ J Ahavial TUAM S] from cs z showing ida pression d on inner surface 0 surface with i impression of cotyledon areola, 1110 Journal of the Botanical R h Institute of Texas 1(2) Seed externally with prominent hilum, anterior micropyle and posterior lens. Hilum oval ca 0.8x1.2 mm; surrounded by a slightly raised rim. Rim-aril tan, mostly of ca 0.2 mm raised bead, notched anterior; with hilar tongue ca 0.9 mm wide by 0.8 mm long attached along entire lateral side of hilum and distal end extended anterior. Hilar tongue on left in about half the seeds; on right in about half the seeds. Coun- ter-palisade surface covered with white cellular funicular remnants, especially adjacent to the rim-aril. Vascular bundle scar between posterior end of hilar groove and rim-aril; white; raised; prominent. Hilar groove visible; when open, allows view of white tracheid bar below. Micropyle outside rim-aril, ca 0.2 mm long; Y-shaped when dry, with arms of Y towards hilum. Lens slightly perceptible as ca 0.4 mm raised area posterior to hilum. Epitesta deposits wanting. Seed in transverse section through the middle of the hilum with testa, endosperm, and two cotyledons of the embryo. Testa 0.1 mm thick dorsally to 0.4 mm thick at the hilum. Palisade (Malpighian) layer of one cell-layer thick; thinner dorsally; light line immediately under the outer surface except at the hilum where it shifts inward. Hourglass layer immediately interior; of a single cell-layer; imperceptible dorsally and later- ally, thicker ventrally, disappearing under the hilum. Parenchyma layer imperceptible dorsally and laterally with 50x optics; increasingly thicker on ventral half. Hilum testa much thicker and more differentiated than dorsal and lateral testa. Rim-aril and extension into hilar tongue top outside edge of counter-palisade and hilar rim. Funicular cell remnants cover counter-palisade, absent from hilar groove area. Hilum palisade and counter-palisade curved upwards towards hilar groove. Light line in hilum palisade about one-tenth down from outer surface; touching each other at the lips of the hilar groove when groove closed. Tracheid bar white; ovate; immediately below hilar groove; sometimes minutely cleft ventrally; no dorsal extension. Sclerenchyma below hilum dark. Parenchyma below hilar rim white, with columns of loosely joined cells with large air spaces. Endosperm thin over entire inside of testa; laterally forming two slightly thicker areas adjacent to cotyledon areoles. Cotyledons tan-yellow; adaxial mesophyll of palisade parenchyma; abaxial mesophyll slightly palisade-like; interior mesophyll spongy; starch test negative; vascular bundles about 5, central one slightly larger. Seed in longitudinal section reveals the same basic testa, endosperm, and embryo structure. Testa palisade tissue thinner in lens area. Vascular bundle from funiculus joins the posterior end of the tracheid bar and extends into the raphe vascular bundle. Lens cells between Malpighian layer and raphe vascular bundle dark or light vertical cells. Endosperm adnate to and lines the entire inner surface of the testa; slightly transparent and dark, allowing partially obscured view of inner surface of testa through endosperm; when dry forms an impression of every feature of the embryo, down to the cellular level; thicker areas especially between radicle and cotyledon, and in a lateral thickening both left and right; cotyledon areole impres- sion from embryo on each lateral endosperm thickening; swells on addition of water, especially hot water, becoming gelatinous, and embryo impression structure lost. Embryo consists of two cotyledons, radicle, and minute plumule between cotyledons. Radicle oriented towards micropyle. Cotyledon areole ovate, medial to basal-medial, ca 0.9x0.4, and immediately internal to lateral endosperm thickening, stains less readily than surrounding cells with toluidine blue. Cells arranged more regularly in cotyledon areole than surrounding epidermal cells (Fig. 3n). Clitoria (Neurocarpum) rubiginosa. Seed globose; ca 4.0x4.0x4.0 mm; brown with tan and brown hilum (Fig. le); epitesta irregularly covers most of seed; sink when forced under water. Seed externally with prominent hilum, anterior micropyle and posterior lens. Hilum oval ca 0.9x1.1 mm; surrounded by a raised rim. Rim-aril tan, mostly of ca 0.1 mm raised bead; with hilar tongue ca 0.5 mm wide at base by 0.8 mm long positioned from slightly anterior of mid-hilum extending toward posterior end; a similar tongue, about half as wide and quarter as long, on opposite side of hilar tongue on posterior quarter. Larger hilar tongue on left in about half the seeds; on right in about half the seeds. Counter-palisade surface covered with white Tp funicular remnants. Vascular bundle scar between posterior end of hilar groove and rim-aril; white; raised; prominent. Hilar groove visible. Air bubble forms above counter-palisade and below hilar tongue upon immersion. Micropyle outside rim-aril, ca 0.1 mm long; Y-shaped when dry, Lackey, Cotyledon areoles of Clitoriinae 1111 with arms of Y towards hilum. Lens slightly perceptible as furrow immediately posterior to hilum. Epitesta deposits covering most testa surface except in hilum, lens, and micropyle area, for seeds in pods or freshly removed from pods; embedded with endocarp cell remnants, extraneous plant parts and foreign material, and sporadic epitesta deposits on hilum, lens, and micropyle area for seeds exposed to storage or handling; absolutely clear and colorless; of endocarp origin; strongly adhesive and plastic when warm, spreading and becoming a viscous liquid at ca 55? C; non-adhesive and firm, almost brittle, when cool; of strongly hydro- phobic material, when immersed in water, large air bubbles form on surface. Seed in transverse section through the middle of the hilum with testa, endosperm, and two cotyledons of the embryo. Testa 0.08 mm thick dorsally to 0.4 mm thick at the hilum. Palisade (Malpighian) layer of one cell-layer; thinner dorsally; light line immediately under the outer surface. Hourglass layer immediately interior; of a single cell-layer; imperceptible dorsally, thin laterally, thicker ventrally, disappearing under the hilum. Parenchyma layer imperceptible dorsally and ventrally with 50x optics. Hilum testa much thicker and more differentiated than dorsal and lateral testa. Rim-aril and extension into hilar tongue and second tongue top counter-palisade at outside edge and on hilar rim. Funicular cell remnants on counter- palisade. Hilum palisade and counter-palisade curved upwards towards hilar groove. Light line in hilum palisade about one-twentieth down from outer surface; touching each other at the lips of the hilar groove when groove closed. Tracheid bar white; ovate; immediately below hilar groove; sometimes minutely cleft ventrally; spanning most of the testa, with no dorsal extension, always with dark sclerenchyma between tracheid bar and endosperm. Cells below hilum of dark sclerenchyma. Endosperm lining testa, scarcely perceptible. Cotyledons white to yellow; mesophyll of spongy parenchyma; starch test positive; vascular bundles 5, faintly perceptible, middle one largest. Seed in longitudinal section reveals the same basic testa, endosperm, and embryo structure. Testa palisade tissue about equal thickness throughout. Vascular bundle from funiculus joins the posterior end of the tracheid bar and extends into the raphe vascular bundle. Lens tissue between raphe vascular bundle and Malpighian layer of dark vertically oriented sclerenchyma. Endosperm adnate to and lines the entire inner surface of the testa; thin and transparent, view of inner surface of testa through endosperm; thicker areas especially in slit between radicle and cotyledons, and as a ridge formed at juncture of cotyledon margins. Embryo consists of two cotyledons, radicle, and minute plumule between cotyledons. Radicle oriented towards micropyle. Cotyledon areole wanting; small medial patch of endosperm often adherent to cotyledon (Fig. 3m). A patch of light-colored epidermal cells apically. Clitoria (Neurocarpum) laurifolia. Seed as in Clitoria rubiginosa. Seed subglobose; ca 4.0x4.0x4.7 mm; brown with tan hilum; epitesta irregularly covers most of seed. Cotyledon areole wanting. Clitoria (Neurocarpum) mariana. Seed as in Clitoria rubiginosa. Seed subglobose; ca 4.4x3.8x4.0 mm; brown to brown-black with tan hilum; thin epitesta irregularly covers most of seed. Second hilar tongue small to scarcely perceptible. Small patch of endosperm adjacent to cotyledon areole. Starch absent. Cotyledon areole circular, basal-medial, ca 0.3x0.3. Cotyledon mesophyll spongy, perhaps slightly palisade-like adaxially. Clitoria (Bractearia) amazonum. Seed as in Clitoria rubiginosa. Seed circular, compressed, ca 7.4x3.4x7.5 mm; dark brown with tan hilum (Fig. 1d). Epitesta wanting (Fig. 3k). Hilar tongue attached to entire side of hilum. Second tongue smaller, attached on posterior hilum half (Fig. 2j). Testa venation massive (Fig. 2i); forms impression on cotyledons (Fig. 3j). Starch test positive. Cotyledon areole absent. Plumule unknown because of inadequate specimen Clitoria (Bractearia) brachycalyx. Seed as in Clitoria rubiginosa. Seed circular, compressed, ca 11.4x2.8x11.2 mm; dark brown with tan hilum. Epitesta wanting. Hilar tongue attached to entire side of hilum. Second tongue about a third as long, attached on entire opposite side. Testa covered with minute punctae. Testa venation massive; forms impression on cotyledons. Starch and cotyledon mesophyll unknown because of inadequate specimen. Cotyledon areole absent. Plumule unknown because of inadequate specimen. Barbieria pinnata. Seed reniform-oval to cylindrical; ca 5.0x1.8x3.0 mm; brown with brown hilum; epitesta irregularly covers most of seed; sink when forced under water (Fig. 1f). 1112 Journal of the Botanical R h Institute of Texas 1(2) Seed externally with prominent hilum, anterior micropyle and posterior lens. Hilum oval ca 0.7x1.5 mm; surrounded by a raised rim. Rim-aril tan, mostly of ca 0.1 mm raised bead; with hilar tongue ca 0.7 mm wide by 0.6 mm long positioned from mid-hilum extending toward anterior end; a similar tongue, about equally wide and half as long, on opposite side of hilar tongue and slightly posterior. Hilar tongue on left in about half the seeds; on right in about half the seeds. Counter-palisade surface mostly naked; covered with white cellular funicular remnants adjacent to the rim-aril. Vascular bundle scar between posterior end of hilar groove and rim-aril; white; raised; prominent. Hilar groove visible; when open, allows view of white tracheid bar below; air bubble often forms on hilum upon immersion in water. Micropyle outside rim-aril, ca 0.05 mm long; Y-shaped when dry, with arms of Y towards hilum. Lens slightly perceptible as ca 0.4 mm irregular crack; 0.4 mm posterior to hilum. Epitesta deposits covering most testa surface except in hilum, lens, and micropyle area, for seeds in pods or freshly removed from pods; embedded with extraneous plant parts and foreign material, and sporadic epitesta deposits on hilum, lens, and micropyle area for seeds exposed to storage or handling; absolutely clear and colorless; of endocarp origin; absent from immature seeds and some other seeds and some specimens; strongly adhesive and plastic when warm; non-adhesive and firm when cool; of strongly hydrophobic material, when immersed in water, large air bubbles form on surface. Seed in transverse section through the middle of the hilum with testa, endosperm, and two cotyledons of the embryo. Testa 0.08 mm thick dorsally to 0.5 mm thick at the hilum. Palisade (Malpighian) layer of one cell layer; thinner dorsally; light line immediately under the outer surface except at the hilum where it shifts inward. Hourglass layer immediately interior; of a single cell-layer; imperceptible dorsally, thin later- ally, thicker ventrally, disappearing under the hilum. Parenchyma layer imperceptible dorsally and laterally with 50x optics. Hilum testa much thicker and more differentiated than dorsal and lateral testa. Rim-aril and extension into hilar tongue and second tongue top counter-palisade at outside edge. Funicular cell rem- nants few; adjacent to rim-aril on counter-palisade. Hilum palisade and counter-palisade curved upwards towards hilar groove. Light line in hilum palisade about one-tenth down from outer surface; touching each other at the lips of the hilar groove when groove closed. Tracheid bar white; ovate; immediately below hilar groove; sometimes minutely cleft ventrally; with no dorsal extension. Parenchyma below hilum of columns of loosely joined dark cells with large air spaces. Endosperm dorsally and ventrally thin; laterally forming two thick pockets 0.07x1.3 mm, about equal to the thickness of the testa, aleurone layer imperceptible, middle and inner layers tan, vitreous. Cotyledons tan-yellow; concave on part of abaxial face to match expanded endosperm; mesophyll of palisade parenchyma; starch test negative; vascular bundles not seen. Seed in longitudinal section reveals the same basic testa, endosperm, and embryo structure. Testa palisade tissue about equal thickness throughout. Vascular bundle from funiculus joins the posterior end of the tracheid bar and extends into the raphe vascular bundle. Lens tissue between raphe vascular bundle and Malpighian layer of dark sclerenchyma. Endosperm adnate to and lines the entire inner surface of the testa; slightly transparent and dark, allowing partially obscured view of inner surface of testa through en- dosperm; when dry forms an impression of every feature of the embryo, down to the cellular level; thicker areas especially between radicle and cotyledon, and in a lateral thickening both left and right; cotyledon areole impression from embryo on each lateral endosperm thickening; swells on addition of water. Embryo consists of two cotyledons, radicle, and minute plumule between cotyledons. Radicle oriented towards micropyle. Cotyledon areole ovate, medial to basal-medial, ca 0.9x0.4, and immediately internal to lateral endosperm thickening, stains less readily than surrounding cells with toluidine blue. Cells arranged more regularly in cotyledon areole than surrounding epidermal cells. DISCUSSION Clitoriinae seeds show a diversity of seed characters. Cotyledon areole presence All studied specimens have cotyledon areoles except Clitoria subgenus Bractearia and two species of Clitoria subgenus Neurocarpum. The current study gives first reports of cotyledon areole presence in the genera Lackey, Cotyledon areoles of Clitoriinae 1113 Periandra and Barbieria as well as several named species of Centrosema, Clitoria (Neurocarpum) mariana, and Clitoria (Clitoria) ternatea. Absence in Clitoria subgenus Bractearia and C. laurifolia and C. rubiginosa seems to be confirmed by diligent searches with light microscopy and SEM. In C. rubiginosa, small endosperm remnants were often found in a medial position on the cotyledon (Fig. 3m), which has been found to be an occasional indicator of cotyledon areole presence in some scant endosperm specimens of some tribe Phaseoleae (unpublished results); however several attempts did not show clear cotyledon areoles or any other clear indication of cotyledon areoles in C. rubiginosa. When present, cotyledon areoles were always circular to ovate and in a medial to basal-medial or slightly medial-apical position, above the midvein, and opposite a local enlargement of endosperm. Some cotyledon areoles were small, especially in Periandra, and difficult to detect initially. In Barbieria, cotyledon areoles were the largest in the subtribe and conspicuous (Fig. 2n). No seeds were available of Clitoriopsis mollis, the single species of the remaining genus in the subtribe. inet o no of cotyledon areole position and appearance in these plants are in accord with ns for other papilionoid 1 legumes (Beck 1878; Dzikowskil936, 1937; Miksche 1961; Yaklich et cal 1984, 1986, 1987, 1989, 1992, 1995, 1996, 1998; Baker & Minor 1987; Endo & Ohashi 1997, 1998a, 1998b, 1999a, 1999b; Matarese & Fasci 2002; Ma et al. 2004). The most immediately obvious indication of cotyledon areole presence is an area of epidermal cells with “granular projections” (Endo & Ohashi 1998a), in other words, projecting external cell walls. All cotyledon areoles were identifiable by these projections, and cotyledon areoles seen here matched the previously reported characteristics of an unbranched single spot on the abaxial surface of each cotyledon, above the midvein, opposite an enlarged middle endosperm layer which takes a mirror-image impression of the cotyledon areole (Fig. 3c-d, e-f), and with cells of dif- ferent size and shape than other cotyledon epidermal cells. Endo and Ohashi (1999a) reported one Clitoria sp. specimen and one Centrosema sp. specimen with medial circular to oval cotyledon areoles. Their two observations are confirmed by material seen here. Cotyledon areole correlation with other seed characters Presence and position of cotyledon areoles is always associated with an enlarged area of middle endosperm. And the size of the cotyledon areole seems to be roughly aes with EIE size of the middle endosperm ll, and so are the middle layer. For instance, cotyledon areoles in most Centrosema and P endosperm layers, which can sometimes be seen as a small ne on n testa ER (Fig. 2b). Barbieria, and to a lesser degree, Clitoria (Clitoria) ternatea, have larger cotyledon areoles, and these are associated with larger, visible endosperm. In Clitoria subgenera Neurocarpum and Bractearia neither cotyledon areoles nor discernible endosperm is found, except for a small visible patch of endosperm and associated cotyledon areole in Clitoria (Neurocarpum) mariana; endosperm was also noted in this species by Pammel (1899). In addition, absence of cotyledon starch is correlated with presence of cotyledon areoles and endosperm, with the realization that the starch test employed here is relatively crude. Nadelmann (1890) and Bailey (1971) also mention a negative correlation between cotyledon starch and endosperm galactomann reserves. No correlation could be found to exist between presence of canavanine, restricted to seeds of Centrosema (Lackey 19773), and any other seed character observed here. This study expands upon Smith's (1981, 1983) findings, supported by earlier Pammel (1899) findings, that Clitoria (Clitoria) ternatea fit Smith's form 1 and Centrosema molle (as C. pubescens) and Clitoria (Neurocarpum) mariana fit form 4 cotyledon and seed arrangement. Form 1 seeds, found in about 58% of legume species, have leaf-like photosynthetic cotyledons, adaxial palisade tissue, endosperm, and are epigeal. A remaining 3096 (form 4) of legumes have fleshy, storage cotyledons, no adaxial palisade tissue, no obvious endosperm, and are either hypogeal or epigeal. The final 12% (forms 2 and 3) are intermediate or anomalous forms. In the current study, C. ternatea and Barbieria are shown to be form 1 seeds. Both have palisade mesophyll tissue, and conspicuous endosperm. Clitoria (Clitoria) ternatea is epigeal (Candolle 1825; Lubbock 1892; Baudet 1974; Yeh et al. 1987). Germination of Barbieria is unknown, but can be presumed to be epigeal because of the conspicuous palisade cotyledon mesophyll. Clitoria subgenus Bractearia and most subgenus Neurocarpum species do not have cotyledon palisade mesophyll, lack obvious endosperm, do have cotyledon 1114 Journal of the Botanical R h Institute of Texas 1(2) starch storage, are epigeal or hypogeal, and fit Smith's form 4 seeds. Clitoria (Neurocarpum) mariana seems to be an intermediate form: no true palisade mesophyll is formed, the cotyledons are thick storage structures, a small amount of endosperm is present, germination is epigeal (Holm 1891), and a starch test is negative. = Although Smith considered Centrosema molle to be type 4, species of Centrosema and Periandra did show some tendency for palisade-like adaxial cotyledon mesophyll, had small amounts of endosperm, no obvious starch idered intermediate forms. All of these storage, and are epigeal or hypogeal. They also could therefore be structural adaptations and observations, endosperm, cotyledon starch, cotyledon mesophyll palisade tissue, (Smith 1983). As suggested here by correlation in Clitoriinae, cotyledon areoles also seem part of this character group. and germination epigeal or hypogeal, seem to be various adaptations to germination [m Systematic value of seed characters The survey given here, although limited in number of species and intensity, l other structures may be of taxonomic and morphological interest, and deserve more study. Two characters distinguish species of Centrosema and Periandra from species of other genera. The first is the peculiar lens cell type (Fig. 2c, d, g, 3b, h) which forms a white tissue of cells similar to the distinc- tive papilionoid tracheid bar cells (Lersten 1982), but the lens cells have scalariform thickenings, and are oriented parallel to the testa surface as well as the immediately dorsal raphe vascular bundle; cells of the tracheid bar have pits or reticulate thickenings, and are oriented perpendicular to the testa surface. In other genera of Clitoriinae, the lens cells do not appear to consist of white tracheoids or have such thickenings, and are unexceptional relative to other papilionoid legumes. The second distinguishing character is a dorsal wing or mere extension of the tracheid bar which appears at low magnification to project the tracheid bar to span the entire testa thickness and contact, or almost contact, the innermost boundary of the testa. Some specimens or species in genera other than Centrosema and Periandra approach this tracheid bar condition; in Clitoria (Neurocarpum) laurifolia and C. (Neurocarpum) rubiginosa the tracheid bar spans most of the testa, but always there is a layer of dark testa cells between the tracheid bar and the innermost layer of testa. A distinct cotyledon palisade mesophyll is found only in Clitoria subgenus Clitoria and in Barbieria. In these genera, about three prominent adaxial layers and several smaller abaxial layers surround a central spongy layer. Mesophyll entirely of spongy tissue was in Clitoria subgenus Bractearia and most Clitoria sub- genus Neurocarpum. In Centrosema and Periandra, and possibly Clitoria (Neurocarpum) mariana, mesophyll was spongy and no true palisade tissue exists, but some mesophyll cells, particularly adaxial mesophyll cells, had one or two cell layers of somewhat palisade-like orientation (Fig. 2h). Current findings concur with Pammel's (1899) report of palisade mesophyll in Clitoria (Clitoria) ternatea and no palisade mesophyll in C. mariana and Smith's (1983) report of palisade tissue in C. ternatea and only spongy mesophyll in Centrosema molle (as C. pubescens). In Barbieria and to a lesser degree in Clitoria subgenera Neurocarpum and Bractearia, the hilum bears the common papilionoid hilar tongue on the left or right, and a second smaller hilar tongue on the opposite side (Fig. 2k, m, o). In some specimens of Clitoria (Neurocarpum) mariana the second tongue is very small or essentially missing. Two hilar tongues are uncommon in other Phaseoleae (Fig. 2e). To my knowledge, it occurs conspicuously in the Phaseoleae only elsewhere in Amphicarpaea, Dumasia, Cologania, and Dysolobium; however, slight and barely perceptible suggestions of a second hilar tongue or minute hilar tongue pair (as in Galactia spp.) may occur in other genera (unpublished results). Berg (1979) described the development of the common single hilar tongue in Phaseoleae subtribe Kennediinae and the possible role in seed abscission The function of a second hilar tongue is unknown. An odd viscid thermoplastic epitesta occurs in Barbieria and Clitoria subgenus Neurocarpum (Fig. 2k, l, m, o, 3D. Noted at least since Bentham (1858) as a “peculiar viscid exudation” or “glanduloso-viscosa” in Clitoria subgenus Neurocarpum, in contrast to other species with “smooth” or “laevia” seeds. Fantz (1996b) and Kirkbride et al. (2003) additionally reported this epitesta in Barbieria. The term “smooth,” as used here, is not appropriately the opposite of viscid. In Lersten's (1981) survey of legume testa patterns, the term smooth, or levigate, is one of nine testa patterns under SEM. Presence of epitestal deposits was considered a character distinct from these nine patterns. He reported testa patterns in Centrosema, Clitoria, and Barbieria Lackey, Cotyledon areoles of Clitoriinae 1115 were rugulate to irregular papillose. None were considered smooth or levigate. The current study concurs with Lersten that the testa (Fig. 3k) is patterned, not smooth, under low magnification ESEM. It would, therefore, seem more appropriate to follow Lersten, and restrict the use of the term smooth for true testa patterns, and separately describe seeds as epitestal or non-epitestal. Js The epitesta seen here, differs from the epitesta common in Glycininae (Schleiden & Vogel 1839; Newell & Hymowitz 1978; Lackey 1981; Gijzen et al. 1999), even though both are of endocarp origin. The epitesta of Glycininae is dark, opaque, in cell-like or other rough irregular patterns, dusty or friable, and is unaffected by temperatures to about 60? C. Gijzen et al. (1999) showed that this epitesta in soybean was a protein, which, when sequenced, exhibits a short hydrophilic end and a longer hydrophobic end. The epitesta in Barbieria and Clitoria subgenus Neurocarpum is much different. As with Glycininae, it also covers most testa surfaces except the hilum, lens, and micropyle area for seeds in pods or freshly removed from pods, but it differs in spreading upon warm contact to other areas of the testa and other objects, and becomes embedded with extraneous plant parts and foreign matter. It is absent from some immature seeds and some other seeds and some specimens. It is absolutely clear and colorless, strongly adhesive and plastic when warm, and non-adhesive and firm, almost brittle, when cool. When these testa adherent deposits are cool and broken, they form curved fracture planes and peel away from the testa, exposing an inner epitestal surface with an impression of testa patterns (Fig. 31). Above 55? C, the epitesta becomes a viscous, spreading, sticky liquid, which cools to a smooth, shiny, surface (Fig. 2m). It is of strongly hydrophobic material. When immersed in water, large air bubbles form on the surface. The functional significance of these deposits must await further study. In Clitoria subgenus Bractearia, testa venation from the two recurrent vascular bundles is especially prominent (Fig. 2i), to a degree that impressions are formed on the abaxial cotyledon surface (Fig. 3j). No other specimens exhibited such a tendency. The functional need for such massive veins is unknown. Generic circumscription (Fantz 1996b) in what is now considered Clitoriinae has remained reasonably stable since Bentham (1858, 1865), who recognized four genera: Centrosema, Periandra, Clitoria, and Barbie- ria. Clitoria was further divided into three sections. Barbieria was distanced from the remaining members and allied with Tephrosia (Bentham 1865). Wilczek (1954) added the monotypic genus Clitoriopsis. Recent taxonomic work, aside from the revisions of Centrosema (Barbosa-Fevereiro 1977) and Brazilian Periandra (Funch & Barroso 1999), has been dominated by the works of Fantz (1976, 1979a, 1979b, 1988, 1991, 1996a, 1996b, 2000, 2001, et lit. infra), which focused on Clitoria, and presented the first serious study since Ben- tham (1858). Generic limits remained as with Bentham, with the addition of Clitoriopsis, acknowledgment of Barbieria as a distinct genus near Clitoria, and recognition of Bentham's Clitoria sections as subgenera. The question of recognition of Clitoria as a single genus, or two or three genera, and the affinity of Barbieria with these has long been a source of ambiguity. Bentham (1858) merged Neurocarpum, which he formerly held to be generically distinct, with Clitoria, and formed another section, now known as subge- nus Bractearia. Lackey (1977b) was inclined to divide Clitoria to the status before Bentham's merger. Fantz (19962) recognized that one could argue for three distinct genera, but that the flowers are unique within the papilionoid legumes, and therefore maintained a single genus. Barbieria has, likewise, always caused much trouble. Originally described as a species of Galactia in 1807, of Clitoria in 1811, and then transferred to the new genus Barbieria in 1825 (Fantz 19963), it had been aligned with several legume groups before realignment with and returned as a species of Clitoria (Lackey 1981). The realignment was suggested by Rob Geesink (personal correspondence 1979), who thought Barbieria better placed close to Centrosema or Clitoria. Fantz (19962) used extensive morphological evidence to argue for a resegregation of Barbieria. Studies using molecular techniques do not clarify much at this time. Doyle and Doyle (1993), used cpDNA inverted repeat regions, to study phylogeny in 43 genera of tribe Phaseoleae. They only sampled Centrosema virginianum and Clitoria (Clitoria) ternatea, and found these two formed a close pair relative to other samples. Kajita et al. (2001), using rcbL sequences and the same two species, found essentially the same thing, but as part of a broader survey. The generic and subtribal structure has not been studied by molecular techniques. 1116 Journal of the Botanical R h Institute of Texas 1(2) Results of the current limited study of seeds, admittedly in need of much broader survey for taxonomic purposes, suggest a few affinities within the Clitoriinae. Despite individual species variation, Centrosema and Periandra seeds are essentially all the same in basic form, and distinct from anything else in the subtribe. Aside from consistent medial cotyledon areoles with an adjacent patch of endosperm from an expanded middle endosperm layer (often externally visible as a lateral protrusion of the testa), there is the peculiar and consistent lens structure of white tracheoids. In addition, the tracheid bar extension to the full thickness of the testa sets these two genera apart. The three subgenera of Clitoria have distinctive seeds. Subgenus Clitoria, only represented here by a single species, has cotyledon areoles, adjacent endosperm enlargement, palisade cotyledon mesophyll, nega- tive starch test, inconspicuous testa venation, a single hilar tongue, and no epitesta. Subgenus Bractearia has no cotyledon areoles, no visible endosperm enlargement, only spongy cotyledon mesophyll, positive starch test, conspicuous testa venation, a second hilar tongue, and no epitesta. Subgenus Neurocarpum usually has no cotyledon areoles, no visible endosperm enlargement, only spongy cotyledon mesophyll, positive starch test, inconspicuous testa venation, a second hilar tongue, and the peculiar viscid thermoplastic epitesta. Clitoria (Neurocarpum) mariana does not fit the pattern for the remainder of the subgenus. It differs by pres- ence of cotyledon areoles, endosperm presence, and negative starch test. It would be enlightening to see if a broader review would maintain these distinctions or clarify the relationships of the subgenera. Barbieria has equally distinctive seeds. With Clitoria subgenus Neurocarpum it shares the peculiar epitesta and second hilar tongue, but almost no other seed characters, with the exception of Clitoria (Neurocarpum) mariana. Barbieria has prominent cotyledon areoles, conspicuous endosperm enlargement, prominent pali- ade cotyledon mesophyll, and a negative starch test. These characters place Barbieria in a peculiar position. If one regards the endospermic seed with palisade cotyledon mesophyll as basic and the non-endospermic seed with spongy cotyledon mesophyll as derived (Smith 1983), then Clitoria subgenus Clitoria and Barbieria are basic for this condition, and most members of the other subgenera of Clitoria are derived. On the other hand, one could assume that the peculiar epitesta is a derived character, and therefore unites Clitoria section Neurocarpum and Barbieria, even though they are currently placed in two distinct genera. A good deal more information on the diverse seed anatomy of Clitoriinae is needed for further taxonomic interpretation. ACKNOWLEDGMENTS Thanks go to Joe Kirkbride for thoughtful discussion and use of facilities and collections at BARC. Scott Whittaker and Stanley Yankowski were most helpful with ESEM and anatomical work. John Kress and War- ren Wagner graciously made available the collections and facilities of the Smithsonian Institution. REFERENCES BaiLev, RW, 1971. Polysaccharides in the Leguminosae. In: Harborne, J.B., D Boulter, and B.L. Turner. Chemotax- onomy of the Leguminosae. Academic Press, London and New York. Pp. 503-541. Baker, D.M. and H.C. Minor. 1987. 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Ethnobotany of Clitoria (Leguminosae). Econ. Bot. 45:511-520. Fantz, PR. 1996a. Taxonomic notes on the Centrosema pubescens Bentham complex in Central America (Legu- minosae: Phaseoleae: Clitoriinae). Sida 17:321-332. FANTZ, PR. 1996b. Resegregation of Barbieria from Clitoria (Leguminosae: Phaseoleae: Clitoriinae). Sida 17: 55-68. Fantz, PR. 2000. Nomenclatural notes on the genus Clitoria for the Flora North American project. Castanea 65: FANTZ, PR. 2001. Barbieria, Centrosema, Clitoria. In: Tellez, O. Flora of Nicaragua. Fabaceae Lindl. Pp. 961-962, 9 m FUNCH, L.S. and G.M. Barroso. 1999. Rivisáo taxonómica do género Periandra Mart. ex Benth. (Leguminosae, Papilionoideae, Phaseoleae). Rev. Brasil. Bot., Sao Paulo 22:339-356. Guzen, M., S.S. MILLER, K. KurLu, R.I. BuzzeLL, and B.L.A. Miki. 1999. Hydrophobic protein synthesized in the pod en- docarp adheres to the seed surface. Pl. Physiol. 120:951-959. GUNN, C.R. 1981. Seeds of Leguminosae. In: Polhill, RM. and PH. Raven. Advances in legume systematics. Part 1. Royal Botanic Gardens, Kew. Pp. 913-923. Horm. 1891. Contributions to the knowledge of the germination of some North American plants. Mem. Torrey Bot. Club 2:57-108. Kasra, T., H. OHASHI, Y. TATEISHI, C.D. Bai ev, and J.J. Dove. 2001. rbcL and legume phylogeny, with particular reference to Phaseoleae, Millettieae, and allies. Syst. Bot. 26:515-536. KIRKBRIDE, J.H., Jr, C.R. GUNN, and A.L. Weitzman. 2003. Fruits and seeds of genera in the subfamily Faboideae (Faba- ceae). USDA ARS Tech. Bull. 1890:v + 1-1208. £L Dag PRA D hi Air 1118 Journal of f Texas 1(2) LAcKEY, J.A. 1977a. A revised classification of the tribe Phaseoleae (Leguminosae: Papilionoideae), and its relation to canavanine distribution. Bot. J. Linn. Soc. 74:163-178 Lackey, J.A. 1977b. A synopsis of Phaseoleae. Ph.D. Dissertation. lowa State University, Ames. Lackey, J.A. 1981. Phaseoleae. In: Polhill, RM. and PH. Raven. Advances in legume systematics. Part 1. Royal Botanic Gardens, Kew. Pp. 301-327. Lersten, N.R. 1981. Testa topography in Leguminosae, subfamily Papilionoideae. Proc. Ilowa Acad. Sci 88: 180-191. LERSTEN, N.R. 1982. Tracheid bar and vestured pits in legume seeds (Leguminosae: Papilionoideae). Amer. J. Bot. 69:98-107. LuBBOCK, J. 1892. Contribution to our knowledge of seedlings. 2 vols. Kegan Paul, Trench, Trubner, London. Ma, F., C.A. Peterson, and M. Guzen. 2004. Reassessment of the pits and antipits in soybean seeds. Canad. J. Bot. 82:654-662. Maatin, A.C. 1946. The comparative internal morphology of seeds. Amer. Midland Nat. 36:513-660. MATARESE PALMIERI, R. and A. Fasa. 2002. Le areole cotiledonari in alcune Leguminosae. In: 98 Congresso Nazionale SBI Catania 24-26 September. P. 24. MikschE, J.P. 1961. Developmental vegetative morphology of Glycine max. Agronomy J. 53:121-128. NADELMANN, H. 1890. Ueber die Schleimendosperme der Leguminosen. Prings. Jarhb. Wissen. Bot. 21:609-691 Oe: NeweLL, C.A. and T. Hvwowrrz. 1978. Seed coat variation in Glycine Willd Brittonia 30:76-88. PAMMEL, L.H. 1899. Anatomical characters of the seeds of Leguminosae, chiefly genera of Gray's manual. Trans. Acad. Sci. St. Louis 9:91-273 + xxxv ic. SCHLEIDEN, M.J. and J.R.T. VoceL. 1839. Über das Albumen, insbesondere der Leguminosen (nebst einem Anhange). Nova Acta Acad. Caes. Leop.-Carol. 19(1):53-96 + 6 ic. ScHRIRE, B.D. 2005. Phaseoleae. In: Lewis, G., Schrire, B., Mackinder, B., and Lock, M. Legumes of the world. Royal Botanic Gardens, Kew. Pp. 393-432. Sum, D.L. 1981. Cotyledons of the Leguminosae. In: Polhill, R.M. and PH. Raven. Advances in legume systematics. Part 2. Royal Botanic Gardens, Kew. Pp. 927-940. Smr, D.L. 1983. Cotyledon anatomy of the Leguminosae. Bot. J. Linn. Soc. 86:325-355. STERLING, C. 1954. Development of the seed coat of lima bean (Phaseolus lunatus L.). Bull. Torrey Bot. Club 81: 2 "i Glycine (Leguminosae) by SEM. WILCZEK, R. 1954. Groupes nouveaux des Phaseoleae-Phaseolinae du Congo Belge et du Ruanda-Urundi. Bull. Jard. Bot. Etat Brux. 24:405-450 YaKLICH, R.W, E.L. Visit, and W.P. Werain. 1984. Scanning electron microscopy of soybean seed coat. Scan. Electron Microscop. 3:991-1000. YakucH, R.W, E.L. Vici, and W.P. Wercin. 1986. Special secretory cells in the soybean seed coat. Protoplasma 134: 78-87. YAKLICH, R.W., E.L. Vici, and W.P. Werain. 1987. Changes in structure of pit and antipit in soybean seeds and seedling development. J. Seed Technol. 11:151-157 YakLicH, R.W, E.L. Via, and W.P. Wera. 1989. The pit and antipit in the genus Glycine. Crop Sci. 29:1304-1309. YaKLicH, R.W, E.L. Via, and W.P. Werain. 1992. The fine structure of aleurone cells in the soybean seed coat. Pro- toplasma 167:108-119. YaKLICH, R.W., E.L. Vici, and W.P. Werain. 1995. Morphological and fine structural characteristics of aerenchyma cells in the soybean seed coat. Seed Sci. Technol. 23:321-330. YakucH, R.W., W.P. Werain, and E.F. Ene. 1996. Observation of unique structures between the endosperm and embryo in seeds of Glycine max. Seed Sci. Res. 6:183-189. YaKLicH, R.W., E.L. Vici, E.F. Erge, and W.P. Werain. 1998. Developmental changes of antipit cell ultrastructure in the soybean seed coat during seed maturation. Scan iatis eR 12523-532. Yen, M.S., H. Yuasa, and K. KyoDa. 1987. Seedli logy of the tribe Phaseoleae (Papilionoideae, Leguminosae) as an aid to their taxonomy. J. Taiwan Mus. 40: 19- 28. REDISCOVERY OF ELEOCHARIS KLEINII (CYPERACEAB), AN OVERĽOOKED SPECIES FROM THE HIGHLANDS OF SOUTH BRAZIL R. Trevisan D.J. Rosen Programa de Pós-Graduação em Botânica S.M. Tracy Herbarium A Federal do Rio do Sul Department of e e a iS nto Goncalves Porto Alegre, ds pau do Sul, i P 970, BRASIL College Station, Texas 7843-2 126, U.S.A. rftrevisan@yahoo.co l.l. Boldrini Departamento de Botânica U En do Rio o do Sul to Gonçalve Porto Alegre, E. es do Sul, » D -970, BRASIL ilsi.boldrini@ufrgs.br ABSTRACT Dr. Manuel B lescribed Eleocharis kleinii in 1966 based | llection from S Catarina state, Brazil. The species is closely related to E. liesneri Pins in pun of the culms, floral scales, dd bristles, and achene surface sculpturing. An expanded description, not additional f Santa Catarina and Rio Grande do Sul states, a key to FEN E. kleinii from the similar species, and TEM are presented. RESUMO Dr. Manoel Barros descreveu Eleocharis kleinii em 1966 tendo como base um único material coletado no estado de Santa Catarina, 1 Brasil. E. kleinii é semelhante à E. liesneri diferindo desta pelas características do escapo, glumas, o superfície do aquénio. Este trabalho consiste de uma nud n p da a nota lógi tatus de conservação, citação i S 1 pa A , uma m dicotómica para distinguir E kleinii de espécies semelhantes e ilustrações. Eleocharis R. Br. is a worldwide genus that includes ca. 200 species with a remarkable richness in tropical and subtropical America (González-Elizondo & Tena-Flores 2000). Distinctive features, such as unbranched aerial culms, leaves reduced to tubular sheaths, an inflorescence constituted of one spikelet on the apex of culms without involucral bracts, and the stylopodium enlarged and persistent on the achene are uniformly — found in this genus. Almost all species of Eleocharis are restricted to wetlands, often in muddy soils that are temporarily wet or inundated. The genus has not been treated comprehensively since the seminal work of Svenson (1929, 1932, 1934, 1937, 1939). Others (Blake 1939; Barros 1947, 1960; González-Elizondo 1994; Faria 1998; González-Elizondo & Reznicek 1998; Smith et al. 2002; Gil 2007; Trevisan 2005) have provided more regional treatments. Trevisan and Boldrini (2005) published records of Eleocharis ochrostachys Steud. from Rio Grande do Sul and Santa Catarina States, Brazil. However, additional study of the specimens cited, including comparison to types of E. ochrostachys, led us to consider that these collections p represented an undescribed species of Eleocharis. Recent correspondence with Drs. Rosa Guaglia SD and Socorro González-Elizondo (CIIDIR) has brought to our attention that one of the specimens studied by the first author (Klein 3718a ) while visiting Herbário Barbosa Rodrigues (HBR) in September, 2006, is the holotype of a poorly known species from the highlands of South Brazil, Eleocharis k as a type, hence our earlier opinion that it was an undescribed species. We fond no reference to this species pads einii Barros. The holotype bears no annotation as either E. kleinii, or in checklists or floristic treatments since the protologue, although it is to be included in the forthcoming Catalogo de la Flora del Cono Sur (pers. comm. Drs. Rosa Guaglianone and Socorro González-Elizondo). J. Bot. Res. Inst. Texas 1(2): 1119 — 1124. 2007 1120 Journal of the Botanical R h Institute of Texas 1(2) The protologue of Eleocharis kleinii provides only a brief description based on a single specimen from Santa Catarina state with no illustration. The purpose of this manuscript is to improve the knowledge of E. kleinii by providing an expanded description, ecological information and conservation status, additional specimen citations, a key to distinguish E. kleinii from the similar species, and illustration. Eleocharis kleinii Barros, Sellowia 18:49. 1966. (Fig. 1 a—e). Ter: BRAZIL: Santa Catarina, Irineópolis (-Valóes), 10 Dec R.M. Klein 3718a (HOLOTYPE: HBR!; isotype: SI!). Perennial herb; roots coarse, fibrous, mostly drab-brown; rhizomes elongated, 2-3 mm thick, dark maroon, the scales 1.2-1.5 cm long. Culms obtusely trigonous distally, terete proximally, 30-70 cm x 1.3-2 mm, soft, internally spongy with incomplete transverse septa, smooth to finely longitudinally striate, light green (grayish green when dry). Leaves 2, reduced to sheaths, membranous, loose, friable, proximally venose, upper sheath 7-18 cm, apex narrowly acuminate and extended into a soft awn, proximally dark maroon, becoming chestnut to tawny distally. Spikelets cylindric, acute, 12-32 x 2.5-4 mm, two times wider than culms, 16- to 30-flowered; proximal scale empty (rarely with a flower), amplexicaulous and appearing as a continuation of the culm; remaining fertile floral scales conspicuously arranged in 4 spiraled rows, weakly imbricate, somewhat appressed to loose at maturity, ovate, 5-6.2 x 2-3.5 mm, medially cartilaginous, marginally and distally translucent hyaline-erose, central area slightly round-keeled, apex acute, abaxiall veins slightly raised to cellular-lineate (the centrally located veins sometimes reddish), only mid-vein dis- tinguishable in adaxial view, medially greenish to stramineous, laterally reddish. Flowers with 6 perianth bristles, slender distally, becoming slightly coarser proximally, sub-equal, exceeding the stylopodium, densely retrorsely spinulose to below the middle with fine spinules, white to yellowish white; stamens 3; anthers 1.9-2.1 mm long, yellow to stramineous; style bifid or trifid. Achenes biconvex, obovoid to obpyriform, 2-2.2 mm x 1.3-1.4 (1.9) mm, with 20-28 longitudinal rows of slightly transversely elongated polygonal cells, finely sculptured to nearly smooth at 20x, dull buff yellow maturing to olivaceous, apex constricted to a distinct neck. Stylopodium triangularly elongate, dorsoventrally compressed with a thickened basal rim, somewhat curvate, 0.7—1.5 x 0.5-0.6 mm, dark brown. Distribution.—South America: Brazil (Rio Grande do Sul and Santa Catarina states). Endemic to the highlands of South Brazil, this species has been endangered by the silviculture of Pinus species. Some wetlands have been dried to cultivate this exotic species on a large scale. The species fits in category of Vulnerable in level D (UICN 2001). Habitat.—The species grows in permanently flooded environments in Du of South Brazil. The 1 21. plants generally have culms partially submerged, forming clonal masses (Fig. 2). Eleochai inii occurs above 30? latitude S, between 500-1400 m above sea. These high regions are the coldest in South Brazil, with an average temperature of ca. 17?C and winter occurrences of frost or snow (Boldrini 1997). The pluviometry is 1500 to 2000 mm a year with rains distributed throughout the year (Moreno 1961). Other Specimens Examined. BRAZIL. Rio Grande do Sul: Cambará do Sul, Faxinal, Dec 1983, M. Sobral & J. R. Stehmann 2717 M ICN), Cambará do Sul, Feb 1948, B. Rambo 36659 (PACA); Campestre da Serra, BR 116, km 70, 03 Nov 2003, Trevisan, R. Lúdtke, Vignoli-Silva & L. Mentz 133 (ICN); São Francisco de Paula, 1996, H.M. Longhi-Wagner (ICN 129292); São Francisco de Paula, 29°17'57" S 50°20'08.8" W, 28 Nov 2003, Trevisan & Boldrini 309 (ICN, MO, TAES); Sao José dos Ausentes, Serra da Rocinha, 03 Feb 1953, B. Rambo 53898 (HBR, PACA). Santa Catarina: Bom Jardim da Serra, desfiladeiro do Funi, Feb 1989, M. Sobral et al. 6480 (ICN); Bom Jardim da Serra, Serra do Oratorio, 15 Dec 1958, R. Reitz & R. Klein 7991 (HBR); Matos Costa, SC 302, km 31, 26°27'41" S 51°07'32,1" W, 21 Dec 2006, R. Trevisan 804 (ICN). Barros (1966) correctly placed Eleocharis kleinii in subgen. Limnochloa (P. Beauv. ex Lestib.) Torr. This species is included in this subgenus on m basis of the following features: indurate floral scales with a distinct hyaline margin; proximal scal g continuation of the culm; achenes ornamented with longitudinal rows of polygonal cells (oa Elizondo & Peterson 1997). Barros (1966) indicated that E. kleinii was very close to E. elongata Chapm., sharing with it slender culms and aquatic habitat. However, we believe that E. kleinii is most closely related to E. liesneri S. González & Reznicek (Fig. 1 f-g), another species from the highlands of South America (González-Elizondo & Reznicek 1998; Rosen 2006). Eleocharis kleinii differs from E. liesneri 1121 Trevisan et al., Rediscovery of Eleocharis kleinii Nikutic Neva a. Detail ofachene. Eleocharis Fi. 1. El i. Detail of achene. D £ (Lindl. & Nees) Steud. h. Detail Fertile floral scale. e. Detail of achene. El 1122 Journal of the Botanical R h Institute of Texas 1(2) Fic. 2. Habitat of Eleocharis kleinii B PI h taken in Santa Catarina state, Matos Costa, SC 302, km 31, 26927'41" S 51°07'32,1" W in 21 Dec in its obtusely trigonous culms, narrowly acuminate leaf sheaths extended into a soft awn, floral scales me- dially greenish to stramineous and laterally dark reddish, finely sculptured to nearly smooth achenes, and less coarse perianth bristles. Eleocharis kleinii also appears to be closely related to E. obtusetrigona (Lindl. & Nees) Steud. (Fig. 1 h-i), differing in its shorter and narrower culms, narrowly acuminate leaf sheath apex, fewer flowered spikelets, longer, less imbricate and conspicuously laterally reddish floral scales, relatively longer and less coarse perianth bristles, and achenes that are finely sculptured to nearly smooth with more numerous longitudinal rows of epidermal cells. The occurrence in E. kleinii of floral scales with medially greenish to stramineous and laterally reddish coloration is unique among currently known New World spe- cies of subg. Limnochloa. The following key will help distinguish between these three taxa. KEY TO SEPARATE E. KLEINII, E. LIESNERI, AND E. OBTUSETRIGONA je Aen ME Scales ROT UE mus dan 12- pos oca pao Ao over 2-times wn with 2 f slightly transversely elongated polygonal cells. ulms 1.3-2 mm wide, obtusely trigonous distally; upper into a e awn; PER bristles relatively fine; achene surface finely sculptured to nearly smooth; floral scales greenish to stramineous, laterally dark reddis E. kleinii Culms 2-4 mm wide, terete (obscurely 3-angled); upper sheath apex acute, not extended into a soft awn; perianth bristles pas coarse; achene surface coarsely Sce floral scales medially greenish to stramineous with a su al dark brownish splotch, laterally stramineous E. liesneri 1. Exposed portion of floral pe about as long as wide; spikelet 40-100-flowered; perianth bristles less n | “ A dan IS =Z Trevisan et al., Rediscovery of Eleocharis kleinii 1123 than 2-times achene length; mature achenes dark green splotched with amber or rarely entirely amber with 11-18 longitudinal rows of transversely reniform (at least the most centrally located) polygonal cells E. obtusetrigona Faria (1998) cited Eleocharis laxiflora (Thwaites) H. Pfeiff. (a currently accepted synonym of E. ochrostachys) from Sáo Paulo state, however, the description and illustrations are close to E. kleinii and perhaps those materials studied belong instead to this species. The superficial resemblance of E. kleinii to the Old World E. ochrostachys is due to the occurrence in both species of spikelets wider than the slender culms and is imbricate floral scales. Eleocharis kleinii differs markedly from E. ochrostachys in the nature of the constri between the stylopodium and achene. In E. kleinii, the stylopodium is well diferentiated from the achene apex, which is markedly constricted to a short neck. In E. ochrostachys, the stylopodium arises from the central sunken region of an annular thickening at the achene apex. Eleocharis ochrostachys also differs from E. kleinii in having usually distally terete to weakly-angled culms, acute leaf sheaths, and red-maculate floral scales with wider marginally and apically translucent hyaline-erose regions. ACKNOWLEDGMENTS The authors are grateful to curators of the herbaria indicated for loan of specimens. The first author thanks the Coordenacáo de Aperfeicoamento de Pessoal de Nível Superior (Capes) for scholarship support. Special thanks to Neva Mikulicz for preparing the illustration and to Drs. Rosa Guaglianone and Socorro González- Elizondo for their help and suggestions. REFERENCES Barros, M. 1947. Cyperaceae. In: H. Descole, ed. Genera et Species Plantarum Argentinarum 4(1):1—243. Barros, M. 1960. Las Ciperáceas del Estado de Santa Catalina. Sellowia 12:181—448. Barros, M. 1966. Uma nova Ciperácea em Santa Catarina. Sellowia 18:49. BLAKE, S.T. 1939. A monograph of the genus Eleocharis in Australia and New Zealand. Proc. Roy. Soc. Queensland 50:88-132. BoLDRINI, I.I. 1997. Campos do Rio Grande do Sul: Caracterização Fisionómica e Problemática Ocupacional. Bol. Inst. Biociências: Ecologia 56:1-33. Faria, A.D. 1998. O género Eleocharis R. Br. (Cyperaceae) no Estado de Sao Paulo. Dissertacáo de Mestrado. Uni- versidade Estadual de Campinas, Sáo Paulo. Git, A.S.B and C.P. Bove. 2007. Eleocharis R. Br. (Cyperaceae) no Estado do Rio de Janeiro, Brasil. Biota Neotrop. 7(1): http://www.biotaneotropica.org.br/v7n1/pt/abstract? Taxonomicreview+bn0050701 2007 ISSN 1676-0603. GONZALEZ-ELIZONDO, M.S. and A.A. Reznicek. 1998. Eleocharis R. Br. In: PE. Berry, B.K. Holst, and K. Yatskievych, eds. Flora of the Venezuelan Guayana, Vol. 4. Caesalpiniaceae-Ericaceae. Missouri Botanical Garden Press, St. Louis. Pp. 548-561. GONZALEZ-ELIZONDO, M.S. 1994. Eleocharis R. Br. (Cyperaceae). In: G. Davidse, M. Souza, and A.O. Chater, eds. Flora Mesoamericana, Vol. 6. Alismataceae a Cyperaceae. Universidad Nacional Autónoma de Mexico, Mexico, D.F; Missouri Botanical Garden, St. Louis; The Natural History Museum, London. Pp. 458-464. GONZALEZ-ELIZONDO, M.S. and P.M. Peterson. 1997. A classification of and key to the supraspecific taxa in Eleocharis (Cyperaceae). Taxon 46:433-449. GONZALEZ-ELIZONDO, M.S. and J.A. Tena-FLORES. 2000. Eleocharis (Cyperaceae) in the New World. In: K.L. Wilson and D.A. Morrison, eds. Monocots: Systematics and Evolution. A Publishing, Melbourne. Pp. 637-643 Moreno, J.A. 1961. Clima do Rio Grande do Sul. Boletim Geográfico do RS, Secretaria da Agricultura, Porto Alegre, Ano 6. v.11, 134p. Rosen, D.J. 2006. New and disjunct records of Eleocharis liesneri (Cyperaceae) from South America. Sida 22: 781-786 SMITH, S.G., J.J. BRUHL, M.S.GONZALEZ-ELIZONDO, and F.J. MENAPACE. 2002. Eleocharis R. Br. In: Flora of North America Editorial Committee, eds. Flora of North America North of Mexico, Vol. 23. Oxford Univ. Press, New York. Pp. 60-120. — 1124 Journal of the Botanical R h Institute of Texas 1(2) Svenson, H.K. 1929. Monographic studies in the genus Eleocharis |. Rhodora 31:121-135, 152-163, 167-191, 199-219, 224-242. Svenson, H.K. 1932. Monographic studies in the genus Eleocharis Il. Rhodora 34:193 -203, 215-227. Svenson, H.K. 1934. Monographic studies in the genus Eleocharis Il. Rhodora 36:377-389. Svenson, H.K. 1937. Monographic studies in the genus Eleocharis IV. Rhodora 39:210-231, 236-273. Svenson, H.K. 1939. Monographic studies in the genus Eleocharis V. Rhodora 41:1-19, 43-77, 90-110. Trevisan, R. 2005. O género Eleocharis R. Br. (Cyperaceae) no Rio Grande do Sul, Brasil. Dissertação de Mestrado: Universidade Federal do Rio Grande do Sul. Trevisan, R. and l.l. BoLbrin1. 2005. Eleocharis oct hys Steud. (Cyperaceae), nova ocorrénia para os Estados do Rio Grande do Sul e Santa Catarina, Brasil. Acta Bot. Brasilica 19:695-699. UICN. 2001. Categorías y Criterios de la Lista Roja de la UICN: Versión 3.1. Comisión de Supervivencia de Especies de la UICN. UICN, Gland, Suiza y Cambridge, Reino Unido. DISTRIBUTION OF GAMOCHAFETA (ASTERACEAE: GNAPHALIEAE) IN TEXAS, OKLAHOMA, ARKANSAS, AND LOUISIANA Guy L. Nesom Botanical Research Institute of Texas 509 Pecan Street Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT Dietribiti f a f ] lat ty level in Texas, Oklahoma, Arkansas, and Louisiana: G. be G. argyrinea, G. E G. chionesthes, G. coarctata, G ana and G. purpurea. pre-1900 collect . Gamochaeta coarctata and G. chionesthes are known only from collections made after 1967; the T apparently is spreading rapidly but has not reached Texas and Oklahoma. RESUMEN c (PM PENNE icc] ] ld dad Texas, Oklahoma, Arkansas, y Louisiana: G. antillana, G. inca G. calviceps, G. PORT G. coarctata, G. db rn. y G. m Ganocaci da aia es us única e conocida B colecciones anteriores a 1900. Gamochaeta coarctata y G. ch 67; la pero no ha alcanzado Texas y Oklahoma. Species of Gamochaeta Wedd. (cudweeds) are common colonizers of roadsides and other disturbed sites in the eastern United States, especially in the Southeast. While several of the species may be native to North America, others probably are native to South America and Central America and recently adventive in our area, rapidly increasing their range (Nesom 2004a, 2004b). In fact, the ubiquity of most of them, within the bounds of their range, is remarkable. The present study details geographic distributions in Texas, Oklahoma, Arkansas, and Louisiana, where most cudweeds are at the eastern or southeastern extremity of their North American range. Symbols in the maps for Texas, Oklahoma, and Arkansas indicate the date of earliest collection in a county and suggest when the species may have arrived in a local region. Three time classes are recognized: pre-1941, 1941-1950, and post-1950. Collections have been studied from BRIT-SMU, LSU, NLU, OKL, TENN, TEX-LL, UARK, and VDB. Descriptions of Gamochaeta species and a key are provided in the recent FNA overview of the genus (Nesom 2006). County distributions in the following paragraphs refer to Texas. Gamochaeta antillana (Urb.) Anderb. (Fig. 1). 1903-1940 Denton, Galveston, Hidalgo, Travis counties. 1941-1950 Burnet, Hardin, Kenedy, Leon, Palo Pinto, Robertson, San Patricio, Smith counties. Gamochaeta argyrinea Nesom (Fig. 2). The only known collections from Kansas and Missouri are ap e in Fig. 2 (see Nesom 2004a). The apparent sharp distributional boundary suggests that a corres sharp ecological limiting factor exists. 1905-1940 Austin, Bastrop, Chambers, Grayson, Hardin, Henderson, Jefferson, Liberty, Polk, Trinity, Walker counties. Older collections were made from near the coast, suggesting that the species may have been spreading northward. 1941-1950 Brazos, Camp, Cass, Delta, Fayette, Harris, Hopkins, Jasper, Montgomery, Newton, Orange, Panola, Robertson, Sabine, Shelby, Smith Titus, Van Zandt, Wood counties. Gamochaeta calviceps (Fern.) Cabrera (Fig. 3). 1931-1940 Houston, Van Zandt, Walker counties. 1941-1950 Fannin Co. J. Bot. Res. Inst. Texas 1(2): 1125 — 1130. 2007 nee Journa l of the Bot IR L jm is — Ree TJ NIEHS L| E E B LI CURS H ices oe DM wh Tie ERES ] — TN AHE ay 40 aa aa TSS EEN es ee Se a H us Y e "TTA RA cbe EC Ie H a e @ 1941-1950 Fic. 1. Distributi fG haet. tillana in T: Oklahoma, Arkansas, an d Louisiana | [Lr SIS E ARTO "E eeu ROI puits REN xi fede Te EHET » PEER Me? e Ar PE ae d) LU "n ay LA 7 an LE EN F a TH pum E- en e us ae = "ed s RA wank M EN E zt Cg e jeje o e RE Susita E c | 4 | Y] e AN | | sae EARS MR CUN OPES " 5% SY. y " Gamochaeta argyrinea CHTSS ^ Q 1905-1940 W @ 1941-1950 3l e 1951-present MILES o 1 26 a | A q 95° Fic. 2. Distributi fG haet. j in T Oklahoma, Arkansas, and Louisiana. TI ly | SF E are mapped. Nesom, Distribution of G haeta in Texas, Oklahoma, Arkansas, and Louisiana 1127 p ma E ECRIRE E [| NEAR BSES: a H mE VEURE NE qr H m i ALT "rp LPS Ee ce Suns fae ius A D l R EE E r DUI SA P = Nu ee | E CN EN x (P E d d A calviceps > 1931-1940 1-1950 e 1951—present MILES (0) 100 200 LLLLL LL LL LL Ln Ln 105: 959 d | Fic. 3. Distributi fG h lvii ps in T , Oklahoma, Arkansas, and Louisiana Gamochaeta chionesthes Nesom (Fig. 4). This species is known by relatively few collections from north-central Louisiana and south-central Arkansas the earliest made in 1968. Of all known collections from over its North American range, the earliest was made in Florida in 1957 (Nesom 2004a for citations). It appears to be rapidly expanding in range In the original description, Gamochaeta chionesthes was known in Arkansas and Louisiana only from three collections. Others are cited here. ARKANSAS. Calhoun Co.: clearcut pine woods beside Ark. 160 ca. 2 mi from the Bradley Co. line and 5 mi SE of Harrell, 19 May 1993, Thomas 133,914 (NLU). Cleveland Co.: Greenwood ND E of US 79 and SW of Ark 35 at Rison, 19 May1993, Thomas 133,983 (UARK). Grant Co.: Brush Creek Cemetery beside Ark. 222 W of Ark. 229 N of Leola and 1.9 mi E of Hot Spring Co. line, 12 May 1996, Thomas 149,010 (NLU). Hot Spring Co.: Malvern P.O., Magnet Cove, rocky ridges, 600 ft, 24 Apr 1973, Demaree 66292 (MO). Perry Co.: Hwy 247 on north Cove Mountain, 1.5 mi N ae ae iui 155, 13 air mi SE of Russellville, [mixed with G. purpurea], 30 Apr 2003, Nunn 7680 (UARK). LOUISIANA. B ill woods W of La. 154 at Bonnie and Clyde Historical Marker on W side of La. 154, 3 mi S of La. 793 and Pleasant Hill Church SW ot Mt. Lebanon, 17 Jun 1987, Thomas 100,399 (NLU). Claiborne Par.: roadbank of La. 520 in Kisatchie Natl. Forest, 7 mi from Homer, 13 Apr 1975, Thomas 43,230 (NLU); Hurricane Church in town of Hurricane, on La. 159 SE of Athens, 21 Apr 1977, Lewis 345 (NLU). Grant Par.: Camp Beauregard, near de j icc ado: and Tank Trail, 2.1 km NW of S end of Twin Lake, 7 km W of Flagon Bayou, upland longleaf pine forest, 8 Apr 2002, Douglas Hastings 8 (LSU); Camp Beauregard, N of Mayhaw Road, upland longleaf pine forest, 22 Apr 1993, McInnis 4240 (LSU). LaSalle Par.: Tullos, 1/2 mi S of jct US 165 and US 84, under telephone line, 16 May 1968, Haan, Fife, et al. 139 (TENN). Lincoln Par.: Woodlawn Park in eastern Ruston, roadside of La. Hwy 146, 12 May 1985, David 128 (LSU); [Ruston], Illinois Central Railway right of way, ni soil, 3 May 1984, Wise 39 (DOV). Natchitoches Par.: W of Natchitoches near jct of Interstate Hwy 49 and La Hwy 6, hard-packed sandy soil in front of gas station complex, past flower and fruit, 5 Jul 2004, Nesom GA04--63 (BRIT); gas line along paved road S of Goldonna, 22 Apr 1977, horas 51,639 (NLU). Tangipahoa Par.: beside La. 10 between Wilmer and Washington Par. line, 9 May 1970, Thomas 18,589 (NLU); roadbank of Interstate 55, 2 mi S of Fluker exit (La. 10), 29 Apr 1979, Thomas 64,021 (NLU). Union Par.: hills 3 mi NW of Spencer, hills, 10 May 1969, Thomas 13, 471 (NLU, on sheet with G. purpurea and G. argyrinea). 1128 Journal of the Botanical R h Institute of Texas 1(2) Co E E F m "Hs y £s a sor x E LUCES | TEE dr ECC E EET Ke mer it ERES m NE RANA OH OER Sai D Y E Ns Boers E ESA Gamochaeta chionesthes Gamochaeta coarctata (Willd.) Kerg. (Fig. 5). The earliest Texas collection of this species was made in 1969. The earliest in Louisiana were made in 1970. The earliest known North American collections were made in 1949 and 1950 in the vicinity of Wilmington, North Carolina (see Nesom 2004b). Gamochaeta pensylvanica (Willd.) Cabrera (Fig. 6). 1903-1940 Bastrop, Bell, Burleson, Cameron, Galveston, Goliad, Gonzales, Harris, Kleberg, Llano, Nacog- doches, Travis, Waller, Washington counties. 1941-1950 Aransas, Brazos, Denton, Erath, Hardin, Hidalgo, Jack, Jim Wells, Lee, Leon, Robertson, Somervell, Tarrant counties. Eight collections of Gamochaeta pensylvanica from Louisiana (7 parishes) were made prior to 1922. One collection was made before 1900 (New Orleans, Coliseum Street, 31 Mar 1871, Dr. Joor s.n. LSU). Gamochaeta purpurea (L.) Cabrera (Fig. 7). Gamochaeta purpurea has the widest distribution of the cudweed species occurring in North America, and itis known from the earliest collections. It is the only one of the Texas species collected prior to 1900. 1843 Harris Co. (probably), collected by Lindheimer. 1890 [ca.] Dallas Co., collected by Reverchon. 1905-1940 Burnet, Calhoun, Erath, Jefferson, Llano, Washington, Wilson counties. 1941-1950 Bastrop, Brazos, Cass, Denton, Franklin, Grayson, Hardin, Harrison, Jefferson, Red River, Smith, Tarrant, Titus, Van Zandt counties. A “race” of Gamochaeta purpurea apparently occurs in east-central Louisiana: Caldwell, Franklin, Pointe Coupee, St. Helena, and West Feliciana parishes (LSU). The plants have densely leafy stems, densely brac- Nesom, Distributi fG haeta in T Oklahoma, Arkansas, an d Louisiana HERE oP Eom mu gra HH (UE LER RO = E Brn Sone e RARUS, S Pe o ANS EERE ERN K ERE RA V S Ni ochaeta coarc ata l oH $ 7 Ey i E A " | " 1129 Fic. 5. Distributi fG h in T Arkansas, and Louisiana. All collecti le after 1967. CHE wee Seer Seige) Ee ideo WI |] H V Gamochaeta pensylvanica @ 1903-1940 © 1871 O 1941-1950 [e Y) e 1951-present MILES nn o 100 200 3 le: Losa LL b c Lu 105° 100° QV 95? 90° 1130 Dat e ID hi - 3 £f T, J Infl JOUTTIal OF I — ae TERR H LATTA ie m ln [| C] UE Hr J ie oH d E OA E ERA TRY DU eed | INASRE En 3 o ALETA ume tees e A UNE As A O 1343-1890 ae aw. TRANS Gamochaeta purpurea ^ ~ v @ 1905-1940 @ 1941-1950 e 1951-present MILES o 100 200 3 LLLA LLL LLL a 105° 95° 90° | Texas 1 (2) ^ Y Fic. 7. Distributi in T Oklahoma, Arkansas, and Louisiana teate capitulescences, abaxial leaf surfaces hirsute-pilose with hairs with prominently (atypically) swollen bases, and a tendency toward perenniality. ACKNOWLEDGMENTS Iam grateful for loans of specimens from LSU, OKL, and UARK, comments on the manuscript by an anony- mous reviewer, and help from staff at TENN, TEX-LL, and NLU while studying in those herbaria. REFERENCES Nesom, G.L. 2004a. New species of Gamochaeta (Asteraceae: Gnaphalieae) from the eastern United States and comments on similar species. Sida 21:717-742. Nesom, G.L. 2004b. New distribution records for Gamochaeta (Asteraceae: Gnaphalieae) in the United States. Sida 21:1175-1186. Nesom, G.L. 2006. Gamochaeta (Gnaphalieae). In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. Vol. 19:431—438. Oxford University Press, New York and Oxford. FLOW DIAGRAMS FOR PLANT SUCCESSION IN THE MIDDLE TENNESSEE CEDAR GLADES J.M. Baskin El illa peed mont P. ark University of Kentucl Nashville pM 3721 E Qm Lexinaton, Kentucl 40506, USA, C.C. Baskin Department of Biology UI versity of f E Xtucky Lexington, Kentucky 40506, U.S.A. Department of Plant and Soil Sciences Ul iversity of! El itucky Lexington, Kentucky 40546, U.S.A. ABSTRACT Elsie Quarterman’s comprehensive study of cedar glade vegetation in the Inner Central (Nashville) Basin of Tennessee did not include flow diagrams for any of the multiple pathways of succession she described in her Ph.D. thesis sa ru or in her major publication on cedar glades (Quarterman 1950a) Thus, our pr imary objecti ve t Į diagrams) for the various (inferred) successional pathways from bare limestone bedrock to subclimax redcedar, pr hardwood forests, based primarily on Quarterman’s study. A second objective was to discuss how other q isi a in the Inner Basin fit into dos S schemes. qc E along with how other ipn ue with them, Tennessee cedar limax oak-hickorv and mixed titati tudi n ceda ar E E glade vegetation complex and thus be an aid t ion pl ing in the Nashville Ban ABSTRACT El dio exl ivo de Elsi t de la vegetación de los “cedar od en e cuenca interior central (Nashville) de Tennessee no luyó di de flujo de ni lel áltiples vías d Quarterman Pi ensu mc f het; DJELU más a sobre ise ‘cedar ladles (uu TM 19502). Así pues ius de due de las varias (inferidas) vías de sucesión desde la caliza desnuda a P subclímax de cedro je preclimax roble- nogal y ł de madera dura, basados principalmente en el estudio de Quarterman. Un segundo obj r como otros dee cuantitativos sobre MM de los “cedar glade” de la cuenca interior ee con los Se " Quarterman. las diferentes Estos ee. oes con | tudios, harán más fácil l omunidad d tación) ] vegetación de "cedar EE de Tennessee central y de este modo ser una ayuda en la planifi ación de | rvació la cuenca ee Nashville INTRODUCTION The Inner Central (Nashville) Basin is well known for its cedar glade vegetation (e.g., Harper 1926; Free- man 1933; Quarterman 1950 a, b; Kuchler 1964; Baskin & Baskin 2004). Historically, in the Central Basin the term “cedar glades” (sensu lato) has been used to include both the natural rocky limestone openings Cglades") and the adjacent redcedar-redcedar/hardwood forest complex, and Galloway (1919) even recog- nized “massive rock or hardwood glades” (Baskin & Baskin 2004). More recently, however, botanists have used “cedar glades” (sensu stricto) in reference to the rocky openings only, i.e., “glades” or “open glades” sensu Quarterman (Baskin & Baskin 2004). Thus, cedar or limestone glades in the Inner Basin are open areas of rock pavement, gravel, flagstone and/or shallow soil in which occur natural, long-persisting (edaphic climax) plant communities dominated by herbaceous angiosperms and/or cryptogams (Baskin & Baskin 1985; Quarterman et al. 1993; Ware 2002). They may, or may not, be surrounded by forest (Galloway 1919). me established in deep soil-filled cracks Cedar glades may support low densities of woody plants, which b in the bedrock [e.g., Picklesimer (1927; see Fig. 1 in Baskin @ Baskin 1996a); Quarterman 1950a]. The J. Bot. Res. Inst. Texas 1(2): 1131 — 1140. 2007 1132 Journal of the Botanical R h Institute of Texas 1(2) dominant plants are C, summer annual grasses (primarily Sporobolus vaginiflorus but also Panicum capillare and P. flexile), C; winter annual, summer annual and/or perennial herbaceous dicots; mosses (primarily Pleurochaete squarrosa); the cyanobacterium Nostoc commune; and crustose, foliose and fruticose lichens [Picklesimer (1927; see Baskin & Baskin 1996a); Freeman 1933; Quarterman 1950a,b; Somers et al. 1986; Mahr & Mathis 1981; Dubois 1993; Baskin & Baskin 1996b; Rollins 1997]. The most complete, comprehensive study of cedar glade vegetation in the Central Basin (as well as in the southeastern USA in general) was done by Elsie Quarterman (1948, 19502, b). She described (apparent) successional pathways that theoretically represent the various stages in vegetation development from bare rock to oak-hickory forest or to a subclimax redcedar (Juniperus virginiana) forest. Quarterman determined constancy of plant species in 22 open glades and in 10 glade woods in the Inner Basin (sensu Edwards et al. 1974; also see DeSelm 1959), and she used quadrats to sample intensively the following stages in the (apparent) ional sequence of plant communities: gravel glade, grass glade, glade-shrub, shrub-cedar, 88-year-old cedar forest, 103-year-old cedar forest and cedar-hardwood forest. She referred to gravel and grass glades, along with rock glades, which she did not quantitatively sample, as open glades. These open glades are the *cedar glades" sensu stricto defined above. Although Quarterman described succession from bare rock to redcedar and redcedar/hardwood for- ests in considerable detail, she did not conceptualize, via flow diagrams, the various pathways and their relationship to each other. Since Quarterman's (19502) classic publication, two flow diagrams of vegetation development in the cedar glades of the Central Basin have been published (Quarterman et al. 1993; Baskin & Baskin 2004). However, neither of these diagrams shows the complexity of successional pathways that lead from bare rock to redcedar and hardwood forests (Quarterman 19502). Thus, our objectives were to (1) construct detailed diagrams showing vegetation development in the cedar glades (sensu lato) of the Central Basin of Tennessee, based primarily on Quarterman (1948, 19503), and Q) discuss how other quantitative studies on cedar glade vegetation in the Inner Nashville Basin fit into Quarterman's schemes. The initial work on constructing these diagrams began when Jerry and Carol Baskin were graduate students at Van- derbilt University in the 1960s and were beginning to learn about the middle Tennessee cedar glades from Professor Elsie Quarterman. Although the primary sources for the successional pathways and seral stages is Quarterman (1948, 19502), other sources include Picklesimer (1927; see Baskin & Baskin, 19963), Free- man (1933), McKinney € Hemmerly (1984), Crites and Clebsch (1986), Somers et al. (1986), DeSelm (1989, 1992), Baskin and Baskin (1996b), Rollins (1997) and numerous unpublished observations by Jerry and Carol Baskin in the middle Tennessee cedar glades. Nomenclature primarily follows that used by Baskin and Baskin (2003). SUCCESIONAL PATHWAYS In actuality, Quarterman's presumed successional pathways may represent vegetational sequences along xeric to submesic environmental gradients that are unrelated to succession. In any case, whether the vegetation mosaic seral stages or vegetation unrelated to succession, the cedar glades support a complex LE of plant groupings ("associations") that are difficult to comprehend via written descriptions, even when the text is supplemented with photographs and a considerable amount of data on community composition and structure, as is the case with Quarterman's study. Thus, thinking of these various communities in a succes- sional or developmental context helps to clarify the structural relationships among them. Limestone without cracks Primary succession on limestone without cracks is shown in Figure 1. There are two starting points: one in slight depressions (with small amounts of mineral soil and gravel) on bare rock pavement of thick beds primarily of the Middle Ordovician Ridley (Ord) and Lebanon (Olb), but also on Pierce and Murfreesboro (Opm, mapped as a unit) (e.g. Ware 1969; Baskin and Quarterman 1970) limestones, and the other on flag- stone/gravel glades weathered from thin beds of the Lebanon [see photographs (Fig. 12.1) in Baskin & Baskin 1999], on which most of the cedar glades in the Central Basin have developed (Safford 1851; see Baskin & 1133 gau T. IW "d " Y MVA J ñ ywak-cni E E ( 4 \ el I". "ll IL “TF td Fr Fr FU SY CUL pue pjo OOV-UJUL OVULI) = (9 S) 11*(2—1) si9quunu Áq umoys are (20561 “8r6L) VOM rs . I . 23 ~I LV AN J HESY " pol 6G " h D'a ‘sjenuue Ja} UIM “YM .(")) od m PER ad ñ [AT MODUIED CofE] Vo J[eruugadgu J m Ļ Vins LE] Juar Mo E bu i 2) . 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No mats form directly on the rock surface, although Nost d Grimmia apocarpa may spread to bare rock. Mats of herbaceous vegetation formed during succession on rock glades coalesce into gravel glades. Then, further on in the sequence, the two pathways of succession join to form the grass-glade stage, which is followed by the glade-shrub, shrub-redcedar, redcedar forest and redcedar-hardwood forest seral stages, and finally by an oak-hickory preclimax forest. The last stage of mat formation (rock glades) may proceed directly to the glade-shrub stage, when woody plants become established in deeper parts of the mat and/or vertical crevices in the bedrock, thus by-passing the gravel and grass stages. Also shown in Figure 1 is the relationship between the redcedar forest stage of succession and the formation of rock barrens (sensu DeSelm 1992; = xeric limestone prairies sensu Baskin et al. 1994; Baskin & Baskin 2000, 2004; Lawless et al. 2006 [see photographs (Fig. 3) in Baskin € Baskin 2004]. According to DeSelm (1992), “Environmentally and vegetationally these barrens ‘fit’ between cedar glades and cedar or oak forests with deeper soil." Further, *Barrens in the Basin occur on the soil thickness/available moisture gradient between cedar glades and cedar forests. Today, they probably occur chiefly on sites where the cedar has been removed. Maintenance of such openings against cedar and hardwood successional pressure has been accomplished by grazing, fire and today perhaps by bushhogging.” Limestone with vertical cracks The multiple pathways on limestone with vertical cracks, as discussed by Quarterman (19502), are dia- grammed in Figure 2. In cracks that are widely-spaced and parallel to each other, succession leads to an open glade with lines of trees and shrubs. Between the cracks, the successional sequence is similar to that shown in Figure 1, i.e., from the gravel glade stage to the grass glade stage and on to oak-hickory forest. In cracks that are close together and intersecting, the successional sequence is from herbs to shrubs to redcedar. In the redcedar forest, a moss mat forms on small ledges and sides of cracks in the bedrock, which is invaded by herbs, ferns and lichens. On large flat blocks of limestone between cracks in the redcedar forest, succession in partial shade begins with the moss Orthotrichum strangulatum or the cyanobacterium Nostoc commune and then proceeds to the moss Pleurochaete squarrosa, which may be long-persisting, to herbs and finally to shrubs. On flat blocks of limestone in shade, succession begins with Orthotrichum strangulatum and ends with a Climacium americanum (moss)-fern-herb stage. These four successional pathways in areas with cracks that are close together and intersecting result in a redcedar subclimax forest with shrub, moss and fern-herb layers beneath the canopy of Juniperus virginiana trees. The bark of redcedar trees forms a substrate for establishment of corticolous bryophytes, the mats of which may also include species of cyanobacteria, fungi and lichens (Quarterman 1949). A successional sequence begins on the smooth bark of young (single, unshaded) redcedar tree trunks with the mosses Frullania and Orthotrichum, proceeds to a Leucodon-Cryphaea-Chasmatodon stage (woods margin, generally older trees than in first stage) and then to a Leucodon-Porella (leafy liverwort) stage (on older trees within redcedar forests). Porella (only on trees in forest) sometimes forms pure colonies, which may be invaded by the fern Polypodium polypodioides (Quarterman 1949). Thus, in general, the majority of moss mats on young redcedar trees represents early stages of succession and the majority of those on old redcedar trees later stages of succession. However, new surfaces created by the shedding of bark, even on old trees, are invaded by the same pioneers (Fig. 2, in REDCEDAR SUBCLIMAX) that occur on young trees, providing *...evidence that a rather definite scheme of succession occurs on bark of cedar trees" (Quarterman 1949). QUARTERMAN'S SCHEMES IN RELATION TO OTHER STUDIES Picklesimer (1927; see Baskin and Baskin 19962) counted all individuals along a 404 ft (123 m) x 1 ft (0.3 m) transect in a open glade apparently on a thick-bed (see her Fig. 4) of the Lebanon limestone (Inner Basin). The transect was divided into 21 zones: 16 in open glades and five in glade woods. Forty-seven percent of the 42,207 individuals recorded along a transect by Picklesimer in the open glade were plants of Sedum pulchellum, and 30.496 were plants of Arenaria (Minuartia) patula. Sedum was the most abundant plant in eight of 16 open glade zones and Arenaria in five. Panicum capillare, Sporobolus vaginiflorus and Baskin et al., Plant ion in the Middle T dar glad 1135 LIMESTONE WITH VERTICAL CRACKS cracks widely spaced (parallel) cracks close together (Intersect at right angles) | | | In cracks between cracks i k large flat blocks of limestone between lines di of mature trees In REDCEDAR FOREST 1 partial shade ——p shade v v (crown cover somewhat broken) Sporobolus! Dalea, 3 herbs dic. herbs smail ledges and sides E m r—-—p of vertical cracks in D 2 REDCEDAR FOREST Nostoc ; 2 Orthotrichum (in slight ASS STAG! depresslons) y E cracks widen & sprite = i n g g filt with soil gra e a soll B E TortellalGrimmialOrthotrichum SHRUB THICKET $ shrubs --- -p Forestier Rhus! shrubs N Plewrochaste e Symphoricarpos Yo 5 (rows of pa A EN 3 Ulmus, & C ` z become sd JuniperuslUlmus! ! Arena Waada HIE 3 Celtis (with shrubs) Anomodon andother he Thuidium r trees JuniperusiUlmusi Celtis 1 (with shrubs) Fi : RENFENAR CIRC! IMAY t REDCEDAR SUBCLIMAX mature Entodon Thuidium shrubs Climaciumt Juniperus forest ^ — ( ferns/herbs Quercus!Carya cma herbs, & lichen: (preclimax) n moss mats in shade) succession on redcedar trees (bark) MIXED HARDWOOD FOREST B Erullaniali Inh » POE AM: di Y dan/Darall dioid (ctimatic climax) ioii d T Vice iia i ? OAK-HICKORY FOREST (preclimax) Erigeron strigosus [= E. strigosus Muhl. ex Willd. var. calcicola J. Allison (Allison and Stevens 2001; Noyes and Allison, 2005)] ranked third, fourth and fifth, respectively, in order of abundance. Plants of each of the cedar glade endemics Leavenworthia torulosa, Lobelia appendiculata var. gattingeri, Phacelia dubia var. interior and Talinum calcaricum were present in several of the zones. Redcedar (Juniperus virginiana) was by far the dominant tree in the glade woods with Adelia (Forestiera) ligustrina (shrub), Ulmus alata and Celtis missis- sippiensis (2 C. laevigata) second, third and fourth, respectively, in abundance. The vegetation of the open glade Sand by Picklesimer fits primarily in the rock glade category (Fig. 1) and that of the glade woods in the redcedar forest (Juniperus/Ulmus/Celtis/Forestiera) (Fig. 1, 2; see Fig. 1, 2 in Baskin & Baskin 19963). Picklesimer proposed that the successional pathway in cedar glades begins with crustose lichens (pioneers) and ends with a redcedar climax forest. See Baskin and Baskin (19962) for updating of nomenclature used by Picklesimer (1927). Freeman (1933) used 1 ft? (0.09 m?) quadrats to determine the relative abundance of species in five spring and 10 summer aspect societies in an open glade in Davidson County, Tennessee, near LaVergne (In- ner Basin). The 15 aspect societies were (1) Dalea gattingeri (not D. foliosa as indicated by Freeman, i.e., see his figure 18, which is a photograph of D. gattingeri and not D. foliosa)-Sporobolus vaginiflorus; (2) Hypericum sphaerocarpum-Dalea gattingeri; (3) Croton monanthogynus-Cocculus carolinus; (4) Isanthus brachiatus-Hypericum sphaerocarpum; (5) Croton capitatus; (6) Croton monanthogynus; (7) Erigeron strigosus-Lespedeza striata (not native); (8) Diodia teres- Croton capitatus; (9) Hypericum sphaerocarpum; (10) Sedum pulchellum; (11) Arenaria patula; (12) 1136 Journal of the Botanical R h Institute of Texas 1(2) Leavenworthia stylosa; (13) Leavenworthia stylosa-Ophioglossum engelmannii; (14) Delphinium carolinianum [ = D ciphilum Warnock; Warnock (1990, 1997)]; and (15) Isoetes butleri. However, we do not consider the Croton monanthogynus-Cocculus carolinus or the Erigeron strigosus-Lespedeza striata societies = carolinianum Walter subsp. ca to have been on open cedar glades since the soil was up to 61 cm deep (see Baskin € Baskin 2004 Overall, species with the highest % importance values (% IVs) [(relative frequency + relative density)/2] x 100 calculated from Freeman’s data on frequency and density of taxa in the 13 societies on open cedar glades were (in decreasing order): Minuartia patula, Sporobolus vaginiflorus, Sedum pulchellum, Leavenworthia stylosa, Hypericum sphaerocarpum, Croton capitatus, Isoetes butleri, Dalea gattingeri, Croton monthogynus, Isan- thus brachiatus and Panicum capillare. Sporobolus vaginiflorus ranked first in the D. gattingeri-S. vaginiflorus; H. sphaerocarpum-D. gattingeri; I. brachiatus-H. sphaerocarpum; C. monthogynus; D. teres-C. capitatus; and H. sphaerocarpum-S. pulchellum societies and second in the C. capitatus society. This species was not present in the Sedum, Arenaria, L. stylosa or L. stylosa-O. engelmannii societies. Except for D. carolinianum and I. butleri, no data on species composition/abundance are given for these two societies. Seemingly, the seven societies in which S. vaginiflorus is important represent gravel and/or grassy glades, while the S. pulchellum, A. patula, L. stylosa and L. stylosa-O. engelmannii societies belong to rock glades (Fig. 1, 2). In a study of the ecological distribution of Talinum calcaricum on rock (Opm) glades near Murfrees- boro, Tennessee (Inner Basin), Ware (1969) recognized four vegetation zones in the *ecotone" between bare rock and grass-dominated glades (0-10 cm soil depth): Nostoc-Talinum; Sedum-Nostoc-Talinum; Arenaria- Talinum-moss; and Pleurochaete (moss)-Talinum. Ware's first and second zones appear to correspond to the Nostoc/Talinum/Panicum/ wA/ bryophyte and SEI Talinum/Panicum/Nostoc/wA zones, respectively, and his third and fourth zones to the Dalea/Erigeron/S lus/Pleurochaeta/wA zone (Fig. 1). He did not include a Nostoc/lichens zone (Fig. 1) in the ecotone Büscen bare rock and grass-dominated glades. Later on in her career, Quarterman (1973, 1989, 1993) recognized two characteristic zones or com- munities of open cedar glades in the Inner Central Basin: Zone I (soil depth 0-5 cm; = gravel glades of Quarterman et al. 1993); and Zone II (soil depth ca. 5-20 cm; = grassy glades of Quarterman et al. 1993). The shallow portions of Zone I are dominated by the winter annuals Minuartia patula, Leavenworthia spp. and Sedum pulchellum; and in summer by the summer annuals Cyperus squarrosus (= C. aristatus, C. inflexus) and the small succulent polycarpic perennial Talinum calcaricum. Nostoc commune grows on bare rock. In deeper soil of this zone, the dominant plants are the long-lived polycarpic perennial Dalea gattingeri, the biennial Erigeron stigosus and the moss Pleurochaete squarrosa. Zone Il is dominated by the C, summer annual grasses S. vaginiflorus, Panicum capillare and Aristida longespica (occasional) and the moss P. squarrosa; many other species occur in this zone. Somers et al. (1986) quantitatively sampled 10 open glades in the Inner Central Basin and identified four xeric (soil « 5 cm deep) - Panicum capillare; foliose lichens, Nostoc commune-Sporobolus vaginiflorus and Dalea gattingeri; and three subxeric (soil » 5 cm deep) - Sporobolus vaginiflorus, Pleurochaete squarrosa and Panicum flexile-Pleurochaete squarrosa-S. vaginiflorus plant communities. Quarterman's zones I and II, which she equated with rock-gravel glades and grassy glades, respectively (Quarterman et al. 1993), correspond closely to Somers et al 's xeric and subxeric plant communities, respectively (Somers et al. 1986; Quarterman et al. 1989). Using combined plot data from five sites on open cedar glades in the Inner Basin that supported popu- lations of the federally -endangered species Echinacea tennesseensis, Drew (1991; also see Drew and Clebsch 1995) identified six plant communities: Echinacea/Ruellia; Houstonia/ Sporobolus; Sporobolus/Petalostemon (Dalea); Echinacea; Schizachyrium/ Houstonia; and Echinacea/Schizachyrium. These community types differ somewhat from the vegetation zones (community types) recognized by Quarterman (19502) and also from those recognized by Somers et al. (1986) and Rollins (1997). Thus, for example, whereas Echinacea ten- nesseensis and/or Schizachyrium are important components of four of Drew's across-site community types, neither species was present in any of the vegetation zones of open cedar glades recognized by Quarterman (Fig l 2). Baskin et al., Plant ion in the Middle T dar glad 1137 On the other hand, 96 importance values (96 IVs) [(relative cover + relative frequency)/2] x 100 calculated for taxa at each of the five glade sites sampled by Drew showed that Sporobolus vaginiflorus had the highest idi across-site 96 IV. Other important plants of open Lebanon limestone glades in Drew's study included II Hedyot ricans, Ruellia humilis, Grindelia lanceolata, Dalea gattingeri, Hypericum sphaerocarpum and Isanthus Tom (Fig. 1, 2). Further, as pointed out by Baskin and Baskin (2004), Drew misinterpreted Quarterman's and Hal DeSelm's (see Quarterman 1989; Quarterman et al. 1993; DeSelm 1992) definition of glades («5096 perennial grass cover) and barrens (2 5096 perennial grass cover), and as a result he assigned two of his six community types to barrens, based on 1996 and 1596 cover of the (only) perennial grass, Schizachyrium scoparium, in the two community types. However, using the criteria of Quarterman and DeSelm for glades vs. barrens, all five open glade sites quantitatively sampled by Drew (1991; Drew and Clebsch 1995) are subxeric open cedar glades. Baskin and Baskin (1996b) described a Grindelia lanceolata plant community type on the (mostly) thin-bedded Lebanon limestone about 7.0 km north of LaVergne (Rutherford County) in the Inner Basin. Grindelia lanceolata (10096 frequency), Sporobolus vaginiflorus (96.596) and Dalea gattingeri (91.896) had the highest frequencies of the 51 native and 3 nonnative species present in 85 1-m? sample plots. This commu- nity and NatureServe's (2005) Sporobolus (neglectus, vaginiflorus)-Aristida longespica-Panicum flexile-Panicum capillare Herbaceous Vegetation Association of the Sporobolus (neglectus, vaginiflorus) Herbaceous Alliance fit the subxeric portion of the open glades category described by Quarterman (Fig. 1, 2). Rollins (1997) sampled 10 open glades in the Inner Central Basin and delineated 10 community types at the 10-m? scale and 14 at the 0.1-m? scale. She assigned names to them based on the two to six and two to four most important species, respectively, in order of decreasing cover and constancy. Species included in names of 10 community types identified, with the total number of times that species is used (base number) and number of times it ranked 1, 2, 3, 4, 5 and/or 6 (10-m? scale) or 1, 2, 3 and/or 4 (0.1-m? scale) (exponents of base number) are as follows: for communities identified at the 10-m? scale — Sporobolus ) vaginiflorus, 10 ©®, Dalea gattingeri 5 © ^ * D, Pleurochaeta squarrosa 6 4 ?, Andropogon virginicus 4 © > l Nothoscordum bivalve 2 9.9. 2. Grindelia lanceolata var. lanceolata 2 9 99. 2. Hedyotis nigricans var. nigricans 2 (0,0,0,1, D. Eleocharis compressa 1 ©), Carex crawei 10,0, D, Aristida longespica 1 © è V, Pediomelum subacaule 0, 0, 0, 0, 1 €69,90. Ruellia humilis 1 9.9.9.9. D and Senecio anonymous 1 € D: and for communities identified at the 0.1-m? scale - Sporobolus vaginiflorus, 10 99:2, Dalea gattingeri 4 2D, Grindelia lanceolata var. lanceolata 3 (1,1,0,D, Nothoscordum bivalve 3 œ * D, Nostoc commune 3 0,2, Ð, Ruellia humilis 2 ©% D, Hedyotis nigricans var. nigricans 2 ®, Sedum pulchellum 1 ®, Carex crawei 1 ®, Ratibida pinnata 1 ®, Eleocharis compressa 1 ®, Andropogon virginicus 1”, Pediomelum subacaule 1 © ), Aristida longespica 1 © Y, Manfreda virginica 1 © P, Minuartia patula 10% Y and Calmintha glabella 1 ©% D. Rollins used Pleurochaete squarrosa as a substrate type in sampling at the 0.1-m? scale, and thus did not include its cover or constancy in her samples, which explains why there are no data for this moss in her community types identified at the 0.1-m? scale. Rollins community types differ somewhat from those recognized by Quarterman. Thus, whereas Andropogon virginicus and wet-loving plants such as Carex, Eleocharis and Nothoscordum (see above) were important components of the open glades sampled by Rollins, they were not important in those sampled by Quarterman (Fig. 1, 2). However, Sporobolus vaginiflorus, the overall dominant in open cedar glades of the Central Basin (see Baskin and Baskin 1999), also was the most important species in Rollins’ study. Rollins states that her study agrees with those of Quarterman in dividing open cedar glades into xeric (gravel) and 1 J subxeric (grassy) zones with “... the only exception being low of Pleurochaete and foliose lichens in the ‘xeric’ zone communities described here.” The overall vegetation of the five forest stands sampled by McKinney and Hemmerly (1984) on Cart- ers and Ridley limestones in the Inner Basin was a mixed hardwood (oak-hickory-maple-ash-elm) forest. However, four of the five stands sampled by these authors were on the thick-bedded Carters Limestone, which overlies the Lebanon and generally is not a glade-forming limestone. Their other stand, which was on the thick-bedded Ridley, a glade-forming limestone, was dominated by Quercus-Carya (%IV, 67.2). Thus, 1138 Journal of the Botanical R h Institute of Texas 1(2) it is an example of the oak-hickory stage shown in Figures 1 and 2. In the seemingly most mesic stand (number 3) sampled by McKinney and Hemmerly (1984), Acer saccharum had the highest % IV (37.9) fol- lowed by Quercus spp. (32.4), Carya spp. Q3.2) and Fraxinus spp. (17.3), a mixed hardwood forest (Fig. 1,2). The overall upland forest vegetation on Ridley (redcedar-sugar maple-hickory-oak-ash-elm) and Lebanon (redcedar-oak-hickory-ash-elm) limestones in the Inner Basin sampled by Crites and Clebsch (1986) are examples of redcedar/hardwood forests in Figure 1. Seemingly, the most mesic stand sampled by these au- thors was a Acer saccharum/Fraxinus americana/Ostrya virginiana plant community type on Ridley Limestone, another example of mixed hardwood forest in the Inner Basin. NatureServe's (2005) Juniperus virginiana var. virginiana-Forestiera ligustrina-Rhus aromatica-Hypericum frondosum Shrubland Association of the Juniperus virginiana-Rhus aromatica Alliance fits well with the glade-shrub and redcedar stages, and its Juniperus vir- giniana var. virginiana-Fraxinus quadrangulata/Polymnia canadensis (Astranthium integrifolium) Woodland of the Fraxinus quadrangulata —(Juniperus virginiana) Woodland Alliance more or less with the redcedar and redcedar/hardwood forests (Fig. 1, 2) Finally, NatureServe's (2005) Quercus muehlenbergii-Juniperus virginiana Wooded Herbaceous Association of the (Juniperus virginiana/Schizachyrium scoparium-Bouteloua curtipendula) Wooded Herbaceous Alliance, the little bluestem-dominated glades described by Baskin and Baskin (1977; see Fig. 1 in this paper) and the four barrens on Ridley and Lebanon limestones described by DeSelm (1992) fit what we refer to in Figure l as xeric limestone prairies, an anthropogenically-derived plant community. Not evident from Figure 1 is the fact that successional stages on open glades may be wet enough in spring to support such moisture-loving plants as Isoetes butleri, Carex species (especially C. crawei), Eleocharis bifida S. G. Smith [segregated from E. compressa (Smith, 2001, 2002)], Nothoscordum bivalve and Schoenolirion croceum. However, Quarterman (1950b) did mention a group of species that included Isoetes butleri, Noth- oscordum bivalve, Schoenolieron croceum and Leavenworthia spp, among others, that *...flourishes and blooms in early spring when the ground is saturated." Freeman (1933) recognized an Isoetes butleri aspect on epe cedar glades, and he spun that, *The area occupied by Isoetes was very wet during the early spring." ha JUNTO q Carex crawei, Ele bivalve were used 1 Rollins (1997) in naming three of the 10 open cedar glade NS types she recognized at the 100-m? scale and of four of those at the 0.1-m? scale. This winter/spring-wet stage seems to fit well with the Eleocharis compressa-Schoenolierion croceum- Carex crawei-Allium cernuum Herbaceous Mi idu Association of the Eleocharis compressa-Nothoscordium bivalve Saturated Herbaceous Alliance of NatureServe's (2005) International Vegetation Classification System. NatureServe (2005) also recognizes a Dalea foliosa-Mecardonia-Mitreola petiolata Herbaceous Association in thin soil along streams and a Sedum pulchellum-Talinum calcaricum-Leavenworthia spp./Nostoc commune (in depressions on limestone that hold water in winter and early spring) Association as wet community types on open glades in the Central Basin. CONCLUDING REMARKS We believe that the successional diagrams (Fig. 1, 2) will make it easier to comprehend the multiple starting points and pathways of succession in the middle Tennessee cedar glade vegetation complex, including how the various plant “associations” are related to each other. Further, these diagrams complement Quarterman's (1948, 19502) detailed description of the apparent successional pathways in cedar glades (sensu lato) of the Central Basin of Tennessee, and they serve as a framework for other quantitative studies that have been (or will be) done on cedar glade vegetation in this physiographic region. Finally, they should be an aid to conservation planning in middle Tennessee, where human population growth and urbanization/industri- alization is rapidly destroying much of the cedar glade habitat. ACKNOWLEDGMENTS The authors wish to thank Edward W. Chester and an anonymous reviewer for helpful suggestions to im- prove the manuscript. Baskin et al., Plant ion in the Middle T dar glad 1139 gau REFERENCES ALLISON, J.R. and T.E. Stevens. 2001. Vascular flora of Ketona dolomite outcrops in Bibb County, Alabama. Castanea 66:154-205. Baskin, J.M. and C.C. Baskin. 1985. Life cycle ecology of annual plant species of cedar glades of southeastern United Sates. 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In: Flora of North America Editorial Committee. Flora of North America (north of Mexico). Volume 3. Magnoliophyta: Magnoliidae and Hamamelidae. Oxford University, New York. Pp. 196-240. OBSERVATION ON VEGETATION CHANGES IN CAJUN PRAIRIE, A COASTAL PRAIRIE FLORA IN SOUTHWEST LOUISIANA Charles Allen and Sara Thames Colorado State le Fort Polk Station 23H St. Fort Polk, eae 459, USA. native@camtel.net ABSTRACT ae Prairie once ica 2:5 Me acres in southwest Louisiana bu has been reduced D E Lane (plowing) to less +1 ] f way. Based on ob d of regrowth in pone inremnants, the pus ofvegetation es is iia The early successional dominants in the Cajun Prairie habitat e mainly annual I location, and history of agricultural use. Most annuals disap- pear in less than two years but goatw ET (Crot Y itatus) of i f three to four years. The second id of succession includes ids rud T perennial o native non- ake introduced, and Cajun Prairie. With time, the Cajun P abundan g two groups. The introduced Brazilian vervain CHA brasiliensis) and vasey g Paspal villei) are very i lers in Cajun Prairie but disappear with time. Typically, the first native species to disappear is late fl ing tl gl Euf i tinum) and the last to be lost is common goldenrod (Solidago altissima). RESUMEN La pradera Caj l ez cubrió 2.5 mill d ] te de Louisi E I lucido por las practi gricol turación) a menos Ea 100 acres en bandas pequeñas, aisladas a lo largo del f ilalad ha del ino. Basad b enl ión d | ] j lizado el patrón d bio de la vegetación. Las dominantes en el inicio de la sucesión en la pradera MUI son UNE t l variación en la cod específica debido a la humedad, localización, e hist I 1 roton capitatu que frecuentemente persiste durante tres o cuatro años. El segundo estadio de la sucesión A tres EDO de ai a nativas no de pradera, introducidas, y de la pradera Cajun. Con el tiempo, | la disminución correspondiente de la abundancia de m otros dos ud La m de Brasil (Verbena brasil a 7 d urvillei son invasoras id comunes en la pradera Cajun p Típicamente, | NO D de E E nM. L is INTRODUCTION Cajun Prairie developed in southwestern Louisiana between the Atchafalaya and Sabine rivers where most of the European Settlers were Cajuns (Allen et al. 2001). The flora is best described as a coastal plain flora overlaid with a major midwestern prairie component. This prairie once occupied 2.5 million acres but most has been destroyed. The remaining Cajun Prairie is reduced to 100 acres or less and is in small, isolated remnant strips along railroad rights of way. There are ongoing Cajun Prairie restoration projects with the oldest being the ten acre site in Eunice (Vidrine et al. 2001) The Cajun Prairie flora was presented at the 17'^ North American Prairie Conference by Allen et al. (2001). These report included 513 specific and sub-specific taxa in 92 families and 277 genera. An under- standing of vegetation changes in Cajun Prairie is important in future restoration and preservation efforts. The present report documents vegetation change in four restorations: Eunice ten-acre restoration; Duralde Wildlife Refuge; Eunice 2.5-acre mitigation restoration; and Acadia Power Partner Prairie. Observa- tions on vegetation change after disturbance in remnants are also included. The goals in these restoration efforts were to recrea l ly as possible the Cajun Prairie as it exists in the remnants and to place Cajun Prairie plants into protected areas to ensure the survival of this threatened and vanishing genome. All four restoration efforts have used transplanted plugs and hand gathered seeds. Due to the high variability in the number and kinds of plant propagules within each plug, no specific data are available on the numbers and kinds of species in each restoration. The senior author has been involved in all four restoration efforts. J. Bot. Res. Inst. Texas 1(2): 1141 — 1147. 2007 1142 Journal of the Botanical R h Institute of Texas 1(2) RESTORATION SITES 1. The Eunice restoration site is a ten-acre site located in the city of Eunice in St. Landry Parish, Louisiana with restoration beginning in 1988 and represents the oldest Cajun Prairie restoration effort. Prior to the restoration effort, the site was an abandoned fallow area with a pasture like appearance. The soil type at this restoration site is Crowley silt loam (Murphy et al. 1986). For a complete description of this restoration effort, refer to Allen and Vidrine (1989, 2004) and Vidrine et al. (2001). 2. The Duralde Wildlife Refuge is a federally owned 334-acre site in Evangeline Parish, Louisiana that has a patchwork of age and restoration methods scattered across its landscape (Allen and Grafe 2004). Res- toration was begun in 1995 and restoration continues today. The site was an agricultural field, mainly rice, that was abandoned a few years earlier and just prior to restoration was dominated by Chinese tallow trees (Triadica sebiferum). The soils at this site are mostly of the Crowley-Vidrine complex with limited amounts of Mowata silt loam and Mamou silt loam (Touchet et al. 1974). 3. The Eunice 2.5 acre mitigation restoration was begun in 1999 and is just north of the ten-acre Eu- nice site. Prior to restoration, this area was a fallow area maintained by periodic mowing and had a pasture appearance. The soils in the northern two-thirds of this site are Mamou silt loam with the southern third containing Crowley silt loam (Murphy et al. 1986). 4. Acadia Power Partner Prairie is a fourteen-acre site with approximately four-acres of the area under- going restoration efforts in late 2003 and early 2004. This area was an agricultural field, mainly rice, and is adjacent to and part of a power plant in Acadia Parish, Louisiana. The soils at this site are Crowley silt loam (Clark et al. 1962). In all four of these restorations plus in the remnants, each area contained one or more wet areas in addition to mesic ones so all contained wet, mesic and intermediate zones. METHODS During the past 15 years, observations were made and recorded by the senior author on speci mposition in all stages of succession in all four restorations plus after disturbance in the remnants. Ten remnants were walked every two weeks from February to November in 1987 and all species present were recorded and the abundance of each determined visually and with 100 permanent plots. Some plots were sampled for species composition during the early stages of the Duralde and Eunice restoration. Notes on which species is found in which wetness zone were also made and recorded. Nomenclature and information on nativity are from USDA Plants Database (USDA, NRCS 2004). The native non-prairie and Cajun Prairie distinction was made by the senior author based on more than 20 years of experience in Cajun Prairie Habitat and more than 30 years of work with the Louisiana Flora. The senior author contributed similar information to the floristic quality assessment for the coastal prairie species (Allain et al. 2006). RESULTS The vegetational changes in prairie restoration and remnant recovery are continuous but can be artificially divided into stages. The initial dominants after disturbance in the Cajun Prairie habitat are annuals; this has been observed in the first years of all four restoration efforts plus the vegetational changes associated with disturbances in the remnants. A list of annual species observed during these early stages in restoration or disturbance recovery in Cajun Prairie is in Table 1. This table includes 140 species; 42 were observed in Cajun Prairie restoration on land that was in agricultural use and 118 on land that was fallow-pasture. Annual species almost always observed in prairie restoration after agricultural field crops include Ambrosia spp., Bromus ca- tharticus, Caperonia palustris, Digitaria spp., Iva annua, Melochia corchorifolia, Sesbania herbacea, Sida rhombifolia and Urochloa platyphylla. Annual species usually associated with prairie restoration after pasture or fallow fields include Agalinis spp., Aristida spp., Conyza canadensis, Croton capitatus, Diodia teres, and Hypericum spp. qos SA Twenty-four of these species are introduced. In Table 1, the species are also identified as to their moisture Allen and Thames, Vegetation chang Cajun P | t Louisiana 1143 Tagle 1. List of annual species observed ir un Prairi tl pos ane An * before the name indicates an introduced species The first letter(s) in OMS E after the name indicates th d condition for the taxon prior to its appearance in early stage Cajun Prairie the name is the moisture preference for the taxon where M = mesic, W = wet, and | = intermediate m moisture requiremen ts. where A = agricultural and F = fallow- S The second letter i in parenthesis after Apiaceae Chaerophyllum tainturieri Hook. (F,M) o Um. digitatum DC. (FM) Ptilimnium capillaceum (Michx.) Raf. (FI) Spermolepis echinata (Nutt. ex DC.) Heller (EM) Asteraceae Ambrosia artemisiifolia L. (AFM) Ambrosia bidentata Michx. (AF,M) Ambrosia psilostachya DC. (AFM) Ambrosia trifida L. (AFM Bidens aristosa (Michx.) Britt. (FI) Conyza canadensis (L.) Cronq. (AFM) Coreopsis tinctoria Nutt. (F.M) Erechtites hieraciifolia (L.) Raf. ex DC. (EM) Erigeron annuus (L.) Pers. (EM) Erigeron strigosus Muhl. ex Willd. (F,M) amochaeta purpurea (L.) Cabrera (F,M) *Helenium amarum (Raf) H. Rock (AFM) Iva annua L. (Al Krigia caespitosa (Raf) Chambers (F.M) Krigia virginica (L.) Willd. (F,M) Lactuca canadensis L. (FM) Lactuca floridana (L.) Gaertn. (F.M) Packera glabella (Poit ) E Jeffery (Fl) Pluchea da (L) DC. (EW) Pyrrhopappus ji (Walt.) DC. (F,M) Sonchus asper (L.) Hill (EM) Sonchus oleraceus L. (EM) Boraginaceae Myosotis verna Nutt. (F,M) Brassicace Cardamine n ENEI) Cardamine parviflora L. var. arenicola (Britt) O.E. Schulz (FI) F. Lepidium virginicum . ” ) Buddlejaceae Polypremum procumbens L. (F.M) Callit litrichac ee pina Pursh (EW) Campanulaceae Triodanis perfoliata (L.) Nieuwl. (EM) Caryophyllaceae Cerastium glomeratum Thuill. (F,M) Silene antirrhina L. (F,M) Clusiaceae pericum d Jii (Grev. & Hook.) Torr. & Gray (M) Hypericum gentianoides (L.) B.S.P. (EM) Convolvulaceae Ipomoea lacunosa L. (A,M) Cucurbitaceae *Cucumis melo L. (A,l) Cyperaceae Bulbostylis n (L) Kunth ex C.B. Clarke (EM ) Cyperus acuminatus Torr. & Hook. ex Torr. (EW) Cyperus Se Muhl. (AW *Cyperus iria L. (AFW Eleocharis microcarpa Torr. (AFW) Eleocharis montana (Kunth) Roemer & J.A. Schultes (EW) Eleocharis obtusa (Willd.) J.A. Schultes (AFW) Fimbristylis autumnalis (L.) Roemer & J.A. Schultes (Fl) Fimbristylis miliacea (L.) Vahl (Fl) Isolepis carinata Hook. & Arn. ex Torr. (Fl) Kyllinga RA Vahl (FI) Euphorbiaceae Acalypha gracilens Gray (F,M) ad palustris (L) St.-Hil. (AI) Chamaesyce humistrata (Engelm.) Small (FM) Chamaesyce maculata (L.) Small (EM) *Chamaesyce nutans (Lag.) Small (A,M) Croton capitatus Michx. (AFM) Croton E 5 (AF,M) Idenov Webster (EM) eee a E (EM) WS Fabaceae Aeschynomene indica L. (AI) Chamaecrista fasciculata (Michx.) Greene (F.M) Glottidium vesicarium (Jacq.) Harper (A,I) *Kummerowia striata (Thunb.) Schindl. (A,M) *Medicago lupulina L. (EM *Medicago polymorpha L. (F,M) *Melilotus indicus (L.) All. (FM) ae bejariense Moric. (F.M) Trifolium dubium Sibthorp (F.M) *Trifolium resupinatum L. (F,M) Vicia ludoviciana Nutt. (F.M) I Gentianaceae *Centaurium pulchellum (Sw.) Druce (FM) Sabatia campestris Nutt. (F,M) — ll Geraniaceae f "n L. (EM) Iridaceae Sisyrinchium rosulatum Bickn. (F,M) Lamiaceae *Lamium amplexicaule L. (EM Monarda punctata L. (F,M) Stachys crenata Raf. (FM) — oa 1144 Journal of the Botanical R h Institute of Texas 1(2) TABLE 1. continued. Linaceae Polygalaceae Linum medium (Planch.) Britt. var. texanum (Planch.) Polygala leptocaulis Torr. & Gray (FI) Fern.(F,M) Polygala mariana P. Mill. (F.M) Linum sulcatum Riddell (FM) timulstese Loganiaceae *Anagallis arvensis L. (F,M) Mitreola petiolata (J.F. Gmel.) Torr. & Gray (F,M) Anagallis minima (L.) Krause (FI) Mitreola sessilifolia (J.F. Gmel.) G. Don (EM) Ranunculaceae Malvaceae Ranunculus laxicaulis (Torr. & Gray) Darby (AFW) Modiola caroliniana (L) G. Don (FM) *Ranunculus muricatus L. (AFW) Sida rhombifolia L. (A,M) Ranunculus pusillus Poir. (AFW) Molluginaceae Rubiaceae *Mollugo verticillata L. (AF,M) Diodia teres Walt. (AF,M) Onagraceae Pell aparine L. md - —— Gaura longiflora Spach (F,M) oustonia micrantha (Shinners) Terrell (F.M) Ludwigia decurrens Walt. (FW) eee Ludwigia leptocarpa (Nutt.) Hara (FW) ermum halicacabum L. (A\l) Oenothera laciniata Hill (F,M) Oenothera linifolia Nutt. (EM) axifragac a d UU Ell. (EM) Plantaginaceae Plantago aristata Michx. (F,;M) Plantago heterophylla Nutt. (F,M) Plantago virginica L. (F.M) Scrophulariaceae Agalinis fasciculata (Ell.) Raf. (FM) Agalinis heterophylla (Nutt.) Small ex Britt. (F,M) e oligophylla Pennell (F,M) x Poaceae galinis skinneriana (Wood) Britt. (FM) Alopecurus carolinianus Walt. (A\l) joris viridis (Small) Pennell (E.M) Aristida longispica Poir. var. longispica (F,M) Buchnera americana L. (EM Aristida oligantha Michx. (EM Gratiola neglecta Torr. (F.) *Briza minor L. (F.M) Gratiola virginiana L. (FW *Bromus catharticus Vahl (A,M) Lindernia dubia (L.) Pennell var. dubia (FI) Digitaria ciliaris (Retz.) Koel. (A,M) Nuttallanthus canadensis (L.) D.A. Sutton (FM) n igi itaria ischaemum (Schreb.) Schreb. ex Muhl. (A,M) Nuttallanthus texanus (Scheele) D.A. Sutton (F,M) git violascens Link (A,M) Veronica arvensis L. (F,M) E crus-galli (L.) Beauv. (AW) Veronica peregrina L. (F,M) Limnodea arkansana (Nutt) LH. Dewey (F.M) *Lolium opu L. (AM Phalaris angusta Nees ex Trin. (AI) Phalaris san Walt. (A,I) Solanaceae Physalis angulata L. (AF, M) olanum americanum P. Mill. (FM) == Setaria pumila (Poit 9| Roemer & J.A. Schultes (AFM) Sterculiacea Sphenopholis obtusata (Michx) Scribn. (FM) *Melochia p UTE L. (AD *Sporobolus indicus (L.) R. Br. (AF,M) Valerianaceae Urochloa platyphylla (Munro ex Wright) R. Webster (A,W) Valerianella radiata (L) Dufr. (EM) Vulpia octoflora (Walt.) Rydb. (EM) Verbenaceae Glandularia pulchella (Sweet) Troncoso (F,M) preference and include 100 in mesic condition, 16 in wet conditions, and 24 in intermediate conditions. Some annuals usually found in wet to moist areas during early restoration include Cyperus spp., Ranunculus spp., Sesbania herbacea, and Urochloa platyphylla. Most annuals disappear in a year or two but Agalinis spp., Croton capitatus, Chamaecrista fasciculata, and Sabatia campestris may persist for three to four years or even longer. Some aggressive perennials may also appear during the first year and include Axonopus fissifolius, Eupatorium serotinum, Paspalum dilatatum, Paspalum plicatulum, Paspalum urvillei, and Verbena brasiliensis A second stage of vegetation change includes three groups of perennial and biennial species: native non-prairie, introduced, and Cajun Prairie (Table 2). A complete listing of Cajun Prairie species can be found Y J y Allen and Thames, Vegetati in Caj irie i t Louisiana 1145 TABLE 2. List of perennial/biennial species observed in Cajun Prairie Restorations in Southwest Louisiana. An * before the name indicates an introduced species. The first letter(s) in parenthesis after the name indicates the observed condition for the taxon Pal io its Ln in early E e Prairie dE A= pl and F = fallow-pasture. The second letter in forthe taxon v 'esic, W = wet, and | z intermediate moisture requirements. The superscripted €, if present indicates a Cajun Baie: species. Amaranthaceae Carex frankii Kunthb (FW) *Alternanthera philoxeroides (Mart) Griseb. (AFW) Carex vulpinoidea Michx. (EW) ! G us croceus Vahl (EM) ain haspan L. (EW) Cyperus pseudovegetus Steud. (AW) Apiaceae Cicuta maculata L. (EW) Asclepiadac Cyperus retrorsus Chapman (F,M) aaa m Ell. (E, MC) *Cyperus rotundus L. (Al) viridis Walt. (FMS) Cyperus strigosus L. (A,l) QUU E laeve (Michx.) Pers. (EM) Cyperus virens Michx. (AW) steraceae Eleocharis quadrangulata (Michx.) Roemer & J.A. Schultes Baccharis halimifolia L. (FI9 FW) Boltonia asteroides (L) U'Hér. (ES Eleocharis tuberculosa (Michx.) Roemer & J.A. Schultesb Boltonia diffusa Ell. (EWS) (FW) Chromolaena ivifolia (L) King. & H.E. Robins. (EMS) Kyllinga brevifolia Rottb. (FM) Cirsium horridulum Michx. (EM) Rhynchospora corniculata (Lam.) Gray (A,W9) Coreopsis lanceolata L. (FMS) Euphorbiacea e *Triadica sebifera (L.) Small (AFI) — Coreopsis pubescens Ell. (F,M* Erigeron philadelphicus L. (F,M) Eupatorium capillifolium (Lam.) Small (AFM) Eupatorium serotinum Michx. EM Euthamia leptocephala (Torr. & Gray) Greene (F,MS) Hydrophyllaceae Euthamia tenuifolia (Pursh) Nutt. (EMS) Hydrolea ovata Nutt. ex Choisy (FW) Helenium flexuosum Raf. (E 19) Helianthus angustifolius L. (F,15) Krigia dandelion (L.) Nutt. (EMS) Ud scandens (L.) Willd. (E, M) > Haloragaceae Proserpinaca palustris L. (FWS Iridaceae Sisyrinchium angustifolium P. Mill. (MS) Sisyrinchium atlanticum Bickn. (EMS) Sisyrinchium langloisii Greene (F,M*) kera COPIER a (Micl IX.) € Jeffery (FIS) Pluchea a ne (EW) uncaceae ey (EW) Juncus brachycarpus Engelm. (FW) m n ME L (EM) Juncus effusus L. (FW) Solidago sempervirens L. var. mexicana (L.) Fern. (Fl) Juncus marginatus Rostk. (F.M) Juncus nodatus Coville (FW) Caprifoliaceae Juncus polycephalus Michx. (EW) "Lonicera japonica Thunb. (FM) Juncus tenuis Willd. (FM) Cistaceae Juncus validus Coville (EW) Lechea mucronata Raf. (EMS) Lechea tenuifolia Michx. (EMS) Lamiaceae *Stachys floridana Shuttlw. ex Benth (F,M) Clusiac Teucrium canadense L. (FM) ls ron Engelm. & Gray (FI) AMES Commelinaceae lium canaden ar. cahadense (F M ) Commelina erecta L. (A,M) Nothoscordum BN (L.) Britt. (EMS) Hu ae pne pud Lythraceae Iradescantia virginiana L. (EM) Cuphea glutinosa Cham. & Schlecht. (FI) Convolvulaceae Lythrum alatum var. (Ell.) Torr & Gray ex Rothrock Dichondra li is Michx. (EM) (EW) ucurbita Melastomataceae ade Mur L. (A,M) Rhexia mariana L. (F9) Cyperaceae Onagraceae Carex alata Torr. (EW) Ludwigia glandulosa Walt. (FW) Carex poen Schwein. (FW) 1146 Journal of the Botanical R h Institute of Texas 1(2) TasLe 2. continued. Ludwigia linearis Walt. (F9) Panicum rigidulum Bosc ex Nees var. rigidulum (F9) Ludwigia palustris (L.) Ell. (FW) *Paspalum dilatatum Poir. (AFM) Oenothera biennis L. (EMS) Paspalum plicatulum Michx. (AFI) Oenothera speciosa Nutt. (F,M) *Paspalum urvillei Steud. (AFI) Orchidaceae a parviflora (Poir.) Kerguélen (AFM) *Sorghum halepense (L.) Pers. (AFM) Spiranthes vernalis Engelm. & Gray (FM*) . Steinchisma hians (Ell.) Nash (FIF) Oxalidaceae Tridens strictus (Nutt.) Nash (FS) Oxalis stricta L. (FM) Pol Passifloraceae Porcia num rd Michx. (AEW) Passiflora incarnata L. (F,M*) Rumex verticillatus L. (AFI) Phytolaccaceae Rosaceae Phytolacca americana L. (FM) Rubus argutus Link (FM) Poaceae Rubus trivialis Michx. (F.M) Agrostis hyemalis (Walt) B.S.P. (A,M*) Rubiaceae Axonopus fissifolius (Raddi) Kuhlm. (AF,I°) irginiana L. (FJ) Bothriochloa exaristata (Nash) Henr. (FM?) M tinctorium L. (EM) *Bothriochloa ischaemum (L) Keng (FM) Oldenlandia boscii (DC) Chapman (F) Bothriochloa longipaniculata (Gould) Allred & Gould (F,MS) *Chloris canterai Arech. (FM) scropnulatia Bacopa e (Michx.) Wettst. (AFW) Dichanthelium A Omin (L Gould var. dichotomum (F,I9) Mecardonia acuminata (Walt) Small (F,M£) Dichanthelium scoparium (Lam.) Gould (FIS Solanaceae Dichanthelium sphaerocarpon (Ell) Gould var. sphaerocarpon Physalis heterophylla Nees (F.M) (FM*) Solanum carolinense L. (FM) Eragrostis bahiensis (Schrad. ex J.A. Schultes) J.A. Schultes Solanum elaeagnifolium Cav. (F,M) (E) Eragrostis elliottii S. Wats. (EMS) Eragrostis hirsuta (Michx.) Nees (F,M‘) ~ yphaceae Typha latifolia L. (EW — Eragrostis lugens Nees (F,M‘) Urticaceae Eragrostis refracta (Muhl) Scribn. (EMS) Boehmeria cylindrica (L.) Sw. (E W^) Eragrostis spectabilis (Pursh) Steud. (EMS) Verbenaceae Leersia hexandra Sw. (AW) Phyla nodiflora (L.) Greene (Fl) Panicum hemitomon J.A. Schultes (AF,W‘) *Verbena brasiliensis Vell. (AF) Verbena halei Small (F,M) in Allen et al. (2001). The total of 122 species include 24 species that were commonly observed with prairie restoration after agricultural field crops and 113 on land that was fallow-pasture areas. The species are also identified as to their moisture preference and include 63 in mesic conditions, 31 in wet conditions, and 28 in intermediate conditions. A total of 12 of these species are introduced, 65 are considered by us as native non-prairie species and the remaining 47 species are early stage Cajun Prairies species. The most commonly observed perennial species include Axonopus fissifolius, Eupatorium serotinum, Paspalum dilatatum, Paspalum plicatulum, Paspalum urvillei, Solidago altissima, and Verbena brasiliensis DISCUSSION The Cajun Prairie vegetation, like other vegetation types, changes dramatically after a disturbance and then continues to change over time. The initial dominants are annuals which are replaced by native and introduced perennial species. With time, the Cajun Prairie species become the dominant species and the introduced species and the native non-prairie species are lost; the mechanism(s) by which the Cajun Prairie species are able to dominate is not known and should be the focus of a future research project. The first Allen and Thames, Vegetation cl in Cajun Prairie i t t Louisiana 1147 1; LN introduced perennial to disappear is Brazilian vervain (Verbenal and the last to disappear is vasey grass (Paspalum urvillei). The first native species to disappear is late flowering thoroughwort (Eupatorium serotinum), followed by dog fennel (Eupatorium capillifolium) and the last to be lost is common goldenrod (Solidago altissima). This change in species composition toward a prairie dominated one is observed in all three moisture regimes. REFERENCES ALLAIN. L., L. SMITH, C. ALLEN, M. VipniNe, and J.B. Grace. 2006. A floristic quality assessment system for the Coastal Prairie of Louisiana. In D. Egan and J. Harrington, eds. Proc. 19^ North Amer. Prairie Conf. University of Wisconsin-Madison, Madison. ALLEN, C. M. and M.F. Viorine. 1989. Wildflowers of the Cajun Prairie. Louisiana Conservationist 41:20-25. ALLEN, C.M. M.F. Viorine, B. Borsari, and L. ALLAIN. 2001. Vascular flora of the Cajun Prairie of southwestern Louisiana. In: N. Bernstein and L. Ostrander, eds. Proc. seventeenth North Amer. Prairie Conf. North lowa Area Com- munity College, Mason City. Auten, C. M. and M.F. Viorine. 2004. The Eunice Cajun Prairie restoration project. In: J. Randall and J. Burns, eds. Proc. Third Eastern Native Grass Symposium. Omnipress, Chapel Hill, N.C. ALLEN, C. M. and V. Gnart. 2004. The Duralde Prairie restoration project, Cajun Prairie on a Federal Refuge. In: J. Randall and J. Burns, eds. Proc. Third Eastern Native Grass Symposium. Omnipress, Chapel Hill, N.C. CLARK, H.L., G.J. Haley, E.J. Hesert, R.M. HoLLIER, and AJ. Roy. 1962. Soil survey Acadia Parish, Louisiana. U.S. Dept. Agric., Soil Conservation Service in cooperation with Louisiana Agric. Exp. Sta. Murphy, KE, J.K. Viorine, D.R. McDaniel, and C.L. Goprnev. 1986. Soil survey of St. Landry Parish, Louisiana. U.S. Dept. Agric., Soil Conservation Service in cooperation with Louisiana Agric. Exp. Sta. and Louisiana State Soil and Water Conservation Committee. ToucHeT, B.A., H.L.CLark, C.M. Rebuic, AJ. Roy, B.L. GrirFIS, S.J. ZAUNBRECHER, and R.M. Hour. 1974. Soil survey of Evangeline Parish, Louisiana. U.S. Dept. Agric., Soil Conservation Service in cooperation with Louisiana Agric. EXpsota: USDA, NRCS. 2004. The PLANTS database (http://plants.usda.gov/plants). National Plant Data Center, Baton Rouge, LA 70874-4490 USA. VIDRINE, M.F., C.M. ALLEN, B. Borsari, L. ALLAIN, and S.R. Johnson. 2001. The Cajun Prairie Restoration Project. In: N. Bernstein and L. Ostrander, eds. Proc. seventeenth North American Prairie Conf. North lowa Area Community College, Mason City. 1148 Journal of the Botanical R h Institute of Texas 1(2) BOOK NOTICES University of California Press MICHAEL G. BARBOUR, Topp KeELER-WOLF, and ALLAN A. SCHOENHERR (eds.). 2007. Terrestrial Vegetation of California. Third Edition. (ISBN 978-0-520-24955-4, hbk.). University of California Press, 2120 Berkeley Way, Berkeley, CA 94704, U.S.A. (Orders: California Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A.; www.ucpress.edu; 609-883-1759; 609-883-7413 fax). $75.00, 730 pages, color and b/w photos, 8 1/2" x 11". A “thoroughly revised, entirely ” past the 2nd edition of 1988, with contributions from 43 authors. Table of Contents.—1. The History of Vegetation Classification and Mapping in California. 2. Climate, Paleoclimate, and Paleovegetation. 3. California Soils and Examples of Ultramafic Vegetation. 4. Nonnative Plants of California. 5. Estuarine Wetlands. 6. Beach and Dune. 7. Northern Coastal Scrub and Coastal Prairie. 8. Sage Scrub. 9, The California Channel Islands. 10. Forests of Northwestern California. 11. Closed-Cone Pine and Cypress Forests. 12. Oak Woodlands and Forests. 13. Chaparral. 14. Valley Grassland. 15. Vernal Pools. 16. Riparian Vegetation of the Great Valley. 17. Montane and Subalpine Vegetation of the Sierra Nevada and Cascade Ranges. 18. Southern California Conifer Forests. 19. Alpine Vegetation. 20. Transmontane Coniferous Vegetation. 21. Sagebrush Steppe. 22. Mojave Desert Scrub. 23. Colorado Desert Vegetation. Species Index, General Index Two Books in the California Natural History Guide Series Joan Easton Lentz, Dow DesJARDIN (Prtincipal Photographer), and PETER Garpr (Illustrator). 2006. Introduc- tion to Birds of the Southern California Coast. (ISBN 0-520-23780-3, pbk.). California Natural History Guide Series No. 84. University of California Press, 2120 Berkeley Way, Berkeley, CA 94704, U.S.A. (Orders: California Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A.; ww press.edu; 609-883-1759; 609-883-7413 fax). $19.95, 316 pages, 120 color, 51 line illustrations, 16 maps, 4 1/2" x 7 1/4". Publisher ee mation from back cover.—An excellent introduction and a perfect a for excursions ranging from the Mexican Obispo County, this indispensable guide will familiarize bird-w s, hikers, naturalists, residents, and travelers with the appearance and behavior of 120 of southern California's most common coastal birds. ARTHUR R. KRUCKEBERG. 2006. Introduction to California Soils and Plants: Serpentine, Vernal Pools, and Other Geobotanical Wonders. (ISBN 978-0-520-23372-0, pbk.). California Natural History Guide Series No. 86. University of California Press, 2120 Berkeley Way, Berkeley, CA 94704, U.S.A. (Orders: California Princeton Fulfillment Services, 1445 Lower Ferry Road, Ewing, NJ 08618, U.S.A.; www.ucpress.edu; 609-883-1759; 609-883-7413 fax). $18.95, 280 pages, 181 b/w and color photos, ZA x TE Publisher information from bach cover. —The definitive resource for exploring California's spectacular landscape and abundant flora and fauna. 1) 148 photographs illustrate California's diverse flora and rich geologic diversity, 2) Provides selected trip itineraries for view- ing the state's geobotanical wonders, 3) Includes information on human influences on the California landscape from the early Spanish explorers through the gold rush and to the present. J. Bot. Res. Inst. Texas 1(2): 1148. 2007 PHYTOGEOGRAPHY OF THE BIG LHICKET, EAST TEXAS Michael H. MacRoberts and Barbara R. MacRoberts Bog Research 740 Columbia, Shreveport, Louisiana 71104, U.S.A. an Herbarium, Museum of Life Sciences Louisiana State University in Shreveport, Shreveport, Louisiana, 71115, U.S.A. ABSTRACT M ined the floristic relationship | he Big Thicket of east T ltl inder of North America. We found that floristi- cally the Big Thicket i helmingl ] bly southeastern. There is very little western influence on it. RESUMEN Hem aminado las relaciones florísticas entre el Big Thicket del este de Texas y el resto de Norte América. Hemos encontrado qu florísticamente el Big Thicket I ] i l bl del sur Hay muy poca influencia occidental en él INTRODUCTION Extraordinary claims have been made regarding the ecology of the Big Thicket of east Texas. It has been described as exceptionally species rich, unusually diverse in plant communities, and a biological “crossroads” between eastern and western floras (see Cozine 2004; MacRoberts & MacRoberts 2004; Diggs et al. 2006 for reviews and literature). On the other hand, recent examinations of these claims have led to the conclu- sion that the Big Thicket is not a crossroads but is southeastern with levels of habitat diversity and species richness comparable to other areas of the southeast (MacRoberts & MacRoberts 2004; Diggs et al. 2006; MacRoberts et al. 2007). As Diggs et al. (2006:166) conclude *these conflicting views need examination." However, such an examination has been hampered by a lack of adequate floristic data. Until recently, the Big Thicket flora has been understudied and undocumented (MacRoberts et al. 2002b), a problem now largely overcome by a floristic inventory of the major part of the Big Thicket National Preserve (MacRoberts et al. 2002b; Brown et al. 2005, 2006a, 2006b, in prep.), by a reexamination of earlier plant collections from the Big Thicket National Preserve (L.E. Brown pers. comm.; Warren Pruess pers. comm.), and by the publica- tion of Atlas of the Vascular Plants of Texas (Turner et al. 2003) and Illustrated Flora of East Texas (Diggs et al. 2006). Additionally, much better distribution information on the flora of North America is now available (e.g., Kartesz & Meacham 1999; USDA 2007; Flora of North America 1993-2009). Figure 1 shows the location of the Big Thicket. While the exact geographic boundaries of the Big Thicket are ill-defined, it is generally considered to consist at least of Newton, Jasper, Hardin, Tyler, Polk, and northern Liberty county, or an area of approximately 13,500 square kilometers (see Diggs et al. [2006] for a discussion of the Big Thicket location and a description of its ecology). Within this area is located the Big Thicket National Preserve, with 12 units ranging from 222 ha to 10,100 ha and totaling about 34,000 ha. Since the units were originally selected to represent the Big Thicket community structure and flora (Peacock 1994; Cozine 2004; Diggs et al. 2006), a floristic inventory of the units should constitute a fair sample of the area. As part of a continuing study of the flora of the West Gulf Coastal Plain, in this paper we examine the floristic relationship between the Big Thicket and the remainder of North America north of Mexico. METHODS 1. Using the vouchered floristic lists of MacRoberts et al. 2002b), Brown et al. (2005, 2006a, 2006b, in prep, Warren Pruess (pers. comm.) for the Big Thicket National Preserve, we developed a master list of the J. Bot. Res. Inst. Texas 1(2): 1149 — 1155. 2007 1150 Journal of the Botanical R h Institute of Texas 1(2) | | ET $1 1 mui: — i [zx LES RE i: p: » O B n Hi E, ze ESA my SEE, e LM PUE 3 ES | | NN NEN HH TA A ss Mane: - SUE e rd Ae m M = == AN em iE EE = M A | | | | : Ne ve es oz a T $ Lo ma Z er, a i E Pis e $ - d^ z = aH Os DAI Ji d SSS p TEES S Din Thiel Fic. 1. Location of Big vascular plant species occurring in the Big Thicket region of east Texas. This list consists of 1076 species. Using Kartesz and Meacham (1999) and Flora of North America (1993-2006), we plotted the North American distribution of these species by state, province, and region. 2. Because Texas is a large state and because there is a major floristic ecotone across its center where east and west meet (MacRoberts & MacRoberts 2003; Diggs et al. 2006), we divided the state into 24 areas approximately 180 km on a side and using Turner et al. (2003) and Diggs et al. (2006), we plotted the dis- tribution of the species where possible in our master list (see method 1 above) across Texas to determine the floristic relationship between the Big Thicket and other areas of Texas. This list consists of 1035 species. 3. Using Matos and Rudolph (1985), Brown et al. (2002), Turner et al. (2003) and Diggs et al. (2006), we located all species from the Big Thicket area (Newton, Jasper, Tyler, Hardin, Polk, and northern Liberty county) not in our master list (see method 1 above). We combined this list with our previous list making a sample of 1462 species. Using Kartesz and Meacham (1999) and the Flora of North America (1993-2006), we determined their North American distribution. We then classified all species into one of five categories: 1) western: species that occur exclusively or predominantly in the west (the floristic divide between east and west is about 97 or 98 west longitude [MacRoberts et al. 2003]) (e.g., Loeflingia squarrosa Nutt.), 2) central: species that are predominantly in the central plains area (e.g., Tradescantia occidentalis (Britt.) Smyth), 3) MacRoberts and MacRoberts, Phytogeography of the Big Thicket, East Texas 1151 Pz RA pe " . 4 PA E PCT i vi Fic. 2. D [ f Bia Thicket fl f | [ reaion, or province. = » d r universal: species that occur in virtually all states and provinces in North America (e.g., Athyrium felix-femina (L.) Roth), 4) eastern-western: species that are not universals but which occur equally in both the east and in the west (e.g., Evolvulus sericeus Sw., Phyla nodiflora (L.) Greene), 5) eastern: species that are either exclusively found in the east or are predominantly in the east (e.g., Xyris spp.)(this category includes West Gulf Coastal Plain endemics [MacRoberts et al. 2002a]). Our purpose was to determine the influence of different regions on the flora of the Big Thicket. 4. Using MacRoberts et al. (2002b), Brown et al. (2005, 2006a, 2006b, in prep.), Turner et al. (2003) and Diggs et al. (2006), we located all species from the Big Thicket area (as in method 3) that also occurred in south Texas (Kenedy, Brooks, Jim Hogg, Zapata, Starr, Hidalgo, Willacy, Cameron counties) and deter- mined their North American distribution. Our purpose was to determine the distributional characteristics of species in common between the Big Thicket and outlying regions to see what their commonality was. The two areas have 303 species in common. 5. In order to develop a comparative sample, using MacRoberts and MacRoberts (2006) we applied methods 1 and 3 to the flora of Caddo Parish, Louisiana. Caddo Parish is located about 200 km north of the Big Thicket in the northwestern corner of Louisiana, in the center of the West Gulf Coastal Plain (Fig. 1). We used a random sample of 417 species of the Caddo Parish flora as in method 1, and 1168 species as in method 3. Exotic species are excluded from all of our data sets. RESULTS Figure 2 shows the distribution of Big Thicket flora as percent of total species found in each state, province, or region. Figure 3 shows the distribution of Big Thicket flora as percentage across Texas. Table 1 shows £L Dat cal D hi PA 1152 Journal of f Texas 1(2) NUS AE NO Fic 3 Distributi £DinThirl f£ T, VAL (C ultf. I Di4; "P 1: i^c Din Thiel R 1: l^ how the regions of North America relate to the Big Thicket and Caddo Parish. Table 2 shows the relationship between the species that occur in common between the Big Thicket and south Texas. Figure 4 shows the distribution of Caddo Parish flora as percent of total species found in each state, province, or region. DISCUSSION The results of method 1 show that the Big Thicket is p hically eastern; notably I 98% of the Big Thicket flora occurs in Louisiana, 91% in OE 8896 in Florida and Georgia, 4896 in New York, but only 1996 in New Mexico. In fact, the Big Thicket has more of its flora in common with Wisconsin, Michigan, Minnesota, Ontario, and New England than it has with any western state or province. Method 2, not surprisingly, shows that within Texas the Big Thicket is floristically most similar to east and central Texas, notably to the West Gulf Coastal Plain region of which it is a part. Method 3 shows that eastern spe- cies dominate the Big Thicket, with 8196 of the total, followed by eastern-western species, with 1096, and universal species, with 596. Western and central species account for slightly less than 496; that is, over 9696 of species are eastern associates (eastern, universal, eastern-western). Method 4 shows that, of the species MacRoberts and MacRoberts, Phytogeography of the Big Thicket, East Texas 1153 TABLE 1. Regional affiliation of Big Thicket and Caddo Parish flora. Big Thicket Caddo Parish Region Number of Percent of Number of Percent of Species Species Species Species Western 19 13 17 14 Central 36 24 5] 44 Universal 77 53 85 73 Eastern-Western 143 9.8 181 15:5 Eastern 1187 81.2 834 714 Total 1462 100.0 1168 100.0 TABLE 2. Regional affiliations of species in common to floras of Big Thicket and south Texas. Region Number of species Percent of Species Western 9 30 Central 12 40 Universal 33 10.9 Eastern-Western 102 334 Eastern 147 48.4 Total 303 100.0 in common between the Big Thicket and south Texas, eastern species dominate, with 48%, followed by eastern-western species, with 34%, and universal species, with 11%; in other words, 93% of the species are eastern associates (eastern, universal, eastern-western). Western and central species account for only 7% of the total. This last analysis also makes clear that the vast majority of species held in common between such areas as south Texas and the Big Thicket are eastern, not western, indicating that the Big Thicket is not an ecotone or “crossroads” between east and west but is located fully in the eastern part of the North Ameri- can floral region (Austroriparan subprovince) (McLaughlin 2007). Almost without exception western and central species that occur in the Big Thicket are very rare there and are usually associated with specialized and uncommon habitat, such as prairies and barrens (e.g., Croptilon rigidifolium (E.B. Smith) E.B. Smith, Hedeoma drummondii Benth.). The Caddo Parish analysis (method 5) reinforces this conclusion. While Caddo Parish has more in com- mon with states and provinces north of it (e.g., Arkansas, Oklahoma) than does the Big Thicket (MacRoberts & MacRoberts 2006, unpublished data), it has approximately the same relationship with New Mexico, New York, Mississippi and Florida as does the Big Thicket. Additionally, while Caddo Parish partakes of the same percentage of western species as does the Big Thicket, it has a higher percentage of central, universal, and east-west species than does the Big Thicket. Clearly, therefore, the Big Thicket is not exceptional in the West Gulf Coastal Plain in its affinities to other regions except perhaps the east and/or southeast. What can be truthfully said about the Big Thicket is that many eastern plants reach their western and southern limit in that region. Some of these species are: Cypripedium kentuckiensis C.F. Reed, Cyrilla racemiflora L., Eriocaulon compressum Lam., Fagus grandiflora Ehrh., Hypericum galioides Lam., Lechea minor L., Leucothoe racemosa (L.) Gray, Rhexia petiolata Walt., Rhododendron canescens (Michx.) Sweet, Stewartia malacodendron L., Triantha racemosa (Walt.) Small, Trillium recurvatum Beck, and Xyris stricta Chapman. Many other eastern species get much farther west than the Big Thicket; examples include: Drosera capillaris Poir., Eriocaulon decangulare L., Liatris pycnostachya Michx., Rhexia mariana L., Sarracenia alata Wood, and Xyris torta Sm. Few truly western or central species reach the Big Thicket; western and central habitat is simply not there or is very limited. Rus 1154 Journal of the Botanical R h Institute of Texas 1(2) Fic. 4. Distributi £fladAn Darich £l | * Lineal: bl region, or province Y ACKNOWLEDGMENTS Warren Pruess provided an updated list (January 20, 2007) of the plants collected from the Big Thicket National Preserve, a project that was supported, in part, by a National Park Service Cooperative Agreement (No. 14001004) to Paul Harcombe, Department of Ecology and Evolutionary Biology, Rice University. Two anonymous reviewers made useful suggestions. REFERENCES Brown, L.E., HILLHOUSE, K., B.R. MAcRoserts, and M.H. MacRoserts. 2002. The vascular flora of Windham Prairie, Polk County, east Texas. Texas J. Sci. 54:227-240. Brown, L.E., B.R. MacRoserts, M.H. MacRoserts, PA. HARCOMBE, W.W. Pruess, 1.5. ELsik, 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. Harcom8E, 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. Pruess, I.S. ELsik, 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, I.S. ELsik, 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. MacRoberts and MacRoberts, Phytogeography of the Big Thicket, East Texas 1155 Cozine, J. 2004. Saving the Big Thicket: From exploration to preservation, 1685-2003. Univ. North Texas Press, Denton. Dicas, G.M., B.L. Liescome, M.D. Reep, and R.J. O'Kennon. 2006. Illustrated flora of East Texas. Sida, Bot. Misc. 26: 1-1594. FLORA or NORTH AMERICA EDITORIAL Committee (eds.). 1993-2006. Flora of North America North of Mexico. Vols. 1, 2, 3, 4,19, 20, 21, 22, 23, 25, 26. Oxford Univ. Press, New York and Oxford. KARTESZ, J.A. and C.A. Meacham. 1999, Synthesis of North American flora. Version 1.0. North Carolina Botanical Garden. Chapel Hill. MacRoserts, M.H., B. R. MacRoserts, B. Sonne, and R. Evans. 2002a. Endemism in the West Gulf Coastal Plain: impor- tance of xeric habitats. Sida 20:767—779. MacRostRrS, B.R., M.H. MacRoserts, and L.E. Brown. 2002b. 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 B.R. MacRoserts. 2003. The east-west transition of flora in Texas: a biogeographical analysis. Sida 20:1693-1700. MacRoserTs, M.H. and B.R. MacRoserts. 2004. The Big Thicket: typical or atypical. East Texas Hist. Assoc. 42:42-51. MacRoberTs, 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 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. Matos, J.A. and D.C. RupoLrH. 1985. The vegetation of the Roy E. Larsen Sandylands Sanctuary in the Big Thicket of Texas. Castanea 50:228-249. McLaucHun, S.P. 2007. Tundra to tropics: the floristic plant geography of North America. Sida, Bot. Misc. 30. Peacock, H.H. 1994. Nature lover's guide to the Big Thicket. Texas A&M Press, College Station. Turner, B.L., H. Nicots, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Sida Bot. Misc. 24: 1-888. Uspa nacs. 2007. The PLANTS Database (http://plants.usda.gov). National Plant Data Center, Baton Rouge, LA 70874-4490. 1156 f the Botanical R hl Journal UI titute of Texas 1(2) BOOK NOTICES Jil Nokes, with Par Jasper, Betty Sue FLowers (Foreword), and Krista WHITSON (Principal photography). 2007. Yard Art and Handmade Places: Extraordinary Expressions of home. (ISBN 978-0-292-71679-7, hbk.). University of Texas Press, PO. Box 7819, Austin, TX 78713-7819, U.S.A. (Orders: http://www. utexas.edu/utpress, 512-471-4032). $29.95, 170 pp., color photographs, 9 1/4" x 8 1/4". This unique book is filled with inspiration, pleasure, and delight! Just read through the contents and you too will want your own personal copy. Table of Contents 1. Places Full of Memory and Meaning 2 toi fh a 3. Cascada de Piedra Pinta 4. A Natural and Peaceful Spirit 5. Little by Little, He's Been Blessing Us 6. Stories from the Valley 7. The Love You Have To Do It 8. Placenames and ie 9. Mango's Jun, 10. The | Virtuosity of C 11. One Big Cactus and Succulent Dish Garden 12. A Surprise Comes Every Day 13. Sculpture You Can Live In 14. The Flower Man 15. The Cathedral of Junk 16. Casa de Azücar 17. Stories from the Panhandle 18. A Great Peaceful Place 19. The Only Trees for Miles Around 20. A Windbreak for the Homeplace 21. God and the Birdhouses Brought Us Together 22. A Lesson in Landmarks 23. This Is How You Stay Alive 24. Desires of the Heart 25. More Than Four Seasons Kiet E. SevERSON and CanoryN HULL Sc. 2006. The Nature of Eastern North Dakota: Pre-1880 Historical Ecology. (ISBN978-0-911042-65-8, pbk.). North Dakota Institute for Regional Studies, North Dakota State University, PO. Box 5075, Fargo, ND 58105, U.S.A. (Orders: http://ndirs.ndsu.nodak.edu, wendy. mcquayOndsu.edu, 701-231-8338, 701-231-1047 fax). $21.95, 308 pp., 18 b/w photos, maps, 10 tables, appendices, 6" x 9". Contents.—1) Introduction, 2) Geology, 3) Climate, 4) People, 5) Fire, 6) Vegetation, 7) Animals, 8) Form and Function, 9) The End. Reference List. J. Bot. Res. Inst. Texas 1(2): 1156. 2007 A FLORISTIC SURVEY OF NATIONAL PARK SERVICE AREAS OF TIMUCUAN ECOLOGICAL AND HISTORIC PRESERVE (INCLUDING FORT CAROLINE NATIONAL MEMORIAL), DUVAL COUNTY, FLORIDA Wendy B. Zomlefer David E. Giannasi E of A Biology un of acu Biology University of Georgia University of Georgia Plant e 2502 Plant sence’ Athens, Georgia 30602-7271, U.S.A. Athens, Georgia 30602-7271, U.S.A. Walter S. Judd TD Depa UTICE OI DUCUTIy isa Gainesville, Florida 32611-8526, U.S.A. ABSTRACT The portion of the Timucuan Ecological and Hi ic P including F aroline National M ial) admini 1 by the National Park Service, U.S. Department of the Interior, comprises 3,723 ha (9, 200 acres ) in a series of disjointed land parcels in eastern Jackson- ville, Duval County, Florida. A floristic survey was conducted to provide Park S l with a hered checklist of vascular Ts plant species, supplemented with such as r d , loca sity data, and community type. Three intensive collecting trips conducted in 2005 yielded 480 taxa (474 species plus 6 varieties) of vascular plants in 318 genera of 122 families. The five largest families are Asteraceae, Poaceae, Cyperaceae, Fabaceae, and Ericaceae. A map, descriptions, and photographs of the various lant communities are also provided RESUMEN La porción de la Reserva Ecológica e Histórica de Timucuan, ae Be el Servicio de Parques Nacionales del Departamento del E Interior, comprende 3,723 ha (9,200 ) de Jacksonville, condado Duval, Florida. Se desarrolló un análisis florístico con el eds de dd A personal im Servicio de Parques Nacionales, de una lista de las especies de plantas vasculares, suplementada iva, localidad y el tipo de comunidad circundante. Se realizaron tres giras de recolección intensiva durante 2005, A un ae de 480 taxa (474 especies y 6 variedades) de plantas vasculares, de 318 géneros y 122 familias. I i I Asteraceae, Poaceae, Cyperaceae, Fabaceae y Ericaceae. De igual forma se aportan mapas, descripciones y fotografías de las diferent les vegetales. INTRODUCTION Study Area The Timucuan Ecological and Historic Preserve is located within the city limits of Jacksonville, Duval County, Florida (Fig. 14), and is the largest cooperative park system in the United States, comprising federal, state, and city park lands, plus properties of private corporations, authorities, and over 300 private landowners (Anderson et al. 1996; Furbish et al. 1996; NPS 20072). The large area, 46,000 acres (18,620 ha), encom- passes the St. Johns River valley between the lower St. Johns and Nassau Rivers (Fig. 1B). These two rivers discharge directly into the Atlantic Ocean and form an extensive estuarine system dominated by salt marsh and coastal hammock habitat with marine to brackish open waters. The preserve itself is over 75 percent wetlands and open water (see Fig. 1B). Dense housing developments (and other scattered pockets of urban growth) within preserve boundaries were established prior to formal land use planning for the area. Preserve upland areas are highly desirable for development due to their aesthetic value and proximity to open water. Industries, such as plant ies and construction companies, are concentrated west of the preserve, and rural residential development is spread out in lower density subdivisions in outlying tracts. Estuarine wetlands and waters within the preserve are claimed under sovereignty of the state of Florida up to the mean tide line, and the city of Jacksonville has J. Bot. Res. Inst. Texas 1(2): 1157 — 1178. 2007 1158 Journal of the Botanical R h Institute of Texas 1(2) 839 82? 81? (b | 9 A GEORGIA FLORIDA Atlantic Ocean Lake City Jacksonville e Beach Ü Butch Bane = Dr. E JAX Internat! Airport Downtown Jacksonville Fe 1: locati UR T S CET: Ecological and Historic P A. Map of nortt n howing location of Ti D (Fig "4D Hm J I g y L ge g R AA I £ I IL HTI n If ty Ļ Š gi fal J JX bounded by St lal INI D; nes TAE " er or mate tape NL o peg MEE CURE ME ME gray I y ):1=Thomas Creek; 2 = Cedar Point; 3 Roosevelt Area (see Fig. 2); 4 = Kingsley Pl ion; 5 = Sohn Purchase. BHI = Black H Island; PP — private property inholdings. A: modified L MID? £, Dal {1007\.D pep IL y MID £ E. lS ul + I 40QL\) INDC (200 4, 2005, 2007a) me Zomlefer et al., Flora of NPS areas of Timucuan Preserve 1159 1 jurisdiction over zoning and land use. The d private citizens work through cooperative partnerships toward common acquisition and Viae den. goals (Anderson et al. 1996). The areas within the preserve controlled by the National Park Service (NPS; U.S. Department of the Interior), comprise ca. 9,200 acres (3,723 ha; NPS 2004) of disjointed land parcels, including Fort Caroline National Memorial, discussed below (NPS 20072). Over 60 percent of this NPS acreage is salt marsh (and submerged lands) bordering brackish waters. To obtain maximum plant diversity over such a broad area within three intense sampling trips, the survey team focused on five NPS land parcels (dark gray areas labeled “1” to “5” in Fig. 1B)—Thomas Creek, Cedar Point, Fort Caroline National Memorial/Theodore Roosevelt Area, Kingsley Plantation, and Sohn Purchase—that encompass a wide range of habitat types (including disturbed areas). Descriptive data on these habitats (discussed briefly below) are summarized in Table 1 1. Thomas Creek Parcel.—This site, bordered along the east by 1-95, is located ca. 3 miles (5 km) northeast of the Jacksonville International Airport. The gated property is accessed by easement through land owned by Castleton Beverage Corporation (a subsidiary of the Bacardi Corporation) at the end of Butch Bane Drive East. To the northeast, the Nassau River splits into Thomas Creek, and a smaller tributary, Seaton Creek, runs through the land more or less parallel to the interstate. Habitats prise hardwood swamp surrounded by pinelands that have been extensively logged; other disturbed areas include a water-filled borrow pit and dike system (from interstate construction). Trespassing poachers, who access the remote property along the interstate border, are a problem for NPS personnel. As of this writing, the future of the easement property, now for sale, is unknown. A Revolutionary War battle may have been fought within property boundaries: the Battle of Thomas Creek (17 May 1777), the final engagement in the second attempt by American forces to invade the British province of East Florida (Boatner 1973). However, the exact location of the battle has not been verified (R. Bryant, pers. comm.). 2. Cedar Point.— Cedar Point is at the end of south Cedar Point Road (ca. 5 mi [8 km] east of New Berlin Road) and comprises extensive salt marsh surrounded by upland maritime hammock and some former pine plantations. This property (and the Sohn Purchase, 5. below) is on the popular Black Ham- mock Island in northeastern Jacksonville, which is bordered by the Nassau River to the north and extensive marshes of the St. Johns River, to the south. (Cedar Point occupies the southwestern tip with wetlands of the St. Johns and Ft. George Rivers.) Black Hammock Island was only sparsely populated until the 1980's when infrastructure for development (paved roads, mail delivery, garbage service) became widely available; the opening of the Dames Point Bridge (over the St. Johns River) in 1989 further stimulated the population boom. The general area is now well known for panoramic views of marshlands and waterways. Cedar Point is a former fish camp (with cleared areas for parking and a boat ramp). Plans by NPS to construct a visitors' center there (with extensive hiking trails) may be delayed due to recent archeological evidence indicating some historical significance to this site (R. Bryant, pers. comm.). Fort Caroline National Memorial/Theodore Roosevelt Area.—This property comprises the Fort Caroline National Memorial/Ribault Monument, NPS headquarters/museum, and the Theodore Roosevelt Area (see detailed map, Fig. 2). Th mbined areas, a significant inholding among very expensive homes and riverfront property, have over 4 mi [6.4 km] of publicly accessible trails through extensive hammock, swamp, and salt marsh. Fort Caroline National Memorial (138 acres [56 ha]; including the fort exhibit, visitors' center, park maintenance buildings, parking area, and boat dock) is located on the north side of Fort Caroline Road (just north of the intersection with Monument Road) and borders the southern shore of the St. Johns River; Ribault Monument is in a small disjunct area (8 acres [3 ha]) about 0.7 mi (1.1 km) further east along the river (see Brief History of these monuments below.) Park headquarters is located about a mile to the south along Mount Pleasant Road, adjacent to the Theodore Roosevelt Area to the north and east. The latter ca. 578 acres 234 ha) of ecologically significant, valuable real estate were bequeathed to the Nature Conservancy by this property's last private owner, Willie Browne (1889-1970), in tribute to President Theodore Roosevelt and his conservation efforts. The NPS acquired the land in 1990. 4. Kingsley Plantation.—The plantation (including NPS offices and parking areas) is located on the northern point of Fort George Island, along the Fort George River. Most of the surrounding habitat (primarily 1160 Journal of the Botanical R h Institute of Texas 1(2) n hahitat £f Tagle 1. Year of acquisition, size, location, significance, access, the five survey sites of Timucuan Ecological and Historic Preserve administered by the National Park Service, Jacksonville, Florida (Fig. 1B). Data from Anderson et al. (1996), a ee NPS (2004), NPS ne cg R. Dan pa comm.). Habitat types: AQ ic habitat (standing freshwater and surrounding margin); DA — dunes n beach; MH = maritime hammock: PF = pine flatwoods; sa = "i It ety a community; sH = shell midden; sm = salt marsh; sw = swamp. Parcel Area Location Historical and/or Ecological Public Habitat [Year Acquired by NPS] ha (acres) Importance Access Types 1. Thomas Creek [1995] 243 (600) | Dead end w Butch Various wetland communities; NO AQ, DA, PF, Bane Dr. East, gated possible site of Revolutionary SW entrance through War "Battle of Thomas Creek" easement Property; bordered by Thomas Creek and I-95 2. Cedar Point [1996] 162(400) Black Hammock Riverfront property, encroaching Yes DA, MH, SM Island: 9023 Cedar development; significant habitats Point Rd.; n of St. including extensive salt Johns and Ft. George marsh; possible archeological Rivers significance 3. Fort Caroline National 293 (724) | 12713 Fort Caroline Valuable real estate, ees Yes AQ, DA, DU, Memorial [1950] & Ribault Rd/13165 Mt. development; national memorial MH, SA, SH, Monument [1958 at its Pleasant Rd. (park sites; significant uen SM, SW present location]/Theodore headquarters; including wetlands Roosevelt Area [1990] Roosevelt Area w of headquarters and s of fort); see Fig. 2 — 1991] 32 (80) Nw Ft. George Island; ^ Valuable real estate; early 19th Yes DA, DU, MH, along me George River cer tury plat tation, including SM planter's residence, slave quarters 4. Kingsley Plantation 5. Sohn Purchase [2001] 85 211) Black Hammock Island: Valuable real estate, encroaching No AQ, DA, SA, gated entrance w of development; significant habitats SW 13501 Saw Pit Rd; including wetlands s of Nassau River, w of Pumpkin Creek hammock) is disturbed. During Florida's plantation period (1763-1865), the isolated Fort George Island was owned by a series of planters, including Zephaniah Kingsley and his wife, Anna (a former slave from Senegal), who lived there from 1814 to 1837 (Schafer 1997; Stowell and Tilford 1998; Mallard 2007). The primary cash crop was sea island cotton (Gossypium barbadense), grown for the exceptionally long fibers spun into fine strong thread. The existing compound comprises the restored planter’s residence (ca. 1798) with heritage gardens, as well as the nearby ruins of the slaves' quarters (arranged in semicircle), constructed of tabby (a mixture of oyster shells, sand, and water). One cabin has been restored to its original appearance for the interpretive display. 5. Sohn Purchase.—The gated property, named after its last private owner, is located along the west side Saw Pit Road, ca. 1.5 mi (2.4 km) north of Cedar Point Road (see 2. Cedar Point above) and is in west- central Black Hammock Island, a rural portion undergoing development. The area was acquired by NPS in 2001 (R. Bryant, pers. comm.). The land is bordered to the west by Pumpkin Hill Creek, a southern branch of Nassau River, which feeds the swamp and wetlands (marsh and open water areas) of the northern, western, and southern parts of the property. Scrubby sandhill (last burned in 1993), densely covering the eastern and central portions, intergrades with the various wetland habitats. The extreme southern end of the property was burned in 1998. Zomlefer et al., Flora of NPS areas of Timucuan Preserve 1161 Fort Caroline st, JOHNS RIVER National Memorial Ta Ribault Monumen O UA KUM um - ) 2s = ats m o Fort Caroline € Mount P, Sant Road v S a $ 5 5 N z | NT nde uan j Theodore t- Le- y NES / Roosevelt " Downtown due Jacksonville Salt marsh VO Vi Area a 14 mi/22 km e 0 0.5 1 Kilometer 0 0.5 1 Mile Fic. 2. Detailed i £L f i Mak TAA enl 1 TL J D Fig. 1B). Modified by WBZ from NPS (2007a). Brief History of Fort Caroline National Memorial and Timucuan Ecological and Historic Preserve In February 1562, an expedition lead by French naval officer Jean Ribault landed on the mouth of the River May (now St. Johns River) and was welcomed by Chief Saturiwas, the head of a Timucua-speaking tribe ie Native Americans. Two years later, René Goulaine de Laudonniére founded the colony ofla Caroline (na for reigning French King Charles IX) on St. Johns Bluff (Bennett 1976, 2001). The Timucuan Indians pelted the settlers build the triangular Fort Caroline (Fort de la Caroline). The colony struggled to survive, and Ribault returned from Europe to take command of the settlement in August 1565. A month later, Don Pe- dro Menéndez de Avilés, governor of Florida for Spain, attacked Fort Caroline. As “heretics” (Huguenots or Protestants), the French were a threat to Catholic Spanish colonization in Florida. Most of the colony and members of Ribault’s fleet were slaughtered (245 French soldiers; see Zomlefer et al. 2004). The massacre ended attempts by the French to colonize Florida. The Spanish abandoned the fort in 1569. Fort Caroline National Memorial (Fig. 2) was established along the lower St. Johns River in 1950 to commemorate sixteenth century French efforts to establish a permanent colony in Florida. The memorial is listed on the National Register of Historic Places under the National Historic Preservation Act of 1966 (NPS 2007b). The park features a visitors’ center to complement a somewhat scaled-down reconstruction £L Dag PRA D hi PEA 1162 Journal of f Texas 1(2) of the fort (ca. 200 x 200 x 300 ft [61 x 61 x 91 m]) based on a sketch by Jacques le Moyne, the colony's mapmaker. Ribault Monument, an obelisk situated atop St. Johns Bluff, commemorates Jean Ribault's land- ing near the mouth of the St. Johns River. The exact site of Fort Caroline (and the settlement) is unknown and was likely destroyed when the river channel was deepened and widened in the 1880's (Anderson et al. 1996; NPS 20072). Fort Caroline National Memorial is specifically designated as a unit within the Timucuan Ecological and Historic Preserve, authorized by Congress in 1988 through Public Law 100-249 (Furbish et al. 1996). The preserve was created to protect and interpret the ecological and historical resources of the area, which include one of the few remaining unspoiled coastal wetlands along the Atlantic. The name honors the ex- tirpated native American Timucua tribes, who made contact with the first European arrivals to the area: “Timucua” actually refers to several culturally diverse tribes, sharing the same language, who inhabited northern Florida and southeastern Georgia (Hann 1996). European diseases devastated the population, and by 1700, less than 550 of these native Americans survived; today, no known indigenous people are Timucua. MATERIALS AND METHODS The first authors, Zomlefer and Giannasi, led three intensive field trips in 2005 (21-24 April, 28 June-1 July, 29 September-2 October; Giannasi & Zomlefer 1184-1573, 1576-1738) to collect vascular plant speci- men vouchers in duplicate using standard field and herbarium techniques (under Ne E permit + TIMU-2003-SCI-0001) with assistance of coauthor Judd and personnel listed in th Plant associations were also assessed. The survey focused on five main land parcels administered by NPS (Fig. 1B; Table 1). The floras of Wunderlin and Hansen (2000, 2003) were primary sources for plant identification, supplemented by Godfrey and Wooten (1979, 1981) and Godfrey (1988). The majority of plants were identi- fied by WBZ and DEG; Stephen Lee Echols identified most grasses and sedges; and coauthor WSJ confirmed other problematic determinations. A complete set of vouchers is deposited at GA. A duplicate set is at the Timucuan Ecological and Historical Preserve Museum (see NPS 2007a); that collection is hereafter designated “TIMU,” a Park Service acronym not registered in Index Herbariorum (Holmgren & Holmgren 2007). RESULTS AND DISCUSSION Floristics The 554 numbered collections comprise 480 taxa (474 species plus 6 varieties) in four major vascular plant groups (see ANNOTATED CHECKLIST OF SPECIES below): lycophytes (1 sp.), monilophytes (“ferns and allies;” 10 spp.), gymnosperms (7 spp.), and angiosperms (256 spp. + 6 vars.). Included in the list are 26 species planted around park headquarter/visitor buildings (Fort Caroline National Memorial, Theodore Roosevelt Area, and St. George Island; indicated as “cuLr”). The largest families are: Asteraceae (62 spp. + ] var.), Poaceae (40 spp. + 3 vars.), Cyperaceae (26 spp.), Fabaceae (21 spp. + 1 var), Ericaceae (16 spp.), Euphorbiaceae (11 spp.), Rubiaceae (11 spp.), Fagaceae (10 spp.), Hypericaceae (10 spp.), Juncaceae (10 spp.), Onagraceae (10 spp.), Lamiaceae (9 spp.), and Plantaginaceae (9 spp.). With completion of this survey, we have vouchered 56 new county records (54 species and two varieties) for Duval County, Florida, according to Wunderlin and Hansen (2007). No state/federally listed endangered plants (Duever 1996; Coile & Garland 2003; U.S. Fish and Wild- life Service 2007) and no UE a were found; however, Liatris tenuifolia var. quadriflora, cited in Wunderlin and Hansen (2003) d to Florida, has been recently vouchered from Cumberland Island Georgia (Zomlefer et al., in E Three state listed threatened species were found: Drosera intermedia (water sundew), Opuntia stricta (erect pricklypear), and Sarracenia minor (hooded pitcherplant); in addition, Os- munda cinnamomea (cinnamon fern), Osmunda regalis var. spectabilis (royal fern), and Rhododendron canescens (mountain azalea) are state listed commercially exploited plants (Coile & Garland 2003). Excluding the 21 cultivated exotics, the remaining 61 introduced species represent 13.596 of the total. Eleven species (including three cultivated plants) are listed as invasive exotics by Florida Exotic Pest Plant Zomlefer et al., Flora of NPS areas of Timucuan Preserve 1163 Council (FLEPPC 2007). Nine of these are ranked as Category I (invasive exo tics altering native plant com- munities by displacing native species, changing community structures/ecological functions, or hybridizing with natives): Asparagus aethiopicus, Cinnamomum camphora, Dioscorea bulbifera, Eichhornia crassipes, Lantana camara, Lonicera japonica, Lygodium japonicum, Macfadyena unguis-cati, and Sapium sebiferum. Two species, Alternanthera philoxeroides and Wisteria sinensis, are in Category II (invasive exotics increasing in abundance/ frequency but not yet altered Florida plant communities to the extent shown by Category I plants). Aspara- gus aethiopicus (Sprenger's asparagus-fern) and Lonicera japonica (Japanese honeysuckle) are planted at the NPS Park offices (Kingsley Plantation, Ft. George Island) and apparently are restricted to their flower bed fad unguis-cati (catclawvine) si plots; however, at the same site Wisteria sinensis (Chinese wisteria) and M are naturalized and spreading. Lantana camara (lantana), represented by a few plants cultivated with L. depressa and L. montevidensis near a boat dock (just east of reconstructed Fort Caroline), has some potential to spread into the disturbed areas surrounding the fort. Lygodium japonicum (Japanese climbing fern) is lo- cally common along the path through the maritime hammock surrounding the fort and should be removed before invading further. Dioscorea bulbifera (air-potato) and Eichhornia crassipes (common water-hyacinth), serious invasives, are common and well established around Spanish Pond. The Thomas Creek Parcel has three invasive species in aquatic habitats where propagules are likely dispersed by water runoff from 1-95: Alternanthera philoxeroides (alligatorweed; infrequent; dike area), Cinnamomum camphora (camphortree; infrequent; swamp), and Sapium sebiferum (popcorntree; locally common; dike area and swamp). Of these, Sapium sebiferum is the most common in this land parcel: the many seedlings, established along the margins of these aquatic habitats and in standing water, have potential to rapidly displace native species, especially in the hardwood swamp. A significant native species is in danger of extinction in the preserve due to a recent invasion of an exotic insect. In late June 2005 coauthors Zomlefer and Giannasi noted dying redbay trees (Persea borbonia), likely infected with redbay wilt, in the maritime forests of Cedar Point after park service personnel had reported similar occurrences at Kingsley Plantation. The presence of this fungal disease carrier, redbay ambrosia beetle (Xyleborus glabratus), was confirmed at the preserve in October of that year (Mayfield & Thomas 2006). Redbay wilt has since spread widely along the coast (R. Bryant, pers. comm.; Zomlefer et al., in prep.). Persea borbonia is a dominant understory component, and large-scale redbay mortality will greatly alter the composition of maritime forests along these coastlines. Plant Communities Nin ity types of the survey area are summarized below and several are depicted in Figure 3: dune; salt marsh; shell midden; maritime hammock; sandhill community; aquatic habitat (standing freshwater and surrounding margin), swamp; pine flatwoods; and disturbed areas. Table 1 includes a listing of habitat types in each of the five survey sites. Habitats often intergrade within a site. These general categories are based upon our field observations and classifications of similar areas by Laessle and Monk (1961), Stalter and Dial (1984), Myers and Ewel (1990), FNAI (1990), Easley and Judd (1993), and Zomlefer et al. (2004). Dunes/Open Beach.—The open beach comprises exposed sandy beach up to the high tide line, and foredune is the zone bordering the open beach, often sparsely vegetated with salt tolerant, pioneer species (Johnson & Barbour 1990; Zomlefer et al. 2004). Limited beach-like plant communities occur on narrow sandy strips (ca. 15 ft [4.6 m] wide) along the river beaches on the northern sides of Kingsley Plantation (St. George River) and the Fort Caroline exhibit (St. Johns River). p beach species found at these locations £ include: Batis maritima, Borrichia frutescens, Cenchrus spinifex, Hyd le bonariensis, Ipomoea alba, Oenothera J laciniata, Physalis walteri, Sarcoconia perennis, Sesuvium portulacastrum, and SONOS helvola. Salt Marsh.—Salt marshes in Florida are coastal communities of salt-tolerant plants pying intertidal zones at least occasionally inundated with salt water, characterized by dense stands of Spartina alterniflora (saltmarsh cordgrass) and/or Juncus roemerianus (black rush; Montague & Wiegert 1990; Easley & Judd 1993; Zomlefer et al. 2004). Extensive expanses of salt marsh comprise Cedar Point, as well as the north- eastern portion of the Theodore Roosevelt Area (Round Marsh, Fig. 3D); smaller areas are located along the western side of Fort Caroline (Calypso Island) and western edge of Kingsley Plantation (Steinway-Rodkin f Texas 1(2) " FE £l D JOurdi OF 1164 s ^ r Si TEE t j ^ dense overstory TL San 2005) D ML Ih: /NDC L J c Lyonia lucida 7 4. genui Fic. 3. E Vucicu Zomlefer et al., Flora of NPS areas of Timucuan Preserve 1165 & Montague 2003). Besides saltmarsh cordgrass and black rush, common salt-tolerant plants within these marshes include: Atriplex cristata, Batis maritima, Chenopodium album, Cynanchum angustifolium, Distichlis spicata, Limonium carolinianum, Sarcocornia perennis, Sesuvium portulacastrum, and Suaeda linearis. These salt marshes are bordered by woody plants such as: Baccharis halimifolia, Borrichia frutescens, Ilex vomitoria, Iva frutescens, Juniperus virginiana, Lycium carolinianum, Myrica cerifera, Opuntia stricta, and Sabal palmetto. Shell Midden (Shell Mound).—This coastal habitat is formed from the activities of native Americans who created elevated mounds of mollusk shell fragments that become mixed with organic matter over time (Fig. 3F). Water drains rapidly through the calcareous substrate and supports a distinctive set of indicator species; a closed hardwood canopy eventually develops (FNAI 1990). Well-developed shell midden habitat occurs around the salt marshes of the Theodore Roosevelt Area (Round Marsh, Fig. 3E), mainly as a sparse shrubby community intergrading with maritime hammock. The scrubby overstory species include: Forestiera segregata, Ilex vomitoria, Iva frutescens, Juniperus virginiana, Myrica cerifera, Quercus geminata, and Sabal pal- metto—with tangled vines of Cynanchum scoparium and Smilax auriculata. Borrichia frutescens, Chiococca alba, Eustachys petraea, Iresine rhizomatosa, Opuntia pusilla, and Psychotria nervosa are common in the understory. Maritime Hammock (Coastal Hammock; Maritime Forest).—This habitat, the terminal stage of suc- cession in coastal areas, is a band of “hardwood” forest just inland of the coastal strand on old, stabilized dunes; the generally mesic conditions are maintained by the dense canopy (Laessle @ Monk 1961; Stalter & Dial 1984; Easley & Judd 1993; Zomlefer et al. 2004). Well established maritime hammock borders the salt marshes of Fort Caroline National Memorial (Fig. 3C)/Theodore Roosevelt Area and Cedar Point. The ] € € branches of dominant species Quercus virginiana and Q. geminata are istically covered with epiphytes Pleopeltis polypodioides, Tillandsia recurvata, and Tillandsia usneoides. Juniperus virginiana, Persea borbonia, Prunus serotina, and Sabal palmetto are principal understory trees; other tree species are common at certain sites: Asimina parviflora and Sapindus saponaria at Theodore Roosevelt Area, and Carya glabra around Cedar Point. Callicarpa americana, Ilex vomitoria, Myrica cerifera, Serenoa repens, and Vaccinium spp. are common understory shrubs (or shrubby trees); Hamamelis virginiana is also ubiquitous around Fort Caroline. In the Roosevelt Area, Psychotia nervosa (a shrub in tropical areas) is a common understory plant but usually much less than 2 ft (0.6 m) tall. Woody vines are prevalent (i.e., Parthenocissus quinquefolia, Smilax spp., Vitis aestivalis, Vitis rotundifolia); common herbaceous understory plants include: Bidens bipinnata, Elephantopus nudatus, Galactia elliottii, Galium hispidulum, Melanthera nivea, Oplismenus hirtellus, Ruellia caroliniensis, Sanicula canadensis, Smallanthus uvedalia, Sporobolus indicus, and. Vernonia gigantea. Sandhill Community.—Sandhill is characterized by rolling hills of deep, relatively sterile sands inhabited usually by widely spaced oaks and pines with a ground cover of characteristic herbs (and some shrubs). Fire is a dominant factor in maintaining this community (FNAI 1990). Sandhill-type associations occur on two survey sites. The eastern and central portions of the Sohn Purchase is a large area of “scrubby sandhill” (Fig. 3A), comprising a relatively dense woody canopy—dominated by Quercus geminata with some Q. chapmanni, Pinus elliottii, and P. taeda—and various shrubs, including: Gaylussacia spp., Ilex vomitoria, Lyonia ferruginea, L. lucida, Myrica cerifera, Serenoa repens, and Vaccinium myrsinites. The sandhill scrub intergrades gradually with more mesic habitats (swamp to the south; marshland to the northwest). A restricted area (ca. 3.7 acres [1.5 hal), enclosed by the circular driveway of NPS headquarters (Mount Pleasant Road, Fig. 2), is typical longleaf pine-turkey oak sandhill (Pinus palustris- Quercus laevis) also dominated by Q. geminata (Fig. 3B); JD: O 1n £1 fl I mn J C. Ion ES £ A J OT — Area, Oct 2005 of Carphept ymt i ). C. Maritime I k (Fort Caroline National Memorial, Oct 2005)—overstory: Quercus virginiana with Vitis rotudifolia ] j ( ), Sanicul l j (l ft fi q 1), IS f (f g 1) Salt marsh ound Marn, Theodore Roosevelt Area, Oct 2005) tidal creek bounded by Spartina alterniflora (bright green) and J ianus (dark grayist g g ae n g e E I I. 4 Ẹ CL W ete | [D JAA AM. TL J D 1 Area. is 2005) —on yerstory J i rgi j (left f g 1) 1 E, ; greg (ba kg und) with Smil p : (rigt £ ); { Y F rq! II HA P het ta f I ry EY Hu | L ll £ P J Pe i I ++ ) ( ) g g 6. A esi Creek Parcel, Apr a) overstory ith expanding | (background): Acer rubrum, Liquid iflua, and Nyssa sylvatica v Ha Fr Y Dash att £, X and TL I PE I PR TO E £, A Photo pen W. B. Zomlefer. J M m” J IF J 1166 Journal of the Botanical R h Institute of Texas 1(2) the sparser shrub layer has most of the species present at the Sohn Purchase, with the notable addition of Ceratiola ericoides. Common understory sandhill plants of these two sites include Andropogon spp., Aristida spiciformis, Balduina angustifolia, Berlandiera pumila, Callisia graminea, Carphephorus odoratissimus, Crotalaria rotundifolia, Eragrostis elliottii, Eryngium aromaticum, Gratiola hispida, Lechea e Liatris Ern ia, Licania michauxii, Palafoxia integrifolia, Pityopsis graminifolia, Pteridium aquilinum, S 1riculata, E Stillingia sylvatica, and Stylisma patens. Aquatic Habitats.—This general habitat category designates areas of open freshwater, including the surrounding margin of often woody vegetation (FNAI 1990). The survey included several ponds in the Fort Caroline National Memorial/Theodore Roosevelt Area (including Spanish Pond; 4 acres [1.6 ha]); expanses of standing water of marshlands in the Sohn Purchase; and the water-filled borrow pit and dikes of Cedar Creek as well as higher waters bordering the swamp there. Floating aquatic species include: Eichhornia crassipes, Landoltia punctata, Lemna aequinoctialis, Limnobium spongia, Nymphaea odorata, Sagittaria filiformis, and Salvinia minima; common emergent wetland plants are: Cladium jamaicense, Eleocharis flavescens, Gratiola ramosa, i dpa mutilum n effusus, Ludwigia spp., Panicum hemitomon, P. verrucosum, Pluchea rosea, Polygonum spp., Pontederia cordata, Rhynchospora fascicularis, Sagittaria graminea, S. lancifolia, Setaria parviflora, and Spartina bakeri. Cephalanthus occidentalis, Diospyros virginiana, Ilex glabra, Lyonia fruitcosa, Pinus serotina, and Salix caroliniana are often components of the shrubby border of these wetlands. Swamp (Lowland Hardwood Forest).—This habitat occurs in low-lying, periodically flooded areas (often bordering stream banks); the soil is a mixture of clay and organic matter (Ewel 1990; FNAI 1990). Swamp forest is characterized as a closed-canopy of tall deciduous hardwoods with dense shrubby un- derstory. Ferns and herbs also occur in more open areas. The southeastern portion of Thomas Creek has substantial swamp (Fig. 3G) that intergrades with flatwoods; the mesic hardwoods of the southern Sohn property gradually transitions into scrubby sandhill. Mixed swamp-maritime hammock vegetation also surrounds Spanish Pond in the Fort Caroline National Memorial/Theodore Roosevelt Area. The following canopy trees are typical for Thomas Creek (and many also occur at the other swamp sites): Acer rubrum, Fraxinus caroliniana, Liquidambar styraciflua, Magnolia virginiana, Nyssa sylvatica var. biflora, Persea palustris, Platanus occidentalis, Quercus laurifolia, Q. nigra, Q. virginiana, and Ulmus americana. Woody understory species include: Carpinus caroliniana, Ilex cassine, Myrica cerifera, Smilax laurifolia, Rubus argutus, and Toxicodendron radicans. Examples of herbaceous plants that flourish under open areas of the canopy (often in standing water) include: Boehmeria cylindrica, Carex vulpinoidea, Chasmanthium laxum, Iris hexagona, Juncus effusus, Osmunda cinnamomea, Paspalum urvillei, Pontederia cordata, Rhynchospora spp., Saururus cernuus, Thelypteris kunthii, and Woodwardia areolata. Pine Flatwoods (Mesic Flatwoods; Pine Savannah).—These relatively flat areas are characterized by poorly drained terrain composed of up to 3 ft (ca. 1 m) of sand overlying organic/clayey hardpan that impedes water percolation, so that this habitat is frequently inundated. The overstory is an open canopy forest of widely spaced pines with little understory and a dense ground cover of shrubs and herbaceous plants (Abrahamson & Hartnett 1990; FNAI 1990). Like sandhill, pine flatwoods are also maintained by fire. Disturbed flatwood habitat is located in the central portion of the Thomas Creek parcel. This logged area, once a pine plantation, grades into the lower lying hardwood swamp to the south. Pinus taeda (loblolly pine) and P. elliottii (slash pine) are the dominant overstory species. Shrubs, scattered throughout the wetland and also along its margins, include: Bejaria racemosa, Cyrilla racemiflora (not vouchered), Ilex spp., Lyonia lucida, Lyonia ligustrina var. foliosiflora (not vouchered), Lyonia mariana, Myrica cerifera, Quercus elliottii, Q. minima, Q. myrtifolia, Rhus copallinum, Serenoa repens, and Vaccinium corymbosum. Shrubby and herbaceous species of Hypericum are common, e.g., H. brachyphyllum, H. fasciculatum H. gentianoides, H. hypericoides, H. mutilum, and H. tetrapetalum. The diverse herbaceous flora (many aquatic species in depressions) also = \ includes: Andropogon spp., Balduina uniflora, Bidens mitis, Bigelowia nudata, Carphephorus odoratissimus, E eocharis spp., eines o a Euthamia caroliniana, Fuirena breviseta, Helianthus angustifolius, Juncus "n effusus, I achnocaulon anceps, Liatris spp., Linum monum Tuc spp., Polygala lutea, P. C nana, Rhexia alifanus, iw n aris, dud brevifolia, Sarracenia minor, Scleria ciliata, Sorghastrum secundum, Typha latifolia, Utricularia subulata, and Xyris spp. Zomlefer et al., Flora of NPS areas of Timucuan Preserve 1167 Disturbed Areas (Ruderal Community). —Disturbed habitats have developed around public-access areas of Cedar Point, Kingsley Plantation, and especially within and around the reconstructed Fort Caroline and visitors' center and along the south side 1 Ft. uc: Road ae: Spanish PM Common weedy Ara] Amhy a artemiciifolia Rnevhav diffu E ? $ plants occurring in these areas include: ij a, Cerastium glomeratum, Chamaesyce spp., Dactyloctenium n Desmodium incanum, Digitaria Alans Diodia teres, Eleusine indica, Gamochaeta antillana, Lepidium virginicum, Paspalum notatum, Phyla nodiflora, Phyllanthus spp., Plantago virginica, Rubus trivialis, nigra, Sida rhombifolia, Sonchus asper, Stellaria media, and Triodanis perfoliata. (m 1 ANNOTATED CHECKLIST OF VASCULAR PLANT TAXA A list of 474 vascular plant species representing 318 genera in 122 families is here compiled from Giannasi & Zomlefer specimens (collection numbers in italic) in alphabetical order by family within four major groups (lycophytes, monilophytes [“ferns and allies”], gymnosperms, and angiosperms). Genera, species, and infraspecific taxa are alphabetical within each family. Scientific nomenclature and common names follow Wunderlin and Hansen (2003); exceptions are vernacular names of a few horticultural plants (not included in their flora) that conform to Huxley (1992). Family circumscriptions for lycophytes and monilophytes follow Smith et al. (2006); for gymnosperms, FNA (1993); and for angiosperms, APG (2003) Underlined taxa = new vouchered Duval County records according to on-line species list by Wunder- lin and Hansen (2007); * = exotic (Wunderlin & Hansen 2003); invasive exotics (FLEPPC 2007): [car 1] = Category I; [car u] = Category II; cuit = cultivated (i.e., planted on park grounds); [uni] = unicate speci- men (i.e., at GA, no duplicate at TIMU). Locality data: (1) = Thomas Creek Parcel; (2) = Cedar Point; (3) = Fort Caroline National Memorial/Theodore Roosevelt Area; (4) = Kingsley Plantation (Ft. George Island); (5) = Sohn Purchase. Habitat data: AQ = aquatic habitat (standing freshwater and surrounding margin); DA = disturbed area; pu = dunes/open beach; mn = maritime hammock; rr = pine flatwoods; sa = sandhill community; sH = shell midden; sm = salt marsh; sw = swamp. Relative abundance: c = common (generally abundant throughout a particular habitat; species easily found); o = occasional (locally common and/or several individuals distributed within a habitat; species not too difficult to locate); i = infrequent (sporadic occurrence of a small number of individuals; species relatively scarce and not easily found); r = rare (very few individuals encountered). LYCOPHYTES Osmundaceae Osmunda cinnamomea L., Cinnamon fern, (3)/(5), ^o/wH/sw, c-o, 1296, 1621 Osmunda regalis L. var. spectabilis (Willd.) A. Gray, Royal fern, Lycopodiaceae Lycopodiella alopecuroides (L.) Cranfill, Foxtail club-moss, (5), SA/sw, i, 1458 Da] MONILOPHYTES ("FERNS and allies") Blechnaceae Woodwardia areolata (L.) T. Moore, Netted chain fern, (3), 5 AO/MH/SW, C, 1 Woodwardia virginica (L.) Sm. Virginia chain fern, (1), Ao/pa, c, 1369 Dennstaedtiaceae iim aquilinum (L) Kuhn var. Cus (Clute e ex A. Heller, Bracken fern, (5), SA, C, — Lomariopsidaceae (Dryopteridaceae) *Nephrolepis exaltata (L.) Schott, Wild Boston fern, (3), DA 4 ! yogane n ONE Rn jodium um (Thunb.) Sw., Japanese climbing fern, J ap CAT I], ro C, 1484 (3), = (3), MH, C, 1719 Polypodiaceae on a (L.) E.G. Andrews & Windham var. ichau a (Weath.) E.G. Andrews € Windham, Resur- rection Pun (3), MH, C, 1473 Salviniaceae PP minima Baker, Water spangles, (1), ^o/ba, i, 1710 [UNI] lypteridaceae Pasión kunthii (Desv.) S.V. Morton, Widespread maiden fern, (1), sw, c, 1490 GYMNOSPERMS Cupressaceae Jul perus virgil Nana Es: Red cedar, (1 (3), AO/ DA/ MH, CO; 1233, 1364 Cycadaceae *Cycas revoluta Thunb., Sago palm, (3), cuLT, 1562 1168 Pinaceae Pinus elliottii Engelm., Slash pine, (5), sa, o, 1640 Pinus palustris Mill, Longleaf pine, (3), sa, c, 1303 Pinus serotina Michx., Pond pine, (1), aQ/Da, i, 1362 Pinus taeda L., Loblolly pine, (5), sa/sw, o, 1641 Podocarpaceae *Podocarpus dec (Thunb) D. Don, Japanese podo- berry, (3), CULT, ANGIOSPERMS Acanthaceae Elytraria caroliniensis (J.F. Gmel.) Pers. var. caroliniensis, Carolina scalystem, (1), sw, O, *Justicia brandegeana Wassh. & L.B r, 1268 [Escaped from cultivation and persisting; may be Sm., Shrimpplant, (3), Da, naturalized.] Ruellia PET (J.F. Gmel) Steud., Carolina wild petunia, , MH, C, 1386 *Odontonema cuspidatum (Nees) Kuntze, Firespike, (3), cuLt, 1725 xaceae AM nigra L. subsp. canadensis (L.) R. Bolli, American elder, (3), Da, o, 1280 *Viburnum odoratissimum Ker-Gawl., T, 1270, 1722 Sweet viburnum, (3), Agavaceae Yucca aloifolia L., Spanish bayonet, (3), cuir, 1592 Aizoaceae Sesuvium portulacastrum (L.) L., Shoreline seapurslane, (4), Su *Tetragonia tetragonioides (Pall) Kuntze, New Zealand spinach (4), DU/SM, C, 1529 a ao SL E E Sm, TI Jj E | | (5) Qi 1652 Sagittaria nd Michx. var. graminea, Grassy SOUS (5 ), AQ Q, O, Sagittaria peus L. var. lancifolia, Bulltongue arrowhead, , AQ/DA, O, 1 aceae Allium canadense L. var. canadense, Meadow garlic, (1), DA/sw, r, 1315 Nothoscordum bivalve (L.) Britton, Crowpoison, (4), Da, r, 1376 [UNI] Altingia iub dd uud EE Sweetgum, (1), sw, c, 1492 Amaranthaceae *Alternanthera philoxeroides (Mart.) Griseb., Alligatorweed, (1), AT Il], AQ/DA, i, 1716 [UNI] Atriplex cristata Humb. & Bonpl. ex Willd., Crested saltbush, , SM, O, 15. *Chenopodium album L., Lamb's-quarters, (2), sm, o, 1536 *Chenopodium ambrosioides L., Mexican tea, (4), DU/SM, C, 1670 ~ Rootstock bloodleaf, (3), sh, c, Iresine rhizomatosa Standl., L737 £L Dag PRA D hi Air Journal of f Texas 1(2) Sarcocornia perennis (Mill.) A.J. Scott, Perennial glasswort, (4), DU/SM, C, bum den Mill: dne m of tl Salicornia ma render Sarcocornia paraphyletic (Kandereit et al. Sa and Salicornia s.l. (including both perennial and annual pee is likely hyletic on the ud cU nd pubescent membranous testa (Judd & Ferguson 1 999).] Elliott) Moq., Sea blite, (2), sm, O, 1537 Amaryllidaceae *Crinum asiaticum L., Poisonbulb, (4), cur, 1672 Anacardiaceae hus copallinum L, Winged sumac, (1)/(5), DA/PE/sa/sw, c-o, 1636, 1695 Toxicodendron radicans (L.) Kuntze, Eastern poison ivy, (3), DA/MH/SW, C, 1726 Annonaceae Asimina incana (W. Bartram) Exell, Woolly pawpaw, (5), Sa, o, 1203 Asimina parviflora (Michx.) Dunal, Smallflower pawpaw, (3), H, o, 1390 Asimina pygmea (W. Bartram) Dunal, Dwarf pawpaw, (1), AO/DA, C, 1325 Apiace E asiatica (L) Urb., Spadeleaf, (3), DA, C, 1419 Cicuta maculata L., Spotted water hemlock, (1), AQ/DA, r, *Cyclospermum leptophyllum (Pers) Sprague ex Britton & P. Wilson, Marsh parsley, (1), Da/sw, i, 1307 Eryngium aoa Baldwin, Fragrant eryngo, (3), Sa, O, 17632. Eryngium baldwinii Spreng., Baldwin's eryngo, (5), DA/SA, O, 1469 Oxypolis filiformis (Walter) Britton subsp. filiformis, Water cowbane, (1), Ao/Pr, r, 1682 Sanicula canadensis L., Canadian blacksnakeroot, (3), MH, C, Spermolepis divaricata (Walter) Raf, Roughfruit scaleseed, (3), DA, C, 1266 cynaceae Asclepias tomentosa Elliott, Velvetleaf milkweed, (3), DA/MH, r, 1560 [uni] ynanchum angustifolium Pers., Gulf coast swallowwort, (2), SM, O, 154 Cynanchum scoparium Nutt., Leafless swallowwort, (3), su, G1738 Aquifoliaceae llex ambigua (Michx.) Torr. var. ambigua, Carolina holly, (3), MH, O, 3, Ilex cassine L. var. cassine, Dahoon, (5), sA/sw, r, 1634 Ilex cassine = var. ue (Walter) Sarg., Myrtle dahoon, (1), DA/PF, llex ic: Al ‘ ^ Gray, Inkberry, (1)/(3), Aa/DA/MH/PF, o, 1294, 1328, 1366 llex opaca Aiton var. opaca, American holly, (1), DA/PF, o-r, 1337 Zomlefer et al., Flora of NPS areas of Timucuan Preserve vomitoria Aiton, Yaupon, (1), AQ/DA, C, 1357 Araceae *Landoltia punctata (G. Mey.) Les & D. J. Crawford, Dotted duckweed, (1), a0/Da, c, 1382 Lemna aequinoctialis Welw., Lesser duckweed, (5) 1209 , AQ, C, raliaceae js MM Devil's walkingstick, » ), MH, C, 1566 e a English din i CULT, 1269 x Lam., Largeleaf marshpen- Teu nywort, (3), DA/DU, C, 2 Hydrocotyle umbellata L., Manyflower marshpennywort, (3), A, O, 1 Arecaceae Sabal palmetto (Walter) | MH, C, 1439 Serenoa repens (W. Bartram) Small, Saw palmetto, (3), MH, C, 1438 dd. ex Schult. & Schult. f, Cabbage pe ay Aristolochiaceae Aristolochia serpentaria L., Virginia snakeroot, (3), wu, r, 1565 [UNI] Asparagaceae *Asparagus aethiopicus L., Sprenger's asparagus-fern, (4), [cat 1], CULT, 1374 Astera finis e. L., Common ragweed, (2), DA/SM, C, 1539 "us halimifolia L., Groundsel tree, (3), sm, c, 160 Balduin e ien B.L. Rob., Coastalplain honey- head, (3), sa, r, 173 mu uniflora Nutt., Oneflower honeycombhead, (1), rr, o, 1688 Berlandiera pumila (Michx.) Nutt., Soft greeneyes, (3), sa, C, 1298 Bidens alba (L) DC., Beggarticks, (3), DA/MH, O, ao Bidens bipinnata L., Spanish needles, (3), M e mitis (Michx.) Sherff, Smallfruit aA ee (1), DA/PF, 1701 j : MA) DC. subsp. nudata, Pineland rayless goldenrod, (1), PF, c, 1687 Borrichia frutescens (L) DC., Bushy seaside oxeye, (3), DU/DA, C, 1478 Carphephorus corymbosus (Nutt) Torr. & A. Gray, Coastalplain chaffhead, (3), sa, c, 1734 Carphephorus odoratissimus (J.F. Gmel.) H. Herbert var. odora- a rd ), DA/PF, O, 1696 Cirsium x. Purple thistle, (1), aq/Da, O, 1370 Cirsium nuttallii DC., er thistle, (1), Da/sw, o, 1498 noclinium coelestinum (L.) DC., Blue mistflower, (3), DA/MH, o, 1480 Conyza canadensis (L.) p var. canadensis, Canadian ed, (3), MH, i, 157. un basalis (A. A e F. Blake, Goldenmane tickseed, TAT UNI] Eclipta ade (L) L., False daisy, (3), ^o/Mu/sw, i, 1431 Elephantopus elatus Bertol., Tall elephantsfoot, (5), sa, c, 1655 1169 Elephantopus nudatus A. Gray, Smooth elephantsfoot, (3)/(5 a SA, €, 1579, 1609 Erechtites hieraciifolius (L.) Raf. ex DC., Fireweed, (5), ao, i, 1447 Erigeron quercifolius Poir., Oakleaf fleabane, (3), DA/MH, c, 1218 Erigeron vernus (L.) Torr. & A. Gray, Early whitetop fleabane, (1), DA/PE, r, 1706 Eupatorium album L.. White thoroughwort, (3), sa, r, 1731 UNI Eupatorium capillifolium (Larn.) Small ex Porter & Britton, gfennel, (5), sa, o, 1615 Eupatorium leptophyllum DC., Falsefennel, (5), sa, o, 1624 Eupatorium mohrii Greene, Mohr's thoroughwort, (1), DA/PE, ETATS. Fupatorium rotundifolium L., Roundleaf thoroughwort, (5), Sw, O, 1465, 1466 Euthamia caroliniana (L. Greene ex Porter & Britton, Slender flarr IA p. flatt Idenrod, PF, C, 16 Gaillardia pulchella Foug, Firewheel, (3), pA/cu.??, c, 1477 [Likely lanted with associated Lantana spp. near dock] Gamochaeta antillana (Urban) Anderberg, Caribbean purple everlasting, (3), Da, c, 1241 [Gamochaeta falcata (Lam.) abrera, misapplied, as listed in Wunderlin and Hansen (2003); see Nesom (20063) and Wunderlin and Hansen (2007).] Gamochaeta pensylvanica ns Cabrera, Pennsylvania everlasting, (3), DA/MH, O, 1237 Helianthus angustifolius L., Narrowleaf sunflower, (1), DA/PF, C, 1694 Í iff H (lam) Britt rn 2 Riich amphorweed, 4), DA, C, 1674 *Hypochaeris brasiliensis (Less.) Hook. & Arn. var. tweedii (Hook. rn.) Baker, Tweed's catsear, (3), DA, i, 1415 Iva ee ee sumpweed, H ), sm, C, 1599 Iva microcephala Nutt., Piedmont marshelder, (5), Ao, o, 1649 Krigia virginica (L e Virginia e (1), Da, O, 1329 es (Greene) K.Schum leaf gayfeather, (1), DA/PF, 1702 [= eu graminifolia Willd. var. d inW lansen ( UU), see NO (2006b) e Wunderlin and Hansen (2007).] Liatris spicata (L.) Willd., Dense gayfeather, (1), PF, c, 1685 Liatris tenuifolia Nutt. var. quadriflora Chapm., Shortleaf gay- feather, (5), sa, c, 1605 [Listed in Wunderlin and Hansen (2003) as endemic to Florida, but this variety has been recently vouchered from Cumberland Island, Georgia mlefer etal. in prep.).] Liatris tenuifolia Nutt. var. tenuifolia, Shortleaf gayfeather, (3), — SA, C, 1735 Lygodesmia aphylla (Nutt.) DC., Rose-rush, (3), sa, r, 1300 Melanthera nivea (L.) Small, Snow squarestem, (3), MH, C, 1476 Mikania scandens (L.) Willd., Climbing hempvine, (3), MH, C, 1717 Palafoxia integrifolia (Nutt) Torr. & A. Gray, Coastalplain palafox, (3), sa, i, 1729 Pityopsis graminifolia (Michx.) Nutt., Narrowleaf silkgrass, (3)/(5), Sa, C, 1607, 17. 1170 ron foetida (L) DC., Stinking camphorweed, (1), DA/PF, 693 pue K. Godfrey, Rosy camphorweed, (5), Ao/sA/sw, o-i, 1443, 1467 Pterocaulon pycnostachyum (Michx.) Elliott, Blackroot, (5), SA/SW, O, 1 Pyrri linianus (Walter) DC, Carolina desertchi a ), DA, O, 7 Src tortifolius (Michx.) Nees, Whitetop aster, (3), Sa, i, NU uvedalia (L) Mack. ex Small, Hairy leafcup, Q), A/MH, C, 1534 Saidaga oe Mill, Pinebarren goldenrod, (1), DA/PF, C, a odora Aiton var. chapmanii (A. Gray) Cronquist, 6 Solidago sempervirens L., Seaside goldenrod, (2), sm, o, 1657 *Sonchus asper (L) Hill, Spiny sowthistle, (3), Da, i, 1414 *Sonchus oleraceus L., Common sowthistle, (3), DA/MH, O, 1239 Symphyotrichum carolinianum (Walter) Wunderlin € B. F. Hansen, Climbing aster, (1), aQ/Da, r, 1715 Symphyotrichum tenuifolium (L.) G. L. Nesom, Perennial salt- marsh aster, (1), Ao/DA, r, 1711 id officinale weber ex F. H. Wigg., Common dande- lion, (3), DA/MH, O, 1 "s. Mea E Peer] (2)/(4), DU/MH, o-r, 1664 [UNI], 1671 Vernonia gigantea (Walter) Trel. ex Branner & Coville, Giant ironweed, (3), DA/MH, i, 1393 DC., Oriental false hawksbeard, (3 Es wo *Youngia japonica (L A/MH, C, 1 Bataceae Batis maritima L. Saltwort, (4), DU/sM, C, 1524 Betulaceae Carpinus caroliniana Walter, American hornbeam, (1), sw, C, 1337, 1489 Bignoniaceae Bignonia capreolata L., Crossvine, (3), DA/MH, i, 1285 Campsis radicans (L) Seemann, Trumpet creeper, (3), DA, O, 1424 *Macfadyena unguis-cati (L.) A. H. Gentry, Catclawvine, (4), [CAT 1], DA, O, 1373 Brassi RUE m L., Hairy bittercress, (3), Ao/MH/Ssw, o, 1290 Descurainia pinnata (Walter) Britton, Western tansymustard, (3), DA, C, 1265 Lepidium virginicum L., Virginia pepperweed, (3), DA, i, 1231 Bromeliaceae Tillandsia recurvata (L.) L, Ballmoss, (3), DA/MH, C, 1252 Tillandsia usneoides (L.) L, Spanish moss, (1), DA/PF, c, 1354 — Cactaceae puntia pusilla (Haw.) Haw., Cockspur pricklypear, (3), sh, c, 1385 Opuntia stricta (Haw.) Haw., Erect pricklypear, (2), sm, o, 1549 Ll Dat PRA D hi PEA Journal of f Texas 1(2) Campanulaceae Lobelia glandulosa Walter, Glade lobelia, (1), Ao/Pr, r, 1683 Triodanis perfoliata (L.) Nieuwl., Clasping Venus lookingglass, (3), Da, C, 1264 *Wahlenbergia marginata (Thunb.) A. DC., Southern rockbell, (1), DA, C, 1330 Cannabaceae Celtis laevigata Willd., Hackberry, (4), Du, i, 1526 Caprifoliaceae *Lonicera japonica Thunb., Japanese honeysuckle, (4), [cat 1], CULT, 1377 Lonicera sempervirens L, Coral honeysuckle, (3), DA, O, 1272 Caryophyllaceae *Cerastium glomeratum Thuill, Mouse-ear chickweed, (3), DA, C, 121 Silene antirrhina L., Sleepy catchfly, (3), Da, i, 1276 Spergularia marina (L.) Griseb., Salt sandspurry, (4), DA, C, 1378 *Stellaria media (L.) Vill., Common chickweed, (3), Da, c, 1213 Celastraceae uonymus americanus L., American strawberrybush, (3), MH, C, 1582 Chrysobalanaceae Licania michauxii Prance, Gopher apple, (3), sa, C, 1550 Cistaceae Lechea torreyi (Chapm.) Legg. ex Britton, Piedmont pinweed, (5) SA GST Commelinaceae Callisia graminea (Small) G. C. Tucker, Grassleaf roseling, (3), SA, O, 1551 *Commelina caroliniana Walter, Carolina dayflower, (3), Q/MH/SW, O, 128 ie erecta L, Whitemouth dayflower, (3), Da, c, 1401 s pellucida (M. Martens & Galeotti) D. R. Hunt, Tahitian bridalveil, (4), Da, o, 1372 [Escaped from cultivation; may be locally naturalized.] Tradescantia ohiensis Raf., Bluejacket, (3), Da, o, 1271 Convol ceae DT carolinensis Michx., Carolina ponysfoot, (3), DA, o, 1256 Ipomoea alba L., Moonflower, (3), DA/Du, o, 1593 Ipomoea cordatotriloba Dennst., Tievine, (3), MH, o, 1580 Ipomoea pandurata (L.) G. Mey., Man-of-the-earth, (2), 0A/MH, 159 Ipomoea sagittata Poir., Saltmarsh morning-glory, (1), AQ/DA, SS Stylisma patens (Desr.) Myint, Coastalplain dawnflower, (3), A, O, 1556 Cor jm BN Marshall var. biflora (Walter) Sarg., Swamp tupelo, (1), Ao/bA/sw, C—O, 1338, 1356, 1367 Cucurbitac Melo dcn NE Creeping cucumber, (2)/(3), DA/MH, O-r, 1661 [UN], 1720 Zomlefer et al., Flora of NPS areas of Timucuan Preserve Cyperaceae Carexalbicans Willd. ex c var. australis (L. H. Bailey) Rettig, Whitetinge sedge, (3), DA, r, 1413 Carex PR . H. Bailey RT capillacea (L. H. Bailey) eznicek, Prickly bog sedge, (1), sw, i, 1311 Carex glaucescens Elliott, Clustered sedge, (1), AQ/DA/PF, O, 1516 Carex striata Michx., Walter's sedge, (1), DA/PF, o, 1351 Carex vulpinoidea Michx., Fox sedge, (1), DA/sw, o, 1318 ladium jamaicense Crantz, Jamaica swamp sawgrass, (1), Q/DA, C, 1544 Cyperus croceus Vahl, Baldwin's flatsedge, (3), Da, o-i, 1395, 1403 *Cyperus esculentus L., Yellow nutgrass, (3), Aa/MH/sw, O, 1434 Cyperus odoratus L., Fragrant flatsedge, (3), MH, o, 1585 diia retrorsus Chapm., Pinebarren flatsedge, (5), sa/sw, O, 614, 1620 mn baldwinii (Torr.) Chapm., Baldwin's spikerush, (1), /PF, i, 1704 [UNI] Eleocharis flavescens (Poir) Urb., Yellow spikerush, (1), AQ/DA, O, 1368 ink, Viviparot IS spikert ish, (5), A 1653 oe non (L) Roem. & Schult, o fimbry, (5), na o, 1618 Full ta (Coville) Coville, Saltmarsh umbrellasedge, C », AOPE, C , 1684 Fi hx, Soutl brellasedge, (5), Ao/sA, o-i, 1200, 1204 Rhynchospora caduca Elliott, Anglestem beaksedge, (1), AQ/SW, O, 1547 Rhynchospora colorata (L.) H. Pfeiff, Starrush whitetop, (1), A O, 1360 Rhynchospora fascicularis (Michx.) Vahl, Fascicled beaksedge, (1)/(3)/(5), Aa/DA/MH/PF/sw, C—O, 1355, 1435, 1444, 1512, 1546 Rhynchospora inundata (Oakes) Fernald, Narrowfruit horned beaksedge, (1)/(5), ^o/sw, c-o, 1548, 1647 Rhynchospora megalocarpa A. Gray, Sandyfield beaksedge, (5), sa/sw, i, 1459 Rhynchospora microcephala (Britton) Britton ex Small, Bunched beaksedge, (1), sw, i, 1545 Rhynchospora plumosa Elliott, Plumed beaksedge, (1)/(3), DA/PF/5A, i, 1297, 1326 Scleria ciliata Michx. var. ciliata, Fringed nutrush, (1)/(5), DA/PF/sA, r, 1191, 1384 Scleria oligantha Michx., Littlehead nutrush, (1), Da, c, 1324 Scleria reticularis Michx. Netted nutrush, (5), AQ, O, 1441 Dioscoreaceae *Dioscorea bulbifera L., Air-potato, (3), [car 1], DA, C, 1426 Droseraceae Drosera intermedia Hayne, Water sundew, (5), ao, O, 1207 Ebenaceae Diospyros virginiana L., Common persimmon, (5), AQ, O, 1440 Ericaceae Bejaria racemosa Vent, Tarflower, (1), DA/PF, o, 1345 Ceratiola ericoides Michx. Florida rosemary, (3), sa, O, 1727 1171 fy ni Gaylussacia d ber ery. ( sh o nana i Gray) Small, Glaucous huckleberry, (5), sa 98 [= x i MOROA ) Torr. & ds d ex on var. tomentosa A. Gra oy a nana is o E tomentosa and from a (J. Kenn.) Torr. & A. Gray, Dwarf huckle- Wu der WEAS d Hans ; diagnosable, and differ in chromosome number. Gaylussacia frondosa is also morphologically distinct and occurs disjunctly in Florida only in Santa Rosa County; see Duncan & Brittain (1966) and Luteyn et al. (1996).] Gaylussacia tomentosa (A. Gray) Small, Southern huckleberry (5), sa, o, 1201 [= Gaylussacia frondosa (L.) Torr. & A. Gray ex Torr. var. tomentosa A. Gray, as listed in Wunderlin and Hd (2003). See annotation above py G. nana. Kalmia hirsuta Walter, Hairy laurel, (5), sa Lyonia ei (Walter) Nutt., Rusty o (5), s Dons Hm (Michx.) G. S. Torr, Coastalplain staggerbush, DA, C, 1323 Lyonia mariana (L) " Don, Piedmont staggerbush, (3)/(5 AQ/MH/SA, i, 1287, 1650 Lyonia Meda (BEN ^ Koch, Fetterbush, (5), sa, c, 1197 Rhododendron canescens (Michx.) Sweet, Mountain azalea, (1), DA/PF, O, 1212 een simsii Planch., Formosa azalea, (3), cuv, 1302 Vaccinium arboreum Marshall, Sparkleberry, (3), DA/MH, O, Mu we 1286 Vaccinium corymbosum L., Highbush blueberry, (1)/(3), a0/DA/ MH/PF, C-i, 1210, 1234, 1321 Vaccinium myrsinites Lam., Shiny blueberry, (5), sa, c, 1189 Vaccinium stamineum L., Deerberry, (3), MH, o, 1292 Eriocaulaceae Eriocaulon decangulare L., Tenangle pipewort, (1), DA/PF, C, I achn aron (Walter) Morong, Whitehead bogbut- ton, (1), Ao/bA/Pr, o-r, 1343, 1507 Euphorbia Acalypha gal: A. Gray, Slender threeseed mercury, (3), o e (L.) Millsp., Pillpod sandmat, (3), Da, o, 1402 Chamaesyce hyssopifolia (L) Small, Hyssopleaf sandmat, (3), 5 Chamaesyce maculata (L) Small, Spotted sandrnat, (3), DA, r, 1262 [UNI Cnidoscolus stimulosus (Michx.) Engelm. & A. Gray, Tread softly, (3), sa, O, 1299 Croton ye L. var. glandulosus, Vente conmigo, (3), Poinsettia on (Murray) Bartl., Paintedleaf, (3), DA/MH, i, 1238 m heterophylla (L.) Klotzsch & Garcke ex Klotzsch, Fiddler's spurge, (3), Da, o, 1417 *Sapium sebiferum (L) Roxb., Popcorntree, (1), [car 1], AQ/DA, E 1 1172 Stillingia sylvatica L., Queensdelight, (3), sa, i, 1552 Tragia urens L., Wavyleaf noseburn, (3), MH, o, 1564 Fabaceae Amorpha fruticosa L, Bastard false indigobush, (1), a0/DA, i, 1521 Amphicarpaea bracteata (L.) Fernald, American hogpeanut, (3), MH, O, 1588 Centrosema virginianum (L.) Benth., Spurred butterfly pea, (4), DA, O, 1531 Cercis canadensis L., Eastern redbud, (3), cuLT, 1563 Chamaecrista fasciculata (Michx.) Greene, Partridge pea, (4 A DA, O, 16 Chamaecrista nictitans (L) Moench var. aspera (Muhl. ex Elliott) H. S. Irwin & Barneby, Sensitive pea, (5), sa/sw, o, 1608 Chamaecrista nictitans (L) Moench var. nictitans, Sensitive pea, (1), o^/sw, c, 1708 Crotalaria rotundifolia J.F. Gmel., Rabbitbells, (3), sa, o, 1301 Desmodium glabellum (Michx.) DC., Dillenius' ticktrefoil, (3 591 *Desmodium incanum DC., Zarzabacoa comun, (3), DA, C, X_ 00 Desmodium paniculatum (L.) DC., Panicled ticktrefoil, (2), A/SM, O, 1656 Erythrina herbacea L., Coralbean, (3), sa, €, 1559 Galactia an cdas milkpea, (3), DA/MH, C, 1423 n, Downy milkpea, (3), SH, o, 1569 *Indigofera E ES ae ), DA/PF, O, 1698 *Indigofera tinctoria L., True indigo,(4), cuLt, 1676 Lupinus diffusus Nutt., Skyblue lupine, (5), sa, r, 1612 *Medicago lupulina L., Black medick, (3), p4, o, 1250 *Melilotus indicus (L) All., Indian sweetclover, (3), DA, o, 1220 Sesbania vesicaria (Jacq) Elliott, Bladderpod, (1), sw, o, 1689 Strophostyles helvola (L.) Elliott, Trailing fuzzybean, (3), DA/DU, I Galactia volubi lis is (L) C, 1479 *Trifolium repens L., White clover, (3), DA, C, 1217 *Wisteria sinensis (Sims) Sweet, Chinese wisteria, (4), [CAT 11 DA, C, 15. == Fagaceae Quercus chapmanii Sarg, Chapman's oak, (5), sa, O, 1616 Quercus elliottii Wilbur, Running oak, (1), DA/PE, o, 1327 Quercus geminata Small, Sand live oak, (5), sa, c, 1199 Quercus hemisphaerica Bartr. ex Willd., Darlington oak, (4), DA/MH, i, 1666 [Included as a synonym of Q. laurifolia Michx. in Wunderlin and Hansen (2003); see Jensen (1997), who treats these as two species. In our area they are reproduc- tively isolated by ecology and phenology.] Quercus laevis Walter, Turkey oak, (3), sa, c, 1736b Quercus laurifolia Michx., Laurel oak, (1)/(5), Da/PE/sw, O, 1347, 1631 [Considered here as Q. laurifolia s.s. sensu Jensen (1997); see annotation under 0) hemisphaerica above.] Quercus minima (Sarg.) Small, Dwarf live oak, (1), DA/PF, O, 83 bai E Myrtle oak, (1), pA/r, i, 1348 Quercus nigra L., Water oak, (1), Da/PF, c, 1500 dm virginiana m Live oak, (1), AQ/DA, O, 1363 £L Dat PRA D hi PEA Journal UI f Texas 1(2) Gelsemiaceae Gelsemium sempervirens (L.) W.T. Aiton, Yellow jessamine, (5), SA/SW, O, 1635 Gentianaceae Sabatia brevifolia Raf., Shortleaf rosegentian, (1), DA/PF, o, 1707 Sabatia stellaris Pursh, Rose-of-plymouth, (5), DA/sa, o, 1470 Geraniaceae . nm L., Carolina cranesbill, (3), Da, c, 1249 aemodora pande a (Lam.) Dandy, Carolina redroot, (1), AO/DA/PF, C, 150 Haloragace o pem Lam., Combleaf mermaidweed, (1), AQ/DA/PF, C, 1349 Hamamelidaceae Hamamelis virginiana L., American witchhazel, (3), MH, C, Hydrocharitaceae jin spongia (Bosc) Rich. ex Steud., Frog's-bit, (3), AQ, i, 1284 Hypericaceae Hypericum brachyphyllum (Spach) Steud., Coastalplain St. John's-wort, (1), per, c, 1508 Hypericum cistifolium Lam., Roundpod St. John's-wort, (5), AQ/SA/SW, O, 1456 Hypericum fasciculatum Lam., Peelbark St. Johr's-wort, (1), AO/DA/PF, O, 1211 Hypericum gentianoides (L.) Britton et al., Pineweeds, (1), DA/PF, C, 1505 Hypericum hypericoides (L a St. Andrew's-cross, (1)/(3), DA/PF/sA/sw, O-i, 1317, 1346, 1728 Hypericum mutilum L., Dort St. John's-wort, (1 1503 Ww DA/PF, O, Hypericum myrtifolium Lam., Myrtleleaf St. John's-wort, (5), AQ, C, 1445 Hypericum reductum (Svenson) W. P. Adams, Atlantic St. John's- ort, (5), SA, C, 145i a jd d Lam., Fourpetal St. John's-wort, (1)/(5), t^ E192, 1193, 1 Rin M (L) Raf., Virginia marsh St. John's-wort, (5), ^o/sw, i, 1627 Hypoxidaceae Hypoxis curtissii Rose, Common yellow stargrass, (1), sw, O, B12 Illiciaceae lllicium parviflorum Michx. ex Vent., Star anise, (3), cuLT, 1561 Iridaceae Iris he agona Walter, Dixie i iris, (1 jr SW C, 1304 Sisyrinchium angustifolium Mill., Narrowleaf blue-eyed grass, (3), DA, O, 1247 ti E. P. Bicknell, Eastern blue-eyed grass, (1), Da/sw, c, 1306 [Included as a synonym of S. angustifo- lium s.l. in Wunderlin and Hansen (2003); see Cholewa and Zomlefer et al., Flora of NPS areas of Timucuan Preserve Henderson Ame who treat these as two species based on h and foliage color] E rosulatum E. P. Bicknell, Annual blue-eyed grass, (3), DA, i, 1223 Juglandaceae Carya glabra (Mill) Sweet, Pignut hickory, (3), DA/MH, o, 1398 Juncaceae Juncus acuminatus Michx., Tapertip rush, (5), ^o/sA/sw, O, 1460 Juncus coriaceus Mack, Leathery rush, (1), oA/sw, C Juncus dichotomus Elliott, Forked rush, (1), Da/sw, c, 1319 Juncus effusus L. subsp. solutus P MN Hámet- ti, Soft rush, (1), aQ/DA/PE, c, 1334, Juncus elliottii Chapm., Bog rush, (1), PAR c-o, 1332, 1358 Juncus marginatus Rostk., Shore rush, (5), ^o/sA/sw, i, 1464 e x P A. Curtis, Bighead rush, (1), AQ/DA, ne om Michx., Manyhead rush, (5), AQ, C, 1446 Juncus roemerianus Scheele, Black rush, (3), sm, c, 1571 Juncus scirpoides Lam., Needlepod rush, (5), Aa/sw, o, 1625 2 Lamia caia c americana L., American beautyberry, (3), DA/MH, "pts L (Rich.) Briq., Tropical bushmint, (3), Da, C, 594 Era rubellus Moench, Taperleaf Waterhorehound, (1), o, 1690 narda punctata L., Spotted beebalm, (4), DA, O, 1675 nnn erm à (W. Bartram ex Benth.) Raf, Wild pennyroyal, (5 ), SA A, f, 194 Salvia lyrata if Lyreleaf sage, (3), DA/MH, C, 1245 Scutellaria integrifolia L, Helmet skullcap, (1), a0/Da, O, 1517 Stachys floridana JL ex Benth., Florida hedgenettle, = + M + + Trichostema ee L., Forked bluecurls, (1), PF, o, 1681 Lauraceae *Cinnamomum camphora (L.) J. Presl, Camphortree, (1), [car i], sw, i, 1493 *Persea americana Mill., Avocado, (4), cut, 1530 Persea e (L.) Spreng. var. borbonia, Red bay, (2)/(3), E G P 2,1 3 Persea ue (Raf) Sarg, Swamp bay, (1), ao/Da, i, 1320 entibulariac a prine L., Zigzag bladderwort, (1), oA/F, o, 1336 inaceae Linum medium (Planch.) Britton var. texanum (Plat c Ve) Fert ald, Stiff yellow flax, (1), DA/PF, c, 1502, 1504 Lythraceae *Lagerstroemia indica L., Crapemyrtle, (3), cut, 1573 Magnoliaceae oe grandiflora L., Southern magnolia, (3), MH, o, 1587 Magnolia virginiana L., Sweetbay, (5), sw, i, 1638 Malvaceae *Gossypium barbadense L., Sea island cotton, (4), cur, 1677 1173 Kosteletzkya virginica (L.) C. Presl ex A. Gray, Virginia saltmarsh mallow, i ) pone A lalvavi reus Cav. var. drummondii (Torr. & A. Gray) MM waxmallow, (4), cult, 1527 Sida acuta Burm. f, Common wireweed, (4), Du, o, 1680 Sida rhombifolia L., Cuban jute, (1)/(3), DA, C-0, 1427, 1495 No * => Melastomataceae Rhexia alifanus Walter, S | (1), Ao/DA/PF, o, 1510 Rhexia mariana L., Pale meadowbeauty, (5), sa/sw, i, 1453 Rhexia petiolata Walter, Fringed meadowbeauty, (5), sa/sw, Menispermaceae Cocculus carolinus (L.) DC., Carolina coralbead, (2 1533 — DA/MH, C, Moraceae *Fatoua villosa (Thunb.) Nakai, Hairy crabweed, (3), MH, O, 1589 *Ficus pumila L., Climbing fig, (3), cur, 1273 Morus rubra L., Red mulberry, (3), DA/MH, O, 1255 _ Myricaceae Myrica cerifera L, Wax myrtle, (5), SA, C, 1 — 95 Nyctaginaceae Boerhavia diffusa L., Red spiderling, (3), Da, o, 1405 Nymphae Nymphaea iem Sol., American white waterlily, (5), Ao, i, 1651 Oleaceae Forestiera segregata (Jacq) Krug & Urb., Florida swampprivet, (3), SH, c, 1570 Fraxinus caroliniana Mill., Water ash, (1), sw, o, 1491 Edi um japonicum Thunb., Japanese privet, (4), CULT, 1523 nanthus americanus (L) Benth. & Hook. f. ex A. Gray, Wild e (3), SA, i, Z Onagraceae Gaura angustifolia Michx., Southern beeblossum, (5), DA/SA, Ww , 1468 Ludwigia linearis Walter, Narrowleaf primerosewillow, (1 F, O, Ludwigia linifolia Poir., Southeastern primrosewillow, (1), /PF, C, 1511 Nod maritima R. M. Harper, Seaside primrosewillow, (5), 54 , O, Yi F microcarpa Michx., Smallfruit primrosewillow, (5 A, O, *Ludwigia peruviana (L.) H. Hara, Peruvian primrosewillow, 3), ^o/MH/SW, O, 1 Ludwigia pilosa Walter, Hairy primrosewillow, (1), DA/PF, C, co 1699 Ludwigia sphaerocarpa Elliott, Globefruit primrosewillow, (5), AQ/Ssw, O, 1628 Ludwigia suffruticosa Walter, Shrubby primrosewillow, (5), AQ, C, 1648 Oenothera laciniata Hill, Cutleaf eveningprimrose, (3)/(5), DA/DU/SA, O, 1186, 1222, 1225 1174 Orobanchaceae dd fasciculata (Elliott) Raf., Beach false foxglove, (5), , O, m setacea (J.F. Gmel.) Raf, Threadleaf false foxglove, (5), sa, i, 161 Seymeria pectinata Pursh, Piedmont blacksenna, (5), SA, O, 1606, 1626 Oxalidaceae Oxalis corniculata L, Common yellow woodsorrel, (3), DA, i, 1228 *Oxalis debilis Kunth, Pink woodsorrel, (3), Da, C, 1248 Passifloraceae Passiflora incarnata L., Purple passionflower, (3), DA, i, Passiflora lutea L., Yellow passionflower, (3), MH, i, 1583 1430 Phyllant ae i s pure Baill, Drummond's leafflower, (3), DA ET tenellus Roxb., Mascarene Island leafflower, (3), DA, C, 1406 *Phyllanthus urinaria L., Chamber bitter, (3), Da, C, 1418 Phytolaccaceae Phytolacca americana L. var. rigida (Small) Caulkins & Wy- att, American pokeweed, (3), D 1425 [This distinct geographical race, previously included in the flora by Wunderlin (19 ot in the recent edition (Wun- derlin & Hansen 2003), merits recognition (see Caulkins & Wyatt 1990).] Plantaginaceae (Veronicaceae) Bacopa monnieri (L) Pennell, Herb-of-grace, (3), ^o/MH/sW, Zz Gratiola hispida (Benth. ex Lindl.) Pollard, Rough hedgehyssop, (5), sa/sw, O, 1451 Gratiola pilosa Michx., Shaggy hedgehyssop, (1)/(5), DA/PE/sa/ SW, O, 1455, 1506 Gratiola ramosa Walter, Branched hedgehyssop, (5), AQ, C, Linaria canadensis (L.) Chaz., Canada toadflax, (5), sa, o, 1185 *Lindernia crustacea (L.) F. Muell., Malaysian false pimpernel, ,DA, C, 1724 Plantago virginica L. Virginia plantain, (3), Da, c, 1219 Scoparia dulcis L., Sweetbroom, (5), sa, i, 1448 *Veronica arvensis L., Corn speedwell, (3), DA, C, 1263 Platanaceae Platanus occidentalis L., American sycamore, (1), sw, o, 1486 Plumbaginaceae Limonium carolinianum (Walter) Britton, Carolina sealavender, , 0, Poaceae Agrostis hyemalis (Walter) Britton et al., Ticklegrass, (1)/(3), —o, 1229, 1361 Andropogon glomeratus (Walter) Britton et al. var. glomeratus, Bushy bluestem, (5), s4, o, 1645 Andropogon glomeratus (Walter) Britton et al. var. pumilus (Vasey) Vasey ex L. H. Dewey, Bushy bluestem, (5), sa, 0, 1643 £L Dag PRA D hi PEA Journal UI f Texas 1(2) Andropogon virginicus L. var. glaucus Hack., Chalky bluestem, (5), SA, C, 1642 Andropogon es L. var. virginicus, Broomsedge bluestem, (5), SA, O, 644 is iom Elliott, Bottlebrush threeawn, (5), SA, C, visio stricta Michx. var. Ai (Trin. & Rupr.) D. B. Ward, ass, (5), SA, O, 16 Bromus catharticus D EN EUN (1), DA/MH, O, 1310 a s Nash, Slender sandbur, (3), sa, r, 1554 Cenchrus spinifex Cav., Coastal sandspur, (3), Da/Du, C, 1596 Chasmanthium laxum (L.) Yates var. sessiliflorum can Wipff & - D. Jones, Longleaf woodoats, (1)/(3), Da/MH/sw, C-i, 1392, 1488 *Dactyloctenium aegyptium (L.) Willd. ex Asch. €: Schweinf., Durban crowfootgrass, (3), DA, c, 1602 Dichanthelium aciculare (Desv. ex Poir.) Gould & C. A. Clark, Needleleaf witchgrass, (1)/(5), Da/PE/sa, c-o, 1187, 1188, 1350 Dichanthelium commutatum (Schult) Gould, Variable witch- SS, (3), DA/MH, C, 1235 Dichanthelium laxiflorum (Lam.) Gould, Openflower witch- grass, (1), pA/sw, C, 1 Digitaria ciliaris (Retz.) Koeler, Southern crabgrass, (3), Da, Ci, 399, 148 Distichlis spicata (L.) Greene, Saltgrass, (3), sm, €, 1597 *Fleusine indica (L.) Gaertn., Indian goosegrass, (3), DA, C, 1428 Elymus virginicus L., Virginia wildrye, (3), DA, C, 1412 Eragrostis elliottii S. Watson, Elliott's lovegrass, (5), Sa, C, 1617, 1633 doc ophiuroides (Munro) Hack., Centipedegrass, (4), 673 pac petraea (Sw.) Desv., Pinewoods fingergrass, (3), sh, Meis mutica S Twoflower melicgrass, (2)/(3), DA/sM, |; + Oplismenus Pm ) Beauv., Woodsgrass, (3), MH, O, 1587 Pani Michx., Beaked panicum, (2)/(5), DA/MH/SA/SW, C—O, 1619, 1663 Pani / j Schult., Maidencane, (1)/(5), A0/DA/PE//sw, ,1630, 1703 Panicum verrucosum Muhl., Warty panicgrass, (1)/(5), A0/DA/sw, —o, 1629, 1632, 1654, 1709 Panicum virgatum L., Switchgrass, (1)/(3), aQ/DA/sm, C-O, 1543, 3 *Paspalum notatum Flüggé var. notatum, Bahiagrass, (3), DA, o, 1429 *Paspalum notatum Flüggé var. saurae Parodi, Bahiagrass, (3), DA, o, 1404 Paspalum setaceum Michx., Thin paspalum, (3), MH, o, 1590 e urvillei Steud., Vaseygrass, (3)/(5), ae O, 9 = a avenaceum (L.) Parodi, Blackseed needlegrass, (3), MH, O, 1389 *Poa annua L., Annual bluegrass, (3), DA, c, 1216 *Polypogon monspeliensis (L.) Desf., Rabbitsfootgrass, (1), DA/SW, r, 1315 Zomlefer et al., Flora of NPS areas of Timucuan Preserve Setaria parviflora (Poir.) Kerguélen, Yellow bristlegrass, (1), AQ/DA, r, 1522 Sorghastrum secundum (Elliott) Nash, Lopsided Indiangrass, 1), DA/PF, O, 1714 *Sorghum bicolor (L) Moench, Grain sorghum, (4), cult, 1678, 1679 [UNI] Spartina alterniflora Loisel., Saltmarsh cordgrass, (2), sm, C, 1660 Spartina bakeri Merr., Sand cordgrass, (5), AQ, c, 1202 i "n (Aiton) Muhl., TE cordgrass, (3), sm, NUN obtusata (Michx) Scribn., Prairie wedgescale, (1)/(3), DA, o-r, 1230, 1267, 1316 *Sporobolus indicus (L.) R. Br. var. indicus, Smutgrass, (3), MH, O, 1471 Stenotaphrum secundatum (Walter) Kuntze, St. Augustine- grass, 6), DA, O, 1421 au. common lawn grass is listed in M UNT] tour collection likely represents an escape from cultivation. The native plants, characteristic of coastal habitats, are diploid, and the cultivated turf plants are polyploi Tridens flavus (L.) Hitchc. var. flavus, a tridens, (3), MH, r, 1564 *Vulpia myuros (L) C. C. Gmel., Rattail fescue, (1), AQ/DA/PF, eT Polemoniaceae *Phlox drummondii Hook., Annual phlox, (3), Da, c, 1275 lygalaceae Pola ee Orange milkwort, (1), DA/PF, c, 1340 Polygala nana (Michx.) DC., Candyroot, (1), DA/PF, o, 1341 Polygonaceae Polygonum hirsutum Walter, Hairy smartweed, (5), AQ, O, 1442 Polygonum Dae ee Michx., Swamp smartweed, (3), "OL /sw, i, 1433 EI D J |, (3), AQ/MH/SW, r, 1432 ee Rumex hastatulus Baldwin, Heartwing dock, (4), DA/DU/SM, PELS Pontederiaceae *Eichhornia crassipes (Mart.) Solms, Common water-hyacinth, (3), [Car 1], AQ, C, 1721 Pontederia cordata L., Pickerelweed, (1), sw, o, 1499 Portulacaceae Portulaca pilosa L., Pink purslane, (3), DA, o, 1601 osaceae Prunus caroliniana Marshall, American plum, (3), Da, o, 1281 Prunus serotina Ehrh. var. serotina, Black cherry, (4), DA/MH, o, 1380 *Rosa laevigata Michx., Cherokee rose, (4), CULT, 1669 Rubus argutus Link, Sawtooth blackberry, (1), DA/PF, c, 1333 Rubus cuneifolius Pursh, Sand blackberry, (1), sw, c, 1305 Rubus trivialis Michx., Southern dewberry, (3), D v. 1282 iaceae Cephalanthus occidentalis L., Common buttonbush, (3), AQ/DA, O, 1416 1175 Chiococca alba (L.) Hitchc., Snowberry, (3), su, o, 1568 Diodia teres Walter, Rough buttonweed, (3 D DA, O, 1604 Diodia virginiana L., Virginia buttonweed, (1), ^o/DA, O, 1519 Galium hispidulum Michx., Coastal bedstraw, (3), MH/SA, C, ! Galium tinctoriun Stiff marsh bedstraw, (3), AQ/DA, O, 1283 Houstonia procumbens (J.F. Gmel) Standl, Roundleaf bluet, , DA, O, puri corymbosa L., Flattop mille graines, (4), Da, C, MLS OT X mom uniflora L., Clustered mille graines, (1)/(5), Da/PE/ 5 Psychotria nervosa Sw., Wild coffee, (3), MH, c, 1388, 1567 *Richardia brasiliensis Gomes, Tropical Mexican clover, (3), DA, r, 1257 Ruscaceae *Aspidistra elatior Blume, Cast iron plant, (3), MH, r, 1391 [Non-reproductive plants persisting from cultivation; not naturalized.] *Liriope spicata Lour., Monkey-grass, (2), DA/MH, r, 1532 [Persist- ing from cultivation; probably not naturalized.] Rutace "Gir xaurantium L., Citrus hybrid, (3), DA/MH, r, 1397 inia buxifolia (Poir) Ten., Chinese boxorange, (3), CULT?, 1258 [Likely planted and persisting; not naturalized.] alicaceae Salix caroliniana Michx., Carolina willow, (3), Ao/DA, C, 1279 Santalac Phoradendron leucarpum p Reveal & M. C. Johnst., Oak stletoe, (4), DA/MH, O Sapindaceae Acer rubrum L., Red maple, (1), o4/Pr, o, 1501 Sapindus saponaria L., Soapberry, (3), MH, C, 1566 Sapotaceae Sideroxylon tenax L., Tough bully, (3), ba/mh, i, 1396 Sarraceniaceae Sarracenia minor Walter, Hooded pitcherplant, (1), AQ/DA/PF, 41335 Saururaceae Saururus cernuus L., Lizard's tail, (1), Ao/sw, o, 1314 Smilacaceae Smilax auriculata Walter, Earleaf greenbrier, (3), sa, C, 1555 Smilax bona-nox L., Saw greenbrier, (3), DA/MH, i, 1254 Smilax glauca Walter, Cat greenbrier, (3), MH, c, 1291 Smilax laurifolia L., Laurel greenbrier, (5), sa/sw, C, 1462 Smilax pumila Walter, Sarsaparilla vine, (3), MH/sa, C-i, 1475, NI] 1558 lu Smilax tamnoides L., Bristly greenbrier, (2), MH, O, 1665 Solanaceae Lycium ea Walter, Christmasberry, (2), sm, C-O, 1542, 165 s ante Walters groundcherry, (3), DA/DU, C, 1227 m americanum Mill, American black rightsaade (3), m i, 1240 1176 Journal of the Botanical R h Institute of Texas 1(2) Tetrachondraceae Phyla nodiflora (L.) Greene, Turkey tangle fogfruit, (3), DA, O, Polypremum procumbens L., Rustweed, (5), sa/sw, o, 1452 1251 Theaceae Verbena brasiliensis Vell., Brazilian vervain, (1), Da/sw, o, 1487 Gordonia lasianthus (L.) J. Ellis, Loblolly bay, (1), a0/Da, 0, 1322 Violaceae Typhaceae (ur pid Pe e Mas (5), AQ, O, 1208 Typha latifolia L., Broadleaf cattail, (1), Ao/»r, i, 1686 YO d mad Lu OO 6), Pae Oe Viola sororia Willd., Common blue violet, (3), DA, o, 1278 Vitaceae Ulmaceae Ulmus alata Michx., Winged elm, (1), sw, r, 1485 Ampelopsis arborea (L.) Koehne, Peppervine, (3), AQ/DA, O, Ulmus americana L., American elm, (1), sw, c, 1309 1422 Urticaceae Parthenocissus quinquefolia (L) Planch., Virginia creeper, (3), Boehmeria cylindrica (L.) Sw., False nettle, (1), sw, c, 1494 AQ/DA, O, 1410 Parietaria praetermissa Hinton, Clustered pellitory, (3), oa, r, Vitis aestivalis Michx., Summer grape, (3), DA/MH, o, 1259 Vitis rotundifolia Michx., Muscadine, (3), DA/MH, C, 1253 1224 Pilea microphylla (L) Liebm., Artillery plant, (3), DA, C, 1226 Xyridaceae Verbenaceae Xyris ambigua Beyr. ex Kunth, Coastalplain y ll VEY d grass, *Lantana camara L., Lantana, (3), [car 1], cULT?, 1244 (1), DA/PF, o, 1697 „cut, *Xyris jupicai Rich. Richard's yelloweyed grass, (1), DA/PF, c, _ Lantana depressa Small, Rockland shrubverbena, (3 1243 1342 *Lantana montevidensis (Spreng) Brig, Trailing shrubverbena, Xyris platylepis Chapm., Tall yelloweyed grass, (5), aa/sa/sw, (3), CULT, 1242 O, 1463 ACKNOWLEDGMENTS We thank Kelly A. Bettinger, Galen Burke, T. Brett Miller, and Adam Smith for enthusiastic assistance in the field; Kelly also carefully maintained specimen lists, organized field supplies, databased label information, and supervised mounting of specimens. We greatly appreciate the invaluable support and cooperation of National Park Service personnel: Joe DeVivo, Network Coordinator for the Southeast Coast Network; Rich- ard Bryant, Chief of Resources Stewardships at Fort Caroline National Memorial and Timucuan Ecological and Historic Preserve; and Anne Lewellen, Museum Curator at Timucuan Ecological and Historic Preserve. Stephen Lee Echols expertly identified many sedge and grass species; Kent D. Perkins and Richard P. Wun- derlin provided careful reviews of the manuscript; and Eric Fuchs translated the abstract into Spanish. We are also grateful to James L. Hamrick for the loan of his field vehicle, and the Department of Plant Biology, University of Georgia, for the use of a departmental vehicle for transporting supplemental field supplies and personnel. This survey was funded by National Park Service contract agreement J2114-03-0006 (PI D.E. Giannasi & coPI W.B. Zomlefer). REFERENCES ANDERSON, A., L. DAHL, J. GARRETT, D. STRAH, D. VANA-MILLER, S.A. BROWN, S. Lewis, C. SHELDON, and R. Sussman. 1996. General management plan, development concept plans, Timucuan Ecological and Historic Preserve, Florida [unpub- lished report]. Report D-7B. National Park Service, Denver Service Center, Denver, Colorado. ABRAHAMSON, W. G. and D. C. Hartnett. 1990. Pine flatwoods and dry prairies. In: Ecosystems of Florida, R. L. Myers and John J. Ewel, eds. University of Central Florida Press, Orlando, Florida. Pp. 103-149. ANGIOSPERM PHYLOGENY GrouP [APG]. 2003. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Bot. J. Linn. Soc. 141:399-436. Bennett, C.E. 1976. Fort Caroline and its leader. University Presses of Florida, Gainesville, Florida. Bennett, C.E. 2001. Laudonniére & Fort Caroline: history and documents. University of Alabama Press, Tuscaloosa, Alabama. BoarNER, M.M. 1973. Landmarks of the American Revolution: a guide to locating and knowing what happened at the sites of independence, Stackpole Books, Harrisburg, Pennsylvania. CauLkINs, D.B. and R. Wyatt. 1990. Variation and taxonomy of Phytolacca americana and P rigida in the southeastern United States. Bull. Torrey Bot. Club 17:357-367. Zomlefer et al., Flora of NPS areas of Timucuan Preserve 1177 CHoLEWA, A.F. and D.M. HENDERSON. 2002. Sisyrinchium. In: Flora North America, vol. 26, Magnoliophyta: Liliidae: Liliales and Orchidales, Flora North America Editorial Committee, eds. Oxford University Press, New York. Pp. 351-371. Cotte, N.C. and M.A. GARLAND, 2003. Notes on Florida's endangered and threatened plants. Botany Contribution 38, ed. 4. Florida Department of Agriculture and Consumer Services, Division of Plant Industry, Gainesville, Florida. URL: http://www.virtualherbarium.org/EPAC/default.htm/. DeLorme. 1997. Florida atlas & gazetteer, ed. 4. DeLorme, Yarmouth, Maine. Duever, L.C. 1996. Rare plant survey of the Timucuan Ecological and Historic Preserve [unpublished report avail- able from NPS]. Conway Conservation, Micanopy, Florida. Duncan, W.H. and N.E. Britain. 1966. The genus Gaylussacia (Ericaceae) in Georgia. Bull. Georgia Acad. Sci. 24:13-26. Easley, M.C. and W.S. Jupp. 1993. Vascular flora of Little Talbot Island, Duval County, Florida. Castanea 58:162- 15227; Ewe, K.C. 1990. Swamps. In: Ecosystems of Florida, R. L. Myers and John J. Ewel, eds. University of Central Florida Press, Orlando, Florida. Pp. 281-323. FLORA or NORTH AMERICA EDITORIAL Committee [FNA]. 1993. 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National Park Service: national register of historic places. URL: http://www. cr.nps.gov/nr/index.htm/. Nesom, G. L. 2006a. Gamochaeta. In: Flora North America, Vol. 19, Magnoliophyta: Asteridae (in part): Asteraceae, part 1, Flora North America Editorial Committee, eds. Oxford University Press, New York. Pp. 431-438 . Nesom, G. L. 2006b. Liatris. In: Flora North America, Vol. 21, Magnoliophyta: Asteridae (in part): Asteraceae, part 3, Flora North America Editorial Committee, eds. Oxford University Press, New York. Pp. 512—535. ScHaFER, D.L. 1997. Anna Kingsley, revised and expanded edition. St. Augustine Historical Society, St. Augustine, Florida. SmitH_ , A.R., K.M. Prver; E. ScHuETIPELZ; P. KoraALL; H. SCHNEIDER; and PG. Worr. 2006. A classification of extant ferns. Taxon 55:705- 731. STALTER, R. and S.C. Diat. 1984. Hammock vegetation of Little Talbot Island State Park, Florida. Bull. Torrey Bot. Club 111:494-497. STEINWAY-RODKIN, C.M. and C.L. MoNrAGuE. 2003. 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URL: http://www. pone ius ZOMLEFER, W.B, D.E. Giannası, W.S. Jupp, L.M. Kruse, and K.A. BETINGER. 2004. A floristic survey of Fort M tional Monument, St. Johns County, Florida. Sida 21:1081-1106. ZOMLEFER W.B., D.E. GiANNAsI, K.A. BETTINGER, S.L. ECHOLS, and L.M. Kruse. (in prep.). Floristic survey of Cumberland Island National Seashore, Camden County, Georgia. To be submitted to Castanea. THE VASCULAR FLORA OF THE CHUNKY RIVER (MISSISSIPPI) Lucas C. Majure! Department of Biological Sciences Mississippi State University, P. O. Box GY Mississippi State, Mississippi 39762, U.S.A. / ICEEIUT 1@Msstate.edu ABSTRACT A floristic survey of a portion of the Chunky River, situated in east-central Mississippi (SE USA), dertal determine the level of species richness within an area recently set aside as a scenic stream of Mississippi. Rare, uncommon, and disjunct species’ popula- tions were investigated and quantified. Invasive and exotic species were evaluated within the habitats in which they were found to subjectively determine possible deleterious DUE that could be observed through their possible expansion. A total of 805 taxa were recorded (including species, infraspecific taxa and 1 hybrid), encompassing 422 genera and 140 families. The famili ntaining the highest number of taxa were: Poaceae (120), Asteraceae (83), Cyperaceae (80), and Fabaceae (56). Nine tracking list species, according to the Mississippi Natural s Program, were encountered which Edi Callirhoe P Carex tenax, Clematis glaucophylla, Ilex montana, P. ] lius, Rhapidophyllum hystrix, RI } bul var. pinetorum, Scl and Staphylea trifolia Ten of the state's wach list d Carex P ell erciol ium, Eleocharis baldwinii, Fuirena pumila, eae Sean Panicum s brachyanthum, Ptelea trifoliata, RI also were found. T : records were located for Clarke Caine 272 new county recor ds were located for Lauderdale County, and 56 new county records were found for Newton County. Carex dasycarpa was collected representing the first specimen of this species found within the state of Mis- sissippi in 112 years and 2 new county records. In total, 79 exotic/invasive species were found during this stu RESUMEN Se hizo una bn pd ids de una mo del Río re ns jue qued l cent te del estado de Mississippi (sureste de los EE.UU.) para l de riqueza I designad énico de Mississippi. Fueron investigadas y cuantificadas poblaciones de especies raras, no comunes y disjuntas. Fueron evaluada ti te especi Sti x 1 d xL pa | pa ra a| 1 dict H E s a e 1 i y | I 8 J que p expansión. Un total de 805 taxa f itad incl peci inf ifi un híbrido) ab d 2 géneros y 140 familias. Las famili ti l ámero de taxa f las: Poaceae a 20), Asteraceae (83) ROA (80) y las Fabaceae (56). Nueve especies de la lista de plantas raras, según el programa de Patrimonio Natural de MS, fueron tre las que se incluyen: Callirhoe Mara Carex tenax, DR iiu cid Ilex montana, Panax dd ia, QU ck Be ee globularis var. pinetorum, S UN queque es 1 rar Carex picta no E Dichantl tifoli Oxalis grandis, Pani bracl } Ptelea ae Rhyncho- Trid lini Se encontraron 13 l larke, 272 para el spora mixta, AA HA b y con derdal ado Lauderdale, y 5 ndado Newton. Se colectó Carex poma por pe vez en e estado de MS desde i 112 años y se representan dos nuevas citas del condado. En total, 7 INTRODUCTION Historical Significance Although the name Chunky sounds as if it pertains to a derogatory term, it is actually the English derivative of a name used for a game played by the Choctaw Indians called “Chunka” at Chunky Shoals (Breiger 1997), where Boyette's Fish Camp sits along the Chunky River. The early Native Americans heavily used this area for its stone. The Tallahatta Formation, located throughout the extent of the Chunky River area that was surveyed, contains an easily fracturable rock known as tallahatta quartzite (Lowe 1915). This material (a combination of quartz sand and glauconite solidified by silica) is capable of forming concoidial fractures, which lends the material useful for making highly sharpened, well-crafted, stone tools. This stone, which can be found throughout certain parts of the river where it outcrops, was mined by the aboriginal peoples of this area for thousands of years. The east-central part of Mississippi therefore became an area of heavy trade and resource availability for the Native Americans throughout parts of the southeast. This is evident 1 jd Florida M fN | Hi University of Florida, P.0. Box 117800, Gainesville, Florida 32611, U.S.A.; Imaj fl.ed ? 7 y J. Bot. Res. Inst. Texas 1(2): 1179 — 1202. 2007 1180 Journal of the Botanical R h Institute of Texas 1(2) by tallahatta quartzite artifacts that are found around many parts of the southeast. Also, stones from other regions (e.g., Fort Payne Chert from Tennessee) eventually made their way to the Chunky River region through trade (McGahey et al. 1992). Tallahatta quartzite is still used by many modern-day flintknappers (Majure pers. exp.). Stuckey's Bridge is an important historical structure located along a portion of the survey area in Lauderdale County (Fig. 1). It was built by the Virginia Bridge and Iron Company in 1901 and was placed on the National Register for Historic Places in 1988 (Historic Bridges of the United States 2006). The man for which it is named is said to have owned a hotel near the bridge and would murder patrons for money and other valuables, then would bury their bodies along the banks of the Chunky River. After local people discovered what was happening, *Stuckey" was hung from the bridge. His spirit is said to still haunt the bridge (local legend). Dunn's Falls is another area of historical significance located along the Chunky River (Fig. 1). This was the state's second natural area, developed in 1981. A total of 27.9 ha around the mill were bought by the Mis- sissippi Wildlife Heritage Committee, as well as a 4.05 ha easement (Anonymous 1981;, Mississippi Natural Heritage Program 2002). Dunn's Falls was originally owned by Irish immigrant John Dunn who settled there in 1854 to establish a cotton mill. This soon was overtaken by the confederate government after the onset of the Civil War. The mill was then used to make blankets, hats, and knives for the Confederate Army. After the war, the mill was used to make flour, corn meal, hats, and money. Eventually, the mill was reclaimed by the forces of nature and carried off by the river. The mill is now represented by an 1857 Carroll-Richardson gristmill and water wheel taken from Cave Springs, Georgia (Pat Harrison Waterway District 2002). Survey Significance The Chunky River is important in that it forms part of the upper Pascagoula River System, which is *the last unimpeded major river system in the lower 48 states.” It forms, along with Okatibbe Creek, a “sub- basin" of the overall Pascagoula River basin. The Pascagoula River Basin drains about 15,446.4 km of land. The system therefore is important in providing nutrients and sediments necessary for salt marsh stability (8,498.4 hectares of coastal marsh exist within the basin), which then *feed" the Mississippi Sound estuary (Mississippi Department of Environmental Quality 2001). In 2003 the Mississippi Legislature designated the Chunky River as a scenic stream under the Missis- sippi Scenic Streams Stewardship Program supported by the Mississippi Department of Wildlife Fisheries and Parks (Andrew Whitehurst Mississippi Museum of Natural Science pers. comm.). However, before a river can be evaluated for nomination as a scenic stream, there are several requirements that it must meet (Andrew Whitehurst Mississippi Museum of Natural Science pers. comm .). First and foremost, any river system under scrutiny has to be on the State of Mississippi's list of *Public Waters." It then has to pass at least one of five puc Listed publication by a government agency, or by id tion group, prof l tion or a or, aa Traverses or i tate or national park, forest, refuge, wildlife management area, monument or other major public land. Exhibits e or potential use for recreational boating (motorized boating, rowing, canoeing, kayaking or tubing Possess accessible unique scenic or historical sites. Supports plants and/or animals that are unique or of special concern Once one of these requirements is met, the p area is subjected to an evaluation through a series of c t criteria (Assessment Criteria for Mississippi's Stewardship Program 2002). These criteria are composed of biological, physical, human interaction and historical elements. As a result of the high scoring of the Chunky River through several areas of testing (Assessment Criteria for Mississippi's Scenic Streams Stewardship Program 2002), the House Bill 952 “designated a certain portion of Chunky Creek in Newton County and the Chunky River in Newton, Lauderdale and Clarke counties, as eligible for nomination to the State Scenic Streams Stewardship Program" (Mississippi Legislature 2002). Of all the related elements subjected to testing for the proposed nomination of the Chunky River, the Majure, Flora of Chunky River in Mississippi 1181 Chunky River Survey Area Site Locations Dunn's Falls E <= m e * Hwy. 11 takeout Newton County A Lauderdale Co. bridge [5] Newton Co. bridge % Gtuckey's Lauderdale County Primary Roads == 20 514 i Clarke County COL E e — mnt ¡lali Fic. 1. The tri y area in which tl y lucted is sl ling from tt f Chunky (N Co.) tott f Enterprise (Clarl Co) I J Jai f 41 L HI G > J L . > £ “sl ed I * JJ , » * flora was not studied. Considering the geological makeup of the area, the wide array of habitats within this system, and the fact that the area had not been studied in detail previously (e.g., only few collections had been made previously by Sydney McDaniel, Ken Rogers, Fanney Cook, Richard Carter, Charles Bryson and a few other botanists), this was an intriguing area in which to perform a floristic study. Recording the flora of this area not only helps conservation efforts but also contributes valuable information and reference material for understanding floral assemblages and plant distributions within this understudied region of Mississippi. Geography, Geology and Climate The Chunky River begins at the confluence of Okahatta and Chunky creeks in east-central (Newton County) Mississippi and ends at the formation of the Chickasawhay River by the junction of the Chunky River and Okatibbee Creek in Clarke County, MS. It is situated in the physiographic region known as the North Central Plateau (Lowe 1921, Fenneman 1938) within the upper coastal plain land resource unit (Vanderford 1962). The Burhstone/Lime Hills is the main ecoregion covering the area studied (Chapman et al. 2004). This £L Dag PRA D hi PEA 1182 Journal of f Texas 1(2) ecoregion is a result of part of the Claiborne group that outcrops in the area (Lowe 1915) known as the Burhstone Cuesta (Fenneman 1938) or Tallahatta Formation and the Enterprise Green Marl (Lowe 1915). Elevations range from 76-201 m. Hard substrates (outcrops and underlying material) consist of Tertiary (Eocene) claystone, siltstone, clay, sandy clay, sandstone, limestone and marl (Lowe 1915; Chapman et al. 2004) which were derived mainly from marine deposits (Lowe 1915). Soil types along the extent of the survey generally are moderately to excessively well-drained sandy loams and are mildly to very strongly acidic. Most of the soil types within the survey area are low in organic matter (Murphree 1957; Neal et al. 1961; Allgood 1983). The majority of the floodplain soils are of the Bigbee-Bibb soil association, where Bigbee soils are medium to strongly acidic and well drained. Bibb soils are acidic or very strongly acidic and poorly drained. Upland soil associations are of Arundel-Lauderdale, Arundel-Sweatman-Smithdale, Arundel-Smithdale-Williamsville, and Lucy-Smithdale soils, which are all well drained and mildly to strongly acidic. Arundel soils are underlain by stratified sandstone, siltstone, and burhstone. Smithdale and Willliamsville soils are underlain by thick, sandy, marine sediments (Allgood 1983). m summers and cool winters are typical. Thirty year annual averages for temperature and precipi- tation (from 1971-2000) are 18.2?C and 149 cm/yr, respectively. Maximum and minimum annual average temperatures for this time period are 24.9°C and 11.3?C, respectively (National Weather Service 2007). The original forest type for this physiographic region consisted of mixed pine and deciduous forests, generally dominated by pine (Lowe 1915). Other than remaining mixed pine and deciduous forests and scrub-shrub forests mainly bordering the river, cutover quite common in many parts of the survey area. These are typically made into loblolly pine plantations except where native vegetation is allowed to grow back naturally. MATERIALS AND METHODS Survey Methods This survey of the Chunky River was performed from March 2004 through early May 2007. The area in- vestigated included sites along roughly 34 km of the Chunky River (32.3264%N, 88.9209°W to 32.1761°N, 88.8198°W) in Newton, Lauderdale and Clarke counties (Fig. 1). Topographic, road, and soil maps of the area were used to choose sites that could be surveyed and re-surveyed over the collection period to account for temporal variation in floral communities. These sites were chosen to account for the heterogeneity of habitats and geology that were observed along the extent of the survey area. The area chosen for study was that extent of the Chunky River that is most heavily used which presumably would suffer the greatest amount of anthropogenic disturbance. Surveying was conducted by canoe in order to search sandbars, rockbars, and river edges and by foot in order to extensively investigate boat ramps, river margins and trails, and roadsides within 50 m of the river at selected sites. About 30 trips were made to the river over the course of this study. Rare plant populations were heavily studied using guidelines set up by the Mississippi Natural Heritage Program to estimate, from an observational perspective, the plant population health, size, and bili These data were supplied to the Mississippi Natural Heritage Program in order to aid in the conservation of these populations through their tracking and watch list program. Collections Currently there is no flora for Mississippi, so collections were identified using various keys for the southeast- ern United States and other regions which include: Bailey (1949), FNA (1993+), Godfrey & Wooten (1979 & 1981), Hitchcock (1971), Radford et al. (1968), Steyermark (1963), Weakly (2003), and Wunderlin (1998). Questionable specimens were compared with collections at the Mississippi Museum of Natural Science Herbarium (MMNS) and the Mississippi State University Herbarium (MISSA). Species within the Cypera- ceae were reviewed by Dr. Charles T. Bryson at the USDA experiment station in Stoneville, MS. Collections made during this survey were vouchered at MMNS in Jackson, MS. Duplicates made during the collection Majure, Flora of Chunky River in Mississippi 1183 process were donated to MISSA in Starkville, MS and the Stoneville Research Station Herbarium (SWSL) in Stoneville, MS. Several primary state herbaria; IBE, MISS, MISSA, MMNS, and USMS were reviewed to check for new county records that were potentially found during collection. The Carex from SWSL were checked for any new county records within that genus. A comprehensive list of the plant species found during the survey is provided (Appendix 1). Species names are given along with the collection number, a general habitat description and the frequency in which they were encountered. The plants are listed by class, family, genus, and species. Nomenclature follows The Plants Database (2007) and FNA (1993+), unless otherwise noted. Family names and their respective genera follow the Angiosperm Phylogeny Website (2007). Any relatively recent changes are detailed in the appendix (e.g., Orobanchaceae (Scrophulariaceae, in part)). Author abbreviations follow IPNI (2007). Rare plants that were found during the survey are supplied with their state ranking given by the Mis- sissippi Natural Heritage Program (2006). Rankings include S1, S2 and S3 for plants on the state tracking list. Higher ranks (S3/S4) are designated for plants only on the state watch list. S-ranks are determined by the abundance and self-maintenance of certain species. The S-1 ranking signifies that the species is under extreme peril in Mississippi through rareness or through some other factor constraining its reproduction or sustained existence. The S-2 ranking is given to denote moderate rarity or some other reproductive or life sustaining, limiting factor. The S-3 ranking means that a species is simply rare or infrequent in the state, and an S-4 ranking means a species is abundant and widespread, but there is reason for future concern relating to this species' continued survival in its present state. An S-5 ranking signifies that the species is relatively common throughout the state and is not apparently threatened in any way (Mississippis Comprehensive Wildlife Conservation Strategy 2004). Invasive and/or exotic species collected during the survey are noted in Appendix 1. These are species deemed non-native (exotic) or non-native and invasive by The PLANTS Database (USDA 2007). Regional introductions of plants are not > considered when designating a species as indigenous or non-indig- enous to an area; therefore, some species deserving of such a listing might not be represented as introduced (e.g., Digitaria horizontalis). Also, native species e are considered to be invasive were not denoted as such in the species list. RESULTS Floral Richness In total 805 taxa, 422 genera, and 140 families were recorded. Families with the highest number of taxa were the Poaceae (120), Asteraceae (83), Cyperaceae (80), and Fabaceae (56). The Poaceae (21) and Fabaceae (13) had the highest number of exotic and/or invasive taxa. The Cyperaceae had the highest number of track- ing and watch list taxa recorded for the survey (7). Thriteen new county records were recorded for Clarke County, 272 for Lauderdale County and 56 for Newton County. Floral Community Heterogeneity A total of 10 different community types were observed. These are river channel, river bank, floodplain forests scrub-shrub, xeric openings, sloughs, spring seeps, upland forests, mesic forests, and anthropogenically disturbed areas. Floodplain forests, scrub-shrub, xeric openings, sloughs, spring seeps and some anthropo- genically disturbed areas are all within the floodplain. Rocky, mixed uplands and mesic forests fall within the floodplain where the Burhstone Cuesta erupts or grade directly into the river. The River Channel (RC) community is consistently disturbed by moderate fluctuations in water levels. The most common constituents of this community are the river channel itself, sandbars, rockbars, snags, stumps, and the river margin (where the bank i the river channel), and the most frequently encoun- E 1 tered species are ruderal. Most species are annuals such as Amaranthus spinosus, Cl iana, Cyperus erythrorhizos, C. iria, C. polystachyos, Eragrostis hypnoides, Fimbristylis autumnalis, F. miliacea, Kyllingia pumila, Lindernia dubia, Ludwigia palustris, Oldenlandia boscii, Paspalum boscianum, P. fluitans, Physalis pubescens and Sesbania herbacea. Perennial species are either on fairly stable sandbars or along the river margins where the 1184 Journal of the Botanical R h Institute of Texas 1(2) substrate is less subject to dislodging. Specific examples are Alternanthera philoxeroides, Brunnichia ovata, Cyperus esculentus, Itea virginica, Nyssa aquatica, Onoclea sensibilis, Osmunda regalis, Planera aquatica, Polygo- num hydropiperoides, Rubus trivialis, Sesbania punicea, Steinchisma hians, Taxodium distichum, Trachelospermum difforme, and Triadenum walteri. Podostemon ceratophyllum is a major component of the actual channel where it grows submerged from early fall through early summer. In early fall small plantlets can be found attached to hard submerged Sunc These continue to grow until early summer when plants can be up to 0.5 m or longer and form xtensive colonies in fast-flowing water. As the water level falls during the first dry periods of early summer, this species can be found slightly emergent on rocks, logs or any other hard substrate barely above the water level. a this point the plant begins to bloom, fruit, and eventually senesce. The water column in drier years (e.g., 2006) is then invaded by a diverse group of filamentous algae that occupies these areas through the rest of the summer or until water levels increase. The River Bank (RB) community consists of those intermediate areas between the river channel and the floodplain. The RB community substrate is composed of either sandy, acidic soils, like those of the adjacent floodplains, or of rock outcrops where the Burhstone Cuesta erupts or has been uncovered by the river chan- nel. Most of the RB flora is perennial, but ruderal annuals also are found in these areas frequently disturbed by annual flooding. The most common species in these areas are Agrostis perennans, Alnus serrulata, Apios americana, Mo iden UE Chasmanthium latifolium DOTT gallioides, Ilex decidua, Ipomoea pandurata, 1 anescens, Solidag a phiotrichum lateriflorum and Viburnum dentatum. Along rock outcrops where spring seeps occur, Selaginella apoda is found along with other pteridophytes and a very interesting assortment of bryophytes (hornworts, liverworts and mosses). Floodplain forests (FF) are scattered throughout the extent of the survey area. In many areas they have been decimated by timber harvesting practices, but some relatively large stands of forest still remain. Soils within these areas are generally well drained to sometimes poorly drained, sandy, and acidic with low organic content (Murphree 1957, Neal et al. nie Allgood 1983). Typical tree species are Acer barbatum, Ty. d 41 M AT [e] e» d Carya pallida, C. glabra Ostrya virginiana, Pinus glabra, Quercus austrina, O. hemisphaerica, Q. michauxii, EE Q. d UMS shrubby species are Callicarpa americana, Ditrysinia fruticosa, Halesia carolina, Hypericum frondosum, Illicium floridanum, Lindera benzoin, Symplocos tinctoria and Vaccinium elliottii. Ariasaema dracontium, A. triphyllum, Carex abscondita, C. digitalis, Chasmanthium laxum, C. sessiliflorum, Dichanthelium commutatum, D. dichotomum, Hexastylis arifolia, Mitchella repens, and Trillium stamineum are common herbaceous species within these areas. A species of Hymenocallis Salisb. was seen within the floodplain forest; however, it was not collected for the lack of flowering throughout the survey period. Backwater slough (S) communities permeate the floodplain forests and may be ephemerally wet or inundated throughout the year. Slough wetlands are host to a variety of species not found in faster flow- ing currents. Species like Brasenia schreberi, Myriophyllum aquaticum, M. heterophyllum, Nymphaea odorata, Potamogeton pulcher and Ceratophyllum demersum occupy the pools. Other wetland species like Carex joorii, C. intumescens, C. lurida, Iris virginica, Leersia oryzoides, Peltandra virginica, Sabal minor and Sparganium ameri- canum were found along the margins of these sloughs. Taxodium distichum and Nyssa aquatica occasionally form large cypress/tupelo stands within these backwaters. The scrub-shrub (SS) communities are between upland ridge and floodplain forest communities. Soils are well-drained, acidic, sandy soils of the Independence loamy fine sand (Murphree 1957) and Bigbee loamy sand (Allgood 1983). Tree and shrub species in these areas typically are those commonly associated with sandy, dry habitats. Chionanthus virginicus, Crataegus uniflora, Ilex vomitoria, Kalmia latifolia, Osmanthus ameri- canus, Pinus echinata, P. palustris, Ptelea trifoliata, Quercus hemisphaerica, Q. incana, Q. margarettiae, Vaccinium arboreum, and V. stamineum were frequently seen within these areas. Other common species are Areolaria flava, Asclepias variegata, Chasmanthium laxum, C. sessiliflorum, Dichanthelium dd. D. scoparium, Dioclea multiflora, Elephantopus tomentosus, Gelsemium sempervirens, I leza] bens, pumila and Solidago E Majure, Flora of Chunky River in Mississippi 1185 — ulmifolia var. microphylla. Vaccinium darrowii was found in this community type along the river by Dr. Sidney McDaniel in 1970 but was not seen during this survey, although it is quite common in Lauderdale County. Xeric openings (XO) are found among the SS communities. Openings apparently are created by flooding events, moisture constraints on vegetation produced by copious amounts of sand, anthropogenic disturbances, and by beaver activity. Oftentimes, areas are “cleared” by beavers, which are subsequently repopulated by species adapted for open, xeric, habitats. In summer 2004, one area in particular had been recently cleared by beavers. Opuntia pusilla was found in this area but was nearly diminutive from overshading by shrubs. In 2006, when revisiting this site, those same plants were much larger and flowering. Opuntia pusilla is fre- quently found in these areas and therefore typifies these sites, fem an Rui UN pele of view, as xeric. Many other species are associated with and only with these as ta, Bulbostylis dig ciliatifolia var. coarctata, Callirhoe triangulata, Carex tenax, aoo filia iculmis, C. o C a. Digitaria +] cognata, Froelichia floridana, F. gracilis, Seymeria pectinata L tuberosa, Triplasis americana, T. purpurea, and Zornia bracteata. Spring seeps (SPS) are the rarest of the communities seen along the survey area. They can be found in deep ravines of some of the larger ridges or along the upper reaches of some of the slough systems. Soils are poorly drained, acidic soils of Bibb (Allgood 1983) or moderately well drained, acidic soils of the Iuka fine sandy loam (Murphree 1957). Typically the herbaceous vegetation is dominated by Carex leptalea subsp. harperi, Eleocharis tuberculosa, Osmunda cinnamomea, Woodwardia areolata, Xyris laxifolia var. iridifolia and other plants typical of acidic seepages. Overstory vegetation generally consists of Alnus serrulata, Magnolia virginiana, Smilax laurifolia, Toxicodendron vernix, and Viburnum nudum. Rocky, mixed upland (RU) and mesic, deciduous forests (MF) are found in areas with large outcrop- pings from the geological uplifts in the area (Burhstone Cuesta). The two communities are segregated by soil moisture, which is apparent by the vegetation. This mainly is a result of aspect, slope and depth of soil relative to harder substrates (rock outcrops), as most of the soils are typically well drained and moderately to highly acidic (Allgood 1983). However, elevated soil pH often is seen where calcareous portions of the Enterprise Green Marl and Tallahatta Formation (consisting of marine shell fragments, etc.) outcrop (Lowe 1915). This provides areas more suitable for plants commonly found in basic soils (e.g., Heuchera americana, Quercus muehlenbergii; Radford et al. 1968). The RU are generally found on southern, western, or southwestern slopes and typically consist of upland tree species such as Acer barbatum, Magnolia acuminata, M. grandiflora, Pinus echinata, P. palustris, Querucs alba, Q. velutina, and Q. muehlenbergii. Shrubs typical of well drained soils include Asimina parviflora, Hydrangea quercifolia, Ilex montana, Kalmia latifolia, Osmanthus americanus, and Viburnum rufidulum. Herbaceous plants generally are species more typical of well drained soils like Asclepias variegata, Iris cristata, Lilium michauxii, Pedicularis canadensis, Piptochaetium avenaceum, Pteridium aqulinum, and Symphyotrichum cordifolium. Mesic, deciduous forests are more typical of the rich, deciduous forests of the Appalachian foothills, although, many of the overstory components were similar to the RU. The MF communities generally are found on northern, eastern, or northeastern slopes. Common tree species are Acer barbatum, Carya caro- linae-septentrionalis, Fagus grandifolia, Magnolia macrophylla, Quercus alba, Q. muhlenbergii, and Q. pagoda. Herbaceous species include Brachyeletrum erectum, Carex kraliana, C. striatula, Chamaelirium luteum, Desmo- dium nudiflorum, D. pauciflorum, Hepatica nobilis var. obtusa, Panax quinquefolius, Phegopteris hexagonoptera, Sanguinaria canadensis, Smallanthus uvedalius, Tragia cordata, and Trilium stamineum. Anthropogenically disturbed areas (ADA) were heavily influenced through human actions. These locations included boat launches, roadsides, bridges, river trails, ditches, cutovers, and any other sites that 1 might have been impacted by human actions. Many non-native, weedy sp were seen in such areas. Tracking and Watch List Species Nine rare species located on the Mississippi Natural Heritage Program tracking list (2006) were located during this survey. These were Callirhoe triangulata (S1/S2), Carex tenax (S2), Clematis glaucophylla (S1), Ilex 1186 Journal of the Botanical R h Institute of Texas 1(2) montana (S3), Panax quinquefolius (S3), Rhapidophyllum hystrix (S3), Rhynchospora globularis var. pinetorum (S1), Schisandra glabra (S3), and Staphylea trifolia (S3). Also, ten species on the state watch list were found. These include Carex picta (S3/S4), Eleocharis baldwinii (SNA (P)), Fuirena pumila (S3/S4), Dichanthelium erectifolium (S3/S4), Oxalis grandis (S3/S4), Panicum brachyanthum (S3/S4), Ptelea trifoliata (S3/S4), Rhynchospora mixta (S3/S4), Rhynchospora scirpoides (S3/S4) and Tridens carolinianus. Callirhoe triangulata (clustered poppymallow) was found in two different locations within the XO com- munity type. Only five non-reproducing individuals were seen in these small openings within the SS. A more intense investigation of this area for this species could produce larger populations, as this species has been recorded in other areas in Lauderdale County. Carex tenax (wire sedge) was found in virtually all of the XO communities studied along the extent of the survey area. There never are very large populations of this Carex, but it generally is seen to produce large numbers of fruit and apparently thrives in these sandy, xeric sites. Opuntia pusilla tends to be a good indicator species for potentially locating this sedge. 1 1 ped Clematis glaucophylla (whiteleaf leather flower) was seen in many locations the survey area. It is found along the floodplain in the ADA, SS, and XO communities. Most of the areas in which it was recorded consisted of a very sandy, well-drained soil that had been previously disturbed by human actions (cutovers, foot traffic, boat ramps) or by flooding along river margins. It generally was associated with scrubby type species of trees and shrubs. Main associates included Amelanchier arborea, Crataegus uniflora, Ilex decidua, I. glabra, Ptelea trifoliata, Quercus hemisphaerica, Q. incana, Q. margarettiae, Rhododendron canescens, Vaccinium arboreum, V. stamineum, and Viburnum dentatum. Other common herbaceous constituents included Carex dasycarpa, C. tenax, Erythrina herl acea, Heterotheca subaxillaris, Opuntia pusilla, Scleria triglomerata, Sporobolus junceus, Stillingia sylvatica, Tephrosia virginiana, Triplasis americana, and Zornia bracteata. There is a large amount of habitat available for this species’ continued existence along river margins, and increased disturbance seems to be a favorable condition for C. glaucophylla. Although large numbers of this Clematis were not seen in any one location, its distribution seems to be fairly widespread throughout the extent of the main survey area. Ilex montana (mountain holly) was seen in several RU communities throughout portions of the survey area. It is generally found on the slope just below the tops of ridges, so soils typically are fairly dry. The associate flora is that typical of the RU. Mature male and female plants were seen in flower and fruit, so there is reproduction among the populations of this Ilex. Its habitat is relatively stable or free from human disturbance as a result of the topographic nature of the areas. Panax quinquefolius (American ginseng) was located in two different areas of the survey in mesic forests. Associates included a variety of herbaceous and woody species typical of rich, mesic forests. The small colonies of Panax consisted of about 20% mature, reproductive individuals and around 80% mixed immature plants and seedlings. Many plants were seen in the two locations. The populations appear to be self-maintaining and relatively safe from habitat loss because of the rugged topography of the areas in which they are growing. As long as these populations are not overexploited by “ginseng hunters,” which is a common practice (Cruise-Sanders & Hamrick 2004), Panax will most likely maintain populations at these sites. Although Panax was known from Lauderdale County previously, it apparently was not collected, as this collection represents a new county record for the species. Rhapidophy = lum hystrix (blue needle palm) was located along and between backwater sloughs dominated by Taxodium distichum (bald cypress) and Nyssa aquatica (water tupelo). They always were found inhabiting areas that were slightly elevated from the rest of the floodplain along the slough margins. Soils were of the Quitman loam series which are deep, poorly drained soils found on stream terraces that are strongly to very strongly acidic (Allgood 1983). Generally, plants were growing under a dense canopy provided by mature floodplain forests. The dominant canopy species included Acer barbatum, Carya glabra, Liguidambar styraciflua, Quercus hemisphaerica and Q. michauxii. The main understory component was Illicium floridanum. Other understory dominants included Lindera benzoin, Carpinus caroliniana, Ostrya virginiana and Symplocos tinctoria. Majure, Flora of Chunky River in Mississippi 1187 Overall, about 90 individuals among 3 populations were seen throughout the area. Several reproduc- tive palms were seen, as well as seedlings and immature individuals growing among the larger plants. Unfortunately, the area containing the highest number of R. hystrix has recently been heavily logged. Storm damage from Hurricane Katrina apparently prompted salvage logging at the site. Plants were seen that had been completely devastated by logging trails and machinery. It is uncertain what effects this disturbance will have on the overall population. Although, R. hystrix has been reported from Lauderdale County previ- ously (Watson 1972; Clancy & Sullivan 1990), these are the northernmost populations of the palm that have been found in Mississippi and are the largest known populations in the county, as a nearby population along Okatibbee Creek consisted of only four immature individuals (Clancy & Sullivan 1990). Rhynchospora globularis var. pinetorum (globe beaksedge) was found in a cutover within a SS community in moist sandy soils. This variety was rare and was only found in one location along the extent of the survey area. It is a new Lauderdale County record and extends the northern range of this variety in Mississippi. Schisandra glabra (bay starvine) was found in one location in a MF community along a northeast-fac- ing slope. Some of the vines were rather large, exceeding 30 m in height and 5 cm in diameter. There were about 30 stems of not more than a few plants. Most stems were the product of asexual reproduction, which subsequently formed small colonies. No plants were seen in flower or fruit. Common associate species were Podophyllum peltatum, Polystichum acrostichoides, Phegopteris hexagonoptera, Maianthemum racemosum, Liriodendron tulipifera, Quercus alba, Ilex opaca, Fagus grandifolia and Acer barbatum. Human disturbance could eventually play a factor in this species' existence at this site, as nearby trails have been cut along the slope where it grows. Tree-falls have already resulted from those actions. The avail- able habitat for this species directly along the extent of the river surveyed is not very extensive, so this is an important population. Also, this is a new county record for Lauderdale County. Staphylea trifolia (bladdernut) was found at the base of rich mesic slopes within a FF community. Three large groups of plants were seen, each consisting of about 400-500 clones. None of the plants were reproduc- ing sexually, although numerous plants were produced ieu through root sprouts. Common associate species were Aesculus pavia, Calycocarpum lyonii, Carex | tl difolia, Hexastylis arifolia, Magnolia acuminata, Phegopteris hexagonoptera, Polystichum acrostichoides a DON lycioides. Carex picta and Oxalis grandis were found in the RU community type, although, C. picta was seen on a more northerly slope that was subsequently more mesic in nature than the site where O. grandis was located. The population of C. picta consisted of about 100 large clumps, which were growing vigorously. Unfortunately, many of the plants had lost the majority of their floral parts at the time of collection, so it was difficult to determine the quantity of male versus female plants throughout the population. The O. grandis population consisted of about 30 individuals scattered over the mid-upper levels of a northwestern slope. The 2 locali- ties where C. picta and O. grandis were found appear to be relatively stable and free from large-scale human disturbance, because they are close to the river margins along steep slopes. O. grandis is a new county record for Lauderdale County. Ptelea trifoliata and Tridens carolinianus were found in the XO communities. Immature, or sterile P. trifoliata also was seen in the ADA, FF, MF, RB, RU and SS communities but was only seen reproducing in a few xeric and riverbank openings. The fruit of this species seems to be widely dispersed, but without a sufficient opening in the upper canopy, plants generally do not reproduce. This collection is a new county record for Newton County. Tridens carolinianus was seen in one XO community. The small population was made up of about 15 plants. This collection represents a new county record for Lauderdale County and a disjunct population, as the closest location for this species is from Wayne County (MISS) around 50 km south of this location. Fi A, 1 scirpoides were found in Dichanthelium erectifolium, Eleocharis baldwinii, F the RC community. Dichanthelium erectifolium was found Deal along a in the RC community but also was seen in an area adjacent to a slough system with an open canopy in moist, sandy soils. This is a new county record for Clarke County. It was recorded previously from Lauderdale County (Freckmann 1188 Journal of the Botanical R h Institute of Texas 1(2) & Lelong 20033). Eleocharis baldwinnii was at one location forming a small colony along the periphery of a sandbar in moist soil. This collection is a new county record for Clarke County and increases the species' known range to the north. The Mississippi Natural Heritage Program designation for this species is SNA (P), which means that this species is reported from the state but not confirmed. However, this species has been collected previously in various parts of the state and is confirmed from the state by Smith et al. (2002). The Mississippi Natural Heritage Program designation for Eleocharis baldwinnii can now be reconsidered. Fuirena pumila and R. scirpoides were seen occasionally or uncommonly along sandbars intermixed with ruderal species of Cyperus and other vegetation typical of sandy, moist, disturbed areas. These collections represent new county records for Lauderdale County. Although various collections from Mississippi exist for F. pumila (MISS), it is not listed for the state by Kral (2002). The closest record of R. scirpoides is from Forrest County, so this collection is disjunct by about 134 km to the northeast. Panicum brachyanthum was found in one location in an ADA community. Very few individuals were seen along the side of a disturbed, sandy river trail. This habitat seems to be typical for the species (Freckmann & Lelong 2003b). This collection represents a new county record for Lauderdale County. Rhynchospora mixta was at the base of a moderate slope in a SPS community under a moderate to dense canopy of mixed pine and deciduous forest. A robust population of about 200 plants, in flower and/or fruit, was seen at the time of collection growing closely associated with Saururus cernuus and Lycopus virginiana. This represents a new county record for Lauderdale County and a disjunct population of around 134 km to the northeast, as the closest collection known is from Forrest County. Notable collections Although not on the state's tracking or watch list, Carex dasycarpa was found in portions of the FF and SS communities along the extent of the survey area. This represents the first documented specimen of C. dasycarpa collected within the state of Mississippi in 112 years. The last specimen of this species recorded in the state was in 1894 by Tracy and Earl found in Jackson County (C.T. Bryson pers. comm.; specimen at MISSA). Therefore, this not only represents new county records for Lauderdale and Newton counties, but it also presents a widely disjunct population, about 320 km, from the previous location. Until seeing these specimens, Charles Bryson had considered delimiting this species as of “historical occurrence” for the state. Further study will be required to determine the extent of these populations and to decide if this species should be placed on the Mississippi Natural Heritage Program tracking or watch list. Ball (2002), states that this species is rare over most of its range, so it is likely to be rare for Mississippi as well. Digitaria horizontalis is only known from several counties in Mississippi but wasn’t previously known from Newton County. Wipff (2003) mentions that this species is tropical in origin and that; it is *probably a recent introduction to the Flora region." This collection is widely disjunct from previous collections. Wipff (2003) illustrates the closest collections in relation to Newton County as being from a north-central county 1 in Mississippi and from Alabama. One other collection by Tracy (s.n.; 1892) from Horn Island (Jackson Co., MS) was discovered at MISSA while updating the Panicum genus. Exotic/Invasive species All together, 79 species of exotic plants were found (9.896 of taxa recorded). Fifty-six of these species are ones deemed to be invasive by The Plants Database (2007). The most abundant of these invasive species were Chenopodium ambrosioides, Ligustrum sinense, Lonicera japonica, Microstegium vimineum, Pueraria lobata and Triadica sebifera. Chenopodium ambrosioides and Triadica sebifera were most common in the RC commu- nity with populations generally occurring on shifting sandbars and small “islands.” Chenopodium was seen throughout the survey area. Triadica was mainly seen in two areas. It formed a large colony on one small island above Stuckey's Bridge in Lauderdale County, and another large colony was seen below the Highway 11 Bridge north of Enterprise in Clarke County. Seedlings were seen on sandbars downriver from these larger colonies. Ligustrum sinense and Lonicera japonica were prevalent in portions of the ADA and FF communities. Majure, Flora of Chunky River in Mississippi 1189 Lonicera was seen mostly in a low vegetative form where canopy cover was dense, but, in canopy openings more robust, sexually reproductive plants were often found. Ligustrum fruited heavily and reproduced vegetatively by means of roots sprouts. Large colonies were found in various parts of the FF communities especially in young forests close to bridges and other previously disturbed areas. Pueraria lobata was seen in three locations, near Hwy. 11 in Clarke County forming an exten lon behind a residence, below Stuckey's bridge in Lauderdale County along the river margin adjacent to a ene house, and along a roadside and railroad track near the first boat launch in Newton County. Colonies of this species extended to the river's edge in certain instances. Seedlings were seen on several sandbars disjunct from any parental colonies, so kudzu is definitely moving along the main channel of the river. Microstegium vimineum was most prevalent in the FF and RC communities. This invasive grass forms a dense groundcover in the floodplain under heavy canopy cover. It also is frequent along the river margins especially in shaded areas (e.g., rockbars where trees overhang the river channel). The remaining invasive species seen along the survey area were not generally in large numbers except for Imperata cylindrica. However, the population of Imperata that was found around one of the boat launches in Lauderdale County was sprayed in 2005. Subsequently, no regrowth was seen. DISCUSSION Although many botanical collections exist for the state of Mississippi, no all inclusive flora has been com- pleted. Few floristic surveys have been published for the state of Mississippi (Alford 2001; Leidolf et al. 2002) relative to the number of counties (82) and overall area occupied by the state. Floristic surveys of reasonably large areas, such as the Chunky River, add valuable data to the paucity of information for poorly studied regions within the state. These surveys also provide bases for the fulfillment of county-level floras, of which only a few have been completed (e.g., Flint 1882; Ferrari 1970; Morris 1987; Winstead 1990; McDonald 1996; Alford 2001; Leidolf et al. 2002). The number of county records that were produced exemplifies the lack of collecting in this area. Several of these new county records increase the known distributions for rare species like Schisandra glabra and Rhynchospora globularis var. pinetorum. Other species such as Carex dasycarpa, which had not been seen in the state for more that 110 years, can be included in the contemporary flora for the state. Other collections such as Fuirena pumila, which is not recorded for the state of Mississippi in the Flora of North America, can be confirmed for the state. Mississippi plant species distributional information is occasionally overlooked in certain treatments within the Flora of North America, although, collections for a given species exist for the state (e.g., Majure 2007). Floristic surveys and herbaria searches can help alleviate this problem. Also, occurrence and distribution data for potentially problematic, invasive, exotic species is gathered through such surveys. Exotic, invasive species comprised 9.8 % of the flora found on the river. Exotic species are spread easily through highly disturbed areas, and certain species tend to prefer riparian zones for dispersion and resources. River margins and peripheral areas that frequently flood are common places for exotics to be- come established (Hood & Naiman 1999). It is difficult to say what overall effect the river has on propagule dispersal, but apparently invasive species are being moved around to a degree by annual flooding. There are a large number of invasive species that are common to the sandbars of the Chunky River system. Anthropogenic disturbance is a continuous threat to this ecosystem. In certain areas, large tracts of forest have been cut within 3-6 meters of the riverbank. This creates potential problems with erosion, and destroys a great deal of habitat for certain species (e.g., blue needle palm). Direct contamination of the river channel is evident especially during the heavily used summer season. Interestingly, some people who use the river for its scenic beauty also contribute to a great deal of waste (e.g., cans, bottles, garbage bags, and tents). Waste also is thrown into the river channel from bridges. More work should be done to insure the natural beauty of this area not be tainted by carefree outdoor enthusiasts. 1190 Journal of the Botanical R h Institut f Texas 1(2) APPENDIX 1 iff ollection number Plants included in this list were collected by the author and are followed by their sp unless otherwise noted (e.g., McDaniel 13369). State ranks are given for tracking or watch list species (S1- S4). The frequency at which a given species was observed is noted as common, occasional, uncommon or rds are denoted by C for Clarke, L for Lauderdale, and N for Newton County. Exotic species are denoted by * while exotic, invasive species rare. Community type acronyms follow species frequency. New county r are denoted by ** before the species name. FILICOPSIDA ASPLENIACEAE Asplenium platyneuron (L.) Britton, Stearns, & Poggenb. (87) on; FF, RU, ADA BLECHNACEAE Ii dunrdianan lata (L.) T. Moore (124) common; SPS Woodwardia virginica (L.) Sm. (431) occasional; SPS DENNSTAEDTIACEAE Pteridium aquilinum (L.) Kuhn (113) common; RU DRYOPTERIDACEAE h (111) common; MF, RB, S, SPS, Onoclea sensibilis L. (82) common; RC, S, SPS Polystichum acrostichoides (Michx.) Schott (112) common; I EYGODINGERE * Lygodium japonicum (Thunb.) Sw. (372) rare; ADA, RC OPHIOGLOSSACEA UP baies Spreng. (1914) uncommon; FF N Botrychium virginianum (L) Sw. (1 342) common; BF MF OSMUNDACEAE Osmunda cinnamomea L. (109) occasional; SPS Osmunda regalis L. (110) common; RB, RC, SPS POLYPODIACEAE Pleopeltis polypodioides (L.) E.G. Andrews & Windham subsp. michauxiana (Weatherby) E.G. Andrews & Windham (265) occasional; RU Mie Ania. hegopte a (Michx) Fée (96) common; MF E agreed 1o a (Gaudich.) Ching (506) rare; EREL e. kunthii (Desv) CV. Morton (661) rare; MF, RC L LYCOPODIOPSIDA SELAGINELLACEAE Selaginella apoda (L.) Spring (371) uncommon; RB, RC L PINOPSIDA CUPRESSACEAE cb dll L. (640) occasional; FF, SS L m (L.) Rich. (522) common; RB, RC, SL PINACEAE Pinus echinata Mill. (11 nus glabra Walter ( 91) common; RU, SS 259) common; FF, RU, SS Pinus palustris Mill. (1178) rare; RU, XO L Pinus taeda L. (1198) common; ADA, FF, RBL LILIOPSIDA AGAVACEAE Manfreda virginica (L.) Rose (1779) uncommon; RU Yucca filamentosa L. (92) occasional; FF, SS, XO ALISMATACEAE Sagittaria graminea Michx. var. chapmanii J.G. Sm. (1505) on; ADA, SL Sagittaria latifolia Willd. (427) occasional; ADA, S, SPS ALLIACEAE (LILIACEAE, in part) Allium canadense L. (1414) common; ADA, MF Nothoscordum bivalve (L.) Britton (1321) common; ADA, AMARYLLIDACEAE (LILIACEAE, in part) Zephyranthes atamasco (L.) Herb. (1371) uncommon; FF ABACERE (including LEMNACEAE) ch tt (148) " al; MF ree pty (L ) Schott subsp. M (116) com- mon; FF, Arisaema iN (L) Schott subsp. quinatum (Nutt.) Huttl. 521) occasional; MF ** Colocasia esculenta (L.) Schott (1 d 566) rare; REN emna minor L. (1427) common; SL Spirodela polyrrhiza Schleid. (1428) uncommon; SL Peltandra virginica (L) Schott (219) occasional; S L ARECACEAE Rhapidophyllum hystrix (Pursh) H. Wendl. & Drude (89) (S3) occasional; FF Sabal minor (Jacq.) Pers. (523) occasional; FF L BROMELIACEAE Tillandsia usneoides (L) L. (2549) rare; RC on Betula nigra C COLCHIACEAE bcd ic in part) Uvularia grandiflora S asi nal; M Uvularia perfoliata L. us al; MF Uvularia sessilifolia L. (436) EU FE MF L COMMELINACEAE lid diffusa Burm. f. (557) common; RCL mmelina erecta L. var. angustifolia (Michx.) Fernald (1606) uncommon; XO L Commelina virginica E (554) common; le RC, SL Tradescantia virginiana E (3 ) mino, FF, MF Majure, Flora of Chunky River in Mississippi CYPERACEAE Bulbostylis ciliatifolia (Ell.) Fernald var. coarctata occasional; XO L — Ell.) Kral (1192 — Carex abscondita Mack. (1485) occasional; FF, MF Carex albicans Willd. ex uo var. australis (L.H. Bailey) J.H 1338) ianal F, RU L Rettig ( create Schwein. (2497) occasional; FF, SN Carex atlantica L Bailey ate ae: capillacea (L.H. Bailey = Rezn id asional; ^ Carex aureolensis Steud. (1597) occasional; FF, S L Carex basiantha Steud. (1165) common; FF, MF L Carex bromoides Schkuhr ex Willd. (2499) common; FF, S N Carex caroliniana Schwein. (1372) UV REE Carex cephalophora Muhl. ex Willd.(20 mmon; MF, RUL Carex cherokeensis Schwein. (1393) occasi i d L arex crebriflora V d (1 364) uncommon; FF, dasycarpa Muhl. (1403, bd crest A L,N ae debilis Michx. (272) comm SPS Carex digitalis Willd. var. floridana i H. ae Naczi & Bryson (201) common; FF, MF Carex flaccosperma Dewey (1359) common; ADA, FF Carex floridana Schwein. (2502) common; FF, MF N Carex gigantea Rudge (364) rare; ADA, SL Carex intumescens Rudge RN common; ve L Carex joorii L.H. Bailey (618) SPS Carex kraliana Naczi & Bryson (221) Carex NA Wahlenb. subsp. nnum (Fernald) W. Stone (130) rare; SPS mmon: 1 MF L Carex longii ras (2496) common; FF, S Carex louisianica L.H. Bailey (1594) occasional; S L Carex lupulina Muhl. ex Willd. (1595) occasional; S L Carex lurida Wahlenb. (214) common; RC, S Carex muhlenbergii Schkuhr ex Willd. var. muhlenbergii (2503 ncommon; ADA, XO L Carex nigromarginata Schwein. (29, 1337) common; FF, MF, we E RUL, N Carex oxylepis Torr. & Hook. (1365) uncommon; FF L Carex picta Steud. (1373) uncommon; RU (S3/S4) Carex reniformis (L.H. Bailey) Small (2498) occasional; FF, S N Carex socialis Mohlenbr. & Schwegman (1439) rare; MF, RC Carex striatula Michx. (1 384) common Carex tenax Chapm. ex Dewey (336) occasional; XO (S2) ll tribuloides Wahlenb. (271) COMMON; EE: ex umbellata Schkuhr ex Willd. (2229) cou FFL a vulpinoidea Michx. (2362) occasional l RC Cyperus compressus L. (1592, 171 30 common; RCL, N Cyperus croceus Vahl (1196) common; ADA, R Cyperus echinatus (L.) Wood (1710) uncommon; RC Cyperus LOSS Muhl. (2649, 1664) common; ADA, RCL Cyperus DIT L. (518) common; ADA, RCL Cyperus filiculmis Vahl (1782) rare; XO L Cyperus haspan L. (1711) common; RC ** Cyperus iria L. (1720) common; Cyperus lupulinus (Spreng.) Marcks subsp. lupulinus (1767 — rare; XO L Cyperus odoratus L. (1783) common; RC Cyperus plukenetii Fernald (1768) rare; XO 1191 E BOTE DU mmon; RB, RC N Cyperu d. (329) common; RB, REL eee ee asional; ADA, XO ** Cyperus rotundus L. (1556) rare; ADA Cyperus strigosus L. (1719) common; RC N virens Michx. (1657) occasional; RC, SPS L Eleocharis baldwinii (Torr) Chapm. (1542) rare; RC; C SNA (P) Eleocharis microcarpa Torr. (1709) uncommon; REN Eleocharis obtusa (Willd.) Schult. (1570) common; RC Eleocharis tuberculosa (Michx.) Roem. & Schult. (1495) uncom- mon; > SPS L imbrich amitinmnniti yl (L) Roem. € Schult. (1162) common; RCL Fimbristylis decipiens Kral (1712) occasional; RC N Fimbristylis dichotoma (L.) Vahl (1821) uncommon; RCL Fimbristylis miliacea (L.) Vahl (1659) common; RC Fimbristylis tomentosa Vahl (1808) occasional; RC L Fimbristylis vahlii (Lam.) Link (1565) common; RC N Fuirena pumila (Torr.) Spreng. (2550, 1751) occasional; RC C, L (S3/S4) Isolepis carinata Hook. & Arn. ex Torr. (1474) uncommon; ADAL Kyllinga les Rottb. : 390) uncommon; ADA L Kyllinga odorata Vahl (1714) occasional; RC PORA Michx. d common; RC Rhynchospora caduca Ell. (1498) occasional; ADA, RC, S Rhynchospora corniculata te A. Gray (549) common; RC, S Hee meus (Chapm.) Small var. globularis (1602) ional; A TEMO ue (Chapm.) Small var. pinetorum Britton & Small) Gale (340) rare; SS L (S1) Rhynchospora glomerata (L.) Vahl (1648) common; S, SPS L Rhynchospora inexpansa (Michx.) Vahl (1647) common; S ospora mixta Britton (1494) uncommon; SPS L — (S3/S4) Rhynchospora scirpoides (Torr) A. Gray (1809) uncommon; Scirpus cyperinus (L.) Kunth (556) common; RC, S Scleria oligantha Michx. (220) common; MF, RU L Scleria triglomerata Michx. (1726) uncommon; XO L DIOSCOREACEAE Dioscorea villosa L. (121) common; MF ERIOCAULACEAE Lachnocaulon anceps (Walter) Morong (210) rare; ADA, SPS IRIDACEA Iris a Aiton (134) aid RUL Iris virginica L. (278) si Sisyrinchium Het EP. Bicknell (140) common; MF, RUL Sisyrinchium rosulatum E.P. Bicknell (1387) occasional; ADA L JUNCACEAE Juncus bufonius L. (1581) common; ADA, RC, S L Juncus coriaceus Mack. (432) common; ADA, S Juncus dichotomus Ell. (334) occasional; ADA, RC 1192 Juncus diff Buckley (216) common; ADA, RC, SL Juncus effusus L. Ga common; ADA, 2 Juncus elliottii Chapm ADAL Juncus marginatus Rostk. (339) D oT ADA Juncus repens Michx. (1504) uncommon; RC, Juncus scirpoides Lam. (343) occasional; ADA, RC L Juncus tenuis Willd. (337) common; ADA, X Juncus validus Coville (342) occasional; ADA, Luzula echinata (Small) FJ. Herm. (30) common; MF, RB, RU LILIACEAE Lilium michauxii Poir. (228) occasional; RU L MELANTHIACEAE (LILIACEAE, in part) Chamaelirium luteum (L.) A. Gray (1382) ional; MF Trillium stamineum Harb. (39) du FF, MF, RU ORCHIDACEAE orallot / HZG VVIOULCHIGOTIG onrad (38) rare; FFL Rich. (1241) rare; FF L lante s Rat (1762) rare; XO POACEAE ia ug (Walter) Britton, Stearns, & Poggenb. (1441) sional; ADA PUn Qin Nene Tuck. ep common; RB, RCL * Aira elegans Willd. ex Kunth (140 ADA RIOR cara Water eet 2e Sn ADA, RC N Stearns, & Poggenb. ar. r. scabriglumis C.S. Campb. (1907) uncommon; ADA L mue tracyi Nash (1855) uncommon L Andropogon virginicus L. var. decipiens cs n (1879) anth a (L ;ADAL Anthaenantia villosa (Michx.) P. Beauv. (1860) uncommon; XO Aristida dichotoma Michx. var. curtissii A. Gray (1188) com- on; ADA, XO Aristida lanosa Muhl. ex Elliott (1875) rare; XO Aristida longispica Poir. var. geniculata Fernald (1236) common; A, X Aristida purpurascens Poir. var. purpurascens (1187) common; Aristida purpurascens Poir. var. virgata (Trin.) Allred (1876) common; ADA, Arundinaria gigantea (Walter) Muhl. subsp. tecta (Walter) cClure (1340) Axonopus fissifolius (Raddi) Kuhlm. (1724) common; ADA L Brachyelytrum erectum (Schreb.) P. Beauv. (548) common; mmon; MF, RB, RC * Briza minor L. (1418) asional; ADA ** Bromus arvensis L. (1417) common; ADA Cenchrus spinifex Cav. (1658) S ADA, XO es (1610) common; FF Chasmanthium latifolium MEN Yates 435) common; FF, hasmanthitim laxm (EJ Yat B, RC ,SL Chasmanthium sessiliflorum (Poir.) Yates (504) common; FF, MF ** Cynodon dactylon (L) Pers. (1232) common; ADA, RC L Danthonia sericea Nutt. (333) occasional; ADA, SS, XO Dichanthelium EE Rb ex Poir) Gould & C.A. Clark p. aciculare (2 t [5 ADA, SS, XO £L Dat PRA D hi Air Journal of f Texas 1(2) Dichanthelium aciculare (Desv. ex Poir) Gould & C.A. Clark ubsp. angustifolium (1397) uncommon; ADA, SS, XO Dichanthelium acuminatum Gould €: C.A. Clark subsp. lind- heimeri (Nash) Freckmann & Lelong (1718) occasional; ADA, RC Miro boscii (Poir) Gould & C.A. Clark (538) com- N; F F, MF TRUM clandestinum (L.) Gould (1055) common; FF, RC, S Dichanthelium commutatum (Schult.) Gould subsp. joorii (Va mann & Lelong (1056) common; FF, RC, S Dichanthelium dichotomum (L) Gould subsp. dichotomum 45) common; FF, RC (4 Dichanthelium dichotomom (L.) Gould subsp. microcarpon Fre (Na sh) Fr sl elong 1473) O Dichanthelium erectifolium (Nash) Gould & C.A. Clark (2546) are; RC, S C (S3/S4) Dichanthelium laxiflorum (Lam.) Gould (1338) common; ADA, SS common: ; ADA, m 2 Dichanthelium Re pu Gould subsp. oligosan- thes (1 No) i | aa ovale de m & C.A. Clark subsp. villosis- simum (Nash) Freckmann & Lelong (341) common; XO Dichanthelium polyanthes (Schult) Mohlenbr. (1649) com- mon; FF, RB Dichanthelium ravenelii (Scribn. & Merr.)Gould (1337) oc- casional; XO Dichanthelium scoparium (Lam.) Gould (1054) common; RC, A Dichanthelium sphaerocarpon (Ell.) Gould (1465) common; Digitaria bicornis (Lam.) Roem. & Schult. ex Loud. (2536) mmon; RCL Digitaria cognata (Schult.) Pilg. (1654) occasional; XO Digitaria filiformis (L) Koeler var. filiformis (1243) occasional; ADA, XO Digitaria filiformis (L) Koeler var. villosa Fernald (1877) oc- asional; ADA * Digitaria horizontalis Willd. (1884) uncommon; ADA N ** Digitaria ischaemum (Schreb.) Muhl. (1830, 1883) common; RCL, Digitaria sanguinalis (L) Scop. (1164) common; ADA, RC * Digitaria violascens Link (1242) common; ADA L ** Echinochloa colona (L.) Link (1579) common; RC ** Echinochloa crus-galli (L.) P. Beauv. (1813) common; RC Echinochloa walteri (Pursh) in A. Heller (625) common; RCL ** Eleusine indica (L.) Gaertn. (1239) occasional; ADA, RC Elymus canadensis L. (1885) occasional; ADA, FF N Elymus virginicus L. var. virginicus (1488) common; ADA, FF, SL Eragrostis capillaris (L.) Nees (1811) rare; RCL Eragrostis curvula (Schrad.) Nees (1899) rare; ADA L Eragrostis hirsuta (Michx.) Nees (1181) occasional; RB, XO Eragrostis hypnoides (Lam.) Britton, Stearns, & Poggenb. (1668) common; RC Fragrostis pectinacea (Michx.) Nees (1588) common; ADA, RC NA. Eragrostis refracta (Muhl.) Scribn. (511) occasional; ADA, XO Majure, Flora of Chunky River in Mississippi * Glyceria fluitans (L.) R. Br. (1336) rare; ADA L Gymnopogon ambiguus Y Britton, Stearns, & Poggenb. (1757) occasional; X Hordeum pusillum Nutt. (1 m common; ADA ** Imperata cylindrica (L.) P. Beauv. (347) occasional; ADA L Leersia lenticularis Michx. (663) occasional; FF, S L Leersia oryzoides (L.) Sw. DA E RC,S Leersia virginica Willd. (662) common; FF, RB, RC, SL Nis cal ota (. A Hitchc. (2641) rare; RCL Lolium temulentum L. (1416) common; ADA Melica mutica Walter (40) common ** Microstegium vimineum (Trin.) A. Camus (527) common; — A, FF, RC Muhlenbergia schreberi J.F. Gmel. (1228) occasional; ADA L Oplismenus hirtellus (L.) P. Beauv. subsp. setarius (Lam.) Mez 1822) uncommon; FF L Panicum anceps Michx. (1616) common; ADA, XO Panicum brachyanthum Steud. (1725) rare; ADA L (S3/S4) Panicum dichotomiflorum Michx. subsp. dichotomiflorum 83) common; RC, SL Panicum rigidulum Bosc ex Nees subsp. rigidulum (1167) RGS Panicum verrucosum Muhl. (622) common; ADA, FF, RC, S Panicum virgatum L. (1177) common; RB, RC L Paspalum boscianum Flueggé (1752) common; RC ** Paspalum dilatatum Poir. (1788) common; ADA L Paspalum distichum L. (1170) a RC Paspalum fluitans (Ell.) Kunth (2646, 1716) common; RC L, N Paspalum laeve Michx. (1615) occasional; ADA, XO un Nee notatum Flueggé (1193) common; ADA, FF, RB, RC Paspalum setaceum Michx. var. ciliatifolium (Michx.) Vasey C. Doffitt (s.n.) common; ADA, XO Paspalum setaceum ee var, MCN (LeConte) Alph. Wood (1 XO, omm n; I A Paspalum setaceum Michx. var. PE im DJ. Banks (1911) uncommon; ADA inc setaceum Michx. var. stramineum (Nash) DJ. Banks 913) occasional; ADA N i cdi setaceum Hir var. villosissimum (Nash) D.J. Banks (1 A, X a = = T urville as (1573) occasional; ADA, Phanopyrum gymnocarpon (Elliott) Nash (1247) occasional; RC, SL E avenaceum (L.) Parodi (128) occasional; RU L * Poa annua L. (1301) common; A Poa autumnalis Muhl. ex Elliott (330) common; RB Poa chapmaniana Scribn. (1328) common; FF, RU Saccharum baldwinii Spreng. (1829) common; RB L Saccharum giganteum (Walter) Pers. (648) occasional; ADA, S ** Schedonorus phoenix (Scop.) J. Holub (1421) common; ADA Schizachyrium scoparium (Michx.) Nash var. scoparium (1189) com ; ,XO Schizachyrium scoparium (Michx.) Nash var. divergens (Hack) Gould (1 common; ** Secale cereale L. (1673) occasional; ADA L 1193 ** Setaria pumila (Poir J Roem. & Schult subsp pumila (1586) occasional; ADA, RC L d mm Bid (C. Mohr) Nash (701) occasional; RB, ** Sorghum halepense (L.) Pers. (1 en common; ADA, RB, RC Sphenopholis nitida Scribn. (13 Sphenopholis obtusata (Michx.) soba R 1420) common; ADA Sporobolus clandestinus (Biehler) Hitchc. (1226) common; XO Sporobolus junceus (P. Beauv.) Kunth (1185) uncommon; XO Sporobolus indicus (L.) R. Br. var. indicus (1233) occasional; ADAL Steinchisma hians (Ell.) Nash (1463) common; RC, S Stenotaphrum secundatum (Walter) Kuntze (1194) common; ,RBL Tridens carolinianus (Steud.) Henrard (1872) rare; XO L (S3/S4) Tridens flavus (L.) Hitchc. var. flavus (1195) common; ADA, RB Triplasis americana P. Beauv. (1186) rare; XO n du ad (Walter) Chapm. (1769) rare; m m darctvi ides (l EE (368) asional; B, RC Denon platyphylla TOR e Wright) R. D o 169) TN elliotea (Raf) Fernald (1346) common; ADA L Vulpia octoflora (Walter) Rydb. (331) occasional; ADA POTAMOGETONACEAE Potamogeton pulcher Tuck. (1510) common; SL Stuckenia pectinata (L.) Boerner (1506) common; S L RUSCACEAE (LILIACEAE, in part) Maian iE racemosum (L.) Link subsp. racemosum (95) occasi md m (Walter) Ell. (1392) uncommon; MF SMILACACEAE milax bona-nox L. (273) common; ADA, FF, RB, RC, SS Smilax ecirrata (Engelm. ex Kunth) S. Wats. (120) occasional; N MFL Smilax glauca Walter (274) common; ADA, FF, RB, RC, SS Smilax laurifolia L. (1641) rare; SPS Smilax pumila Walter (115) common; FF, RU, SS Smilax rotundifolia L. (267) common; ADA, FF, RB, SS Smilax smallii Morong (360) common; ADA, FF, RB, RC, SS L SPARGANIACEAE Sparganium americanum Nutt. (1534) common; SL TYPHACEAE Typha latifolia L. (1560) common; ADA, S C XYRIDACEAE Xyris jupicai Rich. (1652) occasional; S, SPS L Xyris laxifolia Mart. var. iridifolia (Chapm.) Kral (1642, 2648) uncommon; SPS L, N Xyris torta Sm. (455, 456) uncommon; ADA L MAGNOLIOPSIDA ACANTHACEAE Justicia ovata (Walter) Lindau (236) common; RC, SL 1194 Ruellia caroliniensis (Gmel.) Steud. (417) common; ADA ADOXACEAE (in part, CAPRIFOLIACEAE) dl nigra L. subsp. canadensis (L.) R. Bolli (1467) com- $ $ R rere Henan L. (473) common; FF, RB, SS L Viburnum nudum L. (1640) rare; SPS L Viburnum rufidulum Raf. (223) asi ALTINGIACEAE (in part, HAMAMELIDACEAE) Liquidambar styraciflua L. (263) common; ADA, FF, RB, RC, So: AMARANTHACEAE ** Altornmanthern philo ernide«c (Mart.) Griseb. (351) common; nal: ME RUL RC, Amaranthus spinosus L. (1584) occasional; RC L A coto a L. (1166) common; RCL ) Moq. (1244) rare; XO L Froelichia n NE Mog. (495) rare; ADA, XO L ANACARDIACEAE Rhus scd L. (472) common; ADA, RB, SS, XO Rhus glabra L. (605) common; ADA, RB Toxicodendron pubescens Mill. (474) occasional; SS, XO N Toxicodendron radicans (L) Kuntze (1363) common; ADA, F 459 F roelichia floridana (Nut (L) Kuntze (1662) rare; SPS ANNONACEAE Asimina parviflora (Michx.) Dun. (76) common; FF, MF, RU Asimina triloba (L) Dun. (222) common; FF, MF L APIACEAE Chaerophyllum tainturieri Hook. (1314) common; ADA Cicuta maculata L. var. maculata (1486) common; ADA Eryngium prostratum Nutt. ex DC. e occasional; S P O ranunculoides L. f. (1804) uncomm L H lydrocotyle verticillata Thunb. var. P (1 508) occasio- nal; R Piiimnium capillaceum (Michx. a Raf. (1 ud common; ADA C 37) common; mo A p K.M. Pryer & L. n p (533) oc- casional; M Sanicula pun ee (251) l; MI sl ~- (Nutt. ex DC) ale (1503) uncom- al adet 1515 [E (5 on; AD E ee A. Gray (197) occasional; MF, RU Trepocarpus aethusae Nutt. ex DC. (451) common; ADA APOCYNACEAE (including ASCLEPIADACEAE) Amsonia tabernaemontana Walter (1888) occasional: FF, MFN Asclepias amplexicaulis Sm. (323) rare; ADA, SS, XO Asclepias humistrata Walter (1605) uncommon; XO L Asclepias variegata L. (1502) common; FF, SS, XO Matelea gonocarpos (Walter) Shinners (1460) occasional; ADA, FF, MF L 4 Nal y (241) Para n; ADA, EE ME RC SS AQUIFOLIACEAE llex decidua Walter (136) common; FF, MF, RB, SS, XO £L Dag PRA D hi PEA Journal UI f Texas 1(2) llex glabra (L.) A. Gray (307) rare; FF, RB llex montana Torr. & A. Gray (1383) uncommon; MF, RU (S3) llex opaca Aiton (261) common; FF, MF, 1 91 Aa asi sional; RB, SS, XO N Ilex \ omitoria Aiton ( ARALIACEAE Aralia spinosa L. (1341) common; ADA, MF, RB Panax quinquefolius L. (93) rare; MF L (S3) ARISTOLOCHIACEAE Aristolochia serpentaria L. (198) common; FF, MF, SS, XO Aristolochia tomentosa Sims (498) common; FF, RB Hexastylis arifolia Small (32) common; FF, MF ASTERACEAE A F H iciifinti EN FE CON ; ADA Ambrosia trifida L. (1723) common; ; ADA N Baccharis halimifolia L. (1471) common; ADA Bidens aristosa (Michx.) Britton (647) common; ADA Bidens bipinnata L. (1229) common; roadsides, ADA L Bidens uod L. (283) occasional; RC Boltonia caroliniana (Walter) Fernald (635) common; ADA L Caphephous odoratissimus (J.F. Gmel.) Hebert (1873) rare; XO Pun gossypina (Michx.) Ell. d hyssopifolia (Nutt.) mple ( uncommon; X Chrysopsis mariana (L.) Ell. (643) common; ADA, XO Cirsium horridulum Michx. var. horridulum (1395) uncommon; ADA Conoclinium coelestinum (L) DC. (546) common; ADA, FF, S Conyza canadensis (L.) Cronqist var. canadensis (1614) com- A + Conyza canadensis (L.) Crongist var. pusilla (Nutt. (1747) common; ADA Conyza ramosissima Crongist (1791) common; ADA L Coreopsis lanceolata L. (1399) common; ADA Coreopsis major Walter (311) common; SS, XO N S db due (1161) occasional; RB, SS, XOL Cr (Nutt.) Raf. (598) common; ADA, XO Ela prostrata (L.) L. (1 600) common; ADA, RC, S Elephantopus carolinianus Raeusch. (1665) common; FF, Crono. Elephantopus tomentosus L. (639) common; ADA, FF, RB, Erechtites ES ) Raf. ex DC. (619) common; ADA, RC (L) Pers. (286) common; A o a (1313) common; ADA Erigeron pulchellus Michx. (1532) rare; MF Eupatorium album L. (600) uncommon; ADA, SS Eupatorium Rd (Lam. Small ex Porter & Britton (1859) ommon; Eupatorium FEIN L. var. hyssopifolium (1755) rare; XO Fupatorium perfoliatum L. i 770) rare; SPS Eupatorium rotundifolium L. (601) common; ADA, SS Eupatorium serotinum Michx. (624) common; ADA, RC, S Eurybia hemispherica (Alexander) G.L. Nesom (644) uncom- n; ADA, RU * Facelis retusa (Lam.) Sch. Bip. (1410) common; ADA Majure, Flora of Chunky River in Mississippi Gaillardia aestivalis (Walter) H. Rock (442) common; ADA, RB, XO Gamochaeta americana (Mill. Wedd. (1900) uncommon; ,RC Gamochaeta falcata (Lam.) Cabrera (335) common; ADA ta pensylvanica (Willd.) Cabrera (1422) common; ADAL Helenium amarum (Raf) H. Rock (463) common; ADA Helenium autumnale L. (1823) uncommon; FF Helianthus DAT " (626) common; ADA, SPS Helianthus giganteus L. (447) common; ADA, SS N elianthus dad e Torr. & A. Gray (1793) uncommon; — MF, RU Heterotheca subaxillaris (Lam.) Britton & Rusby (592) com- ; ADA, Hieracium gronovii L. (541) common; RU, SS, XO Hypochaeris glabra L. (1350) common; ADA lonactis linariifolius (L) Greene (1607) uncommon; XO L lva annua L. (1235) common; ADA Krigia caespitosa (Raf) Chambers (1330) common; ADA, RC Krigia virginica (L.) Willd. (1345) common; XO L Lactuca canadensis L. (1564) common; ADA Lactuca floridana (L.) Gaertn. (367) uncommon; ADA, FF Liatris squarrosa (L.) Michx. (1763) ra Melanthera nivea (L) Small (536) castor MF, RU Mikania scandens (L.) Willd. (205) common; ADA, RC Packera glabella (Poir) C. Jeffrey (149) rare ityopsis graminifolia (Michx.) Nutt. var. p (596) common; SS, XO Pluchea camphorata (L.) DC. (623) c AM ADA, RC, S, SPS Prenanthes altissima L. (1745) occasi RUL dognaphalium slum Hilliard & B4 Burtt subsp Wr lium (599) co Pyrrhopappus pur S DC. (462) common; ADA Rudbeckia hirta L. (422) common; xeric, sandy areas; ADA, RB, SS Rudbeckia laciniata L. (1792) rare; FF, RB L Sericocarpus tortifolius (Michx.) Nees (608) occasional; ADA, XO e asteriscus L. var. asteriscus (319, 461) occasional; DA, XO o uvedalius (L) Mack. ex Small (244) occasional; AMM n; , RU idea a (Elliott Torr. & A. Gray (542) common; RB, A] RU, SS, XO Solidago gigantea Aiton (603) common; ADA Solidago nemoralis Aiton (602) common; RB L Solidago odora Aiton (627) common; ; RB, RU, SS, XO Solidago rugosa Mill. (636) common; RB, RU Solidago ulmifolia Muhl. ex Willd. var. microphylla Gray (590) common; RB, SS, X ** Sonchus asper (L.) Hill (1299) common; ADA xdi d cordifolium (L.) G.L. Nesom (1797) occasional; jed Smotyoridum lateriflorum (L.) A. Lóve €: D. Lóve var. lateri- EEI ATES \ È 1195 Symphyotrichum patens (Aiton) G.L. Nesom var. patens (634) CO ; RB, RU, SS Symphyotrichum pilosum (Willd.) G.L. Nesom (1909) com- mon; ADA re occidentalis (L ) Walter (1856) rare; FF rbesina virginica L. var. virginica (1844) rare; n eeu walteri Shinners (558) occasional; FF, Vernonia gigantea Trel. subsp. gigantea (1 e common; FF Xanthium strumarium L. (1825) occasional; RC L BALSAMINACEAE Impatiens capensis Meerb. (2544) common; FF, RC, SC PFRBFRIDA CEAE Nandina domestica Thunb. (1666) uncommon; FF N Podophyllum peltatum L. (36) occasional; MF BETULACEAE Alnus serrulata (Aiton) Willd. (512) common; RB, RC Betula nigra L. (235) common; FF, RB, R Carpinus caroliniana Walter (226) common; FF, RB, RC, S Ostrya virginiana K. Koch (85) common; FF, MF, RU L BIGNONIACEAE Bignonia capreolata L. (143) common; RB, RC Campsis radicans (L.) Seem. (207) common; RB, RC alpa speciosa (Warder) Engelm. vM common; RB, RC L pad elio tropium indicum E (3 n; RC M macrosperma Engelm. (1 366) ¢ common; FF L BRASSICACEAE Cardamine concatenata (Michx.) Sw. (147) uncommon; FF, MFL ** Cardamine hirsuta L. (1326) common; ADA N Cardamine pensylvanica Muhl. ex Willd. (1362) occasional; RC SL Lepidium virginicum L. (361) common; ADA nus raphanistrum L. (281) uncommon; ADA, RC L Rorippa palustris (L.) Besser (208) common; RCL Rorippa sessiliflora (Nutt.) Hitchc. (209) occasional; RC MBACE Brasenia schreberi J.F. Gmel. (1507) common; SL CACTACEAE Opuntia pusilla (Haw.) Haw. (309) occasional; ADA, XO N CAMPANULACEAE Lobelia cardinalis L. (613) common; RC Lobelia puberula Michx. (631) common; ADA, XO Lobelia spicata Lam. (1491) rare; ADA ene perfoliata (L) Nieuwl. (1354) common; ADA Wahlenbergia marginata (Thunb.) A. DC. (418) occasional; ADA L CANNABACEAE (ULMACEAE, in part) Celtis laevigata Willd. (1348) occasional; FF, RB Celtis tenuifolia Nutt. (288) occasional; RU, SS, XO CAPRIFOLIACEAE ** Lonicera japonica Thunb. (502) common; ad FF, RB, SS Lonicera sempervirens L. (1343) 1196 CARYOPHYLLACEAE * Arenaria serpyllifolia L. (1320) occasional; ADA N * Cerastium glomeratum Thuill. (1302) common; ADA decumbens Torr. & Gray (1388) occasional; ADA L Silene antirrhina L. (1351) common; ADA * Stellaria media (L.) Vill. (1332) common; ADA CELASTRACEAE Euonymus americana L. (441) common; FF, MF CERATOPHYLLACEAE o Ihiiy Heormercii m L. (280) common; SL CISTACEAE Lechea mucronata Raf. (425) occasional; ADA, XO L Lechea tenuifolia Michx. (326) occasional; ADA, XO L CLEOMACEAE (CAPPARACEAE, in part) * Cleome hassleriana Chod. (375) common; RCL CONVOLVULACEAE Dichondra li is Michx. (1386) occasional; ADA L Ipomoea cordatotriloba Dennst. var. cordatotriloba (420) common; AD ** Ipomoea hederacea Jacq. (1789) common; ADA Ipomoea lacunosa L. (1816) common; RCL Ipomoea pandurata (L.) G. Mey. (359) common; ADA, FF, RB, Jacquemontia tamnifolia (L) Griseb. (595) common; ADA Stylisma humistrata Chapm. (416) common; SS, XO pei ornus drummondii C.A. Mey. (378) occasional; FF, RB L rin florida L. (250) common; FF, MF, RB CUCURBITACEAE * Cucumis melo L. var. reticulatus Naud. (1557) rare; ADA N (nomenclature follows Bailey 194 Melothria pendula L. (1598) occasional; ADA, FF, SL EBENACEAE Diospyros virginiana L. (242) common; ADA, FF, MF, RB, RU, ERICACEAE Gaylussacia dumosa (Andr.) Torr. & Gray (2542) uncommon; SS Kalmia latifolia L. (150) common; RB, RU Leucothoe racemosa A. Gray (1918) occasional; RC, S N Lyonia lucida K. Koch (78) rare; RB Oxydendrum arboreum (L.) DC. (511) Rhododendron canescens Porter. (77) RU asional; MF, RB, RU common; MF, RB, RC, Vaccinium arboreum Marsh. (316) common; RB, SS, XO Vaccinium darrowii Camp McDaniel (13369) not seen; SS Vaccinium elliottii Chapm. (276) common; FF, RB, SS Vaccinium stamineum L. (315) common; RB, SS, XO mild mds Acalypha gracilens A. Gray (466) DA Acalypha epa Riddell (1553) common; de N Acalypha rhomboidea Raf. (515-aux.) DA, RC Chamaesyce maculata (L.) Small (586) common; y ADA N ommon; A £L Dag PRA D hi PEN Journal UI f Texas 1(2) Chamaesyce prostrata (Aiton) Small (1554) common; ADA, RCN Cnidoscolus stimulosus (Michx) Engelm. & Gray (322) com- Croton capitatus Michx. (594) occasional; ADA Croton glandulosus L. (468) occasional; ADA roton willdenowii G.L. Webster (467) occasional; ADA, XO L Ditrysinia fruticosa (W. Bartram) Govaerts & Frodin (204) com- FF, MF, RB, RU, SS, XO Euphorbia corollata L. (313) common; FF, MF, RB, RU, SS, XO ** Euphorbia dentata Michx. (588) uncommon; oe L Euphorbia pubentissima Michx. (1799) mon; MF, RUL Stillingia sylvatica L. (321) asional; XO N Tragia cordata Michx. (547) rare; cl L Tragia urens L. (325) occasional; A ** Triadica sebifera (L.) Small (515) pan RCL FABACEAE ET ac Durazz. (514) common; RC 9) common; RB, RC Amphicarpaea bracteata (L.) Fernald (638) common; ADA, MF == = Amorpha fruticosa > 15 Apios americana Medik. (496) common; RB, RC Centrosema virginianum (L.) Benth.(327) common; ADA, SS, XO Cercis canadensis L. (255) asional; MF, RB Chamaecrista fasciculata (Michx.) Greene var. fasciculata (589) ommon; ADA Chamaecrista nictitans Moench subsp. nictitans (1746) oc- casional; ADA, RC Clitoria mariana L. (443) occasional; ADA, SS, XO Crotalaria rotundifolia Walter ex J.F. Gmel. (606) rare; ADA, L Crotalaria sagittalis L. (458) common; ADA, SS, XO ** Crotalaria spectabilis Roth (1898) uncommon; ADA C Desmanthus illinoensis MacMill. (426) rare; ADA Desmodium glabellum (Michx.) DC. (633) common; ADA, SSL Desmodium nudiflorum (L.) DC. (530) common; MF L Desmodium obtusum DC. (642) common; ADA, SS L Desmodium paniculatum (L) DC. (1897) uncommon; RC Desmodium pauciflort Im (Nutt.) DE (532) common; MF Desmodium strictum (Pursh) DC. (630) common; ADA, SS L Desmodium viridiflorum (L) DC. (641) common; ADA, SS L Dioclea multiflora C. Mohr (369) common; FF, MF, RB, RU, SS a herbacea L. (256) occasional; MF, RB, RU, SS, XO L Galactia volubilis (L.) Britton (545) common; ADA, SS, XO ** Kummerowia striata (Thunb.) Schindl. (1679) common; ** Lesped uneata G. Don (421) common; ADA fue hirta (L poH (1894) uncommon; ADA, XO Lespedeza [ Michx. (607) common; ADA, XO pedeza stuevei Nutt. (604) |; ADAL pes violacea (L. Pers. (1179) uncommon; ADA, XO virginica (L.) Britton (1756) uncommon; XO nonse um Dryand. (365) occasional; XO Orbexilum pedunculatum (Mill.) Rydb. var. psoralioides (Walter) Isely (1402) uncommon; ADA L2 Majure, Flora of Chunky River in Mississippi Phaseolus polystachios (L) Britton, Stearn, & Poggenb. var. polystachios (1748) uncommon; MF, RU **Pueraria montana (Willd.) Merr. var. lobata (Willd.) Maesen . Almeida (1563) asional; ADA SS Rhynchosia reniformis DC. (1764) uncommon; XO L Rhynchosia tomentosa (L.) Hook. & Arn. (1794) rare; RUL Robinia pseudoacacia L. (108) common; RB L Senna obtusifolia (L) H.S. Irwin €: Barneby (1678) common; ADAL Sesbania herbacea (Mill) McVaugh (1677) common; ADA, RC ** Sesbania punicea (Cav.) Benth. (379) common; ADA, RC L Sesbania vesicaria Ell. (1895) uncommon; ADA C Strophostyles helvula (L.) Ell. (587) common; ADA, RC dn leiosperma (Torr. & A. Gray) Piper (1197) rare; ADA oc biflora 7 ) Britton, Stearn, & Poggenb. (430) oc- casional; ADA, X Tephrosia spicata T Torr. & A. Gray (1240) occasional; ADA, XO Tephrosia virginiana (L.) Pers. (320) sional; SS, XO ** Trifolium arvense L. (1487) common; ADA L ** Trifolium campestre Schreb. (1318) common; ADA ** Trifolium incarnatum L. (1352) common; ADA Vicia caroliniana Walter (1391) occasional; MF * Vicia grandiflora Scop. (1311) common; ADA N ** Vicia sativa L. subsp. nigra (L) ae (1310) common; ADA ** Vicia tetrasperma (L.) Schreb. (1312) common; ADA N ** Vicia villosa Roth (1483) occasional; ADA Wisteria frutescens (L.) Poir. (233) occasional; RB C Zornia bracteata J.F. Gmel. (529) rare; XO L AGACEAE Castanea pumila (L.) Mill. (122) occasional; RU, SS Fagus grandifolia Ehrh. (262) common; FF, MF, RB, RU Quercus alba L. (260) common; FF, MF, RU Quercus austrina Small (508) occasional; FF L Quercus falcata Michx. (440) occasional; RU Quercus hemisphaerica Bartr. (317) common; D RB,SSL Quercus incana Bartr. (314) asional; SS, X Quercus laurifolia Michx. (437); occasional; i L Quercus lyrata Walter (268) occasional; RC, S Quercus margaretta Ashe ex Small (470) occasional; SS, XO Quercus michauxii Nutt. (526) common; FF, MF, RB Quercus muehlenbergii Engelm. (248) uncommon; MF, RUL uda uon L. (499) common; FF, MF, RB, RU, SS Raf. (1749) occasional; FF, MF, RU L d shumardi o (1750) rare; MF L uercus stellata War 19er 1. (2547 ) uncommon; RU L Quercus velutina Lam. (539) common; FF, MF, RU GELSEMIACEAE (LOGANIACEAE, in part) G elsemium sempervirens (L.) J. St. Hil. (45) common; ADA, FF, MF, RB, SS GENTIANACEAE Sabatia campanulata (L.) Torr. (454) rare; ADA L GERANIACEAE Geranium carolinianum L. (1315) common; ADA 1197 ** Geranium dissectum L. (1316) occasional; ADA N idos juaticum (Vell) Verdc. (279) common; S L Myr P hera Michx. (1535) uncommon; S L Proserpinaca palustris L. (1360) common; S L HAMAMELIDACEAE Hamamelis virginiana le (1 18) common; MF, RB HYDRANGEACEAE Decumaria barbara L. (83) mmon: ME RR lydrangea arb ns L. (224) common; MF, RU Hydrangea folia Bartr. (531) common; RB, RU HYDROLEACEAE Hydrolea quadrivalvis Walter (1567) occasional; RC N Hydrolea uniflora Raf. (1575) occasional; RC, SL HYPERICACEAE Hypericum crux-andreae (L.) Crantz (1781) uncommon; XO Hypericum drummondii (Grev. & Hook.) Torr. & Gray (1655) CO ,XOL Hypericum frondosum Michx. (444) common; FF, RBL Hypericum galioides Lam. (1527) common; RB, REL Hypericum gentianoides (L.) Britton, Stearn, & Poggenb. (612) ; ADA, XO Hypericum hypericoides (L.) Crantz (324) occasional; ADA, XO Hypericum mutilum L. (428) common; ADA pu setosum L. (1878) rare; XO L Triadenum walteri (J.F. Gmel) Gleason (1184) common; RC, SPS ILLICIACEAE Illicium floridanum Ellis (42) common; FF, MF ITEACEAE Itea virginica L. (196) common; FF, RB, RC, S JUGLANDACEAE Carya aquatica (Michx. f) Nutt. (1480) uncommon; FF, RB L lis (Ashe) Engl. & Graebn. (1529) HUSO WANI d a sept MF, RU JL Carya 2. i Mill) Sweet (1 e EE (Wangenh.) K. Koch (501 180) common; FF, RB ) uncommon; m p" (Michx. f) Nutt. (1493) occasional; FF L Carya ovalis (Wangenh.) Sarg. (500) common; FF, RB Carya pallida (Ashe) Engl. & Graebn. (1661) occasional; FF, RUL LAMIACEAE Blephilia ciliata (L) Benth. (1526) uncommon; MF os llicarpa americana L. (247) common; FF, MF, SS * Lamium amplexicaule L. (1300) common; ADA Lycopus rubellus Moench (1896) rare; REC ud virginicus L. (1674) common; RC, S narda punctata L. (593) common; ADA, XO Perilla frutescens (L) Britton (1 ps 869) UNCOMMON; ADAL la vulgaris L. (433) rare cnanthemum albescens Torr & A. Gray (559) occasional; 1198 Pycnanthemum tenuifolium Schrad. (543) occasional; ADA Salvia lyrata L. (135) common; ADA, RC Scutellaria elliptica Muhl. ex Spreng. var. elliptica (475) occasional; ADA, SS Scutellaria integrifolia L. (285) common; ADA, SS Scutellaria ovata Hill subsp. ovata (1795) rare; MF L Um ll L. (521) occasional; FF, S L ] um L. (629) common; ADA, XO Who ems setaceum SUE (1861) rare; XO LAURACEAE Lindera benzoin (L) Blume (43) common; FF, MF L Sassafras albidum (Nutt.) Nees (346) common; FF, MF, SS LENTIBULARIACEAE Utricularia gibba L. (1852) occasi LINACEAE Linum medium (Planch.) Britton var. texanum (Planch.) Fern. (382) common; ADA L Linum striatum Walt. (429) common; ADA LINDERNIACEAE (SCROPHULARIACEAE, in part) us dubia (L.) Pennell var. dubia (373) common; ADA, nak SI Mdl, > L Tim dubia (L.) Pennell var. anagallidea (Michx.) Cooper- rider (1558) common; ADA, LOGANIACEAE Mitreola petiolata (J.F. Gmel) Torr. & A. Gray (544) occasional; Spigelia marilandica (L.) L. (243) common; MF LYTHRACEAE Ammannia coccinea Rottb. (1717) common; RC Ammannia latifolia L. (1582) asional; RCL * Cuphea carthagenensis (Jacq) J.F. a (519) common; A RCL Rotala ramosior (L.) Koehne (525) common; RC, S MAGNOLIACEA On ds 15252) E FF, MF (L) E (119 acj al; ME RIJ o grandior L (266) common; FF, M Magnolia macrophylla Michx. (227) a i MF, RU Magnolia virginiana L. (438) lS SPS ALVACEAE Callirhoe triangulata (Leavenw.) A. Gray (1760) rare; XO (S1/S2 Hibiscus moscheutos L. (1552) occasional; ADA, S Sida rhombifolia L. (446) common; ADA, RC L Sida spinosa L. (621) common; ADA, RC L MELASTOMATACEAE Rhexia mariana L. var. mariana e common; ADA, RCL nal; RE, SL RI HEAT virgit HCG [5 MELIACEAE ** Melia azedarach L. (1357) MENISPERMACEAE Calycocarpum lyonii (Pursh) A. Gray. (497) common; i MF aie eu (L.) DC. (234) common; FF, MF, S | PRY E £L Dat PRA D hi PEA Journal UI f Texas 1(2) MOLLUGINACEAE Mollugo verticillata L. (516) common; ADA, RC MONOTROPACEAE Monotropa uniflora L. (1501) uncommon; SS, XO L RACEAE Morus rubra L. (258) common; FF, MF MYRICACEAE Morella cerifera (L.) Small (584) common; RB, RC, SPS NYMPHAEACEAE Nymphaea odorata Aiton (1509) common; S NYSSACEAE Raw aquatica L. (231) common; RC, S sa biflora Walter (51 ) common “RO, SL ae sylvatica Marsh. (1404) common; MF, RU, SS OLEACEAE Chionanthus virginicus L. (138) occasional; RB, RU, SS Fraxinus americana L. (439, 1889) common; FF, MF, RB L, N Fraxinus pennsylvanica Marsh. (232) common; FF, RB ** Ligustrum sinense Lour. (503) common; ADA, FF, RB, SS Osmanthus americanus (L.) Benth. & Hook.f. ex Gray (81) common; RU, SS ONAGRACEAE Gaura Rd od (591) rare; ADA, XO .(552) common; RC, S, SPS ion decurrens Walt. (1864) common; RC, S Ludwigia glandulosa Walt. (553) common; RC, S Ludwigia leptocarpa (Nutt.) Hara (1753) common; RC, S Ludwigia linearis Walt. (1772) uncommon; RC Ludwigia palustris (L.) Ell. (277) common; RC, S i Penes (Kunth) Raven (1805) uncommon; RC common; ADA Oenothera laciniata Hill (349) common; ADA Oenothera speciosa Nutt. (1400) common; ADA OROBANCHACEAE ORO BHU EARIACERE: in part) Agalinis obtusifolia Raf. (1908) uncommon Agalinis setacea (J.F. Gmel.) Raf. (632) ol ADA, XO L Aureolaria flava (L.) Farw. (628) common; RU, SS, XO Fpifagus virginiana (L.) W.P.C. Barton (665) common; MF Pedicularis canadensis L. (94) occasional; RU Seymeria pectinata Pursh (597) common, XO OXALIDACEAE Oxalis dillenii Jacq. (1368) common; ADA N Oxalis grandis Small (1334) uncommon; RU L Small subsp. colorea (1333) rare; RU Oxalis stricta L. (328) common; ADA, RC Oxalis violacea L. (125) common; RU PAPAVERACEAE Sanguinaria canadensis L. (37) occasi PASSIFLORACEAE Passiflora incarnata L. (1528) uncommon; ADA Passiflora lutea L. (137) common; RB, RC, SS PENTHORACEAE Penthorum sedoides L. (2545) common; FF, RC, SC enothera biennis ls (5 ~ xalis vi icede nal ME dl, VU Majure, Flora of Chunky River in Mississippi PHRYMACEAE (SCROPHULARIACEAE, in part) Mimulus alatus Aiton Es ets je » L Phryma leptostachya L. ( al; MF L E (EUPHORBIACEAE, in part) hyllantht Walter (370) occasional; ADA, RCL p e urinaria L. (1230) occasional; ADA, RC L PHYTOLLACACEAE Phytolacca americana L. (1234) occasional; ADA, RB, RC PLANTAGINACEAE (SCROPHULARIACEAE, in part) Gratiola A Torr. o occasional; ADA, RC, SL ADA (A ianal Gratiola pilosa Mich IA. VT a>IUIlal, E 1339) ianal Gratiola VH UH HUTU dolOlldl f RC, Mecardonia acuminata (Walter) Small (610) occasional; ADA, RC e umbrosum (J.F. Gmel.) Blake (1589) occasional; Nuttallanthus canadensis (L) D.A. Sutton (1327) occasional; aristata Michx. (345) common; ADA ** Plantago lanceolata L. (424) common; ADA Plantago rugelii Decne. (452) common; ADA L Plantago virginica L. (1409) common; ADA ** Veronica arvensis L. (1319) common; ADA N Veronica peregrina L. (1329) occasional; RC N PLANTANACEAE Platanus occidentalis L. (44) PODOSTEMACEAE Dod. f, WE, Plantag mmon; RB, RCL hyllum Michx. (1458) common; REN POLEMONIACEAE Phlox pilosa L. subsp. pilosa (114) common; MF POLYGALACEAE Polygala cruciata L. (457) occasional; ADA, SPS L Polygala grandiflora Walter (350) common; ADA, XO Polygala incarnata L. (1482) rare; ADA Polygala mariana Mill. (1672) rare; ADA Polygala nana (Michx.) DC. (194) common; ADA, SS, XO POLYGONACEAE Brunnichia ovata (Walt) Shinners (210) common; RB, RCL Polygonum caespitosum Blume var. longisetum (Bruijn) Stew- ard (1479) common; RCL Polygonum glabrum Willd. (1812) occasional; RCL dul idit Michx. (1464) common; RC, S um L. (620) common; RC, S onum virginianum L. (1824) uncommon; FF x eda crispus L. (282) rare; RC Rumex hastatulus Baldwin (318) common; ADA, XO PORTULACACEAE Claytonia virginica [s (91) common; FFL ** Portulaca oleracea L. (1585) uncommon; RC L RANUNCULACEAE Clematis glaucophylla Small (367) occasional; ADA, SS, XO (S1) Clematis virginiana L. (1497, 1847) occasional; ADA, RB, RC ! 1199 Hepatica nobilis Schreb. var. obtusa (Pursh) (139) occasional; MF, RU Ranunculus abortivus L. (1331) uncommon; RC N Ranunculus bulbosus L. (377) rare; RC Ranunculus pusillus Poir. (1361) rare; RC Ranunculus recurvatus Poir. (1531) rare; MF ** Ranunculus sardous Crantz (1476) occasional; ADA, RC Thalictrum thalictroides (L.) AJ. Eames & B. Boiv. (50) com- mon; M Xanthorhiza simplicissima Marsh. (80) asional; RB RHAMNACEAE Berchemia scandens (Hill) K. Koch (507) common; FF, MF, RB Frangula caroliniana (Walter) A. Gray (245) occasional; MF ROSACEAE Agrimonia microcarpa Wallr. (646) uncommon; ADA, RU Amelanchier arborea (Michx. f) Fernald (117, 513) common; B, RU Crataegus marshallii Eggl. (79) occasional; RB L Crataegus uniflora Münchh. (287) common; SS L * Duchesnea indica (Andrews) Focke (1322) occasional; ADA Photinia pyrifolia (Lam.) K.R. Robertson & J.B. Phipps (614) S Potentilla simplex Michx. (1429) common; ADA Prunus angustifolia Marsh. (1405) uncommon; SS, XOL Prunus caroliniana E ) Alten (46) common; RB L Prunus serotina Ehr FRB SS Prunus umbellata Ell. (90) e le RB, SS Pyrus calleryana Decne. (2548) uncommon; RB € Rosa carolina L. (1415) common; RB, ** Rosa multiflora Thunb. ex Murray (146) occasional; RB L Rubus argutus Link (1440) common; RB, Rubus flagellaris Willd. (332) common; ADA, RC L Rubus trivialis Michx (47) common; ADA, RCL RUBIACEAE phal th is L. (509) common; RC, S Diodia teres Walter (449) common; ADA N Diodia virginiana L. (284) common; ADA, RC Galium aparine L. (1353) common; A Galium circaezans Michx. (253) common; MF calar pilosum Aiton (423) occasional; ADA, i i Galium tinctorium (L.) Scop. (1599) occasior Galium uniflorum Michx. (200) common; MF, - RU, sS Houstonia purpurea L. (229) common; RB Houstonia pusilla Schópf (1325) common; ADA N Mitchella repens L. (195) common; FF, MF Oldenlandia boscii (DC.) Chapm. (465) common; ADA, RC Oldenlandia uniflora L. (1758) common; R Richardia scabra L. (517) common; RC * Sherardia arvensis L. (1411) common; ADA L RUTACEAE Ptelea trifoliata L. (144) occasional; FF, MF, RB, RU, SS, XO N (S3/S4) SALICACEAE dr pO Bartram ex Marshall (471) uncommon; "e aos ies (213) common; RC, S 1200 SANTALACEAE Phoradendron leucarpum (Raf) Reveal & M.C. E (550) occasional; FF parasitic on Acer barbatum SAPINDACEAE (including HIPPOCASTANACEAE) Acer barbatum Michx. (86) common; FF, MF Acer negundo L. (48) common; FF, RB, RC Acer rubrum L. (249) common; FF, MF, S Aesculus pavia L. (26) common; FF, MF, RB SAPOTACEAE Sideroxylon lycioides L. (225) occasional; FF, MF SAURURACEAE Saururus cernuus L. (275) common; S SAXIFRAGACEAE Heuchera americana L. (1490) rare; RU L SOLANACEAE Sd V Mill. (1475) uncommon; ADA, RC lanum e L. (211) common; RC ce Pd L (1807) uncommon; RC L Physalis pubescens L. var. pubescens (1583) uncommon; RC Physalis virginiana Mill. (1530) uncommon; MF L N STAPHYLEACEAE Staphylea trifolia L. (1525) uncommon; FF, MF (S3) £L Dag PRA D hi PEA Journal UI f Texas 1(2) TETRACHONDRACEAE (BUDDLEJACEAE, in part) Poh premumM pro umbens L. (450) common; ADA, RC ACEAE Planera aquatica J.F. Gmel. (145) occasional; RC, SL Ulmus alata Michx. (469) common; FF, MF, RB, SS Ulmus americana L. (528) occasional; FF, RB Ulmus rubra Muhl. (1394) common; FF, MF, RU L URTICACEAE Boehmeria Pos (L.) Sw. (505) common; RC, S Pilea pumila (L) A. Gray (524) occasional; RC, SL Urtica Vienn que Pursh (1461) uncommon; ADA, FF L VALERIANELLACEAE Valerianella radiata (L.) Dufr. (1317) common; ADAN VERRENACPAE brasiliensis Vell. (363) common; ADA o n Small (419) common; AD * Verbena rigida Spreng. (1231) occasional; ADA VIOLACEAE Viola affinis LeConte (27, 31) common; FF, RB, RC N Viola xprimulifolia L. (pro sp.) [lanceolata x macloskeyi] (33) common; RB, RC Viola sororia Willd. x uncommon; pu L Viola walteri House (28) comm F, RU STYRACACEAE Halesia carolina L. (84) common; FF, RB Styrax americanus Lam. (246) occasional; FF, MF, S L Styrax grandifolius Aiton (1492) uncommon; RU SYMPLOCACEAE Symplocos tinctoria (L.) UHér. (88) common; FF, MF, RB VITACEAE Ampelopsis arborea (L.) Koehne (206) common; RB, RC L Parthenocissus quinquefolia (L.) Planch. (257) common; FF, F, RB, Vitis cinerea (Ei gelr n D Millardet (1 457) common; RBL Vitis palmata Vahl (1462) common; FF, RB L Vitis rotundifolia Michx. (254) common; FF, MF, RB, SS, XO ACKNOWLEDGMENTS Partial funding for this project was provided by the Mississippi Museum of Natural Science in Jackson, Mis- sissippi. Terrence C. Majure provided necessary resources and extended a helping hand with some of the field work. I would like to thank Andrew Whitehurst for sharing information about the Mississippi Scenic Streams Program with me. Heather Sullivan (MMNS) helped me locate county level and plant population data, and reviewed some specimen determinations. Chris Doffitt helped search for records at IBE. I would like to thank Charles T. Bryson (SWSL) for examining species within the Cyperaceae collected during this survey and for providing valuable information about the Carex in Mississippi. Sidney McDaniel gladly let me explore IBE in search of records. Ron Altig, Charles T. Bryson, Gary N. Ervin, and another anonymous reviewer provided helpful comments on an earlier draft of this manuscript. I would also like to thank landowners for the use of some of their property for this research. REFERENCES ALFORD, M.H. 2001. The vascular flora of Amite County, Mississippi. Sida 19:645-699. AtLGoop, V.C. 1983. Soil survey of Lauderdale County, Mississippi. United States Department of Agriculture, Soil Conservation Service, Washington, D.C. ANONYMOUS. 1981. State gets newest natural area. Mississippi Outdoors. P. 11 ASSESSMENT CRITERIA FOR Mississippi's SCENIC STREAMS STEWARDSHIP PROGRAM. 2002. Public Notice No. 3661, Mississippi De- partment of Wildlife, Fisheries and Parks, Jackson. Barney, L.H. 1949. Manual of the cultivated plants. McMillan Publishing Company, New York. Majure, Flora of Chunky River in Mississippi 1201 Ball, PW. 2002. Carex (sect. Hallerianae). In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. New York and Oxford. Vol. 23, pp.487-489. BRIEGER, J.F. 1997. Hometown Mississippi. Town Square Books, Inc. Jackson. CHAPMAN, S.S., G.E. GRIFFITH, J.M. OMERNKK, J.A. Comstock, M.C. Belser, and D. Johnson. 2004. Ecoregions of Mississippi. (color poster with map, descriptive text, summary tables and photographs): Reston, VA, U.S Geological Survey (map scale 1:1,000,000). CLANCY, K.E. and M.J. SuLuvAN. 1990. Distribution of the Needle Palm, Rhapidophyllum hystrix. Castanea 55:31-39. CRUISE-SANDERS, J.M. and J.L. Hamrick. 2004. Genetic Diversity in Harvested and Protected Populations of Wild American Ginseng, Panax quinquefolius L. (Araliaceae). Amer. J. Bot. 91:540-548. FENNEMAN, N.M. 1938. Physiography of eastern United States. McGraw-Hill Book Company, Inc. New York. FERRARI, RV. 1970. A survey of the vascular flora of DeSoto County, Mississippi. M.S. Thesis, Memphis State Univ, Memphis. Funt, M.B. 1882. The exogenous flora of Lincoln Co., Mississippi, from October to May. Bot. Gaz. 7:74-76, 79-81. FLORA of NORTH AMERICA EDITORIAL Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. New York and Oxford. FRECKMANN, RW, and M.G. LELowc. 2003a. Dichanthelium. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. New York and Oxford. Vol. 25, pp. 406-451. FRECKMANN, RW, and M.G. LeLonG. 2003b. Panicum. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. New York and Oxford. Vol. 25, pp.450-488. Goprrey, R.K. and J.W. Wooten. 1979. Aquatic and wetland plants of southeastern United States; Monocotyledons. University of Georgia Press, Athens. Goprrey, R.K. and J.W. Wooten. 1981. Aquatic and wetland plants of southeastern United States; Dicotyledons. University of Georgia Press, Athens. Historic BriDGES of the Uniten States. 2002-2006. (http://bridgehunter.com/) HitcHcock, A.S. 1971. Manual of the grasses of the United States. Vols. 1 & 2. Dover Publications, Inc. New York. Hoop, W.G. and R.J. NAIMAN. 1999. Vulnerability of riparian zones to invasion by exotic vascular plants. Plant Ecol- ogy 148:105-114. KraL, R. 2002. Fuirena. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. New York and Oxford. Vol. 23, pp. 32-37. Leipote, A., S. McDanieL, and T. Nume. 2002. The flora of Oktibbeha County, Mississippi. Sida 20:691-765. Lowe, E.N. 1915. 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Soil Survey: Clarke County, Mis- sissippi. United States Department of Agriculture, Soil Conservation Service, Washington, D.C. Par Harrison Waterway District. 2002. Dunn's Falls. (www.phwd.net.) RADFORD, A.E., H.E. AHLes, and C.R. Bett. 1968. Manual of the vascular flora of the Carolinas. The University of North Carolina Press, Chapel Hill. Smith, S.G., J.J. BRUHL, M.S. GonZALEZ-ELIZONDO, and F.J. Menapace. 2002. Eleocharis. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. New York and Oxford. Vol. 23, pp. 60-120. Stevens, PF. (2001 onwards). Angiosperm Phylogeny Website. Version 8, June 2007. http://www.mobot.org/MO- BOT/research/APweb/ STEYERMARK, J.A. 1963. St k's flora of Missouri. Missouri Dept. of Conservation in cooperation with Missouri Botanical Garden Press, Jefferson City. THE INTERNATIONAL PLANT Names INDEX (IPNI). 2007. Published on the Internet http://www.ipni.org September 2007]. USDA, NRCS. 2007. The PLANTS Database (http://plants.usda.gov, 20 September 2007). National Plant Data Center, Baton Rouge, LA 70874-4490 USA. VANDERFORD, H.B. 1962. Soils of Mississippi. Mississippi Agric. Exp. Sta., Mississippi State. WATSON, R. 1972. Rhapidophyllum hystrix in Mississippi. Rhodora 74:525 Weakty, A.S. 2003. Flora of the Carolinas and Virginia: Dicotyledons. Working draft of July 2003. University of North Carolina at Chapel Hill, NC. WINSTEAD, R. 1990. A taxonomic and ecological survey of the plant communities of Attala County, Mississippi. MS. Thesis, Mississippi State Univ., Mississippi State. WUNDERLUIN, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida, Gainesville. accessed 20 — NEW PLANT DISCOVERIES FOR SONORAN ISLANDS, GULF OF CALIFORNIA, MEXICO Benjamin T. Wilder Richard S. Felger Desert Laboratory University of Arizona Herbarium (ARIZ) University of Arizona PO 36, University of Arizona 1675 West Anklam Road, Tucson, Arizona 85745, U.S.A. Tucson, Arizona 85721, U.S.A. ou arizona.edu rfelgereag.arizona.edu Humberto Romero-Morales AONAN a Mascareñas Tribu Seri, Punta Chueca U A Nicolás de Hidal Sonora, MEXICO Morelia, MEXICO ABSTRACT Renewed 1 li in the S islands of the Gulf of California (Islas Tiburón, San Esteban, San Pedro Nolasco, Dátil, and Alcat p D “Botanical pu of Isla Tiburón and Satellite Islands, Gulf of California," has resulted in 61 new ad- ditions and distrib f these islands. These new udi HN increases 2: 16% for Tiburón, 496 for San Esteban, 3.5% for San Pedro Nolasco, 3% for Dátil, and 12% for Al hecklists. We also report three new records for the Sonora mainland. On Isla Tib of the new addi f th the extensive Sierra Kunkaak on the east side of the island, which highligh? the sheltered habitats of the Bona and their accordingly strong connection with more tropically inclined floras. For all islands, the new records expand our understanding of this region and : B sf 1; +1 : 1 A f 1 : ES £1 aer 1 1 € eel dor ; m 1 TUIS da o o E" o TNT D ~ 1 1 ]; (egy IA La ASE BeA pee f+1 e] 5| 11 eL hi : f. : o [e] O Oo o o Lr threats from development and invasive species. RESUMEN Un interés botánico renovado en las islas del Golfo de California en Sonora (Islas Tiburón, San Esteban, San Pedro Nolasco, Dátil, y Alcatraz), como parte del proyecto *Diversidad Botánica de Isla Tiburón e Islas Satélite en el Golfo de California", ha dado como resul- tado 61 nuevas adiciones a la flora de estas islas y varias clarificaciones sobre su distribución. Estos nuevos registros representan un aumento del 1696 para Tiburón, 496 para San Esteban, 396 para Dátil, 1296 para Alcatraz, y 3.596 para San Pedro Nolasco, sobre las listas florísticas más recientemente pido Reportamos también tres nuevos registros m el territorio continental de Sonora. En Isla Tiburón la mayoría de | proviene n de los cañones profundos yp picos de tensa na de la Sierra Kunkaak, al este de la isla, lo que eee los refugios oF habitat 7 la Sierra y su fuerte conexión con floras de afinidad tropical. Para todas las islas, los 2 i > 5 1 4; +] : i d o E especies. El Sed trabajo florístico en las islas de S fade infi ión significati bre | laci biogeográficas de est islas, así como su situación actual dentro de una región que se enfrenta a amenazas i procedentes del d ll Smico y de las especies invasoras. INTRODUCTION The Sonoran islands of the Gulf of California, in the Central Gulf Coast subdivision of the Sonoran Desert (Shreve 1951), support largely undisturbed, unique, and magnificent desert ecosystems. Scientific explora- tion focused on the flora and vegetation of the mainland and islands associated with the Gulf of California began with Edward Palmer's collections (Watson 1889). Principal works focusing on the islands include Johnston (1924), Gentry (1949), Felger and Lowe (1976), Case and Cody (1983), Felger and Moser (1985), Moran (1983), Turner et al. (1995), and Case et al. (2002). The most exhaustive works on the plants of the Gulfislands are: Johnston’s (1924) classic work, produced from three months of collections, field observations, and subsequent herbarium studies; Gentry’s (1949) evaluation and interpretations, based largely on Rempel’s 1937 collections and Dawson’s 1940 collections; Felger and Lowe’s (1976) description and interpretation of the vegetation and flora of the Sonoran islands including a checklist; Case and Cody’s Island Biogeography in the Sea of Cortéz (1983), and Case et al.’s Island Biogeography of the Sea of Cortéz (2002). Rebman et al. (2002) J. Bot. Res. Inst. Texas 1(2): 1203 — 1227. 2007 1204 Journal of the Botanical R h Institute of Texas 1(2) and Moran and Rebman (2002) provide the most recent and comprehensive checklists of vascular plant species of the Gulf islands; it is upon these extensive checklists that we base the new records, clarifications, and interpretations presented here. Recent field work on the Sonoran islands (Table 1), as part of our project “Botanical Diversity of Isla Tiburón and Satellite Islands", has resulted in 48 new plant records and further study has yielded 13 dis- tributional clarifications (Table 2). From largest to smallest, these islands are Tiburón, San Esteban, San Pedro Nolasco, Dátil, and Alcatraz (Figs. 1 and 2). The majority of these records (49) are from Isla Tiburón, representing a 1696 increase over the 298 species reported in Rebman et al. (2002). Felger and Lowe (1976) predicted a 10-15% increase from their report of 286 species for this island. Similar predictions were made for the other islands treated here and are presented in Table 3 along with past and current species totals. While we report 61 distributional records, it is expected that about 1596 the flora of Tiburón is still not known, the majority expected from canyons and high elevations of the Sierra Kunkaak, and further discoveries on other islands are likely especially when fieldwork is conducted in seasons with favorable rains (Table 3). Close to 1096 of the new records reported here are non-native species, all occur on Isla Tiburón and two are also on Isla Alcatraz. West and Nabhan (2002) note the occurrence of the majority of the non-native species on the Midriff islands (islands found in the central part of the Gulf of California, between latitudes 29?40' and 28°20’) and we provide specific details regarding localities, current status, and documentation with voucher specimens. A number of biogeographical patterns are highlighted in the Sonoran island new records. One such connection is with “subtropical” areas. Species showing this pattern are supported by the extensive and relatively sheltered canyons in the Sierra Kunkaak of Isla Tiburón (Fig. 3A), where rainfall is presumably higher than the surrounding more xeric, desert areas. We report four species from the canyons of the Sierra Kunkaak whose nearest known populations are in both Baja California in the Sierra San Francisco ca 160 km to the southwest, and in Sonora in the subtropical canyons of the Sierra el Aguaje in the Guaymas region ca 150 km to the southeast: Celtis reticulata, Lantana velutina, Notholaena lemmonii var. lemmonii, and Rhynchosia precatoria. A relationship is also seen between Isla San Pedro Nolasco and the Sonoran Midriff islands, which are ca 130 km to the north. Two of the records; Bothriochloa barbinodis on San Esteban and Chloris crinita on Tiburón occur on no other Gulf islands besides San Pedro Nolasco, and Notholaena lemmonii var. lemmonii is only found on Isla Cerralvo in the southern Gulf besides on Tiburón and San Pedro Nolasco. This signal is reinforced by the fact that Tiburón and San Pedro Nolasco have a 6896 floristic similarity. Another strong pattern is seen in species with distributions in the Baja California peninsula and the Sonoran mainland that we record for the Sonoran Midriff islands. We present new records for seven such taxa: Ambrosia carduacea, Brahea armata/brandegeei, Dalea bicolor var. orcuttiana, Euphorbia xanti, Sideroxylon leucophyllum, and Tetra- merium fruticosum on Tiburón and Ambrosia divaricata on San Esteban. Taken together and with other taxa showing similar distributions and presence on the Midriff islands, evidence is mounting for plant migrations between the peninsula and mainland via the islands (Cody et al. 1983). We treat each new record individually and note the significance of the record, the nearest population, the geographic range, and cite voucher specimen(s). Additional information for localities is in the gazetteer. Common names are given in the following order if known: Cmiique litom (Seri; in Roman font), local Span- ish (italics), and English (Roman). Non-native species are denoted with an asterisk (*). Voucher specimens are deposited at the University of Arizona herbarium (ARIZ), unless indicated otherwise. Many of the duplicate specimens of our collections are variously deposited at MEXU, SD, and USON, as well as other herbaria in Mexico and the United States. We cite some specimens from Isla Alcatraz from the herbarium of the Prescott College Field Station at Bahía de Kino; photos of these specimens are at ARIZ and data based in SEINET. More detailed information for specimens at ARIZ can be accessed oi via Southwest J / Ju Environmental Information Network database (SEINET: http://seinet.asu jsp) and data for specimens at USON can be found at Plantas Silvestres de Sonora: Un Herbario Electrónico en Línea, Catálogo de Especies (http://herbario.uson.mx:8080/plantson.htm), and the cited herbarium collections. The majority Wilder et al., Plant records for 4l c Is, Gulf of Mexico TABLE 1. Recent Sonoran Island fieldtrips by the authors. Collector Associated Collectors Island Locality Dates Collection Number of New Numbers Records Benjamin Wilder Edward Gilbert, Tiburón Eastern bajada, 29 Dec 2005- 05-01-05- 7 Humberto Romero Pazj Hax waterhole 4 Jan 2006 54 and 06- (Tinaja Anita), north 01-06-52 and central Sierra nkaak David Bertelsen Tiburón Coralitos, Arroyo 16-19 Mar 06-54-06 1 Sauzal, Xapij 2006 : (Sauzal waterhole) Richard Felger, Tiburón Eastern Bajada, 23-25 May 06-143- 2 Humberto Romero Estero San Miguel, 2006 06-169 central Sierra Kunkaak Gloria Guadalup Tiburón Estero San Miguel 2 Sep 2006 06-274- 0 Morales-Figueroa, 06-288 Jesús Sanchez Escalante, Mikhal Gold, Humberto Romero Richard Felger, Tiburón Northern Sierra 23-26 Nov 06-345-22 2 Edward Gilbert, Kunkaak canyons, 2006 06-513 Humberto Romero Zozni Cmiipla Seth Turner San Esteban Arroyo Limantour, 7-9 Mar 2007 07-38-07-93 S entral peak, south- west corner Seth Turner Dátil Central peak and 10-11 Mar 07-94- 3 canyon 2007 07-142 Exequiel Ezcurra, Tiburón Ensenada de Perros 12Apr 2007 07-143- 0 Richard Felger, 7-165 Jesús Ventura Trejo Exequiel Ezcurra, Dátil Central eastern 12 Apr 2007 07-166- 0 Richard Felger, canyon 7-183 Jesús Ventura Trejo Humberto Romero, Tiburón Estero San Miguel, 1-5 May 2007 07-184- 3 Seth Turner opc Hax waterhole, 07-272 rancho Carocol, northern portion of eastern bajada, Tecomate Adrian Quijada- Tiburón Cerro San Miguel 9 Nov 1990 90T002-90T012 2 Mascareñas Gil Gillenwater Tiburón Cerro San Miguel 9 Mar 1991 91T007-90T017 2 Richard Felger Alberto Búrquez San Pedro Northeast side 28 Nov 2006 06-73-06-107 1 ontijo, Florencio Nolasco of island Cota Moreno, Jesús Ventura Trejo of the records introduced here were obtained during Felger and Wilder's fieldwork from 2005 to 2007 and Quijada's field work in 1990 and 1991 (Table 1). However, some records were discovered while searching herbarium specimens, and dates of collection are given for those specimens. 1206 Journal of the Botanical R h Institut f Texas 1(2) A EE a Ss, lc imt If TABLE D New [ECOIU lul f California. New is defined as not included in the checklist of Rebman et al. (2002). Bold = New for the island. * 2 Non-native species. j = Previously cited but not by Rebman et al. 2002. Family Species Tiburón San Nolasco Dátil Alcatraz Esteban Acanthaceae Tetramerium fruticosum TIB Amaranthaceae Suaeda (e TIB Apocynaceae Funastrum muse var. hartwegii TIB Arecacea ¡Brahea a TIB Asteraceae Ambrosia e TIB Asteraceae Ambrosia divaricata TIB EST DAT Asteraceae Gymnosperma glutinosum TIB Asteraceae Heliopsis anomala TIB Asteraceae Pectis pa pposa var. quie TIB ALC Asteraceae CARLA diffus TIB ALC ce Batis maritima TIB ALC ixaceae as palmatifida TIB Convolvulaceae — Cuscu a TIB Convolvulaceae A d TIB Convolvulaceae ¡Ipomoea hederacea TIB Convolvulaceae o agrestis TIB Cucurbitaceae Tumamoca macdougalii TIB modaceae ¡Halodule wrightii TIB Cyperaceae ¡Cyperus elegans TIB NOL Cyperaceae yperus squarrosus NOL Euphorbiaceae Euphorbia abramsiana TIB Euphorbiaceae ¡Euphorbia arizonica TIB Euphorbiaceae Euphorbia eriantha TIB DAT Euphorbiaceae Euphorbia a TIB Euphorbiaceae Euphorbia x TIB Fabaceae Dalea o var. orcuttiana TIB Fabaceae dium procumbens TIB Fabaceae *Pithecellobium dulce TIB Fabaceae ia precatoria TIB Fabac hi I TIB Henn eee praca Pr TIB Malpighiaceae Echinopterys E TIB Malvaceae iAbutilon palmeri TIB Malvaceae Horsfordia new pr TIB EST DAT Myrtaceae *Fucalyptus us camaldulensis TIB Orobanchaceae Orobanche c TIB Poaceae Meus comi var. glabarata TIB Poaceae thri barbinodi EST Poaceae pde SUE TIB Poaceae Ge TIB Poaceae Chori virgat TIB Poaceae neapogon CT TIB Poaceae uns a dubia TIB Poaceae i*Pennisetum ciliare TIB ALC Poaceae pi leucopila TIB EST DAT Poaceae ee virginicus TIB ALC Poaceae Tridens muticus var. muticus TIB Poaceae Urochloa fusca TIB EST Portulacaceae Portulaca oleracea TIB Pteridaceae Notholaena californica ssp. californica TIB EST Pteridaceae Notholaena lemmonii var. lemmonii TIB Pteridaceae UE standleyi TIB Wilder et al., Plant records for the S Islands, Gulf of Mexico 1207 TABLE 2. continued Family Species Tiburón San Nolasco Dátil Alcatraz Esteban Rubiaceae Galium proliferum TIB Rubiaceae Galium stellatum var. eremicum EST Sapotaceae Sideroxylon leucophyllum TIB EST Solanaceae Lycium berlandieri var. longistylum TIB Solanaceae Physalis crassifolia var. versicolor TIB DAT Tamaricaceae ¡Tamarix aphylla TIB Ulmaceae Celtis reticulata TIB Verbenacea Lantana tina TIB Zygophyllaceae *Tribulus terrestris TIB ALC NEW RECORDS ACANTHACEAE Tetramerium fruticosum Brandegee New for Tiburón where it is common in deep, protected canyons in the interior of the Sierra Kunkaak. Herbaceous perennials. Geographic Range.—Widespread in Baja California Sur (Daniel 1997) and known from mainland Sonora (Tinaja Picu, SW of Pitiquito; Daniel 1986, 2004) by a single collection (Wiggins 6055, DS). The Tiburón specimens provide a disjunct, stepping-stone locality. Tiburón.—Base of Sierra Kunkaak, between Sierra Kunkaak Mayor and Sierra Kunkaak Segundo, head of arroyo, Wilder 05-46. Base of canyon on N slope of Sierra Kunkaak, Wilder 06-386. Siimen Hax waterhole, Wilder 06-449. AMARANTHACEAE Suaeda esteroa Ferren & S.A. Whitmore. Sipjó yanéaax; estuary seablite We confirm the presence of this species on Tiburón. We expected and found it at the margins of mangroves at Estero San Miguel, where it is often inundated by the highest tides. The nearest popul are on the op- posite shore of the Canal del Infiernillo. This species and S. nigra (Raf.) J.F. Macbr. are the only Suaeda species on Tiburón, although others are reported (e.g., Moran 1983a; Rebman et al. 2002). Halophytic succulents. Geographic Range.—Mangrove and salt scrub esteros in the Gulf of California and Pacific coast of the Californias. The Gulf of California plants are annuals and perhaps a different taxon from those on the Pacific Coast, which generally are perennials. Tiburón.—Zozni Cmiipla, Wilder 06-362. APOCYNACEAE Funastrum cynanchoides (Decne.) Schltr. var. hartwegii (Vail) Krings [Sarcostemma cynanchoides Decne. subsp. hartwegii (Vail) R.W. Holm; Funastrum heterophyllum (Engelm.) Standl.] Hexe; huirote, hierba lechosa; climbing milkweed New for Tiburón, where we found it in a canyon bottom among dense vegetation near a waterhole in the Sierra Kunkaak. It is widespread and common on the adjacent Sonora mainland and it is surprising that we have only one record for it on the island. Perennial vines, the stems dying back during drought. Flowering at various seasons. Geographic Range.—SW United States to Michoacán and Queretero, and northern Baja California. Tiburón.—Pazj Hax waterhole, wet soil, Wilder 06-10. 1208 Journal of the Botanical R h Institute of Texas 1(2) ESTADOS UNIDOS DE AMÉRICA zZ , Desemboque Isla Angel de la Guarda Isla Patos ò OCEANO PACÍFICO 3 Isla Punta Chueca o . r ¡o Tiburón o e Bahía de los 25 Isla \. Bahía de Kino Hermosillo Á e Angeles Alcatraz”? ` Isla San Lorenzo Sy Y ae Isla D) Isla Dátil Isla San Lorenzo Sur San Esteban Isla - Pedro Mártir Baja CALIFORNIA Guaymas Isla San S Nolasco 0 20 km Fic. 1. Midriff island region. Map by Cathy Moser Marlett. ARECACEAE Brahea armata S. Watson. Zamij cmaam (female palm); palma azul; blue hesper palm The report of a palm on Isla Tiburón had been enigmatic even though a number of Comcáac have known of a palm on the northwest side of the island (Felger & Moser 1985, p. 351). We now document this report with a specimen and additional information. On May 20, 1964, José Juan Moreno brought Mary Beck Moser a single leaf of a palm that he collected in a canyon along the northwest side of the island. Sr. Moreno had been on a fishing trip and knew that Mary Beck and Richard Felger were interested in the *palm from Tiburón" as they were beginning their studies of Seri knowledge and uses of the plant world. Soon thereafter Richard and Alexander “Ike” Russell made a number of searches by airplane along the west side of the island for the elusive palm. The search was futile even though Ike would fly low and close enough that Richard usually could identify all but the smaller plants (Felger 20002, 2002). Sometimes, however, shadows, other trees, and cliffs can make it difficult for plant spotting from the air. On May 4, 2007, Humberto confirmed that he had seen a single palm at this site about 20 years earlier. Additional information from R. James Hills (pers. comm., April 2007) further substantiates the presence of one or more palms on the island. The site is reported to be at or near a fresh water source in a canyon on the seaward side and base of high cliffs on the northwest side of the extremely arid Sierra Menor. However, on a trip to the claimed canyon in September 2007 by Felger, Romero, and Wilder, no palm was found and we believe the lone individual is now deceased. Wilder et al., Plant records for the S Islands, Gulf of Mexico 1209 Zozni Cmiipla Ensenada del Perro 0 Isla Cholludo ya Dátil 0 20 Km a NNI Ex 9 Isla Tiburó I vicinity. Map by Cathy M Geographic Range.—The nearest locality for B. armata is at the SE side of Isla Ángel de la Guarda (John- ston 1924; Moran 1983b; Turner et al. 1995), ca 65 km to the W of the Tiburón site. The vegetation and habitats of the Isla Ángel palm canyon sites and the NW coast of Tiburón are quite similar. The nearest Brahea population from the Tiburón site is on the Sonora mainland about 37.5 km to the NE: *The most isolated and undoubtedly smallest palm population in Sonora occurs on a north-facing slope below Pico de Johnson in the Sierra Seri. There are probably fewer than one dozen of these palms, hidden from view and far from any road" (Felger & Joyal 1999, p. 10). The population on Pico de Johnson seems conspecific 1210 Journal of the Botanical R h Institute of Texas 1(2) Tagle 3. Island flora totals and predictions for additional records. A = Felger and Lowe 1976, B = Moran 1983, C = Rebman et al. 2002, D = Our current information. A B C D Islands # Species Prediction # Species # Species # Species Prediction Tiburón 286 315-329 298 298 340 391 (+15%) San Esteban 107 118 WF 123 125 131 (2-590) San Pedro Nolasco 55 NA 56 57 57 57 Dátil 99 NA 99 99 106 111 (45%) Alcatraz 43 NA 43 43 44 46 (4-596) with the hesper palms in the Sierra el Aguaje, N of San Carlos and Guaymas, which have been treated as conspecific with B. brandegeei (Purpus) H.E. Moore of Baja California Sur (Felger & Joyal 1999; Felger et al. 2001; Henderson et al. 1995). These western Sonora populations, however, appear to be closely related to B. armata of Baja California Norte and Isla Ángel de la Guarda and the distinctions *have not been quantified or studied in depth" (Felger & Joyal 1999, p. 11). Tiburón.— Canyon on NW side of the island, 20 May 1964, José Juan Moreno, s.n. ASTERACEAE Ambrosia carduacea (Greene) W.W. Payne New for Tiburón. It is a fairly common component along the drainageways of the rich canyon desertscrub/ thornscrub community of the Sierra Kunkaak (Fig. 4A), especially east- and north-facing canyons. It also a new record for western mainland Sonora, where it occurs on the north facing slopes of Pico de Johnson on the Sonora mainland opposite Tiburón. Robust shrubs ca 1—2.5 m tall; reproductive October to at least February. Geographic Range.—Widespread in both states of Baja California and on Isla Cerralvo, mostly in Sonoran desertscrub and extending into tropical deciduous forest in the Cape Region. Widely disjunct in E-central Sonora where it is often arborescent to 4 (5) m tall. The new records for Sierra Kunkaak and Sierra Seri nar- row the gaps between the Baja California peninsula and the disjunct Sonora populations along the E side of the desert as mapped by Turner et al. (1995). Tiburón.—Top of San Miguel Peak in a sheltered canyon, 28 Oct 1979, Knight 1019 et al. (UNM). Cerro San Miguel near Sierra Kunkaak, upper bajada, 1 m tall, Quijada-Mascareñas 901007. Head of an arroyo at base of Sierra Kunkaak, between Sierra Kunkaak Mayor and Sierra Kunkaak Segundo, 1,285 ft, Wilder 05- 26. Head of an arroyo at E base of Sierra Kunkaak, 875 ft, Wilder 06-31. Sonora mainland.—N side of Pico de Johnson, Sierra Seri, 30 Dec 1991, Powell B. (Gil) Gillenwater III (photo). Ambrosia divaricata (Brandegee) W.W. Payne. An icoquéete; chicurilla, huizapol New for San Esteban. The nearest populations are on Tiburón and Dátil where it is fairly widespread as shrubs to ca 1 m in height. On San Esteban we know it only from the tall central peak where it is localized as a dwarf shrub tucked into the north-facing slope of the ridge. Geographic Range.—Primarily a Baja California peninsular species with outlying populations along the Sonora coast and Gulf islands. San Esteban.— Central peak, 365 m, steep slopes of ridge top, dwarf shrubs ca 20 cm tall, Wilder 07-59. Gymnosperma glutinosum (Spreng.) Less. Gumhead New for Tiburón, where we found a few plants along the bottom of a canyon in the Sierra Kunkaak. The nearest populations are on the Sonoran mainland where it occurs in scattered localities from the Guaymas region northward (Felger 1999, 2000b). Small shrubs or subshrubs. Wilder et al., Plant records for the S Islands, Gulf of Mexico 1211 Geographic Range.—Arizona and Texas to Guatemala. Tiburón.—Base of N portion of Sierra Kunkaak, ca 0.5 km E of Siimen Hax waterhole, 360 m, broad canyon bottom, subshrub, stems water swept [from a flash flood], Wilder 06-467. Heliopsis anomala (M.E. Jones) B.L. Turner [H. parvifolia A. Gray var. rubra (T.R. Fisher) Wiggins] New for Tiburón; growing in deep canyons of the Sierra Kunkaak where it is localized, usually in shaded places, often among rocks, and occasionally along open washes of canyon drainages. The nearest popula- tion occurs in similar habitat in the Sierra Seri. Mostly perennials, probably short lived, and also flowering in the first year or season. Geographic Range.—W Sonora from the Sierra Bacha (S of Puerto Libertad) to the Guaymas region, and both Baja California states. Tiburón.—Head of an arroyo at E base of Sierra Kunkaak, Wilder 06-32. Canyon bottom at N base of Sierra Kunkaak, Wilder 06-411. Pectis papposa Harv. & A. Gray var. papposa. Caasol heecto, caasol ihasíi quiipe, cacáatajc; mansanilla del coyote; desert chinchweed New for Alcatraz; widespread and common on adjacent mainland and Tiburón. Aromatic hot-weather ephemerals. Geographic Range.—Aridlands of W North America. Alcatraz.—25 Oct 2001, Jones 16 (Prescott College Collection). Xylothamnia diffusa (Benth.) G.L Nesom. Caasol cacat, caasol ziix iic cóihíipe; hierba de pasmo New for Alcatraz; common on the adjacent mainland and on parts of coastal Tiburón. Small shrubs. Geographic Range.—Coastal Sonora from the vicinity of Bahía de Kino to NW Sinaloa, Baja California Norte y Sur, and Islas Ángel de la Guarda and San Lorenzo. Alcatraz.—2 May 2002, Gracida 56 (Prescott College Collection). BATACEAE Batis maritima L. Pajóocsim, xpajóocsim, xpacóocsim; dedito; saltwort New for Alcatraz; common in adjacent mainland and Tiburón coastal wetlands. Halophytic succulent peren- nials locally forming dense ground cover of trailing and scrambling stems. Geographic Range.—Shores of the Americas including the Gulf of California. Alcatraz.—27 May 2002, Gracida 77 (Prescott College Collection). BIXACEAE Amoreuxia palmatifida Sessé & Moc. ex DC. Xoját; saiya Although reported for Tiburón by Felger and Moser (1985), we verify its occurrence here with a specimen from the middle of the island. The roots were an important food resource for the Seris. Perennials from a thick, tuberous root, responding to summer rains and dormant during the rest of the year. It also occurs on the adjacent Sonora mainland. Geographic Range.—S-central Arizona and Sonora to Colombia, and Baja California Sur, the closest peninsular location is in the Sierra San Francisco. Tiburón.—Haap Hill, 8 Sep 1974, Felger T74-2. CONVOLVULACEAE Cuscuta americana L. Hamt itóozj; fideo; dodder New for Tiburón and the first record for a Gulf island. The nearest known localities in Sonora are in the San Carlos-Guaymas region ca 160 km to the southeast; it is widespread elsewhere in Sonora in non-desert regions. Stems moderately thick; on various woody shrubs, especially Colubrina viridis M.E. Jones. Specimens of both Cuscuta were identified by Mihai Costea in 2007. Geographic Range.—Florida; Mexico to South America and the West Indies. 1212 Journal of the Botanical R h Institute of Texas 1(2) Nel f, C; Lu terhole. (R | ACA, nj I. T Fic. 3. Sierra Kunkaak. (A, top) Sheltered, interior canyon, p cany I I L HA | £f c AAS 1 DL L D > > MEIA 25 Nov 2006 b J 7 J Wilder et al., Plant records for the Sonoran Islands, Gulf of Mexico Fic. 4. New records from Sierra Kunkaak. (A, top left) Ambrosia carduacea (Wilder 06-383). (B, bottom) Euphorbia xanti (Wilder 06-453). (C, top right) Echinopterys eglandulosa (Wilder 06-377). Photos by Benjamin Wilder. 1214 Journal of the Botanical R h Institute of Texas 1(2) Tiburón. —Foothills of NE portion of Sierra Kunkaak, Wilder 06-373. Top of Sierra Kunkaak Segundo, Wilder 06-495. Cuscuta desmouliniana Yunck. Hamt itóozj; fideo; dodder New for Tiburón and the first record for a Gulfisland. Widespread on the island and common on the adjacent Sonora mainland. Stems very slender; on various annuals. Geographic Range. —Widespread in W and central Sonora and on the Baja California peninsula. Tiburón.—SE side of Agua Dulce Valley, ca 12 mi S from Tecomate, ca 280 m, 11 Dec 1976, Felger 76- T14. NE base of Sierra Kunkaak, *bench" above arroyo bottom, Wilder 06-381. Ipomoea hederacea Jacq. Hatáaij; trompillo azul; morning glory An unidentified Ipomoea reported for Tiburón (Moran 1983a; Rebman et al. 2002) is undoubtedly this spe- cies, which is the only one known for the island. It is locally common at widely separated localities on the island and may be seasonally more widespread. Warm weather ephemeral vines. Geographic Range. —Widespread in the Sonoran Desert including western Sonora and Baja California Sur with the closest peninsular location being the Sierra San Francisco. North and South America and adventive in the Old World. Tiburón.—Haap Hill, 11 Dec 1976, Felger 76-T23. Vicinity of Zozni Cmiipla, Wilder 06-349, Jacquemontia agrestis (Mart. ex Choisy) Meisn. [J. palmeri S. Watson, Proc. Amer. Acad. Arts 24:63. 1889] New for Tiburón, where it was found in the interior of the island, and new for islands in the Gulf of California. There are a number of records for this species to the east and south of the Sonoran Desert in Sonora but it is apparently rare or poorly known from the desert. Despite more than a century of intensive collecting in the Guaymas region, it has not been recorded there since Palmer's collection in 1887 (Watson 1889). In Arizona itis known only from the Baboquivari Mountains where it has not been recorded since 1945 (Austin 1998). Is its range shrinking in the Sonoran Desert region? Or has it been overlooked? Annuals, generally vining but the Tiburón specimens are stunted summer/fall ephemerals and not vining. Geographic Range.—Arizona through Mexico, uncommon in Baja California Sur (present in the Sierra San Francisco), to Honduras and South America; often a vining weed in cultivated fields. Tiburón.—Haap Hill, 11 Dec 1976, Felger 76-124. CUCURBITACEAE Tumamoca macdougalii Rose. Hatoj caaihjó (“that which makes the eyes red”; Felger & Moser 1991, p. 388; Moser & Marlett 2005); Tumamoc globe-berry New documentation for Tiburón and a new generic record for the Gulf islands. Perennials with a single tuberous root, producing a small, ephemeral vine following summer/fall rains. The small fruit is bright red. The Comcáac know of it on the east side of Tiburón. Paul Knights collection is from the general vicinity of Punta Tormenta on the east shore of the island. The nearest known populations are in the vicinity of Bahía de Kino. Geographic Range.—SW and central Sonora to S-central Arizona. Tiburón.— Growing near the beach, under Frankenia palmeri, single seed per fruit, 24 Oct 1979, Knight 909 et al (UNM). Punta Tormenta, 14 Sep 2007, Romero 07-19. CYMODACEAE Halodule wrightii Asch. Shoal grass New listing for Tiburón, although reported for the Canal del Infiernillo in shallow sea water (15—150 cm below low tide level (Felger 2004; McMillan & Phillips 1979; Meling-López & Ibarra-Obando 1999). Richard observed Halodule in the stomachs of sea turtles (Chelonia) butchered by Seris along the Infiernillo coast of Tiburón and at Punta Chueca in the 1970s and 1980s, but only recently realized that at least much of it was Halodule rather than Ruppia as documented by the specimen cited below. At that time (1970s and 19803) large Wilder et al., Plant records for the S Islands, Gulf of Mexico 1215 and small fragments of Ruppia maritima L. were common in the summertime coastal and beach drift along both coasts of the Infiernillo but the source of rooted plants may have been further south in the Estero de la Cruz at Bahía de Kino (Alf Meling-López, pers. comm., 2006). Halodule is a small, delicate seagrass locally known only from the summer months when the common local seagrass Zostera marina L. is not present. (Felger & Moser 1985; Meling-López & Ibarra-Obando 1999; Torre 2002.) Geographic Range.—Shallow, protected sea water; intermittently on the Pacific coast from Canal del Infiernillo, Sonora, and Sinaloa to Central America, and Atlantic waters from southeastern United States to South America and the West Indies. Canal del Infiernillo.—S end of Infiernillo Channel, S of Punta Chueca, water 26°C, recovered from stomach content of a Chelonia mydas harpooned and butchered at Punta Chueca, stomach full, ca 9996 com- prised of this seagrass, 16 Oct 1973, Felger 21205 et al. Punta Chueca, water depth of 6 in. to 1.5 m, 16?C, 5-7 Jan 1979, Ronald Phillips & Calvin McMillan s.n. Ca. 2 km N of Punta Chueca, 25 Jul 2007, Wilder 07-370. CYPERACEAE Cyperus elegans L. Sticky sedge Documented for Isla San Pedro Nolasco by Gentry (1949) but not included in subsequent checklists. The nearest populations are at waterholes and other wetland sites on the opposite mainland and it also occurs at waterholes on Tiburón. The Nolasco record is enigmatic because there are no wetland habitats on the island, although on occasion, following favorable rains, the steep, north-facing canyon-slopes on the east side of the island can be lush and grassy (Felger & Lowe 1976). The plants are notably viscid, even the spikelets, and birds are presumed dispersers. It has not been documented on the island since 1940, in spite of Richard's several extensive collecting trips and visits by other botanists. Annuals in western Sonora (elsewhere perennials). Geographic Range.—S United States and Baja California Sur to South America. San Pedro Nolasco.—6 Feb. 1940, Dawson 1036 (LCU, NY; verified by Gordon Tucker, pers. comm., 2007, also see Tucker 1994). Cyperus squarrosus L. [C. aristatus Rottb.]. Dwarf sedge New for Nolasco. Documented from a few other islands in the Gulf (San Pedro Martír and Espíritu Santo) and we predict it will be found on other larger islands. Diminutive, short-lived annuals. Geographic Range.— Cosmopolitan and widespread in the Sonoran Desert, including Baja California Norte y Sur and present in the Sierra San Francisco. San Pedro Nolasco.—NE side of island, ca 60 m, open area near canyon bottom, E-facing exposure in local area mostly free of Vaseyanthus with Boerhavia, Coreocarpus, Eragrostis pectinacea, Setaria liebmannii, localized population of several hundred plants, Felger 06-87. EUPHORBIACEAE Euphorbia abramsiana L.C. Wheeler New for Tiburón and not recorded for other Gulf islands except Isla San José off Baja California Sur. Known from widely separated localities in western Sonora. Annuals, mostly growing with summer-fall rains. Geographic Range.—SE California to Baja California Sur and S Arizona to Sinaloa. Tiburón.—Near airstrip [= Palo Fierro landing field at Punta Tormenta], 25 Oct 1979, Knight 906 et al. (UNM). Euphorbia arizonica Engelm. Tomítom hant cocpéetij; golondrina; Arizona spurge This species was listed by Felger and Lowe (1976) and Moran (19833) for Tiburón but not by Rebman et al. (2002). We confirm the earlier listings with specimens from six localities. Non-seasonal ephemerals to small perennials. Geographic Range.—SW United States and NW Mexico including both Baja California states. Tiburón.—SW part of Central Valley, ca 13 mi S of Tecomate, upper rocky slope of mountain bordering 1216 Journal of the Botanical R h Institute of Texas 1(2) valley, 1200-1400 ft, S facing slope, 2 Feb 1965, Felger 12423. Haap Hill, 8 Sep 1974, Felger 74-T40. Between Sopc Hax water hole and Hant Hax camp, 26 Oct 1963, Felger 9339. Cerro San Miguel, Quijada-Mascareñas 91T005. Large arroyo heading to a valley at E base of Sierra Kunkaak, Wilder 06-52. Siimen Hax waterhole, Wilder 448. Euphorbia eriantha Benth. Pteept; beetle spurge New for Dátil; widespread on nearby Tiburón. Non-seasonal annuals. Geographic Range.—Southwestern United States, northern Mexico, and Baja California Norte y Sur. Dátil.—N-facing slopes of the central ridge, common, ca 30 cm tall, Wilder 07-124. S-facing slope of ridge just N of center of island, less common than on N-facing slopes, Wilder 07-140. Euphorbia prostrata Aiton. Tomítom hant cocpéetij; golondrina; ground spurge New for Tiburón where we found it at a single locality along the eastern coast near the old Seri camp Zozni Cmiipla. This is the first record for a Gulf island. Known from widely separated localities in western Sonora, often in disturbed habitats (Steinmann & Felger 1997; Felger 2000b). Warm-weather ephemerals (short-lived perennials in non-desert regions). Geographic Range.—Widespread in the Americas and naturalized in warm regions of the world. Tiburón.—Vicinity of Zozni Cmiipla, sandy soil, Wilder 06-513. Euphorbia xanti Engelm. ex Boiss. Hehe ix cooxp; juemetón New for Tiburón. Common on the bajada of the northeastern side of the island and also documented from a canyon in the Sierra Kunkaak. Also on the opposite Sonoran mainland. Shrubs 1.5—2.5 m tall, often propagating by rhizomes (Fig. 4B). Cody et al. (1983) hypothesized this species migrated from the Baja California peninsula to the Sonoran mainland via the Midriff islands. In addition, many other species show a similar distributional pattern: for example, Steinmann and Felger (1999) list six euphorbs with this pattern. In a thesis on the group (section Alectoroctonum) to which E. xanti belongs, E. peganoides Boiss. is given as the most similar relative of E. xanti (Ramírez 1996). Victor Steinmann (pers. comm., 2007) treats both E. peganoides and E. colletioides Benth. as synonyms of E. cymosa Poir. as eluded to by Steinmann and Felger (1997). Euphorbia xanti is the only member of section Alectoroctonum occurring on the Baja California peninsula, so the relatives of E. xanti are mainland taxa. Geographic Range.—Predominately in Baja California Sur and the S half of Baja California Norte, and sporadically along 215 km of the Sonora coast, from near El Desemboque San Ignacio S to Bahía de Kino and farther S near Guaymas (Turner et al. 1995). Also on Islas Monserrat and Tortuga off Baja California Sur. Tiburón.— Canyon on a N slope of Sierra Kunkaak, vicinity of Siimen Hax waterhole, cluster of five plants on N-facing wall on side of canyon, Wilder 06-453. Lower E bajada, near Valle de Águila, ca 4 km N of Punta Tormenta, common shrub ca 1.25 m tall, Wilder 07-232. FABACEAE Dalea bicolor Humb. & Bonpl. ex Willd. var. orcuttiana Barneby New for Tiburón and the first record for this species for a Gulf island. Found on steep slopes of the Sierra Kunkaak—both island collections are from approximately the same locality. The nearest population occurs on the opposite mainland near the western base of the Sierra Seri range, which is a new record for this variety for mainland Mexico. Shrubs or subshrubs to 1« m tall. Geographic Range.—Variety orcuttiana was previously known only from the Baja California peninsula where it is widespread in both states. Tiburon.—Sierra Kunkaak Mayor, Mar 2007, Romero 07-3. Cerro San Miguel, común en la cima, 8 Mar 1991, Quijada-Mascareñas 911019. Sonora mainland.—Vicinity of Cerro Pelón, 29°34’N, 112?09"W, ca 5 mi SE of Desemboque San Ignacio, 21 Apr 1968, Felger 17936. Wilder et al., Plant records for the S Islands, Gulf of Mexico 1217 Desmodium procumbens (Mill.) Hitchc. Tick clover New for Tiburón where it appears to be widespread. The nearest known populations are in the vicinity of Hermosillo and the Guaymas region but it is expected in the Sierra Seri; the closest peninsular population is in the Sierra San Francisco. Short-lived summer/fall ephemerals on Tiburón and in western Sonora. Geographic Range.—Baja California Sur and S Arizona to South America and the West Indies; also in the Old World where it is probably introduced. Tiburón.—Haap Hill, 8 Sep 1974, Felger 174-38. Hant Hax camp, canyon bottom in lower foothills at base of mountain [Sierra Kunkaak], on way to Sopc Hax from Zozni Cmiipla, 26 Oct 1963, Felger 9348. Canyon bottom at N base of Sierra Kunkaak, between Sierra Kunkaak Mayor and Sierra Kunkaak Segundo, 395 m, Wilder 06-409. *Pithecellobium dulce (Roxb.) Benth. Camótzila; guamúchil; manila tamarind New for Tiburón, where we found one young tree that had been planted at the Sopc Hax waterhole in the Sierra Kunkaak. Also planted in the Comcáac villages on the opposite mainland (Nabhan 2003). Geographic Range.—Large trees native to the hot lowlands of Mexico and northern South America. Widely planted for its edible fruit throughout lowland Sonora but not known to persist untended (without supplemental water) within the Sonoran Desert, or at least in Sonora (Felger et al. 2001). Reid Moran docu- mented a tree on Isla Cerralvo (16 Apr 1962, tree 5 m tall x 10 m, Moran 9511, SD). Tiburón.—Sopc Hax waterhole, small tree ca 3 m tall, Wilder 07-214. Rhynchosia precatoria (Humb. & Bonpl. ex Willd.) DC. Chanate pusi, ojo de pajarito; rosary bean New for Tiburón, where we found it in a deep canyon of the Sierra Kunkaak. This is the first record for it on a Gulf island. The nearest known localities are in the Sierra el Aguaje north of Guaymas (Felger 1999), ca 150 km to the southeast, and ca 160 km to the southwest in Baja California in the Sierra San Francisco. Perennial vines growing over shrubs. Geographic Range.—Extreme S Arizona to northern South America Tiburón.—Deep canyon on N slope of Sierra Kunkaak, SW and up canyon from Siimen Hax waterhole, Wilder 06-480. Tephrosia vicioides Schltdl. New for Tiburon where it is widespread, often growing along sandy-gravelly washes. It also occurs on the opposite mainland in the Sierra Seri. Non-seasonal annuals (or sometimes short-lived perennials?) on Tiburon. Geographic Range.—A somewhat variable species, widespread in the Americas including Baja California Sür. Tiburón.—San Miguel Peak, 27 Oct 1979, Knight 1019 et al. (UNM). Arroyo Sauzal: Ca 2.75 mi inland from shore, Wilder 06-99; Ca 1.25 mi inland from shore, main wash channel, Wilder 06-78. SW part of Central Valley, ca 13 mi S of Tecomate, ca 450 ft, 2 Feb 1965, Felger 12389. Haap Hill, 11 Dec 1976, Felger 76-139. Valley at E base of Sierra Kunkaak, ca 875 ft, Wilder 06-51. Foothills of NE portion of Sierra Kunkaak, Wilder 06-445. HYDROPHYLLACEAE Phacelia affinis A. Gray A single collection from the interior of the island is a new record for Tiburón and the first record for a Gulf island. The nearest known population is ca 150 km to the north in the Sierra del Viejo near Caborca. Cool- season ephemerals. Geographic Range. —SW United States, N Sonora, and both states of Baja California. Tiburón.—SW part of Central Valley, ca 13 mi S of Tecomate, 20 Feb 1968, Felger 17337. MALPIGHIACEAE Echinopterys eglandulosa (Juss.) Small. Hap oáacajam 1218 Journal of the Botanical R h Institute of Texas 1(2) New for Tiburón and the first record for a Gulf island. We encountered a single shrub (conspicuous because of its bright yellow flowers) in a Sierra Kunkaak canyon but it is probably more widespread in these moun- tains. The nearest population is in the Sierra Seri on the opposite Sonoran mainland. Drought-deciduous shrubs. (Fig. 4C). Geographic Range.—Sonora (S from the vicinity of El Desemboque San Ignacio and Altar) to Oaxaca. Tiburón.—NE base of Sierra Kunkaak, S-facing side of arroyo in rocky talus, shrub 1.6 m tall, Wilder 06-37T. MALVACEAE Abutilon palmeri A. Gray. Caatc ipápl; pelotaso; Indian mallow This species was listed by Gentry (1949), Felger and Lowe (1976), and Moran (1983) as occurring on Tiburón and other Gulf islands, but was not included in the checklist by Rebman et al. (2002). We confirm it for the southern and eastern parts of Tiburón. Sparsely branched, open and scarcely woody shrubs or subshrubs. Geographic Range.—SW Arizona to Sinaloa, and SE California to the Cape Region of Baja California Sur, and many islands in the Gulf; disjunct in Tamaulipas. Tiburón.—Arroyo Sauzal, ca 1 mi from shore, Wilder 06-84. Head of arroyo at base of Sierra Kunkaak, between Sierra Kunkaak Mayor and Sierra Kunkaak Segundo, Wilder 05-30. Canyon bottom at N base of Sierra Kunkaak, Wilder 06-413. Horsfordia newberryi (S. Watson) A. Gray. Mariola; yellow felt plant, orange velvet mallow New for Dátil where it is most common on steep slopes and rocky outcrops. It is common on nearby Tiburón and on San Esteban. Slender, erect shrubs with orange flowers. Geographic Range.—Widespread in the Sonoran Desert including coastal Sonora, both states of Baja California, and Isla Ángel de la Guarda. Dátil.—Steep rocky slopes, summit of island, ridge crest, rare, 4 Dec 1965, Felger 13448. NW side of island, 20 Dec 1966, Felger 15343. MYRTACEAE *Eucalyptus camaldulensis Dehnh. Eucalipto; Murray red gum New for Tiburón and a new plant family for the Gulf islands. Planted at Tecomate and persisting near two decaying small buildings. Not reproducing on the island or in the Sonoran Desert. Geographic Range.—Native to Australia; this is the most commonly grown Eucalyptus in the Sonoran Desert and the most wide-ranging Eucalyptus species in Australia. Tiburón.—Tecomate, about 6 trees ca 5 m tall, Wilder 07-264. OROBANCHACEAE Orobanche cooperi (A. Gray) Heller. Matar; flor de tierra; desert broom-rape New for Tiburón and a new family record for the Gulf islands. The nearest record is from the vicinity of El Desemboque San Ignacio, ca 62 km to the north (Felger and Moser 1985), which previously was the south- ernmost record in Sonora. Apparently annuals, appearing in spring; parasitic. Geographic Range.—Deserts and semiarid regions in SW United States and NW Mexico in Baja California Norte and northern Sonora. Tiburón.—Cerro Kunkaak, 700 m, 11 Apr 1979, Scott s.n., et al. (UNM). POACEAE Aristida californica Thurb. ex S. Watson var. glabrata Vasey Recorded for Tiburón by Reeder and Felger (1989) but not listed by Rebman et al. (2002). This interior Ti- burón population is notable for the occurrence of var. glabrata well within the desert in contrast to its general distribution at higher elevation and/or in areas of higher precipitation, and for the most part geographically peripheral to the desert var. californica. Perennials and sometimes reproductive in the first season. Wilder et al., Plant records for the S Islands, Gulf of Mexico 1219 Geographic Range.—Arizona Upland in Arizona and Sonora, and both states of Baja California at the N and S margins of the desert. Also in grassland, oak grassland, and Chihuahuan Desert in S Arizona and N Sonora. Tiburón.— Central Valley, 13 mi S of Tecomate, 20 Feb 1968, Felger 17351. Bothriochloa barbinodis (Lag.) Herter. Zacate popotillo; cane bluestem f ing id n yon on the south- New for San Esteban, where it has been found only in a sheltered northwest west side of the island. Isla San Pedro Nolasco is the only other Gulf island Inm The nearest populations are on the Sonora mainland, mostly at widely scattered sites of higher soil moisture such as north-facing cliffs and slopes, as well as canyons and waterholes especially in mountains. Tufted perennials. Geographic Range.—Widespread in the Americas, mostly in non-desert regions including Baja California Norte and Sur, the closest peninsular location being the Sierra San Francisco. San Esteban.—SW corner of island, sheltered canyon branched off from main drainage, ca 1 m tall, rare, 200 m, Wilder 07-91. *Cenchrus echinatus L. [C. insularis Scribn.]. Guachapori, zacate toboso; southern sandbur New for Tiburón and the first record for this genus on a Gulf island other than the ubiquitous Sonoran Desert endemic C. palmeri Vasey. In our opinion C. echinatus is not native to the Sonoran Desert. It is common in weedy, disturbed places in the Bahía de Kino region and likely reached the island from there. Its occurrence at a fishing camp is strong evidence of the role fishermen or tourist campers (who often spend one to several nights on the islands) in being a vector for the spread of this and other non-native species (plants and animals) to Gulf islands. West and Nabhan (2002) report C. brownii Roem. & Schult. for Tiburón, but it is in fact C. echinatus and their report is based on the specimen cited here. The taxonomic distinction between the two species is subtle (John Reeder, personnel communication 2007). Warm weather annuals. Geographic Range.—Widespread in the Americas and adventive in the Old World; often weedy. Tiburón.—Fishing camp at Ensenada del Perro, just above high tide, 31 Oct 1993, Burillón s.n. Chloris crinata Lag. [Trichloris crinita (Lag.) Parodi]. Zacate escoba; feather fingergrass New for Tiburón, where we found it only on the lower portion of the extensive eastern bajada in a swale supporting dense desertscrub. It was growing through desert shrubs, the stems mostly 1.5—2 m tall, and intermixed with C. virgata, which was decidedly more numerous and extensive. The nearest known popu- lations are in western Sonora near the Arizona border (Felger 2000b), and to the south at Bahía San Pedro (Felger 11620) north of Guaymas and on Isla San Pedro Nolasco. Large, tufted perennials, growing and reproductive during the warmer months. Recent data shows that this species is best treated as Chloris (Travis Columbus, pers. comm., 2007). Geographic Range.—Arizona to Texas and northern Mexico, and disjunct in South America. der 06-512. = Tiburón.—Vicinity of Zozni Cmiipla, Wi Chloris virgata Sw. Zacate lagunero; feather fingergrass New for Tiburon where it was locally abundant at the inland margin of the Frankenia palmeri zone on the lower portion of the eastern bajada. It is common and weedy on the Sonoran mainland in the Bahia de Kino region. Warm-weather annuals, highly variable in size (ca 10—80 cm tall). Geographic Range.—Widespread in the Americas including Baja California Sur where the closest pen- insular location is in the Sierra San Francisco; also introduced in the Old World. Tiburón.—Vicinity of Zozni Cmiipla, abundant in water channels and decreasing in density towards the Frankenia zone (seen from this point southward for at least several km), mostly ca 60—80 cm tall, Wilder 06-353 Enneapogon desvauxii P. Beauv. Zacate lobero; spike pappusgrass New for Tiburón; documented from two inland sites. The nearest known populations are in western Sonora in the vicinity of Hermosillo and near the Arizona border (Felger 2000b) but it can expected closer in coastal mountains. Small, tufted perennials. 1220 Journal of the Botanical R h Institute of Texas 1(2) Geographic Range.—This species is cosmopolitan. The closest peninsular population is in Baja California Norte just north of Calmalli, about 140 km from Tiburón. Tiburón.—Haap Hill, 11 Dec 1976, Felger 76-T17. Deep canyon on N slope of Sierra Kunkaak, SW and up canyon from Siimen Hax waterhole, Wilder 06-464. Leptochloa dubia (Kunth) Nees [Diplachne dubia (Kunth) Scribner]. Green sprangletop New for Tiburón, where we found it near sea level along the eastern shore. It is common on the mainland above the desert and there are but few, scattered records for it at lower elevations within the Sonoran Desert (Felger 2000b). Tufted perennials. Geographic distribution.—SW US, Florida and México including Baja California Sur to South America. Tiburón.—1 km inland from Zozni Quimplla, N side of Punta San Miguel, Wilder 06-371. *Pennisetum ciliare (L.) Link. Oot iconée (coyote's grass); zacate buffel, buffel; buffelgrass New for Tiburón. It has been in the vicinity of Caracol at least since 1998 (West & Nabhan 2002). This is currently the most actively used and disturbed site in the interior of the island, and where Comcáac guides often stay with bighorn sheep hunters during prolonged hunting activities on the island. A second popula- tion was found in 2007 just inland from Estero San Miguel on the eastern bajada. On the Sonoran mainland buffelgrass is extensively planted for cattle grazing after removing the desert vegetation. For example, one large plot was observed in 2007 at the western base of Pico de Johnson of the Sierra Seri. Buffelgrass lines Highway 10 linking Hermosillo and Bahía de Kino, and is especially thick at the entrance to the coastal town. It also is along the length of the road between Kino Nuevo and Punta Chueca and is expanding in this area. Perennials and often reproductive in the first season; growing and reproductive with sufficient soil moisture especially during the warmer months. Buffelgrass was reported for Alcatraz (West & Nabhan 2002; West et al. 2002), where it is said to be eradicated (Tad Pfister, pers. comm., 2006), although there are no voucher specimens for the island. Extensive buffelgrass on the nearby mainland makes continued re-population likely. The Midriff islands are free of significant populations of buffelgrass. Unlike most of the Sonoran Desert, A, a unique opportunity exists to control this invasive species on the Gulf islands before it becomes nearly impossible to thwart its invasion and the subsequent ecosystem transformation associated with this noxious weed (Búrquez et al. 2002; Franklin et al. 2006). This species is one of the most serious conservation threats to Tiburón and other Gulf islands (West & Nabhan 2002). Immediate reporting of new populations followed by control efforts is essential. In May 2007, Humberto asked Steve and Cathy Marlett for suggestions for a Seri name for buffel — so that the people could recognize it by name. The three of them decided to call it oot iconée, because unusual or odd items are said to be belong to the coyote. Humberto and other Comcáac are attempting to eradicate in from Tiburón. Geographic Range.—Native to the Old World and widely introduced for forage and fodder in arid and semiarid regions worldwide, and generally becoming invasive. Tiburón.—Caracol Research Station, E foothills of Sierra Kunkaak, small canyon arroyo: 100 m W of station, scattered plants occur along a 100 m section of the arroyo, with the total population ca 50 plants, 24 May 2006, Wilder 06-151; Population expanded since May 2006, buffel lines arroyo for ca 200 m, all plants pulled, 3 May 2007, Wilder 07-230. Vicinity of Estero San Miguel, ca 0.75 km S of Zozni Cmiipla and ca 400 m inland, scattered populations in the area, 28%57,824'N, 112?13.241'W, in total ca 200 plants, many seedlings, all plants pulled, 4 May 2007, Wilder 07-232. Lower E bajada, ca 3.75 km S of Zozni Cmiipla and ca 500 m inland, along road that leads to Pazj Hax waterhole, 28%56.171'N, 112?13.503'W, single plant seen, pulled, 2 May 2007, Wilder (observation). Setaria leucopila (Scribn. & Merr.) K. Schum. Hasac, xiica quiix; white-haired bristlegrass Recorded as S. macrostachya Kunth for San Esteban by Johnston (1924) and Tiburón, San Esteban, and Dátil by Felger and Lowe (1976) and Moran (1983). The specimens are identifiable as S. leucopila if one is to rec- Wilder et al., Plant records for the S Islands, Gulf of Mexico 1221 ognize it as distinct from S. macrostachya (see McVaugh 1983:361). Rebman et al. (2002) omit S. macrostachya and S. leucopila from the main checklist but report S. macrostachya from Dátil. Here we re-confirm Tiburón and San Esteban island occurrences of S. leucopila with specimens. Densely tufted perennials. Geographic Range.— Central and NW Mexico including Baja California Norte y Sur and SW United States. Tiburón.—Deep canyon on N slope of Sierra Kunkaak, SW and up canyon from Siimen Hax waterhole, Wilder 06-478. Canyon bottom at N base of Sierra Kunkaak, Wilder 06-421. Coralitos, S end of island, just inland from beach, ca 5 m, Wilder 06-55. La Pescadita, S shore of island, 11 Oct 1977, Wilkinson s.n. San Esteban.—Arroyo Limantour, Van Devender 92-482. Arroyo [Limantour], 13 Oct 1977, Wilkinson s.n. Sporobolus virginicus (L.) Kunth. Xojásjc; zacate salado de la playa; seashore dropseed Felger and Moser (1985, p. 316) reported it for Tiburón “along beaches and inland margins of mangroves” but it is not listed for the island by Moran (1983a) and Rebman et al. (2002). We confirm its presence with a specimen. Perennial saltgrass; the Gulf of California plants are notably robust. Geographic Range. —Tropical and subtropical shores worldwide. Tiburón and Alcatraz and the Sonora coast S from near Bahía de Kino, and the Baja California peninsula S from Bahía de San Francisquito. Tiburón.—Zozni Cmiipla at Estero San Miguel, Wilder 06-361. Tridens muticus (Torr.) Nash var. muticus. Slim tridens New for Tiburón. We found it in the interior of Sierra Kunkaak. The nearest known population is 150 km to the north in the Sierra del Viejo southwest of Caborca, which is the southernmost documentation for this grass for mainland Sonora (Felger 20005). Itis one of the most widespread and drought-tolerant perennial grasses of the Sonoran Desert; often in surprisingly harsh, xeric habitats. Tightly clumping small perennials; growth and flowering response apparently non-seasonal depending on soil moisture. Geographic Range.—SW United States and N Mexico (N Sonora, N Baja California Norte, Chihuahua, Coahuila, and Nuevo León); deserts, grasslands, and oak woodland. Tiburón.— Canyon bottom at N base of Sierra Kunkaak, between Sierra Kunkaak Mayor and Sierra Kunkaak Segundo, 405 m, Wilder 06-401. Deep canyon on N slope of Sierra Kunkaak, SW and up canyon from Siimen Hax waterhole, Wilder 06-479. Urochloa fusca (Sw. B.F. Hansen & Wunderlin [Brachiaria fasciculata (Sw.) Parodi] Browntop signal- grass New for Tiburón and San Esteban. It is apparently fairly common locally following times of favorable sum- mer/fall rains. The nearest populations are on the Sonoran mainland at Bahía de Kino (Van Devender 90-519). Summer/fall ephemerals, highly variable in size. Geographic Range.—Widespread in the Americas including Baja California Sur. Tiburón.—Vicinity of Haap Hill, 11 Dec 1976, Felger 76-T10. San Esteban.—Arroyo Limantour, uncommon annual, 15 Sep 1990, Van Devender 90-537. PORTULACACEAE Portulaca oleracea L. Verdolaga; purslane New report for Tiburón. The two widely separated localities on the island indicate it is probably more widely distributed. It is widespread on the adjacent Sonora coast. Warm weather annuals. Geographic Range.—Worldwide in tropical to warm-temperate climates including Baja California Norte y Sur. It is often difficult to determine which populations might be native. Tiburón.—Ensenada de la Cruz, 27 Feb 1965, Felger 12778. Haap Hill, 8 Sep 1974, Felger 174-24. PTERIDACEAE Notholaena californica D.C. Eaton subsp. californica. California cloak fern New for San Esteban. This is the only fern on San Esteban where it occurs in sheltered north-facing canyons 1222 Journal of the Botanical R h Institute of Texas 1(2) in the southern part of the island. On Tiburón and San Esteban it is restricted to seasonally moist niches among rocks. It is the most widely distributed fern on Tiburón and other Gulf islands (Johnston 1924:980), and one of the most xeric-inhabiting ferns in the Sonoran Desert (e.g., Felger 2000b). Small tufted ferns or with short rhizomes. Geographic Range.—S California to the Cape Region of Baja California Sur, S Arizona, and N Sonora to the vicinity of Bahía de Kino (see Felger 2000b). San Esteban.—SW corner of island: Sheltered canyon branched off from main drainage, in sheltered nooks below rocks, extremely common in this side canyon only, Wilder 07-86; W-most N-trending canyon der 07-93. = in the area, in sheltered nooks below rocks, occasional, Wi Notholaena lemmonii D.C. Eaton var. lemmonii New for Tiburón, where it grows in sheltered sites at higher elevation in the Sierra Kunkaak. The nearest known populations are in the Sierra el Aguaje, ca 150 km to the southeast (e.g., Bahía San Pedro; Johnston 1924, p. 980) and on Isla San Pedro Nolasco, ca 140 km to the SE of Tiburón. Small tufted ferns. Geographic Range.—Mountains in Arizona, Chihuahua, much of Sonora, Baja California Sur, and on Isla Cerralvo. Tiburón.—Sierra Caracol [part of Sierra Kunkaak], 27 Oct 1979, Knight 1062 et al. (UNM). Top of Sierra Kunkaak Segundo, ca 450 m, Wilder 06-493. Notholaena standleyi Maxon. Star cloak fern New for Tiburón, found at higher elevations in the Sierra Kunkaak, and the first record for a Gulf island. It occurs in the Sierra Seri in adjacent western Sonora. Small tufted ferns. Ours belong to the western “golden race,” so called because of the golden-colored exudate or farina on the lower surface of the leaf blades. Geographic Range.—SW United States and northern Mexico to Pueblo. Tiburón.— Cerro San Miguel, Quijada-Mascareñas 911013. S slope of Sierra Kunkaak, in sheltered rock crevice, ca 350 m, rare, 1 Jan 2006, Wilder (photo). RUBIACEAE Galium proliferum A. Gray. Desert bedstraw New for Tiburón and the first record for a Gulf island. The nearest known locality is on the Baja California peninsula in the Sierra Asamblea just northwest of Bahía de los Angeles, ca 140 km west from the Sierra Kunkaak. It is also present in Sierra el Aguaje (La Balandrona, Felger 01-650), ca 150 km to the southeast. Delicate, small and often minute winter-spring ephemerals. Geographic Range.—SE California to W Texas and northern Mexico. Tiburón.—Base of the N portion of Sierra Kunkaak, ca 2 km E of Siimen Hax waterhole, 360 m, sand- gravel soil in shaded margin of arroyo bed among leaf litter, Wilder 06-458. Galium stellatum Kellogg var. eremicum Hilend & J.T. Howell. Starry bedstraw New for San Esteban, collected by Frank J. Wegscheider of California State University, Fullerton. The near- est population is on Isla San Lorenzo (Moran 19833). Not recorded from Tiburón, although it occurs on the opposite mainland at Cerro Tepopa. Untidy small shrubs or subshrubs. Geographic Range.—Mostly in desert mountains; SW United States to N Baja California Sur and N Sonora southward in arid, coastal mountains to Cerro Tepopa Q9'21'N). San Esteban.—Inland on NE quadrant of island, (UTM 12 3178413 N, 12 345752 E, NAD 83), 30 Mar 2005, Wegscheider 101 (SD). SAPOTACEAE Sideroxylon leucophyllum S. Watson. Hehe pnaacoj New for Tiburón and the first report for mainland Sonora. On Tiburón we know of it only on the north-face of an eastern peak of the Sierra Kunkaak, where it grows on a steep talus of large boulders (Fig. 3B). Here Wilder et al., Plant records for the S Islands, Gulf of Mexico 1223 it is locally common, developing into large shrubs and small trees to ca 5 m tall with substantial trunks to ca 30 cm in diameter. The peaks of San Esteban and Tiburón are often shrouded in fog clouds—these mountains “make their own microclimate.” The leaves of S. leucophyllum are densely woolly-pubescent and might be efficient in capturing moisture. On San Esteban it occurs only in a few sheltered canyons and ridgecrests. On 4 May, 2007, Benjamin spotted a small population with binoculars on a steep talus slope of Pico de Johnson, on the Sonora mainland opposite Sierra Kunkaak. Who will be the first to confirm this mainland sighting with a specimen? Geographic Range.— Previously known only from Baja California (Norte) and Islas Ángel de la Guarda and San Esteban. Tiburón.— Cerro Kunkaak, 700 m, 11 Apr 1978, Scott s.n., et al. (UNM 53277). Cerro San Miguel, árbol de baja estatura, común en laderas rocosas, principalmente en la ladera norte, 4—5 m, con tronco definido, Quijada-Mascarenas 917009. Top of Sierra Kunkaak Segundo, ca 490 m, N-facing slope among talus boulders, large trunks, Wilder 06-491. SOLANACEAE Lycium berlandieri Dunal var. longistylum C.L. Hitchc. Bachata, salicieso New species record for Tiburón. There are records from the Sierra el Aguaje north of Guaymas, ca 150 km to the southeast, and in the vicinity of Hermosillo, although we expect it in closer mountains. Shrubs with smooth, dark red-brown bark, white or pale yellow-white flowers, and campanulate corollas. Geographic Range.—This species occurs in SW United States and NW Mexico. Var. longistylum ranges from central and SW Arizona southward in W Sonora to the Guaymas region (Chiang-Cabrera 1981). Tiburón.—Base of N portion of Sierra Kunkaak, Siimen Hax waterhole, ca 355 m, Wilder 06-454. Physalis crassifolia Benth. var. versicolor (Rydb.) Waterf. Xtoozp; tomatillo del desierto; desert ground cherry New for Dátil, where it was found on the peaks of the north-central part of the island. It is widespread on nearby Tiburón. Small perennial subshrubs. Geographic Range.—SW United Sates and NW Mexico. Two other varieties occur in the Baja California peninsula and on Gulf islands. Dátil.—Peak of N-central part of island, ca 140 m, Wilder 07-126. TAMARICACEAE *Tamarix aphylla (L.) H. Karst. Hocó hapéc (translated as: any planted tree from outside of the area); pino salado; athel tree, salt cedar West and Nabhan (2002) report it on Tiburón, but it is not listed for the island by Moran (19832) and Reb- man et al. (2002). We confirm its presence with specimens from two localities. Tamarisk trees were planted to a significant extent at the now abandoned small military station at Tecomate and there are also several trees at the active Mexican marine station at Punta Tormenta. These trees were planted several decades ago and have persisted to develop into large, healthy shade trees, but are not reproducing. Geographic Range.—Native to the Old World and extensively planted in the Sonoran Desert including the villages on the Sonoran mainland opposite Tiburón (Felger & Moser 1985). Tiburón.—Palo Fierro [Punta Tormenta], just inland from beach, Wilder 06-143. Tecomate, ca 30 indi- viduals planted in rows, trees ca 6 m tall forming dense cover, a few also near two decaying buildings where a few Eucalyptus are also planted, Wilder 07-250. ULMACEAE Celtis reticulata Torr. Cumero; canyon hackberry, western hackberry New for Tiburón and the first record for a Gulf island. We found it on a high rock talus in the Sierra Kunkaak, a surprising and unusual habitat for this species (it usually grows along riparian or semi-riparian drain- ageways). The nearest Sonoran populations are in deep riparian canyons in the Sierra el Aguaje north of 1224 Journal of the Botanical R h Institute of Texas 1(2) Guaymas, such as Cañón Nacapule (Felger 1999) and the closest peninsular population is in the Sierra San Francisco. Shrubs on Tiburón, trees with smooth, gray bark in west-central and southern Sonora. The type locality is considered to be somewhere in Colorado (James Henrickson, pers. comm., 2006), and trees from that region might not be conspecific with those in southern Sonora. The Arizona and espe- cially the Tiburón and Sonoran populations have relatively thick leaves. The southern Sonora populations are essentially evergreen rather than winter-deciduous like those farther north (Felger et al. 2001). Geographic Range.—Widespread in W North America including Baja California Sur. Tiburón.—Top of Sierra Kunkaak Segundo, 490 m, shrub ca 2 m tall, Wilder 06-486. VERBENACEAE Lantana velutina M. Martens & Galeotti. Confiturilla blanca New for Tiburón. The nearest locality in Sonora is in the Sierra el Aguaje (La Balandrona, Gutiérrez 00-07, USON) and the closest peninsular population is in the Sierra San Francisco. The only other Gulf island record is for Cerralvo (Rebman et al. 2002), the most southern Gulf island. Small shrubs with corollas white to cream. Geographic Range.—Baja California Sur and Sonora to Panama. Tiburón.—Cerro San Miguel, flores de color blanco a crema, de común a abundante en la pediente del cerro, Quijada-Mascareñas 90T011. Top of Sierra Kunkaak Segundo, 425-490 m, common on slope and becoming less so closer to the peak, shrubs ca 1 m tall, flowers bright white, Wilder 06-484. ZYGOPHYLLACEAE *Tribulus terrestris L. Cosi cahóota, cózazni caacól, hee inóosj, heen ilít, hehe ccosyat; torito, toboso; goathead, puncture vine New for Tiburón and Alcatraz. The localities on these islands are along the shore with sand soil and associ- ated with human disturbance. Tribulus is abundant in disturbed habitats along the adjacent Sonora coast. Warm weather ephemerals. Geographic Range.—Native to the Old World, this noxious weed is now widespread in the warmer regions of the world. Tiburón.—Punta Tormenta, ca 1.5 km S of military (marine) station, Wilder 06-345. Alcatraz.—12 Nov 2001, Gracida 35 (Prescott College Collection). Sand flat at E side of island, not com- mon, 8 Oct 1966, Felger 14923. GAZETTEER Key to Abbreviations: BCN =Baja California hii BCS = Baja California Sur, EST = Isla San Esteban, SON = Sonora, TIB = Isla Tiburón. Coordinates are from Google E Arroyo Limantour (EST). Main drainage of the island, terminating on the E shore at 28°41'14.7'N, 1 T oa Arroyo Sauzal (TIB). Major drainage of the S part of the island, 28°47' PN 112°25'28"W at the co Bahia de Kino (SON). Coastal fishing and vacation town; largest ( ation center in the vicinity ie Midriff islands. Bahía de San Francisquito (BCN). NE part of the state. 28°26'24"N, 112°52' 14"W. Bahía San Pedro (SON). Ca 20 km NW of San Carlos and ca 15 km NE of Isla San Pedro Nolasco. 28°03'17"N, 111°14'39"W. Balandrona, La (SON). "ud riparian canyon on the N side of the Sierra Aguaje, canyon mouth in the vicinity of 28°06'N, 11°04'17"W, ca 21 Canal del Infiernillo (son). Shallow channel separating Tiburón from the mainland. N end at 29°14'49"N, 112°15'10"W and S end at 28%57'58"N, 112°10'57"W. Caracol Research Station (TIB). E side and base of Sierra Kunkaak. 29°00'56.7"N, 112°17'36.9"W, 190 m. Central Peak (EST). The major mountain mass, dominating the interior, peak at 28°42'29.02"N, 112*34'31.86"W. Central Valley [= Valle Agua Dulce] (TIB). Expansive valley draining to the N shore, between Sierra Kunkaak on the E and Sierra Menor on the W, running the majority of the island's lengt SED seas Miguel [= Sierra Kunkaak Segundo] (TIB). High eastern a of the Sierra Kunkaak, slightly less than 1,000 m. °58'52.27"N, 112°18'30.96"W ol (TIB). Cove at S shore. 2845! 30"N, 112°19'20"W. Desemboque del Rio San Ignacio, El (SON). Northern of the two Comcáac villages. 29°30'15"N, 112?23'46"W. Wilder et al., Plant records for the S Islands, Gulf of Mexico 1225 Ensenada de la Cruz [=Pescadita, La] (TIB). Bay at S shore. 28?45'54"N, 112%21'4,56"W, Ensenada del Perro (TIB). Tem porani Tnn) camp at SE shore. 28°46' aui 11 21 6 AN | ie NA; | Estero San Miguel (TIB). Tidal wetla ly peninsula (P E shore of island end of Canal del I eS: 28°58'1 14'N, 1 12:12 Q9"W. Haap Hil (IB). SE side of Central Valley, ca 12 mi S from Tecomate. N side and base of basaltic hills, vicinity of former Seri mp Haap Caaizi Quih Yaii, named for gathering of wild tepary beans (Felger & Moser 1985; Moser & Marlett 2005). Vicinity of 28°57'20"N, 112°24.5'W, ca 280 m. Felger collections: 8 Sep 1974 with Cayetano Montaño, Hank Gunn, and Alexander Russell; 11 Dec 1976 with Rosa Flores, Cathy Moser Marlett, and Alexander Russell. Hant Hax (TIB). Fou! Pell camp, at E base of Sierra Kunkaak, between Sopc Hax and Zozni Cmiipla; approximately 28°57'30"N, 112°15 Pescadita, La (TIB). [See Ensenada la Cruz] Palo Fierro (TIB). See Punta Tor Pazj Hax [= Tinaja Anita] A at E base of Sierra Kunkaak. 28°56'25.41"N, 112%16'45.63"W, Pico de Johnson (SON). Highest peak in the Sierra Seri, summit ca 29°02'15.65"N, 112°08'14.66"W. Punta Chueca (SON). Southern of the two Comcáac villages. 29°00'54"N, 112°09'37"W. Punta Tormenta [= Palo Fierro landing field] (TIB). Small Mexican Marine station and Comcáac vehicle-staging area on shore opposite Punta Chueca. 29°01'11"N, 112%11'43"W, Sierra el Aguaje (SON). Extensive coastal mountain mass to the N of San Carlos (N of Guaymas), including Cañón del Na- capule (see Felger 199 Sierra Bacha [= ss See (SON). Granitic ins S of Puerto Libertad (29°50'40"N, 112°38'18"W) and N of El Desem- e del San Sierra P Viejo ou E rge mountain mass ca 45 km SW of Caborca. 30°20'35"N, 112°19'54"W. Sierra Kunkaak (TIB). Largest and highest (1,200+ m) mountains on the island, running along most of the E side of the island. Sierra Kunkaak Mayor (TIB). The highest and most extensive portion of the Sierra Kunkaak, containing deep, sheltered Sierra Kunkaak Segundo (TIB). The E peak of Sierra Kunkaak; see Cerro San Miguel. Sierra San Francisco (BCS). Large volcanic mountain range, ca 1,600 m tall, S of state line separating Baja California and Baja California Sur. The range provides a link between tropical components of the southern peninsula and temperate influences of the north. 27°39'20" N 1 a 54'58"W. (Jon Rebman, pels comm., 2007). Sierra Seri (SO N). E tensive ra nge and ODpos site the Sie Kunkaak (see Pico de Johnson). Siimen Hax (TIB). Waterhole in the E ee 28*58'47 2"N, A 19'30.6"W. Sopc Hax (TIB). Waterhole in the Sierra Kunkaak. 28°57'15.9"N, 112?16'39.3"W. Tecomate MIB Former Comcáac village on the N shore, and more recently site of abandoned small military station. 29*11'12"N, 112?2458"W. Tinaja Picu ee ld site of Ira Wiggins, in the Sierra Picu, along the road from Pitiquito to Puerto Libertad, ca 20 i from the Valle Agua Puls uu See Central Valley. Valle de Águila (TIB). An area of bajada at the NE side of the island. Vicinity of 29*05'28.68"N, 112*15'15.12"W. Zozni Cmiipla (TIB). Historic Comcáac camp near the base of Estero San Miguel on the E side of the island. 28*58'08"N, 1125 12:5 DAN ACKNOWLEDGMENTS Many friends and colleagues have facilitated our work on the islands. Dave Bertelsen, Alberto Bürquez- Montijo, Exequiel Ezcurra, Edward Gilbert, Mikhal Gold, Jesás Sanchez-Escalante, Gloria Guadalupe Mo- rales-Figueroa, Powell B. (Gil) Gillenwater III, Florencio Cota-Moreno, Jesús Ventura-Trejo, and Seth Turner accompanied us in the field—thank you for sharing in the adventure. Ana Luisa Rosa Figueroa-Carranza of CONANP, Larry Johnson at Bahía de Kino, Tad Pfister and the Prescott College Center for Cultural and Ecological Studies at Bahía de Kino, and William J. (Bill) Risner of Tucson provided logistical support and shared their knowledge of the region. A. Elizabeth (Betsy) Arnold, Exequiel Ezcurra, Cathy Moser Marlett, Stephen Marlett, John R. Reeder, and Charlotte Goodding Reeder provided valuable information and assis- tance. Cathy Moser Marlett prepared the maps. Previous floristic work on the Gulf islands, especially that by Reid Moran and Jon P. Rebman, provided the foundation for this publication. Brad Boyle, Phil Jenkins, and Michelle (Shelley) M. McMahon of the University of Arizona herbarium (ARIZ), and Jon P. Rebman and especially Judy Gibson of the herbarium of the San Diego Natural History Museum (SD) provided valuable 1226 Journal of the Botanical R h Institute of Texas 1(2) assistance. Additionally we thank Xavier Basurto, Travis Bean, Julio Betancourt, Janice E. Bowers, Thomas Bowen, Mark Dimmitt, Christopher K. Frazier (UNM), Powell B. (Gil) Gillenwater III, Paul Knight, R. James (Jim) Hills, Paul S. Martin, Jane Mygatt (UNM), Ana Lilia Reina-Guererro, Peter Sherman, Raymond M. Turner, Thomas Van Devender, and Michael Wilson for their help. Collections have been made under Mexican Federal Collecting permit NOM-126-SEMARNAT-2000 obtained with the generous assistance of Exequiel Ezcurra. We appreciate comments by reviewers John L. Anderson and Jon P. Rebman. Funding for this project has been received by Wilder from the University of Arizona Mycological Herbarium, Arizona-Nevada Academy of Science, University of Arizona Honors College Alumni Legacy Grants, The University of Arizona Department of Ecology and Evolutionary Biology's Leslie N. Goodding Memorial Scholarship, and the Research Committee of the Cactus and Succulent Society of America. Felger received support from the Wallace Research Foundation and more recently the World Wildlife Fund in collaboration with El Área de Protección de Flora y Fauna (APFF) “Islas del Golfo de California” en Sonora, de la Comisión Nacional de Áreas Naturales Protegidas (CONANP), de la Secretaría de Medio Ambiente y Recursos Naturales (SEMARNAT). REFERENCES Austin, D.F. 1998. Convolvulaceae, morning glory family [Vascular plants of Arizona]. Arizona-Nevada Acad. Sci. 30:61-83. Burquez-Montuo, A., M.E. MILLER, and A. Martinez-Yrizar. 2002. Mexican grasslands, thornscrub, and the transforma- tion of the Sonoran Desert by invasi ic buffelgrass (Pennisetum ciliare). In: B. Tellman, ed. | ive exotic species in the Sonoran Region. University of Arizona Press and Arizona-Sonora Desert Museum, Tucson. Pp. 126-146. Case, T.J. and M.L. Copy (eds.). 1983. Island Biogeography in the Sea of Cortéz. University of California Press, Berkeley. Case, T.J, M.L. Copy, and E. Ezcurra (eds). 2002. A new island biogeography of the Sea of Cortés. University of California Press, Berkeley. CHIANG-CABRERA, F. 1981. A taxonomic study of the North American species of Lycium (Solanaceae). Ph.D. disserta- tion, University of Texas, Austin. Copy, M.L., R. Moran, and H. THompeson. 1983. The plants. In: T.J. Case and M.L. Cody, eds. Island biogeography in the Sea of Cortéz. 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Seed plants of the Northern Gulf of California. In: R.C. Brusca, E. Kimrey, and W. Moore, eds. A seashore guide to the Northern Gulf of California. Arizona-Sonora Desert Museum, Tucson. Pp. 147-163. FELGER, R.S., M.B. Johnson, and M. F. Witson. 2001. Trees of Sonora, Mexico. Oxford University Press, New York. FELGER, R.S. and E. Joyat. 1999. The Palms (Arecaceae) of Sonora, Mexico. Aliso 18:1-18. FeLGER, R.S. and CH. Lowe. 1976. The island and coastal vegetation and flora of the northern part of the Gulf of California, Mexico. Nat. Hist. Mus. Los Angeles County, Contr. Sci. 285:1—59. Wilder et al., Plant records for the S Islands, Gulf of Mexico 1227 FELGER, R.S. and M.B. Moser. 1985. People of the desert and sea: ethnobotany of the Seri Indians. University of Arizona Press, Tucson. Reprinted 1991. FRANKLIN, K.A., K. Lyons, P.L. NAGLER, D. LAMPKIN, E.P. GLENN, F. MOLINA-FREANER, T. MARKOW, and A.R. Huete. 2006. 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Arts Sci. 24:36-82. — 1228 Journal of the Botanical R h Institute of Texas 1(2) BOOK NOTICES CHRISTINE Finni (ed.). 2006. Annual Plant Reviews, Volume 28: Plant Proteomics. (ISBN 1-4051-4429-7, hbk.). Blackwell Publishing, Ames, IA 50010-0570, U.S.A. (Orders: 800-862-6657, 515-292-0155, www.blackwellprofessional.com). $199.99, 253 pp., color photographs, black-and-white photographs, black-and-white illustrations, 6 1/4" x 9 1/2". The 28'^ volume of Blackwell Publishing's Annual Plant Reviews is edited n Christine Finnie Bi the Technical University of Den- mark and covers the emerging research in the field of Lena proteomics. T t ..all protein species resulting from ..the expression pattern, cellular location, activity, gene expression in a cell, organelle, oteomics is the m Olas regulation, post-translational modifications EU a interactions, three dimensional structures ian functions of each protein in a ther biological system." With the completion of the tees thaliana I , the pr ee d Sales and ae 1 Aat m plants are under investigation. This volume is intended to keep p-t this fascinating field. Chapter 1 covers the challenges of pl i d tl ilabl includi ic datab and online tools. Chapter 2 covers “Proteomic analysis of post-translational modifications by mass peace: ” Chapter 3 covers “Strategies for the investigation of cu -protein interactions in uod e 4 discusses the research on proteins that undergo respectively. Chapter 7 covers “Proteome cysteine and disulfide oe reactions. p 1 E] analysis for the study is a discussion of the pyar sonics of 2 ER — Marissa N. Oppel, MS, Irving, IX, 75060, U.S.A. I ” and Chapter 8 di ll wall proteome. Finally, Chapter 9 SABEEHA MERCHANT, WINSLOW R. Bricas, and Vicki L. CHANDLER (eds). 2007. Annual Review of Plant Biology: Volume 58, 2007. (ISBN 978-0-8243-0658-8, hbk; ISSN 1543-5008). Annual Reviews Inc., 4139 El Camino Way, PO. Box 10139, Palo Alto, CA 94303-0139, U.S.A. (Orders: www.AnnualReviews. org, onlineactivation@annualreviews.org, 800-523-8635, 650-493-4400, 650-424-0910 fax). $212.00 (USA), 501 pp., 7 3/8" x 9 1/4". Contents of Volume 58 of Annual Review of Plant Biology: 1. From Analysis of Mutants t tic Engineering 2. Phototropin Blue-Light Receptors 3. Nutrient Sensing and dnd NPKS 4.1 d Hydrogen PI ion in Oxygenic P ] Organisms 5. Hidden Branches: E in Root System Architecture 6. Leaf Senescence 7. The Biology of Arabinogalactan Proteins 8. Stomatal Development 9. Gibberellin Receptor and Its Role in Gibberellin Signaling in Plants 11. Cyclic Electron Transport Around LEE I: Genetic Approache ight Regulation of Stomatal Movement 13. The Plant H imeric G-Protein ACER 14. Alternative Splicing of Pre-Messenger RNAs in Plants in the G ic Era 15. The Production of Unusual Fatty Acids in Transgenic Plants 16. Tetrapyrrole Biosynthesis in Higher Plants 17. Plant ATP-Binding Cassette Transporters 18. Genetic and Epigenetic Mechanisms for Gene Expression and Phenotypic Variation in Plant Polyploids 19. Tracheary Element Differentiation 20. Populus: A Model System for Plant Biology 21. Oxidative Modific: to Cellular Components in Plants Indexes J. Bot. Res. Inst. Texas 1(2): 1228. 2007 CHECKLIST OF THE VASCULAR PLANTS OF WASHINGTON COUNTY, PENNSYLVANIA Cynthia M. Morton Loree > Section of Bota Section of Bot uL Mee ou r d Mi cun on Rem crei "€ MNA 15213, U.S.A. "T piis A 15213, U.S.A. ABSTRACT Md County has been botanized since the a AE however, a checklist of tl j lar floral previously been ompiled. A search of herbaria augmented by targeted fiel lted in a list of 982 t f nati ] lized pl prising 472 genera and 125 families. The five families, with the N LN S pd; are, Asteracae, Poaceae cC Rosaceae and Fabaceae. Carex, Symphyotrichum, Solidago, Polygonum, and R There are approximately 203 non-native species that have been introduced mainly from Europe and Eurasia. This de recognizes 3 species of Lycophytes, 31 species of Pteridophytes, 10 species of Gymnosperms and 911 species of Angiosperms. Forty-one species have global or state rankin RESUMEN El Condado de Washington, se localiza en el estado de Pennsylvania. Est dad iderado una de las ár ás di estado, sin embargo hasta ahora no ha sid pletad lista de verificación. Esta lista de verificación contiene familias, géneros, y ebbe alfabéticamente dentro de los grupos k ve Vee tales como Pteridofitas, Gimnospermas y tas Esta lista i 1 Condado de Washington. Un total de al menos 125 familias, 472 géneros, y 955 p fueron encontradas. Las cinco m. ru D el número de es- pecies fueron Asteraceae, Poaceae Cyperaceae, R Kosaceae yF abaceae. I gi g p núm p i , fueron Carex, Symphyotrichum, Solidago, Polygonum y Rubus. INTRODUCTION 1 DES INES | 1 E Even though P diverse, Rhoads and Klein (1993) reported 3318 taxa of vascular plants for the state, including 2076 native for most of its counties, the vascular flora known thus far is quite and 1242 introduced. Approximately 26 of Pennsylvania's 67 counties have floras completed, but some are unpublished theses and surveys and therefore unavailable to the public. The exact number of genera and species in the flora of Washington County was not previously known because a checklist had never been compiled. Currently Washington Country is being seriously impacted by coal mining making documentation of the flora all the more urgent. This checklist will be needed when reclamation and revegetation decisions are made of the areas that have been mined for coal. Such checklists provide baseline information that can be used to monitor environmental changes and guide conservation decisions. Conservation of plant biodiversity, including monitoring of native and/or invasive plants, has not been a priority in Washington County. Three county parks, one state park and six game lands have been created to protect and manage biota. The three county parks consist of Cross Creek, Mingo Creek, Ten Mile Creek and the state park, Hillman State Park, is managed for hunting by the Pennsylvania Game Commission. To-date, no publications have been produced documenting native and/or invasive plants for any area of Washington County. Site Description Washington County, Pennsylvania is located in the southwestern corner of the state (Fig. 1). It is bordered by the counties of Beaver and Allegheny to the north, Allegheny, Westmoreland and Fayette to the east, Greene to the south and Marshall, Brooke, and Ohio Counties, West Virginia to the west. Washington County receives total annual precipitation of approximately 38 inches, of which 55 percent usually falls in April through September. Average seasonal snowfall is 31 inches. The last frost of the season J. Bot. Res. Inst. Texas 1(2): 1229 — 1249. 2007 1230 Journal of the Botanical R h Institute of Texas 1(2) Washington County, Pennsylvania Hanover Twp BM X ES SP Burgettstown Gu 3 Sd «McDonald Smith Twp Midway Jefferson T "E Cecil Twp Mt Pleasant Twp Peters Twp Chartiersf Canonsburg SHouston Finleyville Twp ^ North Union Straban Tw e Tw Nottingham New Eagle South P Two Monongahela Strabane LONE ANC A Tw Carroll Twp Donora Somerset Twp ‘Charleroi Fallowfield Twp Sart Bentleyville |Speers or A Ellsworth Long Amwell ¿Bethlehem b Branch Twp West Pike Allen ort T pi TE Run Twp i e es California West Beallsville Bethlehem 1 Wes) C t zu s TWP Deemston VIS Brownsville a East Bethlehem Kilometers Miles is usually around May 18" and the first frost of the season approximately September 25!^. The length of the growing season, from the last killing frost in spring to the first killing frost in autumn, is about 149 days (USDA 1983). Wright’s Woods in Washington County has been reported by Jeffrey D. Wagner of the Western Penn- sylvania Conservancy as an old growth oak forest. Located behind Wright's United Methodist Church in Washington County it is classified as a Mesic central Forest. In Wagner's own words *below the white and red oak (Quercus alba and Q. rubra) canopy is a sub-canopy and shrub layer of predominately sugar maple (Acer saccharum). Basswood (Tilia sp.), hackberry (Celtis occidentalis), slippery elm (Ulmus rubra), white ash (Fraxinus americana) and a number of other tree species grow within the oak-sugar maple forest, as do a number of shrub/trees like and spicebush (Lindera benzoin), common elderberry (Sambucus canadensis), and flowering dogwood (Cornus florida). A rich spring flora of trout lily (Erythronium americanum), spring beauty (Claytonia virginica), false mermaid (Floerkea proserpinacoides), virginia waterleaf (Hydrophyllum virginianum) and many other species blanket the slopes, particularly the lower slopes. To the east, the slope transforms Morton and Speedy, Flora of Washington County, Pennsylvania 1231 to a younger forest of sugar maple and black cherry (Prunus serotina), ending eventually at a driveway and several homes. Confined by an old cemetery and road to the west and by old strip mined land that is now residential development to the south and east, this community stands as a remnant of a forest type that likely once covered large parts of the county. A number of hiking/walking trails run through the forest and some cutting of individual trees (possibly dead or damaged red oaks) has occurred on the lower slopes. The creek itself is open, swampy in places, and appears to have been pastured at one time. Although some invasive exotic plants like garlic mustard (Alliaria officinalis), multiflora rose (Rosa multiflora) and periwinke (Vinca minor) occur densely in areas surrounding the forest, the interior remains relatively free from these plants. Although this community is destined to transform to a sugar maple forest as the old oaks die, this area will remain, if protected, one of the older forest communities in the country. Critical to the maintenance of this community is limiting disturbance within the forest and expanding, wherever possible, the buffer areas surrounding the forest. Motorized vehicles should be restricted and clearing of vegetation or cutting of trees, even dead or downed trees, should be prohibited (Wagner 1994).” We observed Wright's Wood in the spring, summer and fall of 2005. The area appeared heavily browsed by deer. In fact it was the least diverse of all our sites for Washington County. The effects of herbivory have not been well documented for Washington County. This site has been possibly altered by selective deer browsing which has reduced species richness and thus species composition. Whether these changes are permanent will be determined in due time and with additional studies. The soils of the county consist of four main series: Dormont-Culleoka association, Guernsey-Dormont- Culleoka association, Dormont-Culleoka-Newark association, and Udorthents-Culleoka-Dormont association The Dormont-Culleoka association consists of hills with benches and ridges. Dormont soils are found predominantly on hillsides and benches and commonly have slips on them. These soils are deep and mod- erately well drained. They consist of shale, siltstone, limestone and colluvium. The Culleoka soils are mostly on ridges and hilltops but some areas are on hillsides. They consist of limestone, sandstone, siltstone, and shale. Most of this area is wooded or cleared and used for pasture or hay and corn growth. The Guernsey-Dormont-Culleoka association consists of rolling hills and ridges. Dormont and Culleoka soils are as described above. The Guernsey soils are on hillsides and hilltops and commonly have slips on them. These soils are deep and moderately well drained. These areas were formed in residuum of weathered clay shale, siltstone, and limestone. Most of this area is used for cultivated crops or hay. Steeper hillsides are used for pasture. The Dormont-Culleoka-Newark association consists of hills and flood plains. Dormont and Culleoka soils are as described above. The Newark soils are on flood plains. These soils are deep and somewhat poorly drained. These areas were formed from an alluvium derived from limestone, sandstone, siltstone, and shale. The Udorthents-Culleoka-Dormont association consists of hills and areas disturbed by strip mining activities. Dormont and Culleoka soils are as described above. The Udorthents are in the areas used for strip mining. These soils are very shallow to deep and are well drained to somewhat poorly drained. Most of the area is associated with strip-mining or is wooded or reverting to woodland (USDA 1983). Washington County is part of the Allegheny Plateau west of the Appalachian Mountains. The county's terrain consists of rolling hills. In the northeastern part of the county the hills are gently sloping whereas in the southwestern section they become almost rugged. Elevations range from a high of 1,516 feet to a low of 727 feet above sea level. The Monongahela River, one of the most traveled rivers east of the Mississippi, forms an eastern natural boundary to the county. Washington County contains five major watersheds. These major watersheds are grouped into two subbasin catagories including the Monongahela and the Ohio. The Monongahela subbasin has a total drainage area of 2737 square miles. It includes the Pennsylvania portion of the Monongahela River, from West Virginia and Maryland to Pittsburgh. The subbasin encom- 1232 Journal of the Botanical R h Institute of Texas 1(2) passes all of Fayette County, and parts of Greene, Washington, Westmoreland, and Somerset Counties. The Monongahela subbasin in Washington County is divided into the following major watersheds: Middle Monongahela River and Tenmile Creek. The Middle Monongahela River has a total drainage area of 509 square miles and its major streams include the middle portion ofthe Monongahela River, Pigeon Creek and Redstone Creek. The Tenmile Creek has a total drainage area of 388 square miles and its major streams include Tenmile Creek and South Fork Tenmile Creek. The Middle Monongahela and Tenmile Watershed show many extremes in water quality due to both man-made and natural causes. The greatest water problem is the pollution of the water resources by the drainage from coal mining operations, and the hundreds of abandoned oil and gas wells that are not properly plugged. The Ohio subbasin has a total drainage area of 3084 miles. It includes the Pennsylvania portion of the Ohio River, from its beginning at the confluence of the Allegheny and Monongahela Rivers in Pittsburgh to the borders of Ohio and West Virgina. The subbasin encompasses all of Beaver and Lawrence Counties, parts of Allegheny, Bulter, Crawford, Greene, Mercer, Venango and Washington counties. The Ohio subba- sin is divided into the following major watersheds: Chartiers Creek, Wheeling-Buffalo Creeks and Raccoon Creek. Chartiers Creek has a total drainage area of 296 square miles and its major stream is Chartiers Creek. The Wheeling-Buffalo Creek has a total drainage area of 319 square miles and its major stream include Buf- falo Creek, Enlow Fork, and Wheeling Creek. Raccoon Creek has a total drainage area of 327 square miles and the major stream is Raccoon Creek. The upper portion of the watershed has numerous surface mines. The lower and central Raccoon Creek watershed contains some of the most actively farmed land in Wash- ington and Beaver Counties. The major impairment in the Raccoon Creek watershed is abandoned mine drainage. Forty miles of the main stem of Raccoon Creek and 30 to 40 miles of tributaries are degraded by mine drainage. Baseline conditions need to be established for the flora in these watersheds. At this time we can only provide a checklist, however, future watershed conditions need to be determined to detect if the biodiversity is increasing or decreasing. These comparisons will be extremely important because they will reflect the interactions of many of the other indicators, and these comparisons are measurable effects of the manage- ment decisions (Pennsylvania DEP Watershed Notebook 2006). History The first settlers of southwestern Pennsylvania were primarily English and came from Virginia and Maryland following Braddock's Road to Redstone Fort. These settlers were enticed by the Ohio Company to settle and commence trade with the Indians in the early 1760s. After provincial Pennsylvania's land purchase from the Indians in 1769, a migration of primarily Scotch-Irish people began coming from the eastern counties of Chester, Lancaster, York, and Dauphin as well as those of Northern Maryland. By 1773 the Scotch-Irish population was increasing steadily. Between 1771 and 1775 lands west of the Monongahela River were settled. Most of these lands were obtained under Virginia titles because the price was one fourteenth the cost asked by Pennsylvania. After the Revolution most of the settlers were Scotch-Irish mingled with some English and Germans. During the first half of the 19' century scattered settlements grew into boroughs and townships. Washington County was created on March 28, 1781 from part of Westmoreland County. Both the county and the city are named for George Washington, the first President of the United States. The first white travelers found only trails or paths made mainly by animals. George Washington's early years as a surveyor enabled him to see the need for a national road through the Allegheny Mountains con- necting the eastern seaboard centers with the Ohio Valley and the western frontier. Completed in 1818 and still in use today, the National Pike (Route 40) runs through Washington County. Another major turnpike was the Pittsburgh and Steubenville turnpike, chartered in 1818. It was much used by stagecoaches and travelers from the west until 1855 but ceased entirely when the Pennsylvania Railroad was built. The Wash- Morton and Speedy, Flora of Washington County, Pennsylvania 1233 ington and Williamsport Turnpike, which runs from Washington to Monongahela City was completed in 1831 and became a very valuable road for convenience of travel. The first railroad surveys in Washington County were those of the Baltimore and Ohio Railroad. From 1850 to 1901 several major railroads were built and included the Hempfield Railroad, Pittsburgh Southern Railroad, Pittsburgh and Steubenville Railroad, Chartiers Valley Railroad, Pittsburgh, Virginia and Charleston Railroad, and the Waynesburg and Washington Railroad. Early 18% century farmers and settlers carved a living from the rich soil. The discovery of vast coal deposits, coupled with the industrial revolution, transformed economic life in the county. The manufacture of steel and glass, along with the discovery of oil, further stimulated rapid change and development throughout the region. To date Washington County is Pennsylvania's number one coal producer. Wholesale and retail establishments employ more than 16,500 workers, while the service industry provides over 16,200 jobs. The manufacture of steel and steel by-products represents the largest industrial employer with over 5,000 laborers. Forestry, fishing and other agribusiness's play a key role in the county (Forrest 1926). The county is divided into 67 municipalities with the majority of the population residing in the urban and suburban areas. The first census taken in 1790 documented that Washington County had a population of nearly 24,000. As of 2005, the population was 206,406. According to the U.S. Census Bureau, the county has a total area of 2,230 km2 (861 mi2), 2,220 km2 (857 mi2) of it is land and 10 km2 (4 mi2) is water (U.S. Census Bureau 2007). Major Collectors Collecting in Washington County for the Carnegie Museum Herbarium began in the early 1800s and has continued to the present. Most of the collectors have been members of the Botanical Society of Western Pennsylvania and the staff of the Section of Botany at Carnegie Museum. Botanists who have made significant contributions to the knowledge of the Washington County are W.E. Buker, L.K. Henry, O.E. Jennings, L. Speedy, and S.A. Thompson and J.H. Nishida. METHODS This checklist was compiled by searching the herbaria of the Carnegie Museum of Natural History (CM) and Washington and Jefferson College, Biology Department for Washington County specimens. Herbarium collection information was obtained from the Morris Arboretum (MOAR) and the Washington and Jefferson College for 16 samples. Other herbaria that were examined for specimens include the Academy of Natural Sciences of Philadelphia. In addition, fieldwork was conducted targeting underrepresented areas of the county. Most of the collections were made during the last two years, but several specimens date back as far as 1869 and 1885. The majority of the specimens are deposited at the Carnegie Museum of Natural History (CM). Rhoads and Block (2000) was the primary source for plant identification. For generic and species names, we have generally followed the Synthesis of North American Flora (Kartesz 1999). Authorities are abbreviated for the majority of taxa according to Brummitt and Powell (1992). RESULTS AND DISCUSSION This list includes the names of all native and naturalized species known to occur in Washington County. It includes a total of 125 families, 472 genera, and 955 species. The five families with the largest number of species are Asteraceae, Poaceae, Cyperaceae, Rosaceae, and Fabaceae and Carex, Symphyotrichum, Solidago, Polygonum, and Rubus are the largest genera. This checklist recognizes 3 species of Lycophytes, 31 species of Pteridophytes, 10 species of Gymnosperms, and 911 species of Angiosperms. There are approximately 203 non-native species that have been introduced mainly from Europe and Eurasia. Forty-one species have global or state ranking. Of the 41 plants that have global or state ranking only Delphinium exaltatum has a global ranking of G3 or vulnerable status. The remaining 40 plants are either a G4 or G5 status, indicating an apparently secure 1234 Journal of the Botanical R h Institute of Texas 1(2) or secure condition globally. Delphinium exaltatum contain alternate leaves palmately divided into 3-5 parted segments, 15 cm long, silvery-green abaxially and pubescent above and below. The basal lobes of some of the larger leaves are divided again and make the blade appear 5-lobed. The flowers are in terminal racemes. The 4 purplish-blue petals are dimorphic with the upper two petals expanded at the base and forming a spur. The lateral petals are reflexed in the apical half with long white hairs. The stamens are approximately 30 in number with yellow pollen. The 5 sepals are irregular. The species can be confused with Delphinium tricorne however D. tricorne is a smaller plant, which flowers much earlier than D. exaltatum, May to early June. By July, when D. exaltatum is beginning to flower, D. tricorne already has set fruit. Most of the collec- tions rich shaded woods and on rocky limestone bluffs. This species is quite conspicuous and may be subject to casual picking or may be dug for gardens. There are six taxa in the Washington County flora that are listed by the Pennsylvania Department of Agriculture (2007) as noxious weeds. It is therefore illegal to propagate, sell or transport the following taxa in the commonwealth: Cirsium arvense (Canadian thistle), Cirsium vulgare (bull or spear thistle), Datura stramonium (jimsonweed), Lythrum salicaria (purple loosestrife), Polygonum perfoliatum (mile-a-minute vine), and Rosa multiflora (multiflora rosa). Other species considered serious invasives' in our native ecosystems are: Acer platanoides (Norway maple), Alliaria petiolata (garlic mustard), Celastrus orbiculatus (oriental bit- tersweet), Elaeagnus umbellate (autumn olive), Lonicera morrowii (morrow's honeysuckle), Lonicera tartarica (tartarian honeysuckle), Microstegium vimineum (Japanese stilt grass), and Polygonum cuspidatum (Japanese knotweed) (DCNR 2004) While this checklist is probably not all-inclusive of every species in Washington County, it is the most comprehensive list presently available. ANNOTATED CHECKLIST OF THE SPECIES OF WASHINGTON COUNTY, PENSYLVANIA Taxa are listed according to the following format: taxon name, author(s), ((year) collector and number) global: state ranking [Synonyms] and non-native source. We follow the state (S) and global (G) ranking systems developed by The Nature Conservancy (1996 version). The global numbers are designated from 1 (critically imperiled) to 5 (secure). Other notations include SH, which denotes historical occurrence, and SR, which indicates reported without persuasive documentation. Synonyms are included for names not in common usage in the state or regional manuals. In cases where there was more than one specimen present in the collection, recent collections of current collectors for the western Pennsylvania region were cited. Families, genera, and specific and infraspecific taxa are arranged alphabetically within vascular plant groups Angiosperms, Gymnosperms, Lycophytes, and Pteridophytes. ANGIOSPERMS Alismatac Acanthaceae Alisma cordon Raf., 2005 L. Speedy and M. Bowers Justicia americana (L.) Vahl, 2006 L. Speedy LS-06-505 LSB1031 E Ruellia strepens L, 1950 L.K. Henry and WE. Buker s.n. G4G5: Sagittaria latifolia Willd., 1954 L.K. Henry and W.E. Buker s.n. S2 Amaranthaceae Amaranthus albus L., 2006 L. Speedy LS-06-927 Amaranthus retroflexus L., 1984 S.A. Thompson and J.H. Nishida 2097 Tropical America Aceraceae Acer negundo L. var. negundo, 2005 L. Speedy LSB12 Acer nigrum Michx. f., 2005 L. Speedy LSB595 e L., 1985 S.A. Thompson and J.H. Nishidaand Anacardiaceae M. Mac nald 2250 Europe Rhus aromatica Ait. var. aromatica, 2001 J. Polonoli FG73 rubrum e var. rubrum, 2006 L. Speedy LS-06-17 Rhus copallinum L. var. latifolia Engl., 1970 W.E. Buker s.n. ioe saccharinum L., 2005 L. Speedy LSB229-2 Rhus glabra L., 1951 L.K. Henry and F.H. Beer s.n Acer saccharum Marsh. var. saccharum, 2006 L. Speedy LS- Rhus typhina L., 2005 L. Speedy and M. Bowers LSB1350-2 Toxicodendron radicans (L.) Kuntze ssp. radicans, 2005 L. Acoraceae Speedy LSB1661 Acorus calamus L., 2005 L. Speedy LSB1665 Annonaceae Asimina triloba (L) Dunal, 1950 L.K. Henry and F.H. Beer s.n. Morton and Speedy, Flora of Washington County, Pennsylvania Apiaceae Bupleurum rotundifolium L., 1869 S.W. Knipe s.n. Europe and Central Asia ur procumbens (L.) Crantz var. procumbens, 2005 L. Speedy and D. White LSB157 Cicuta mca L., 2005 L. Speedy LSB821 Con n L., 2005 L. Speedy LSB802 Europe Cryptotaenia canadensis (L.) DC., 2005 L. ~ LSB1004 Daucus carota L., 1942 D. Wilson 103 Eur Erigenia bulbosa (Michx.) Nutt., 2006 E T LS-06-21 G5:S2 Heracleum maximum Bartr., 2005 L. Speedy LSB1160-2 Osmorhiza claytonii (Michx.) C.B. Clarke, 2005 L. Speedy LSB584 Osmorhiza longistylis (Torr.) DC., 2005 L. Speedy LSB356 Pastinaca sativa L., 2006 L. Speedy LS-06-380 Eurasia Sanicula canadensis L. var. canadensis, 2005 L. Speedy LSB634 Sanicula marilandica L., 1946 E. Mason s.n. Sanicula odorata (Raf.) K.M. Pryer & L.R. Phillippe, 2005 L. 89 Sanicula trifoliata Bickn., 2005 L. Speedy LSB1145 Taenidia integerrima (L.) Drude, 2005 L. Speedy LSB619 Thaspium barbinode (Michx.) Nutt., 2005 L. Speedy LSB588 Thaspium trifoliatum (L.) Gray var. aureum Britt, 2005 L. Speedy LSB599 Torilis japonica (Houtt.) DC., 2005 L. Speedy LSB1007 Europe Zizia aurea (L.) W.D.J. Koch, 2005 L. Speedy LSB532 Apocynacea ene ean ondescemiolum L., 1929 W.C. Grimm s.n. 005 L. Speedy LSB823 Vinee minor L., 2005 L. a LSB40 Europe Aquifoliaceae llex verticillata (L.) Gray, 1887 C.C. Mellor s.n. Paene na dracontium (L.) Schott, 2005 L. Speedy LSB706 proe triphyllum (L) Schott ssp. pusillum (Peck) Huttleston, 94 Arisaema triphyllum (L.) Schott ssp. stewardsonii (Britt) Hut- tleston, 2005 L. Speedy LSB963 Arisaema triphyllum (L.) Schott ssp. triphyllum, 2005 L. Speedy LSB243 Symplocarpus foetidus (L.) Salisb. ex Nutt., 2005 L. Speedy Araliaceae Aralia nudicaulis L., 2005 L. Speedy and M. Bowers LSB884 Aralia racemosa L. ssp. racemosa, 2005 L. Speedy and S. Speedy and M. Bowers LSB1525-1 Panax quinquefolium L., 1999 J. Polonoli FG24 Aristolochiaceae Aristolochia macrophylla Lam., 1888 A.M. Fulton s.n. Aristolochia serpentaria L., 1947 E. Mason s.n. Asarum canadense L., 2005 L. Speedy LSB596 Asclepiadaceae Asclepias incarnata L. ssp. incarnata, 1999 B.L. Isaac and J.A. Isaac 12075 1235 Asclepias quadrifolia Jacq., 2005 L. Speedy LSB643 Asclepias syriaca L., 1977 ker s.n. Asclepias tuberosa L. ssp. tuberosa, 1967 W.E. Buker s.n. steraceae Achillea millefolium L. var. millefolium, 2005 L. Speedy LSB834 Eurasia Ageratina altissima (L.) King & H.E. Robins. var. altissima, 2005 L. Speedy LSB1181 Ambrosia artemisiifolia L. var. artemisiifolia, 2005 L. Speedy Ambrosia artemisiifolia L. var. elatior (L) Descourtils, 1951 L.K. H nd F.H. Beer s.n. Ambrosia trifida L. var. trifida, 1950 L.K. Henry and W.E. Buker S.N. Antennaria howellii Greene ssp. neodioica (Greene) Bayer, 1996 cand J.A. Isaac 8954 Antennaria howellii Greene ssp. petaloidea (Fern.) Bayer, 1940 Antennaria parlinii Fern. ssp. fallax ( ) Bayer & Stebbins, 952 D. Wilson 32 ea parlinii Fern. ssp. parlinii, 2005 L. Speedy LSB12 Anthemis cotula L., 1889 C.C. Mellor s.n. Europe Arctium minus Bernh., 1995 B.L. Isaac and J.A. Isaac 8340 Eurasia ii ud atriplicifolium (L) H.E. Robins., 1966 W.E. Buker =>: m m muehlenbergii (Schultz-Bip.) H.E. Robins., 1919 O.E. Jennings and Artemisia ludoviciana Nutt. ssp. ludoviciana, 1949 W.E. Buker Artemisia vulgaris L. var. vulgaris, 1940 FH. Bell 389 Eurasia Bidens aristosa (Michx.) Britt., 1965 N.R. Farnsworth WP-189 Bidens bipinnata L., 1950 L.K. Henry de ze E. Buker Bidens cernua L., 2005 L. Speedy LSB Bidens connata Muhl. ex Willd., 1941 = | 678 Bidens frondosa L., 2005 L. Speedy LSB1498 Bidens tripartita L., 2005 L. Speedy LSB1497 Bidens vulgata Greene, 1995 B.L. Isaac and J.A. Isaac 8301 Brickellia eupatorioides (L.) Shinners var. eupatorioides, 1916 ings s.n. Centaurea biebersteinii DC., 2006 L. Speedy and J. Nusser L 4 e Cichorium intybus L., 1950 L.K. Henry and W.E. Buker s.n. Europe Cirsium altissimum (L.) Spreng., 1970 W.E. Buker s.n Cirsium arvense (L.) Scop., 2005 L. Speedy LSB801 Eurasia Cirsium discolor (Muhl. ex Willd.) Spreng., 1940 L.K. Henry S.n. Cirsium muticum Michx., 2005 L. Speedy and M. Bowers LSB1429 Cirsium pumilum (Nutt) Soreng., 1971 W.E. Buker s.n. Cirsium vulgare (Savi) Ten., 1951 L.K. Henry and F.H. Beer s.n. Eurasia Conoclinium coelestinum (L.) DC., 2005 L. Speedy and M. Bow- M. Lange L$81288 G5:S3 Conyza canadensis (L.) Cronq. var. canadensis, 2005 L. Speedy M. Bowers and M. Lange LSB1294 1236 Coreopsis tinctoria Nutt. var. tinctoria, 1901 J.A. Shafer s.n. Coreopsis tripteris L., 1955 W.E. Buker Eclipta prostrata (L) L., 2005 L. us a M. Bowers and M. Erechtites hieraciifolia (L) Raf. ex DC. var. hieraciifolia, 2005 L. Speedy e Frigeron a (L.) Pers., 2005 L. Speedy LSB997 Eriaeron E L. var. philadelphicus, 2005 L. Speedy LSB27 Erigeron dm Michx. var. pulchellus, 1985 S.A. Thompson nd J.H. Nishida 24 Erigeron strigosus Muhl.ex Willd. var. strigosus, Buker s.n. sued rium altissimum L., an L.K Henry and FH. Beer s.n. ) Barratt, 2005 L. Speedy LSB1632 EO pur E var. perfoliatum, 2005 L. Speedy LSB1258 1970 W.E. Eupatorium purpureum L. var. purpureum, 2005 L. Speedy LSB1150 Eupatorium serotinum Michx., 2005 L. Speedy and M. Bowers an ange 3 Eupatorium sessilifolium L. var. brittonianum Porter, 2002 S.P. Grund 2968 Eupatorium sessilifolium L. var. sessilifolium, 2005 L. Speedy 1228 Eurybia divaricata (L.) Nesom, 2005 L. Speedy LSB1438 [Aster divaricata L] o o (L) Cass., 1940 L.K. Henry s.n. [Aster m L] ms c (Nees) Nees, 2005 L. Speedy LSB1010 [Aster schreber, Euthamia o (L) Greene var. graminifolia, 1940 L.K. Henry s.n. Galinsoga quadriradiata Cav., 2005 L. Speedy LSB1165 Central erica Gnaphalium uliginosum L., 2005 L. Speedy and M. Bowers LSB1035 Europe Hasteola suaveolens (L) Pojark., 2005 L. Speedy and M. Bow- ers LSB1393 Helenium autumnale L. var. autumnale, 2005 L. Speedy and M. Bowers LSB1431 Hele fle n Raf., 1966 W.E. Buker s.n. anta annuus E 1965 N.R. Farnsworth WP-1 Helianthus decapetalus L., 2005 L. Speedy and M Bowers LSB1425 Helianthus divaricatus L., 1977 W.E. Buker s.n. Helianthus microcephalus Torr. & Gray, 2005 L. Speedy LSB1238 G5:S3 Helianthus strumosus L., 1950 L.K. Henry and W.E. Buker s.n. Helianthus tuberosus L., 2005 L. Speedy LSB146 Heliopsis helianthoides (L) Sweet var. helianthoides, 2005 L. Speedy LSB1456 Hieracium aurantiacum L. 1972 L.K. Henry and W.E. Buker rope Hieracium caespitosum Dumort., 2006 L. Speedy LS-06-288 Europe Hieracium gronovii L., 1975 W.E. Buker s.n. Hieracium paniculatum L., 2005 L. Speedy LSB1176 Ll Dat PRA D hi Air Journal UI f Texas 1(2) Hieracium piloselloides Vill., 1947 W.E. Buker s.n. Europe Hieracium scabrum Michx. var. scabrum, 2005 L. Speedy LSB1478 Hieracium venosum jn 2005 L. Speedy LSB637 pocha adicata L., 2006 L. Speedy LS-06-907 Eurasia Inula helenium L., 1946 E. Mason s.n. Europe Ms) oe ld Fern., 2006 L. Speedy and M. Bow- ms iM L., 1970 W.E. Buker Lactuca floridana (L) Gaertn. var. mum 1970 W.E. Buker s.n. Lactuca saligna L., 1946 W.E. Buker s.n. dE Lactuca sativa L., 1940 L.K. Henry s.n. Lactuca serriola L., 2006 L. Speedy e pm LS-06-814 Eur m ee vulgare Lam., 2005 L. Speedy LSB923 Europe Liatris spicata (L.) Willd. var. spicata, 1969 W.E. Buker and H. Clark s.n Matricaria recutita L., 1947 L.K. Henry s.n. Europe Packera aurea (L.) A. & D. Love, 1985 S.A. Thompson and JH. Nishida 2312 [Senecio aureus L.] Packera obovata (Muhl. Ex Willd.) W.A Weber & A. Love, 2005 L. Speedy LSB80 [Senecio obovatus Muhl.Ex Willd.] Packera paupercula (Michx.) A. & D. Love, 1950 L.K. Henry and eer s.n. [Senecio paupercula Michx.] emma canadensis L., 2005 L a LSB1144 Prenanthes alba L., 1921 SS. bic Prenanthes altissima L., 2005 L. MUN LSB1635 Prenanthes crepidinea Michx., 2006 L. Speedy LS-06-128 cda m (Cass.) Fern., 1969 W.E. Buker s.n. Pseudognaphalium obtusifolium (L.) Hilliard & Burtt ssp. obtu- sifolium, 2005 L. Speedy LSB1477 Ratibida pinnata (Vent) Barnh., 2006 L. Speedy LS-06-643-2 5:SNA Rudbeckia hirta L. var. hirta, 1924 H. Torp s.n. Rudbeckia hirta L. var. pulcherrima Farw., 1967 W.E. Buker s.n. Rudbeckia laciniata L. var. laciniata, 2005 L. Speedy LSB1278 Rudbeckia triloba L. var. triloba, 1977 W.E. Buker s.n. Senecio vulgaris L., 1987 AW. Cusick 27116 Eurasia Sericocarpus asteroides (L.) B.S.P, 1955 W.E. Buker a trifoliatum L. var. trifoliatum, 2006 L. T LS- 6-742 W uvedalius (L.) Mackenzie ex Small, 2004 R. Coxe Solidago altissima L., 2005 L. Speedy LSB1447 Solidago arguta Ait. var. arguta, 1944 O.E. Jennings s.n. Solidago bicolor L., 2005 L. Speedy LSB1636 Solidago caesia L., 2005 L. Speedy LSB1437 Solidago canadensis L. var. canadensis, 2005 L. Speedy SB1473 d M as & Gray, 1984 S.A. Thompson and J.H. Nishida 5:51 Solidago EN L., 2005 L. Speedy LSB1451 e ii Ait., 2005 L. Speedy and M. Bowers sold re Ait., 2006 L. Speedy LS-06-496 idago nemoralis Ait. var. nemoralis, 2005 L. Speedy LSB1643 Morton and Speedy, Flora of Washington County, Pennsylvania Solidago patula Muhl. ex Willd. var. patula, 2005 L. Speedy 1283- Solidago rugosa P. Mill. ssp. aspera (Ait.) Cronq., 1968 W.E. Buker s.n. Solidago rugosa P. Mill. ssp. rugosa, 2005 L. Speedy and M. Bowers LSB1399 Solidago ulmifolia Muhl. ex Willd. var. ulmifolia, 2005 L. Speedy LSB1225 Sonchus arvensis L. ssp. uliginosus (Bieb.) Nyman, 1952 LK. rope Sonchus oleraceus L., 1951 L.K. Henry and F.H. Beer s.n. Symphyotrichum cordifolium (L.) Nesom, 2005 L. Speedy S 0 [Aster cordifolium L. Symphyotrichum drummondii (Lindl.) Nesom var.drummondii, n is Roe S d ont [Aster drummondii i Lindl.] illd lanceolatum var. interior (Wieg.) Nesom, 2005 L. Speedy pe 1520 [Aster e na. Nel ee) (Wieg.) AG. J LAL Y NI SN a NM var. lanceolatum, 1951 L.K. Henry and F.H. Beer s.n. [Aster lanceolatum Willd.) o lateriflorum (L.) A. & D. Love var. lateriflorum, 2005 L. Speedy LSB1448 [Aster lateriflorum (L.) Britton var. l Le Porter) Nesom, 2005 L. Speedy LS81 1630 [Aster lowrieanum Por Symphyotrichum novae-angliae (L 3 ON 2006 L. Speedy LS-06-909 [Aster novae-angliae L.] Symphyotrichum phlogifolium Muhi ex js Nesom, 1949 ker s.n. [Aster phlogifolium Muhl. ex Willd.] Symphyotrichum pilosum (Willd.) Nesom var. eae 2006 L. Speedy LS-06-917 [Aster pilosus Willd.] Edd pilosum (Willd.) Nesom var. pringlei ee esom, 1949 L.K. Henry s.n. [Aster pilosum Willd. va B cn Blake] Symphyotrichum praealtum (Poir. Nesom var. angustior (Wieg.) Nesom, 1955 W.E. Buker s.n. G5:S3 [Aster praealtum d var. rod on Auhl. ex Willd.) Nesom, 2005 L Speedy LSB1450 [Aster prenanthoides Muhl. ex Willd. Symphyotrichum puniceum (L) A. & D. Love var. puniceum, 2005 L. Speedy LSB1677 [Aster puniceus L.] Symphyotrichum shortii (Lindl.) Nesom, 2005 L. Speedy LSB1519 [Aster shortii Lindl.] diri bi undulatum (L) Nesom, 1968 W.E. Buker s.n. [Aster un ls] DUE urophyllum (Lindl.) Nesom, 1955 E. Mason s.n. [Aster urophyllum Lindl.) Tanacetum vulgare L., 1940 L.K. Henry s.n. Europe Taraxacum laevigatum (Willd.) DC., 2001 A.W. Cusick 35785 ia Europe ee bus G.H. Weber ex Wiggers, 2005 L. Speedy us pons a 2005 L. Speedy LSB236 Eurasia Verbesina alternifolia (L.) Britt. ex Kearney, 2005 L. Speedy Vernonia gigantea (Walt.) Trel., 2005 L. Speedy LSB1470 1237 Xanthium strumarium L. var. canadense (P. Mill.) Torr. & Gray, 005 L. Speedy LSB1494 Xanthium strumarium L. var. glabratum (DC.) Cronq., 1951 L.K. Henry and F.H. Beer s.n. Balsaminaceae Impatiens capensis Meerb., 2005 L. Speedy LSB922 Impatiens pallida Nutt., 2005 L. Speedy LSB919 Berberidaceae Berberis thunbergii DC., 2005 L. Speedy LSB266 Japan Socal Age eas .) Michx., 2005 L. Pond LSB94 la (L.) Pers., 2005 L. Speed Podophyllu P mL, Bae L. Speedy ee Betulaceae Alnus serrulata (Ait) Willd., 2005 L. Speedy and M. Bowers LSB887 Betula lenta L., 2005 L. Speedy and M. Bowers LSB886 Carpinus caroliniana Walt. ssp. caroliniana, 2005 L. Speedy LSB1537 Carpinus caroliniana Walt. ssp. virginiana (Marsh.) Furlow orylus americana Walt., 2005 L. Soeedy LSB1679-1 Ostrya virginiana (P. Mill.) K. Koch var. virginiana, 2005 L. Speedy LSB259-2 Bignoniaceae Campsis radicans (L.) Seem. ex Bureau, 1970 W.E. Buker s.n. Catalpa bignonioides Walt., 1985 S.A. Thompson and J.H. 0 Catalpa speciosa (Warder) Warder ex Engelm., 2006 L. Speedy LS-06-289 Boraginaceae susie arvensis (L.) LM. Johnston, 1940 L.K. Henry s.n. mene officinale L., 1947 E. Mason s.n. Eurasia Cyn said virginianum L. var. virginianum, 2006 L. Speedy LS-06- Echium isi L., 2006 L. Soeedy LS-06-381 Europe Hackelia virginiana (L.) .M. Johnston, 2005 L. Speedy LSB995 Lithospermum latifolium Michx., 2005 L. Speedy LSB594 G4: S4 Mertensia virginica (L.) Pers. ex Link, 2005 L. Speedy LSB247 Myosotis macrosperma Engelm., 1996 B.L. Isaac and J.A. Isaac 8951 Myosotis scorpioides L., 2006 L. Speedy LS-06-237 Europe Onosmodium molle Michx. ssp. ca (Mackenzie) Boivin, 1929 J. Bright 2536 G 4:51 Symphytum officinale L., 1964 Wn T SP-220 Eur- asia Brassicace e um (Bieb) Cavara & Grande, 2005 L. Speedy hos thaliana (L. Heynh., 1913 O.E. c eios and G.K. and A.R eae s.n. Euro 50 DH. pen s.n. Arabis glabra (L.) p 1970 W.E. Buker s.n. Arabis laevigata (Muhl. ex Willd.) Poir. var. laevigata, 2005 L. Speedy LSB909 Arohi bo nci 1238 Barbarea verna (P. Mill.) Aschers., 1972 G.G. Ingotz Europe Barbarea vulgaris Ait.£, 2005 L. Speedy LSB286 Eurasia Brassica juncea (L.) Czern., 1947 W.E. Buker s.n. Eurasia pa nigra (L.) W.D.J. Koch, 1940 L.K. Henry s.n. Eurasia apsella ME (L) Medik., 1951 L.K. Henry and F.H. andate ee O.E. Shulz, 1941 L.K. Henr Cardamine bulbosa (Schreb. ex Muhl.) B.S.P, A 2 Speedy LSB279 Cardamine concatenata (Michx.) Sw., 2005 L. Speedy LSB244 Cardamine diphylla (Michx.) Wood, 2005 L. Speedy LSB269 Cardamine douglassi Britt., 2005 L. Speedy LSB271 Ca pee hirsuta L., 2006 L. Speedy LS-06-5 Europe and mm parviflora L. var. arenicola (Britt) O.E. Schulz, 1951 K d FH. Beer s.n. Kun pensylvanica Muhl.ex Willd., 2005 L. Speedy SB293 uc a Michx., 2005 L. inde LSB378 Draba verna L, J.A. Churchill 198 Euro Erysimum inicr EOS Sle vus and F.H. Beer s.n. Eurasia Hesperis matronalis L., 2005 L. Speedy LSB950 Europe lodanthus pinnatifidus (Michx.) Steud., 2006 L. Speedy LS- 06-286 G5:S1 Lepidium campestre (L.) Ait. f, 2006 L. Speedy LS-06-378 Eurasia Lepidium virginicum L. var. virginicum, 1919 O.E. Jennings s.n. Rorippa nasturtium-aquaticum (L.) Hayek, 2005 L. Speedy LSB827 Rorippa palustris (L.) Bess. ssp. fernaldiana (Butters & Abbe) Jonsell, 1942 O.E. Jennings Rorippa palustris (L.) Bess. ssp Puls 1947 W.E. Buker s.n. Sibara virginica (L.) Rollins, 2001 A.W. Cusick 35789 [Cardamine virginica L. Sisymbrium altissimum L., 1952 D. Wilson 153 Eurasia Sisymbrium officinale (L.) Scop., 1952 D. Wilson 188 Europe Buxaceae Pachysandra terminalis Sieb. & Zucc., 2002 S.P. Grund 2985 Japan Campanulaceae Campanula rapunculoides L., 1971 W.E. Buker s.n. Eurasia Campanulastrum americanum (L.) Small, 2005 L. Speedy LSB1001 Lobelia inflata L., 2005 L. Speedy LSB1490 Lobelia siphilitica L. var. siphilitica, 2005 L. Speedy LSB1444 Lobelia spicata Lam. var. spicata, 1957 W.E. Buker s.n Triodanis perfoliata (L.) Nieuwl., 2006 L. Speedy LS-06-393 Cannabaceae Humulus japonicus Sieb. & Zucc., 1996 S. Grund 1681 Japan Humulus lupulus L. var. lupuloides E. Small, 2006 L. Speedy LS-06-422 Caprifoliaceae Lonicera maackii (Rupr.) Herder, 2006 L. Speedy LS-06-250 Asia £L Dag PRA D hi PEA Journal of f Texas 1(2) Lonicera morrowii Gray, 2005 L. Speedy LSB276 Japan Lonicera tatarica L., 1972 L.K. Henry and W.E. Buker s.n. Eurasia Sambucus nigra L. ssp. canadensis (L) R. Bolli, 1952 J. Wilson 127 Symphoricarpos albus (L) Blake var. albus, 1940 L.K. Henry s.n. Symphoricarpos albus (L.) Blake var. laevigatus (Fern.) Blake, . Jennings s.n. Symphoricarpos orbiculatus Moench, 1949 W.E. Buker s.n. Bickn. var. illinoense e Palmer & Steyermark, 1996 B.L. Isaac and J.A. Isaac 8961 L., 2005 L. Speedy D Viburnum lantana L., 2006 L. Speedy LS-06-245 Europe and IFÁAIYREIACgHIF Viburnum acerifolium Asia Viburnum lentago L., 1887 C.C. Mellor s.n Viburnum opulus L. var. opulus, 2006 L. speedy LS-06-258 Eura \ mn e LE, 2005: L: Speedy LSB284-2 aryophyllaceae € Arenaria serpyllifolia L, 1978 F.H. Utech and W.E. Buker 78- Cerastium o Baumg. ssp. vulgare DD Greuter & B 04 O.E. Jennings s.n. Eura Cerastium pm Thuill., 1946 L.K. s.n. Eurasia Cerastium nutans Raf. var. nutans, 1996 B.L. Isaac and J.A. Isaac 8969 Dianthus armeria L., 2006 L. Speedy LS-06-374 Europe Myosoton aquaticum (L.) Moench, 2005 L. Speedy LSB1486 Europe Paronychia canadensis (L.) Wood, 2005 L. Speedy LSB1216 Paronychia fastigiata (Raf.) Fern. var. fastigiata, 1885 B.H. dicen officinalis L., 2005 L. Speedy and M. Bowers LSB1432 urope Er antirrhina L., 1947 L.K. Henr Silene dichotoma Ehrh., 1952 L.K. e and F.H. Beer s.n. Europe Silene nivea (Nutt.) Muhl. ex Otth, 1869 S.W. Knipe s.n. Silene noctiflora L., 1920 O.E. Jennings s.n. Europe Silene stellata (L.) Ait. f, 1994 J.A. Isaac 6051 Silene virginica L. var. virginica, 2005 L. Speedy LSB600 Silene vulgaris (Moench) Garcke, 1979 P. Singleton and D.E. ufford and EW. Wood 6 Europe Stellaria corei Shinners, 1918 O.E. Jennings and G.K. Jen- nings s.n. Stellaria longifolia Muhl. ex Willd. var. longifolia, 2005 L. Speedy and K. Jurkovic LSB429 Stellaria media (L.) Vill. ssp. media, 2005 L. Speedy LSB239 Furope Stellaria pubera Michx., 1937 C.M. Boardman s.n. elastraceae Celastrus de Thunb., 2005 L. Speedy LSB1666 China and Ja mn p L., 1975 W.E. Buker Euonymus alata (Thunb.) Sieb., 2005 L. pu LSB367 China and Japan Morton and Speedy, Flora of Washington County, Pennsylvania Euonymus atropurpurea Jaca. var. atropurpurea, 1990 B.L. Isaac and J.A. Isaac 3098 Chenopodiac Atriplex patula b a 7 O.E. Jennings et al. s a album L. var. album, on L K. Henry s.n. Eur mm album L. var. missouriense (Aellen) |.). Bassett & C.W. Crompton, 1951 L.K. Henr Chenopodium ambrosioides L. var. ambrosioides, 2005 L. y and M. Bowers and M. Lange LSB1317 Central and South America E berlandieri Mog. var. bushianum (Aellen) Cronq., 194 Chenopodium ban " 1979 D.E. Boufford and N. Foust tei pe m L, 1901 J.A. Shafer s.n. Cistaceae Lechea racemulosa Michx., 1975 W.E. Buker s.n. Clusia Hypericum eer (Grev. & Hook.) Torr. & Gray, 1927 O.E. ae gentianoides ( " : B.S.P, 1892 ET. Aschman s. L. Speedy and M. Bowers n M Lange LSB13 perforatum L., 1965 W.E. Buker s.n. Europe Hypericum prolifi um L., 2006 L. Speedy LS-06-642 Hypericum punctatum Lam., 2005 L. Speedy LSB924 \ peri UM Commelinaceae Commelina communis L. var. ludens (Miq.) C.B. Clarke, 1995 B.L. Isaac and J.A. Isaac 8339 Asia volvulac case "n (Ledeb.) G. Don, 1901 J.A. Shafer s.n. Exotic [Calystegia dahurica (Herb.) Choisy] UN sepium (L) R. Br. ssp. angulata Brummitt, 2005 L. Speedy LS B817 E sepium (L.) R. Br. ssp. erratica Brummitt, 1950 L.K. W.E. Buker s.n fi iflora (Mackenzie & coseno silvatica um ) Griseb. ssp sh) 1940 LK. Henry s.n. mos m (L) Pursh ssp. spithamaea, 2005 L. Speedy LSB635 nvolvulus arvensis L., 1901 J.A. Shafer s.n. Europe Ipomoea hederacea Jacq., 1947 W.E. Buker s.n. Ipomoea pandurata (L) G.FW. Mey., 1953 W.E. Buker s.n. Cornaceae Cornus alternifolia L. f, 2005 L. Speedy LSB639 ornus amomum P. Mill., 2005 L. Speedy LSB813 Cornus florida L., 2005 L. Speedy LSB385 Cornus obliqua Raf., 1951 L.K. Henry and F.H. Beer s.n. [Cornus amomum Mill. subsp. oblique (Raf) J.S.Wilson] Cornus racemosa Lam., 1972 L.K. Henry and W.E. Buker s.n. Nyssa sylvatica Marsh., 1906 O.E. Jennings s.n. Crassulaceae Hylotelephium telephium (L.) H. Ohba. ssp. telephium, 1951 L.K. Henry and F.H. Beer s.n. Exotic 1239 Penthorum sedoides L., 2006 L. Speedy LS-06-481 Sedum sarmentosum Bunge, 1987 S.A. Thompson 3707 China Sedum ternatum Michx., 2005 L. Speedy LSB543 Cucurbitaceae Cucurbita pepo L. var. pepo, 1995 B.L. Isaac and J.A. Isaac 8323 Echinocystis lobata (Michx.) Torr. & Gray, 2005 L. Speedy and i 385 Sicyos angulatus L., 2006 L. Speedy and D. Sofran LS-06-809 Cuscutaceae Cuscuta gronovii oe ex J.A. Shultes var. gronovii, 2005 L. Speedy LSB14 Cyperaceae Bulbostylis capillaris (L) Kunth ex C.B. Clarke ssp. capillaris, 1940 F.H. Bell 400 Carex aggregata Mackenzie, 2005 L. Speedy LSB1152 Carex albicans Willd. ex Spreng. var. albicans, 1921 O.E. Jen- nings s.n. Carex albursina Sheldon, 2002 J. Polonoli FG115A Carex amphibola Steud., 1984 J.H. Nishida and C.W. Bier 726 Carex annectens (Bickn.) Bickn., 2005 L. Speedy and M. Bow- ers LSB1059 Carex appalachica J. Webber & P.W. Ball, 2005 L. Speedy LSB938 Carex blanda Dewey, 2005 L. Speedy LSB946 Carex cephalophora Muhl.ex Willd., 1951 L.K. Henry and FH. Beer s.n. Carex communis Bailey var. communis, 1996 B.L. Isaac and J.A. Isaac 8949 Carex conjuncta Boott, 2005 L. Speedy and M. Bowers LSB864 G4G5:SNR Carex cristatella Britt., 2005 L. Speedy LSB1268 Carex digitalis Willd. var. digitalis, 2006 L. Speedy LS-06-247 Carex emoryi Dewey, 2001 M. Bowers s.n. Carex frankii Kunth, 2005 L. Speedy LSB768 Carex gracilescens Steud., 1951 L.K. Henry and F.H. Beer s.n. Carex granularis Muhl.ex Willd., 2001 M. Bowers s.n. Carex grisea Wahlenb., 1924 J. Bright s.n. Carex gynandra Schwein., 2001 M. Bowers s.n Carex hirsutella Mackenzie, 2006 L. Ta and J. Nusser -06- Carex hirtifolia Mackenzie, 2006 L. Speedy LS-06-386 Carex hystericina Muhl. ex Willd., 1965 W.E. Buker s.n. Carex interior bo 1945 FH. Beer s. Carex jamesii Schwein., 1926 J. B cde E (Kukenth) cee 2006 L. Speedy Carex laxiculmis Schwein. var. laxiculmis, 2005 L. Speedy LSB993 Carex laxiflora Lam. var. laxiflora, 1951 LK. Henry and F.H. Carex lupulina Muhl.ex Willd., 2005 L. Speedy and M. Bowers LSB1062 Carex lurida Wahlenb., 2005 L. Speedy LSB770 Carex molesta Mackenzie ex Bright, 1927 J. Bright s.n. Carex normalis Mackenzie, 2006 L. Speedy LS-06-231 1240 Carex pensylvanica Lam., 2005 L. Speedy pun Carex ME ree nie E r? LSB1 Carex pra / Speedy and (n Society PE PA LS- 06- 267 Carex projecta Mackenzie, 2006 L. Speedy LS-06-395 Carex rosea Schkuhr ex Willd., 2006 L. Speedy LS-06-480 Carex scoparia Schkuhr ex Willd. var. scoparia, 1972 L.K. Henry and W.E. Buker s.n. Carex shortiana Dewey, 2006 L. Speedy LS-06-419 G5:S3 Carex sparganioides Muhl.ex Willd., 1988 K.K. Lutz s.n. Carex squarrosa L., 2005 L. Speedy and M. Bowers LSB1060 Carex stipata Muhl. ex Willd. var. stipata, 2006 L. Speedy 06-486 Pd stricta Lam., 2006 L. o xi bd 399 arex torta Boott ex Tuckerman L. Speedy LSB189 mis Visio var bin 2005 L. Speedy and M. rs LS Carex virescens Muhl.ex Willd., 1923 O.E. Jennings s arex vulpinoidea Michx. var. vulpinoidea, 2006 s Speedy LS-06-398 Carex willdenowii Schkuhr ex Willd., 2002 J. Polonoli FG116A Eua bipartitus Torr., 1993 J.A. Isaac 5403 Cyperus esculentus L., 2005 L. Speedy and M. Bowers Cyperus flavescens L., 1904 O.E. Jennings s.n. Cyperus strigosus L., 2005 L. Speedy LSB1284 Eleocharis acicularis (L.) Roemer €: J.A. Schultes, 1947 W.E. Eleocharis obtusa (Willd.) J.A. Schultes, 2005 L. Speedy LSB1260 Eleocharis palustris (L) Roemer & J.A. Schultes, 1951 L.K. Henry an Eleocharis quadrangulata (Michx.) Roemer & J.A. Schultes, 1951 LK. Henry and FH. Beer s.n. G4:S1 Kyllinga pumila Michx., 1906 R.E. Schuh s.n. Schoenoplectus tabernaemontani (K.C. Gmel.) Palla, 2005 L. peedy LSB8 Scirpus atrovirens Willd., 2005 L. Speedy LSB1277 Scirpus cyperinus (L) Kunth, 2005 L. Speedy and M. Bowers Makino, 2005 L. Speedy LSB805 A ee Fern., 1993 J.A. Isaac 5401 G4:S1 Scirpus polyphyllus Vahl, 2005 L. Speedy LSB1267 Dioscoreaceae Dioscorea villosa L., 2006 L. Speedy LS-06-243 Dipsacaceae Dipsacus fullonum L., 2006 L. Speedy LS-06-649 Exotic Dipsacus laciniatus L., 1989 A.W. Cusick 28431 Ebenaceae Diospyros virginiana L., 1966 W.E. Buker s.n. Elaeegna Elaeagnus eee Thunb. var. parviflora (Royle) Schneid., 2005 L. Speedy LSB1459 Asia ricaceae Epigaea repens L., 2006 L. Speedy and C. Patsche LS-06-74 £L Dag eal D hi PEA Journal UI f Texas 1(2) Gaultheria procumbens L., 2006 L. Speedy and C. Patsche -06- Gaylussacia baccata (Wangenh.) K. Koch, 1908 O.E. Jennings S.N. Kalmia latifolia L., 1970 W.E. Buker s.n. Vaccinium corymbosum L., 2005 L. Speedy LSB299 a Md d 1944 Q.. Jennings s.n. 2005 L. Speedy LSB948 Vaccinium stamineum i 2006 L. Speedy and J. Nusser LS- 06-653 inium nallidum Euphorbia Acalypha D T Raf., 2005 L. Speedy LSB1491 Cham Mes oo eg 2005 L. Soeedy and M. Bow- and M. Lange L EAM nutans pes ; E 2006 L. Speedy and D. Sofran Euphorbia a Engelm., 1946 W.E. Buker s.n. Ed corolata L, 2005 L. Speedy LSB1241 ta Michx. var. dentata, 1970 W.E. Buker s.n. Fabaceae Amphicarpaea bracteata (L.) Fern. var. bracteata, 2005 L. eedy LSB125 Apios americana Medik., 1999 B.L. Isaac and J.A. Isaac 12074 Astragalus canadensis L. var. canadensis, 1948 O.E. Jennings Baptisia australis (L.) R. Br. ex Ait. f. var. australis, 1980 D.E. Boufford G5:S3 Cercis canadensis L. var. canadensis, 2005 L. Speedy LSB615 Coronilla varia L., 2005 L. Soeedy LSB828 Europe Cytisus scoparius (L.) Link var. scoparius, 1888 A.M. Fulton s.n. Europe Desmodium canadense (L.) DC., 1969 W.E. Buker Desmodium canescens (L.) DC., 2006 L. Speedy dar D Sofran Desmodium glutinosum (Muhl ex Willd.) Wood, 2005 L. Spe- edy LSB937 Desmodium nudiflorum (L) DC., 1995 B.L. Isaac and J.A. Isaac 8341 Desmodium paniculatum (L.) DC. var. paniculatum, 1955 W.E. Buker s.n Desmodium perplexum Schub., 1965 W.E. Buker s.n. 1942 OE. Jennings s.n. Gymnocladus dioicus (L.) K. Koch, 1941 W. Harris s.n. Kummerowia stipulacea (Maxim.) Makino, 1951 L.K. Henry an . Asia Lathyrus yest L 2006 L. Speedy and J. Nusser LS-06-609 Afric Lp. : ji | Gleadltsia THACcantnos L., lespedeza de Du Don, 2006 L. Speedy and LS-06-54 Le To rice " pm 2005 L. Speedy LSB1226 Lespedeza hirta (L.) Hornem. ssp. hirta, 1944 O.E. Jennings s.n. Lespedeza PEE EA Michx., 2006 L. Speedy and M. Bow- ers LS-06-891 Lespedeza violacea (L.) Pers., 1939 W.G. Burig s.n. Lotus corniculatus L., 1970 W.E. Buker s.n. Europe Medicago lupulina L., 2005 L. Speedy LSB826 Eurasia Medicago sativa L. ssp. sativa, 1886 J.A. Shafer s.n. Eurasia Morton and Speedy, Flora of Washington County, Pennsylvania n officinalis (L.) Lam., 2006 L. Speedy LS-06-284 ram hi L. var. hispida, 1979 EW. Wood and D.E. Bouf- ford 4 d noU T L., 2006 L. Speedy LS-06-248 marilandica (L.) Link, 1917 B.H. Patterson s.n. G5:S1 E helvula (L.) Ell., 2006 L. Speedy LS-06-883 Thermopsis villosa (Walt) Fern. & Schub., 1962 W.E. Buker s.n. Trifolium arvense L., 1904 Houston, M.E. s.n. Eurasia Trifolium aureum Pollich, 1957 W.E. Buker s.n. Eurasia Trifolium campestre Schreb., 1920 O.F. Jennings s.n. ias Trifolium hybridum L., 1904 O.E. Jennings s.n. Eur Trifolium pratense L., 2005 L. Speedy LSB602 rn Trifolium repens L., 1972 L.K. Henry and W.E. Buker s.n. Europe Vicia caroliniana Walt., 1950 L.K. Henry and F.H. Beer s.n. Vicia sativa L. ssp. nigra (L.) Ehrh., 2005 L. Speedy LSB298 Furope Fagaceae Castanea dentata (Marsh.) Borkh., 1904 O.E. Jennings s.n. Fagus grandifolia Ehrh., 2005 L. Speedy LSB1502 Quercus alba L., 2005 L. Speedy nios 2 Quercus bicolor Willd., 1940 L.K. Henr Quercus coccinea Muenchh. var. coccinea, 1922 O.E. Jen- nings s.n Quercus imbricaria Michx., 2005 L. Speedy LSB1481 Quercus macrocarpa d var. macrocarpa, 2006 L. Speedy and M. Bowers LS-06- Quercus muehlenbergii p 2006 L. Speedy LS-06-746 Quercus palustris Muenchh., 2005 L. ee LSB296-2 Quercus robur L., 1956 R. Cottrell s.n. Eur Quercus rubra L. var. rubra, 2005 L. ey ee 1 Quercus velutina Lam., 2005 L. Speedy LSB1640 Quercus x leana Nutt. (pro sp.), 1944 O.E. Jennings s.n. Quercus x runcinata (A. DC.) Engelm., 1964 J.F. Lewis s.n. Cont ps uA T 2005 L. Speedy LSB249 len die) Walp., 2005 L. Speedy LSB86 Dicena al (L ) a 2005 L. Speedy LSB89 E Gentianaceae Gentiana Dum Griseb. var. d and J. Bright and A.E. McKee "i angularis (L.) Pursh, oo Buker s.n. 917 O.E. Jennings Geraniaceae Erodium cicutarium (L.) UHer. ex Ait. ssp. cicutarium, 2001 A.W. Cusick 35784 Europe Geranium maculatum L., 2005 L. un LSB362 Geranium pratense ES 1962 D. Gondella s. Geranium thunbergii Sieb. & Zucc. ex D 8 Paxton, 2005 L. Speedy LSB1280 Grossulariaceae Ribes cynosbati L., 2005 L. Speedy LSB110-2 Ribes rotundifolium Michx., 1948 E. Mason s.n. Haloragaceae Myriophyllum spicatum L., 2006 L. Speedy LS-06-503 Eurasia 1241 Hamamelidaceae Hamamelis virginiana L., 2005 L. Speedy LSB446 Hippocastanacea (o ia Ait., M L. Speedy LSB274-2 Willd., 2005 L. Speedy LSB131-2 VVA., « Hydrangeaceae Hydrangea arborescens L., 2005 L. Speedy LSB857 Philadelphus pubescens Loisel. var. pubescens, 2006 L. Speedy LS-06-383 aa aes is Michx., 2006 L. Soeedy LS-06-504 Hydrophyllaceae Hydrophyllum appendiculatum Michx., 2005 L. Speedy LSB352 Hydrophyllum canadense L., 2005 L. Speedy LSB1137 Hydrophyllum virginianum L., 2005 L. Speedy and K. Jurkovic LS B433 Phacelia purshii Buckl., 2006 L. Speedy et al., LS-06-228 lridaceae Cieurinrhin M nnnii i tifolium P. Mill., 2005 L. Speedy LSB632 Juglandaceae Carya alba (L.) Nutt. ex Ell., 2005 L. Speedy LSB111 Carya cordiformis (Wangenh.) K. Koch, 2005 L. Speedy LSB1207-1 Carya glabra (P. Mill) Sweet, 2005 L. Speedy LSB1514 Carya laciniosa (Michx. f) G. Don, 1970 A. Liberatore Carya ovalis (Wangenh.) Sarg., 1917 OE. Jennings and J. Bright AE McKee s.n Carya ovata (P. Mill) K. Koch, 2005 L. Speedy LSB1651-1 Juglans cinerea L., 2005 L. Speedy and B. Brown LSB655 Juglans nigra L., 2006 L. Speedy LS-06-262 Juncaceae Juncus articulatus L., 2006 L. Speedy LS-06-497 Juncus effusus L. var. pylaei (Laharpe) Fern. & Wieg., 2006 L. Speedy LS-06-290 Juncus effusus L. var. solutus Fern. & Wieg., 2005 L. Speedy LSB1276 Juncus tenuis Willd., 2005 L. Speedy and M. Bowers LSB1071 Juncus torreyi Coville, 2005 L. Speedy and M. Bowers LSB1066 G5:S2 Luzula multiflora (Ehrh.) Lej. ssp. multiflora var. multiflora, 2005 L. Speedy LSB542 Lam nean ciliata (L.) Benth., 1970 W.E. Buker Blephilia hirsuta (Pursh) Benth. var. RE pu L. Speedy and M. Bowers LSB858 db si dll I S E Speedy LS-06-644 005 L. Soeedy LSB1505-1 chon c E a L. Speedy LSB281 Eurasia pulegioides (L.) Pers., 1957 L.K. Henry and F.H. Beer nin canon, mw brachiatus (L.) B.S.P, 1904 O.E. Jennings s.n. Lamium amplexicaule L., 1940 L.K. Henry s.n. Eurasia Lamium maculatum L., 1940 RV. Clark s.n. Eurasia 1242 Lamium purpureum L. var. purpureum, 2005 L. Speedy LSB187 Eurasia Leonurus cardiaca L. ssp. cardiaca, 1977 W.E. Buker s.n. Asia Lycopus americanus Muhl. ex W. Bart., 2005 L. Speedy LSB1273 Lycopus uniflorus Michx. var. uniflorus, 2005 L. Speedy LSB1521 Lycopus virginicus L., 2005 L. Speedy LSB1201 Marrubium vulgare L., 1901 J.A Shafer s.n. Eurasia Meehania cordata (Nutt) Britt., 2005 L. Speedy and M. Bowers Melissa officinalis L., 1938 H. Scarton s.n. Asia Mentha arvensis L., 2005 L. Speedy and M. Bowers and M. Lange 12 Mentha doen L H sp.), 2005 L. Speedy and M. Bowers LSB1371 Eur Monarda >o L., 2005 L. Speedy and M. Bowers Monarda didyma L., 2005 L. Speedy and M. Bowers LSB881 Monarda media Willd., 2005 L. Speedy and M. Bowers LSB899 Nepeta cataria L., 1970 L.K. Henry and W.E. Buker s.n. Europe Prunella vulgaris L. ssp. lanceolata (W. Bart.) Hulten, 2005 L. Pycnanthemum incanum (L) Michx. var. incanum, 1952 D. Wilson 156 Pycnanthemum tenuifolium Schrad., 1919 O.E. Jennings and AE. T Scutellaria incana Biehler var. incana, 2005 L. Speedy LSB1148 Scutellaria lateriflora L. var. lateriflora, 2005 L. Speedy and M. B 07 pangs nervosa Pursh, 1951 L.K. Henry and FH. Beer s.n. chys germanica L., 1943 E. Mason s.n. Europe e nuttallii Shuttlw. ex Benth., 2004 J.A. Isaac 18573 1 Stachys tenuifolia Willd., 2005 L. Speedy and M. Bowers and M. Lange LSB1303 leucrium canadense L. var. canadense, 2006 L. Speedy and J. Nusser LS-06-611 Lauraceae E benzoin (L.) Blume var. mr (Palmer & Steyer- ark) Rehd., 2005 L. Speedy L a albidum (Nutt.) Nees, ee i Speedy LSB11 Lemnaceae Lemna turionifera Landolt, 1951 L.K. Henry and FH. Beer s.n. G5:SU Liliaceae Allium canadense L. var. canadense, 2005 L. Speedy and B. Brown LSB669 Allium cernuum Roth var. cernuum, 2005 L. Speedy LSB943 Allium tricoccum Ait., 1982 WW. Thomas and S.A. Thompson and M. Ohara 2973 Allium vineale L. ssp. vineale, 1972 L.K. Henry and W.E. Buker s.n. Europe Asparagus officinalis L., 2006 L. Speedy LS-06-249 Europe Camassia scilloides (Raf) Cory, 2006 L. Speedy LS-06-241 £L Dag PRA D hi PEA Journal Ol f Texas 1(2) Chionodoxa luciliae Boiss., 1985 S.A. Thompson and J.H. Nishida and M. Macdonald 2249 Europe Clintonia umbellulata (Michx.) Morong, 1940 L.K. Henry s.n. a rid L., 1940 RV. Clark s.n. Eur m Nutt., 2006 L. Speedy i i 85 G5:S3 ACA americanum Ker-Gawl. ssp. americanum, 2006 L. Speedy LS-06-86 Hemerocallis fulva (L.) L., 1985 S.A. Thompson 2436 Asia Hypoxis hirsuta de .) Coville, 2006 L. Speedy LS-06-339 Lilium canadense L. ssp. canadense, 1929 W.C. Grimm s.n. Lilium pon L. ssp. editorum (Fern.) Wherry, 1967 W.E. Maianthemum canadense Desf., 1970 W.E. Buker s.n. Maianthemum racemosum (L.) Link ssp. racemosum, 1999 J. Polonoli FG1 Narcissus addas D ee W.E. Buker s.n. Europe Ornithogalum un 1L,, 2005 L. Speedy and K. Jurkovic E Europe Polygonatum biflorum (Walt.) Ell, 2005 L. Speedy LSB628 Polygonatum pubescens (Willd.) Pursh, 2005 L. Speedy LSB905 bie gramineum (Ker- aa ) Morong var.gramineum, 1 son s.n. G4G5: Tri de erectum L., 2005 L. P LSB88 Trillium flexipes Raf., 2005 L. Speedy LSB83 G5:S2 Trillium grandiflorum (Michx.) Salisb., 2005 L. Speedy LSB90 Trillium nivale Riddell, 2006 L. Speedy LS-06-1 G4:S3 Trillium sessile L., 2005 L. Speedy and D. White LSB155 Uvularia grandiflora Sm, 2005 L. Speedy LSB477 Uvularia perfoliata L., 2001 J. Polonoli FG81 Limnanthace Floerkea an Willd., 2005 L. Speedy LSB47 Linaceae Linum virginianum L., 2005 L. Speedy LSB912 Lythraceae 1950 L.K. Henry and W.E. Buker s.n. a salicaria L., 2005 L. Speedy and M. Bowers and M. e LSB1325 Europe L $ o UPFICG VISCO5ISSTHTIG Jacq Magnoliaceae Liriodendron vivido L., 2006 L. Speedy LS-06-259 lia (L) L, 1904 OE. Jennings s.n. Malvac vane Peer Medik., 1970 W.E. Buker s.n. Asia Hibiscus trionum L., 1949 L.K. Henry s.n. ded Malva moschata L., 1888 L. Hazzard s.n. Euro Malva neglecta Wallr, 1952 D. Wilson 215 Ad Sida spinosa L., 1892 FT. Aschman s.n. Menispermaceae Menispermum canadense L., 2006 L. Speedy and J. Nusser LS-06-539 Mollugina Mollugo iate L., 1946 H.N. Mozingo s.n. Monotropac Monotropa pe L., 2005 L. Speedy LSB915 Morton and Speedy, Flora of Washington County, Pennsylvania Moraceae Maclura pomifera (Raf) Schneid., 2005 L. Speedy LSB1657-1 Morus alba L., 2006 L. Speedy LS-06-394 Asia leacea En americana L., 1987 S.A. Thompson 3702 Fraxinus nigra 2005 M. Bowers and B. Owen s.n. Fraxinus Mb. Marsh., 2005 L. Speedy LSB1041-2 Ligustrum vulgare L., 2005 L. Speedy LSB1668 Europe nagra Circaea hone E ps USE (L.) Aschers. & Magnus, Speedy LSB iion: EIU "d 2 oe 2005 L. Speedy and M. and M. Lan Eicon coloratum E 2005 L. Speedy LSB12 pilobium parviflorum Schreb., 2006 L. Speedy nm Nusser Gaura biennis L., 2006 L. Speedy LS-06-877 Ludwigia alternifolia L., 2005 L. Speedy and M. Bowers and M. Lange LSB1320 Ludwigia palustris (L.) Ell, 1951 L.K. Henry and FH. Beer s.n. Oenothera biennis L., 1951 L.K. Henry and F.H. B o fruticosa L. ssp. glauca (Michx.) ne Ar OE Ings S A do L., 2005 L. Speedy LSB1656 Oenothera perennis L., 1970 W.E. Buker s.n. rchidaceae Galearis spectabilis (L.) Raf., 1888 A.M. Fulton s.n. Goodyera pubescens (Willd.) R. Br. ex Ait.f., 2006 L. Speedy and 3 Liparis liliifolia (L) L.C. Rich. ex Ker-Gawl, 2006 L. Speedy and J Liparis loeselii (L) L.C. Rich., 1995 J.S. Shriver and A. Shriver n mith 390 Platanthera lacera (Michx.) G. Don, 1970 W.E. Buker s.n. Spiranthes cernua (L.) L.C. Rich., 1994 J.S. Shriver and A. Shriver and C. Smith 345 Sante lacera (Raf.) ipi var. gracilis (Bigelow) Luer, K. Henry and W.E. B T Aden (E p 1968 W.E. Buker s.n. 1950 Orobanchaceae Conopholis americana (L.) Wallr. f, 2005 L. Speedy LSB1540 Epifagus virginiana (L.) W. Bart. 1942 O.E. Jennings s.n. 0 1939 OE. Jennings s.n. robanche uniflora L., Oxalidaceae Oxalis corniculata L., 2005 L. Speedy LSB617 Oxalis grandis Small, 2005 L. Speedy LSB601 Oxalis stricta L., 2005 L. Speedy LSB1489 Oxalis violacea L., 2005 L. Speedy LSB573 Papaveraceae cod majus L. var. majus, 2005 L. Speedy LSB265 rope is L, 2005 L. Speedy LSB92 anguinaria canaden Passifloraceae Passiflora v L., 2004 J.A. Isaac 18574 G5:S1 1243 Phytolaccaceae Phytolacca americana L. var. americana, 2005 L. Speedy LSB958 Ed Ri teglnaceas P. tata Mi hx. 1940 W.G. Burig eee lanceolata L., ~ L. Speedy D. Sofran LS-06- 1031 Europe Plantago major L., 1952 D. Wilson 202 Plantago rugelii Dcne. var. rugelii, 2005 L. Speedy LSB1495 Plantago virginica L., 1913 O.E. Jennings s.n. Platanaceae Pl q $21 talis L., 2006 L. Speedy LS-06-495 Poaceae Agrostis canina L., 2006 L. Speedy LS-06-1020 Agrostis gigantea Roth, 2005 L. Speedy LSB765 um Agrostis hyemalis (Walt.) B.S.P, 1965 W.E. Buker o E (Walt) Tuckerman, x. L Speedy men pu Willd., 1946 R.W. Pohl 5888 Agrostis stolonifera L., 1979 D.E. Boufford and N. Foust 21191 Alopecurus pratensis L., 2001 M. Bowers s.n. Eurasia Andropogon um L. var. virginicus, 1999 B.L. Isaac and J.A. Isaac Antonium ME L. ssp. odoratum, 2005 L. Speedy ls si A ichx., 1960 W.E. Buker s.n. P Pe (L.) Beauv. ex J. & K. Presl var. elatius, 2001 : Brachyelytrum erectum (Schreb. ex Spreng.) Beauv., 2005 L. Speedy LSB990 Bromus Schrad., 2005 L. Speedy and M. Bowers 5 Europe Bromus inermis Leyss. ssp. inermis var. inermis, 1970 W.E. Buker Bromus pubescens Muhl.ex Willd., 1957 W.E. Buker s.n. Bromus secalinus L., 1907 J.A. Shafer s.n. Europe Cinna arundinacea L., 2005 L. Speedy LSB1211 o glomerata L. ssp. ie 1985 S.A. Thompson and H. Nishida 2408 Euro Da omnia spicata (L) Beauv. ex Roemer & J.A. Schultes, 2006 L. Speedy LS-06- s americana Beauv., 2002 B.L. Isaac and J.A. Isaac 5791 G4?:S1 aO E Risa (Sw.) Gould €: C.A. Clark var. 002 J. Polonoli FG123A Dichanthelium acuminatum (Sw.) Gould & C.A. Clark var. fas- ciculatum (Torr.) Freckmann, 1987 S.A. Thompson 3703 Dichanthelium boscii (Poir) Gould & C.A. Clark, 1904 O.E. nnings s.n Dichanthelium clandestinum (L.) Gould, 2005 L. Speedy acuminatum LSB1479 Dichanthelium dichotomum (L.) Gould var. dichotomum, 2005 L. Speedy LSB1177 Dichanthelium latifolium (L) Gould & C.A. Clark, 1957 W.E. ker s.n. Dichanthelium linearifolium (Scribn. ex Nash) Gould, 2006 L. Speedy and J. Nusser LS-06-543 1244 Dichanthelium sphaerocarpon (Ell.) Gould var. sphaerocarpon 1970 W.E. Buker s.n. Digitaria ischaemum (Schreb) Schreb. ex Muhl., 1919 O.E. nni 6272 Eurasia Echinochloa crus-galli (L) Beauv., 2005 L. Speedy and M. Bow- ers LSB1364 Eurasia Echinochloa muricata (Beauv.) Fern. var. muricata, 2005 L. Speedy LSB1261 Eleusine indica (L.) Gaertn., 1919 O.E. Jennings s.n. Old World Tropics Elymus hystrix L. var. hystrix, 2005 L. Speedy LSB932 Elymus riparius Wieg., 2005 L. Speedy LSB1458 Elymus villosus Muhl.ex Willd., 2005 L. Speedy and M. Bowers LSB1024 Elymus virginicus L. var. virginicus, 2005 L. Speedy and M. Bowers LSB1072 Eragrostis cilianensis (All.) Vign. ex Janchen, 1919 O.E. Jennings Eragrostis frankii C.A. Mey. ex Steud., 2005 L. Speedy and M. and M. Lange LSB1312 Eragrostis hypnoides (Lam.) B.S.P, 2005 L. Speedy and M. Bowers LSB1030 Eragrostis minor Host, 1940 FH. Bell 293 Europe Eragrostis pectinacea (Michx.) Nees ex Steud. var. pectinacea, 1919 OE. Jennings s.n Festuca subverticillata (Pers.) Alexeev, 2005 L. Speedy LSB1138 Glyceria striata (Lam.) A.S. Hitchc., 2005 L. Speedy LSB1205 Holcus lanatus L., 2005 L. Speedy LSB918 Europe Leersia oryzoides (L.) Sw., 2005 L. Speedy LSB1264 Leersia virginica Willd., 2005 L. Speedy and M. Bowers Leptochloa fusca (L.) Kunth ssp. fascicularis (Lam.) N. Snow, 1986 AW. Cusick 25725 Lolium sd (Huds) SJ. Darbyshire, 2006 L. Speedy LS- 06-925 Hirst v vimineum (Trin.) A. Camus, 2006 L. Speedy Molte rondo (Poir.) Fern., 2006 L. Speedy and M. Muhlenbergia ni (Muhl. ex Willd.) Trin., 1940 J.M. Fogg Muhlenbergia sylvatica Torr. ex Gray, 2005 L. Speedy LSB1215 Muhlenbergia tenuiflora (Willd.) B.S.P., 2005 L. Speedy LSB1354 Panicum capillare L., 1940 L.K. Henry s.n. Panicum dichotomiflorum Michx. var. dichotomiflorum, 2006 L. E ES 06- 910 Pan tti Nash, 1919 O.E. Jennings s.n. Panicum ROCCO Bernh. ex Trin., 1983 A.W. Cusick Le- == 23226 Panicum rigidulum Bosc ex Nees var. elongatum (Pursh long, 2006 L. Speedy and D. Sofran LS- d 824 Bid. setaceum Michx., 1940 FH. Bell 4 Pennisetum glaucum (L.) R. Br., 1940 LK. Hens China Phalaris arundinacea L., 2005 L. Speedy LSB82 Phleum pratense L., 1952 D. Wilson 161 a Ll Dag PRA D hi Air Journal UI f Texas 1(2) Poa alsodes Gray, 2005 L. Speedy ls K Jurkovic LSB417 Poa annua L., 1947 W.E. Buker s.n. Poa compressa L., 2006 L. Speedy ipt Europe Poa cuspidata Nutt., 2005 L. Speedy LSB74 Poa pratensis L. ssp. pratensis, 1970 W.E. Buker s.n. Poa sylvestris Gray, 2005 L. Speedy LSB359 Poa trivialis L., 2005 L. Speedy LSB969 Europe Puccinellia distans (Jacq.) Parl. ssp. distans, 1984 A.W. Cusick 23492 Schizachyrium scoparium (Michx.) Nash var. scoparium, 1904 nnin Sclerochloa dura (L.) Beauv., 2001 A.W. Cusick 35790 E Setaria faberi Herrm., 2006 L. Speedy LS-06-924 Setaria pumila (Poir. Roemer & J.A. Schultes, dn i Speedy -06-735 Europe Setaria viridis (L.) Beauv. var. viridis, 1 Eurasia ao cdd Bosc ex Link, 2006 L. Speedy and D. Sofran LS-0 1 S.S. Dickey s.n. P intermedia (Rybd.) Rybd., 1980 D.E. Boufford and E.W. Wood and J. Bawa 22157 Sphenopholis nitida (Biehler) Scribn., 1906 O.E. Jennings s.n. Tridens flavus (L.) A.S. Hitchc. var. flavus, 2005 L. Speedy and M. Bowers LSB1376 Polemoniace Phlox a ssp. divaricata, 2005 L. Speedy LSB355 Phlox maculata L. ssp. maculata, 1970 W.E. Buker s.n. Phlox paniculata L., 2005 L. Speedy LSB1163 Polemonium reptans L. var. reptans, 2005 L. Speedy and M. Bowers LSB872 Polygalaceae Polygala sanguinea L., 2006 L. Speedy LS-06-906 Polygala verticillata L. var. verticillata, 1956 E. Mason s.n. Polygonaceae Polygonum aviculare L., 1989 S.A. Thompson 6852 Europe Polygonum caespitosum Blume var. longisetum (de Bruyn) A.N. Steward, 2005 L. Speedy LSB1488 Polygonum convolvulus L. var. convolvulus, 1950 L.K. Henry and PECORI a a & Zucc., 2005 L. Speedy LSB1673 Japan Polygonum erectum L., 1946 W.E. Buker s Polygonum hydropiper L., 1984 P ral pson and J.H. Nishida 2 Furope Polygonum hydropiperoides Michx., 2005 L. Speedy LSB819 pensylvanicum L., 2005 L. Speedy and M. Bowers and M. Lange LSB1327 Polygonum perfoliatum L., 2005 L. Speedy LSB1678 Asia Polygonum persicaria L., 2005 L. md LSB1262 Europe Polygonum punctatum Ell. var. (Meisn.) Fassett, 2005 L. Speedy and M. Bowers LSB1049 Polygonum Ed Ell. var. punctatum, 2006 L. Speedy and D. Sofran LS-06-806 Polygonum pt L., 2005 L. Speedy LSB1252 Polygonum scandens L. var. dumetorum (L.) Gleason, 2005 L. Speedy LSB1493 Polygonum virginianum L., 2005 L. Speedy LSB1501 Polygonum Morton and Speedy, Flora of Washington County, Pennsylvania Rumex acetosella L., 2005 L. Speedy and K. Jurkovic LSB409 Europe Rumex crispus L. ssp. crispus, 2005 L. Speedy LSB675 Europe Rumex obtusifolius L., 2005 L. Speedy LSB809 Europe Portulac Corona. d Michx. var. caroliniana, 1920 O.E. Jen- nings Claytonia Mm L. var. virginica, 2005 L. Speedy LSB91 Portulaca oleracea L., 1952 D. Wilson 205 tamogetonace Potamogeton crispus s 2006 L. Speedy LS-06-251 Europe Primulaceae Anagallis arvensis L. ssp. arvensis, 1901 J.A. Shafer s.n. Eurasia Lysimachia ciliata L., 1984 S.A. Thompson and J.H. Nishida 2114 Lysimachia nummularia L., 2005 L. Speedy LSB804 Europe Lysimachia quadrifolia L., 2005 L. Speedy LSB717 Pyrolaceae B a ) Pursh, 2005 L. Speedy LSB71 Pyrola americana Sweet, 1970 W.E. Buker s.n. Pyrola RE: Nutt., 2006 L. Speedy LS-06-489 Ranunculaceae Actaea pachypoda Ell., 2006 L. Speedy LS-06-254 ctaea racemosa L. var. racemosa, 2005 L. Speedy and M. Bowers LSB874 Anemone canadensis L., 1920 O.E. Jennings s.n. Anemone quinquefolia L. var. quinquefolia, 1918 J. Bright s.n. Anemone virginiana L. var. virginiana, 2005 L. Speedy 4 Aquilegia canadensis L., 2006 L. Speedy and B of Western PA LS-06-269 Caltha palustris L. var. palustris, 2005 L. Speedy LSB52 Clematis virginiana L., 2005 L. Speedy LSB1467 Consolida ajacis (L.) Schur, 1901 J.A. Shafer s.n. Europe Delphinium exaltatum Ait., 2005 L. Speedy LSB1233 G3:S1 Delphinium tricorne Michx., 2005 L. Speedy LSB373 Hepatica nobilis Schreber var. acuta (Pursh) Steyermark, 2005 tanical Society Hepatica nobilis Schreber var. obtusa (Pursh) Steyermark, 2005 is E 2 pios Hy is L., 2006 L. Speedy LS-06-242 RATE abortivus L., i L. Speedy LSB590 Ranunculus dae 5 var. acris, E W.E. Buker s.n. Europe Ranunculus t, 2005 L. Speedy LSB435 Ranunculus E E oo Utech and S.A. Thompson a 82-174 Eur Rann me MU var. t (Chapman) T. Duncan, 2 .Spee B37 ES micranthus dm 2005 L. Speedy LSB591 ees recurvatus Poir. var. recurvatus, 2005 L. Speedy LSB59 Rranrincitltic renens li) 2005 L. Speedy LSB379 Europe dla sceleratus x var. sceleratus, 1993 J.A. Isaac 5398 5 L. Speedy LSB27 Thalictrum DUC sp 2001 J. Polonoli FG86 Thalictrum revolutum DC., 1904 O.E. Jennings s.n. alietrim dioicim 1245 SN Thalictrum thalictroides (L.) Eames €: Boivin, 2006 L. Speedy Rhamnaceae a americanus L., 1957 W.E. Buker s.n hartica L., 1948 M.A. Caoba s.n. Europe hamniı Rosaceae Agrimonia gryposepala Wallr, 1966 W.E. Buker s.n. a parviflora Ait., 2005 L. Speedy LSB1272 Agrimonia pubescens Wallr., 2005 L. Speedy LSB1195 depen arborea (Michx. f) Fern. var. arborea, 2005 L. Speedy LSB1515 Amelanchier humilis Wieg., 1923 O.E. Jennings s.n. G5:S1 Amelanchier stolonifera Wieg., 1923 O.E. Jennings s.n. Aruncus dioicus (Walt) Fern. var. dioicus, 2005 L. Speedy LSB926- E chrysocarpa Ashe var. chrysocarpa, 2005 L. Speedy 0 Pun crus-galli L., 1921 J. Bright s.n Crataegus dissona Sarg., 1907 O.E. ISTIS and G.K. Jen- nings tree Crataegus intricata Lange, 1907 O.E. Jennings tree 5 Crataegus iracunda Beadle, 1907 O.E. Jennings D. G.K. nnings s.n. Crataegus pruinosa (Wendle. f) K. Koch, 1905 O.E. Jennings G.E. Kinzer tree 35 Crataegus punctata Jacq., 2005 L. Speedy LSB847 Duchesnea indica (Andr.) Focke, 2005 L. Speedy LSB289 Asia dd ee Duchesne y poe (Vilm. ex J. Gay) audt, 2006 L. Speedy LS-0 Fragaria virginiana Duchesne ssp. virginiana, 1951 LK. Henry and F.H. Be e gout Jacq., 2005 L. Speedy LSB258 Geum canadense Ja a var. canadense, 2005 L. Speedy and M. Bowers En Geum vernum (Raf.) oe & Gray, 2005 L. Speedy LSB280 Malus baccata (L.) Borkh., 2005 L. Speedy and B. Brown LSB658 Eurasia Malus ~ (L) P. Mill, 2005 L. Speedy and K. Jurkovic LSB3 o — Sieb. exVan Houtte, 2005 L. Speedy LSB1474 "m — P. Mill, 2005 L. Speedy LSB19 1 Eurasia [Pyrus malus L] mu um opulifolius (L) Maxim. var. opulifolius, 2005 L. Speedy LSB640 Porteranthus trifoliatus (L.) Britt. 1977 W.E. Buker s.n. Potentilla arguta Pursh ssp. arguta, 1919 O.E. Jennings and .E. McKee s.n. Potentilla canadensis L. var. canadensis, 1951 L.K. Henry and .H. Beer s.n. Potentilla canadensis L. var. villosissima Fern., 2005 L. Speedy LSB440 Potentilla norvegica L. ssp. monspeliensis (L.) Aschers. & Graebn., 1920 OE. Jennings s.n Potentilla recta L., 1987 S.A. Thompson 3709 Europe Potentilla simplex Michx., 2006 L. Speedy LS-06-255 Prunus americana Marsh., 1978 W.E. Buker s.n. 1246 Prunus avium (L.) L., 2006 L. Speedy LS-06-82 Eurasia Prunus mahaleb L., 2005 L. Speedy LSB716 Europe Prunus serotina Ehrh. var. serotina, 2005 L. Speedy LSB194-1 Prunus virginiana L. var. virginiana, 2005 L. Speedy and K. Jurkovic LSB406 Rhodotypos scandens (Thunb.) Makino, 2005 L. Speedy LSB272 Japan Rosa canina L., 1930 W.C. Grimm s.n. Euro Rosa carolina L. var. M 2005 L. m LSB1223 Asia Rosa eglanteria L., 1904 O.E. Jennings s.n. Europe Rosa multiflora Thunb. ex ma 2005 L. Speedy LSB829 Asia Rosa palustris Marsh., 1972 L.K. Henry and W.E. Buker s.n Rosa setigera Michx. var. tomentosa Torr. & Gray, 2005 L. Speedy Rubus allegheniensis Porter var. allegheniensis, 2005 L. Soeedy Rubus alumnus Bailey, 1958 H.A. Davis and T. Davis 12147 Rubus bellobatus Bailey, 1958 H.A. Davis and T. Davis 12156 Rubus fecundus Bailey, 1958 H.A. Davis pet Fund 2154 Rubus Maus Willd. 2005 L. Speedy LSB4 Rub s L. ssp. strigosus (Michx) cane a L.K. Henry n E M Rubus laudatus Berger, 1958 H.A. Davis and T. Davis 12153 Rubus da (Card ex Bailey) Bailey, 1958 H.A. Davis and T. D 12140 ioi mulier ES ee H.A. Davis and T. ME 10953 talis L., 2005 L. Speedy LSB91 L. var x 2006 L. en LS-06-494 Rubus pensilvanicus Poir., 1961 H.A. Davis and T. Davis 13248 ubus recurvicaulis Blanch., 2006 L. Speedy and J. Nusser Ririh A. + LS-06-552 Rubus roribaccus (Bailey) Rydb., 1958 H.A. Davis and T. Davis 12152 Rubus rosa Bailey, 1958 H.A. Davis and T. Davis 121241 Spiraea alba Du Roi var. ~ (Ait) Dippel, 1919 O.E. Jen- nings and A.E. McKee s.n. Spiraea tomentosa L., 2006 i Speedy LS-06-737 Waldsteinia ee (Michx.) Tratt. ssp. fragarioides, 2006 L. Speedy LS-06-12 Rubiaceae Cephalanthus occidentalis L., 2005 L. Soeedy and M. Bowers LSB1380-2 Galium aparine L., 2005 L. Speedy LSB291 Galium asprellum Michx., 2005 L. Speedy and M. Bowers Galium circaezans Michx. var. circaezans, 1967 W.E. Buker s.n. Galium circaezans Michx. var. hypomalacum Fern., 2005 L. Speedy LSB962 Galium concinnum Torr. & Gray, 1946 E. Mason s.n. Galium mollugo L., 2005 L. Speedy LSB618 Eurasia Galium pilosum Ait. var. pilosum, 1999 J. Polonoli FG4 lium tinctorium (L) Scop, 2005 L. Speedy LSB1263 Gaium um m 2005 L. Speedy LSB 4 6 aL, 2005 L. Speedy LSB5 ou E Willd. ex Roemer & J. A Schultes, 2005 . Speedy LSB612 Houstonia longifolia Gaertn., 1920 O.E. Jennings s.n. Mitchella repens L., 1970 L.K. Henry and W.E. Buker s.n. tonia Or £L Dat PRA D hi PEA Journal UI f Texas 1(2) Rutaceae Zanthoxylum americanum P. Mill, 1940 L.K. Henry s.n. icaceae Populus balsamifera L. ssp. balsamifera, 1906 O.E. Jennings 5:51 Populus grandidentata Michx., 2006 L. Speedy and J. Nusser -06- Salix alba L., 1919 O.E. Jennings and E.M. Gress s.n. Eurasia Salix caroliniana Michx., 1908 O.E. Jennings s.n. G5:S1 Salix discolor Muhl., 1919 O.E. Jennings and E.M. Gress s.n. Salix eriocephala Michx., 1979 D.E. Boufford and E.W. Wood W.E. Buker 20553 Salix fragilis L., 1970 L.K. Henry and W.E. Buker s.n. Europe Salix interior Rowlee, 1978 W.E. Buker s.n. Salix nigra Marsh., 2005 L. Speedy LSB690-2 Salix pupurea L., 2005 L. Speedy LSB691-2 Europe Salix sericea Marsh., 2005 L. Speedy LSB197-2 Saxifragaceae Chrysosplenium americanum Schwein. ex Hook., 1970 W.E. Buker Heuchera americana L. var. americana, 2006 L. Speedy LS- 06-387 Mitella diphylla L., 2005 L. Speedy and K. Jurkovic LSB420 Saxifraga virginiensis Michx. var. virginiensis, 2005 L. Speedy LSB75 Scrophular Agalinis Spi (Vahl) Raf. var. tenuifolia, 2005 L. Speedy SB1641 uo sd minus (L.) Lange, 2006 L. Speedy LS-06-281 NUN m L., 2005 L. Speedy and M. a. LSB1427 Collinsia verna Nutt., 2005 L. Speedy LSB370 Gratiola neglecta Torr., 2005 L. Speedy hw i Bowers LSB880 Linaria vulgaris P. Mill., 1940 L.K. Henry s.n. Eura Lindernia dubia (L) Pennell var. dubia, 2005 L. ae and M. Bowers LSB1033 Mimulus ringens L. var. ringens, 2005 L. Soeedy LSB1257 RP canadensis L. ssp. canadensis, 1970 W.E. Buker s.n. n digitalis Nutt. ex Sims, 2006 L. nd LS-06-420 Penstemon hirsutus (L.) Willd., 1930 W.C. Grim Penstemon laevigatus Ait, 1954 L.K. Henry s.n. ioo. Scrophularia lanceolata Pursh, 1970 W.E. Buker s.n. Scrophularia marilandica L., 2005 L. Speedy LSB1465 Verbascum blattaria L., 2006 L. Speedy LS-06-377 Eurasia Verbascum thapsus L., 1950 L.K. Henry and W.E. Buker s.n. Furasia AS) Veronica americana Schwein. ex Benth., 2006 L. Speedy -06-485 Veronica anagallis-aquatica L., 1952 D. Wilson 101 G5:S1 Veronica arvensis L., 1921 O E. Jennings s.n. Eur Veronica officinalis L. var. officinalis, 2005 L. Sere LSB560 Veronica peregrina L. ssp. peregrina, 2005 L. Speedy LSB608 Veronica persica Poir., 2001 A.W. Cusick 35787 Eurasia Veronica serpyllifolia L. ssp. serpyllifolia, 2006 L. Speedy LS- Europe Veronicastrum virginicum (L.) Farw., 1931 W.C. Grimm s.n. Morton and Speedy, Flora of Washington County, Pennsylvania Simaroubacea Ailanthus WA (P. ne Swingle, 2006 L. Speedy and D. Sofran LS-06-819 Smilacaceae Smilax herbacea L., 2006 L. Speedy and J. Nusser LS-06-537 Smilax tamnoides L., 2005 L. Speedy LSB1676 Solanaceae Datura stramonium L., 1970 W.E. Buker s.n Nicandra physalodes (L.) Gaertn., 1962 M.G. Netting s.n. Peru Petunia x atkinsiana D. Don ex Loud,. 1940 L.K. Henry s.n. Exotic Physalis Mi Nees var. heterophylla, 2005 L. Speedy and M LSB1028 Pa m D var. subglabrata (Mackenzie & eh onq., 2005 L. Speedy and M. Bowers and M. Lange Solanum carolinense L. var. carolinense, 1995 B.L. Isaac and A. Isaac 8338 Solanum dulcamara L. var. dulcamara, 2005 L. Speedy LSB825 Eurasi Solanum pt nthum D Solanum Pca Dunal, ee E o and D. Sofran LS-0 nal] Staphyleac Staphylea ms L., 2005 L. Speedy LSB484 Thymelaeaceae Dirca palustris L., 2002 S.P. Grund 2967 Tiliaceae Tilia americana L. var. americana, 2005 L. Speedy LSB1675 lilia americana L. var. heterophylla (Vent) Loud., 2005 L. Speedy LSB1543 Typhaceae Typha angustifolia L., 1952 D. Wilson 249 Typha latifolia L., 2005 L. Speedy LSB1660 Ulmacea Celtis oM ES d L. Speedy LS-06- mericana L., 2005 L. Speedy and K. er LSB388 Ulmus rubra Muhl., js L. Speedy LS-06-18 Urticaceae Boehmeria cylindrica (L.) Sw., 2005 L. Speedy LSB1439 nidis EA (L.) Weddell, 2005 L. Speedy and M. Bowers LSB1387 m pud Muhl. ex Willd., 1951 L.K. Henry and Pilea pumila (L.) Gray var. pumila, 2005 L. Speedy LSB1508 Urtica dioica L. ssp. dioica, 1952 D. Wilson 122 Europe Urtica dioica L. ssp. gracilis (Ait.) Seland., 2005 L. Speedy LSB844 Valerianaceae Valeriana pauciflora Michx., 2005 L. Speedy LSB592 Valerianella chenopodiifolia (Pursh) DC., 2005 L. Speedy LSB607 Valerianella umbilicata (Sullivant) Wood, 1951 L.K Henry and 1247 Verbenaceae Glandularia canadensis (L) Nutt., 1901 J.A. Shafer s.n. [Glan- NS ue drummondii p ) Small] Phryma leptostachya L., 1954 L.K. Henry and W.E. Buker s.n. e no L. var. es 2005 L. Speedy and M. Bowers .Lange LSB1301 Verbena urticifolia L. var. leiocarpa Perry & Fern,, enry s.n. Verbena urticifolia L. var. urticifolia, 2005 L. Speedy LSB1663 1940 L.K. Violaceae Hybanthus concolor (T.F. Forst.) Spreng., 2005 L. Speedy S LSB593 Viola affinis Le Conte, 1940 L.K. Henry Viola canadensis L. var. canadensis, 2002 S.P. Grund 2982 Viola cucullata Ait., 2005 L. Speedy LSB285 Viola pubescens Ait. var. scabriuscula Schwein.ex Torr. & Gray, 20 y LSB24 Viola sagittata Ait. var. sagittata, 1969 W.E. Buker s.n. Viola sororia Willd., 2005 L. Speedy LSB54 Viola striata Ait., 2005 L. Speedy LSB368 Viola x palmata L (pro sp.), 1951 L.K. Henry and FH. Beer s.n. Vitaceae Parthenocissus quinquefolia (L.) Planch., 2005 L. Speedy LSB294 Vitis aestivalis Michx. var. bicolor Deam, 1951 W.E. Buker s.n. Vitis cinerea (Engelm.) ae var. baileyana (Munson) Co- m 1919 E.M. Gress s Vitis riparia Pond. 2006 L. nue LS-06-492 Vitis vulpina L., 1994 J.A. Isaac 6047 GYMNOSPERMS Cupressaceae aer communis L. var. ca Pursh, 1916 O.E. Jennings ings s. m virginiana Ñ var. virginiana, 2006 L. Speedy LS- 06-911 Pinac d E (Lam.) Carr., 2006 L. Speedy and J. Nusser -06-607-1 Japan Picea glauca (Moench) Voss, 2005 L. Speedy LSB1180 Pinus banksiana Lamb., 2006 L. Speedy and J. Nusser LS- -550-1 Pinus resinosa Ait., 1979 P. Singleton and D.E. Boufford and Pinus strobus L., 1919 OE. Jennings s.n. Pinus virginiana P. Mill., 1913 O.E. Jennings and G.K. Jennings and A.R. Hillard s.n. Tsuga canadensis (L.) Carr., 2006 L. Speedy LS-06-22 aceae Taxus canadensis Marsh., 1919 O.E. Jennings s.n. LYCOPHYTES Lycopodiaceae d digitatum Dill. ex A. Braun, 2006 L. Speedy and J. Nusser LS-06-542 1248 Lycopodium hickeyi W.H. Wagner, Beitel & Moran, 2006 L. Speedy and J. Nusser LS-06-546 Lygodiace Lygodium palmatum (Bernh.) Sw., 1999 B.L. Isaac and J.A. Isaac 1 PTERIDOPHYTES Aspleniaceae E ace x e P. var. platyneuron, 1999 B.L. and J.A. Isaac md END E eos L. Speedy and M. Bowers LSB878 Dennstaedtiaceae Dennstaedtia punctilobula (Michx.) T. Moore, 2005 L. Speedy Pteridium aquilinum (L.) Kuhn var. latiusculum (Desv.) Under- wood ex Heller, 1954 L.K. Henry s.n. Dryopteridaceae Athyrium filix-femina (L.) Roth ae tum (Willd.) Clausen, 2005 L. ae LSB11 Athyrium filix-femina (L.) sen ssp. asplenioides (Michx.) Hulten, 1954 LK. Henry s.n Cystopteris bulbifera (L ) Bernh., 2006 L. Speedy LS-06-493 Cystopteris fragilis (L) Bernh., 2005 L. Speedy and M. Bowers LSB879 Cystopteris protrusa (Weatherby) Blasdell, 2005 L. Speedy LSB273 Cystopteris tenuis (Michx.) Desv., 1996 B.L. Isaac and J.A. Isaac 8921 Deparia acrostichoides (Sw.) M. Kato, 2005 L. Speedy Diplazium pycnocarpon (Spreng.) Broun, 2005 L. Speedy 981 £L Dag PRA D hi Air Journal of f Texas 1(2) Dryopteris goldiana (Hook. ex Goldie) Gray, 2005 L. Speedy SB1161 Dryopteris intermedia (Muhl. ex Willd.) Gray, 2005 L. Speedy SB1507 Dryopteris marginalis (L.) Gray, 2005 L. Speedy LSB1139 uon x triploidea Wherry, 1995 B.L. Isaac and J.A. Isaac Onocko sensibilis L, Ud la M LSB1202 hx.) Schott var. acrostichoides, 2005 L. Speedy LSB902 Woodsia obtusa (Spreng.) Torr. ssp. obtusa, 1940 E.T. Wherry nd J gg Jr Equiseta nes arvense L., 2005 L. Speedy LSB295 Equisetum hyemale L. var. affine (Engelm.) AA. Eat., 2006 L. Speedy LS-06-19 Hymenophyllaceae Trichomanes intricatum Farrar, 1984 J.C. Parks 4381 Ophioglossaceae Botrychium dissectum Spreng., 1995 B.L. Isaac and J.A. Isaac Botrychium virginianum (L.) Sw., 2006 L. Speedy LS-06-238 Osmundaceae Osmunda claytoniana L., 1951 L.K. Henry and F.H. Beer s.n. Polypodiaceae Polypodium appalachianum Haufler € Windham, 1951 L.K. Henry and F.H. Beer s.n. Pteridaceae Adiantum pedatum L., 2005 L. Speedy LSB1142 Thelypteridaceae Phegopteris hexagonoptera (Michx.) Fee, 2006 L. Speedy f . -06-484 Dryopteris carthusiana (Vill) H.P. Fuchs, 2005 L. Speedy Thelypteris noveboracensis (L.) Nieuwl., 2005 L. Speedy LSB945 SB1174 Dryopteris cristata (L) Gray, 2003 S. Ernst and E. Zimmerman Thelypteris palustris Schott var. pubescens (Lawson) Fern., 1951 148 nry and FH. Beer s.n ACKNOWLEDGMENTS Our thanks go to the Wild Resource Conservation Fund of the Pennsylvania Department of Conservation and Natural Resources for partial funding of the field work. We would also like to thank an anonymous reviewer for reviewing the manuscript. We are also thankful to Amanda Juretic for data entry. REFERENCES Brummit, R.K. and C.E. Powe. 1992. Authors of plant names. Royal Botanic Gardens, Kew. DCNR. 2004. Invasive plants of Pennsylvania. http://www.dcnrstate.pa.us/forestry/wildplant/invasive.aspx PENNSYLVANIA DEPARTMENT OF AGRICULTURE. 2007. Pennsylvania noxious weed control list. http://www.agriculture.state pa.us/agriculture/cwp/view.asp?q=127683 PENNSYLVANIA DEP WATERSHED NoteBook. 2006. http://www.dep.state.pa.us/dep/deputate/watermgt/WC Forrest, E.R. 1926. History of Washington County, Pennsylvania. The S.J. Clarke Publishing Company, Chicago, II. Kartesz, J.T. 1999. A synonomized checklist and atlas with biological attributes for the vascular flora of the United States, Canada and Greenland. 1% ed. In: Kartesz, J.T. and C.A. Meacham. Synthesis of the North American flora, Version 1.0. North Carolina Botanical Garden, Chapel Hill. Morton and Speedy, Flora of Washington County, Pennsylvania 1249 RHOADS, A.F. and W.M. Kein, Jr. 1993. The vascular flora of Pennsylvania: annotated checklist and atlas. American Philosophical Society, Philadelphia, Pennsylvania. RHoaps, A.F. and T.A. BLock. 2000. The plants of Pennsylvania. University of Pennsylvania Press, Philadelphia, Pennsylvania. THe Nature CONSERVANCY. 1996. Element ranking list - Pennsylvania Natural Heritage Program. http://www.natul alheritage.state.pa.us/PlantsPage.aspx U.S. Census Bureau. 2007. State and County QuickFacts. http://quickfacts.c gov/qdfd/states/42/42125.html USDA. 1983. Soil survey of Greene and Washington counties Pennsylvania. USDA Soil Conservation Service. US Government Printing Office, Washington, D.C. Wacner, J.D. 1994. Washington County Natural Heritage inventory. Report by the Western Pennsylvania Conservancy. 1250 Journal of the Botanical R h Institute of Texas 1(2) BOOK NOTICES NEAL K. VAN ALFREN, GEORGE BRUENING, and WiLLiam O. Dawson (eds). 2007. Annual Review of Phytopathology: Volume 45, 2007. (ISBN 978-0-8243-1345-6, hbk; ISSN 0066-4286). Annual Reviews Inc., 4139 EI Camino Way, PO. Box 10139, Palo Alto, CA 94303-0139, U.S.A. (Orders: www.AnnualReviews.org, onlineactivation@annualreviews.org, 800-523-8635, 650-493-4400, 650-424-0910 fax). $197.00 (USA), 485 pp., 7 3/8" x 9 1/8". Contents of Volume 45 of Annual Review of Phytopathology: 1. Tell Me Again What It Is That You Do 2. Noel T. Keen—Pioneer Leader in Molecular Plant Pathology 3. Structure and Function of Resistance Proteins in Solanaceous Plants 4. Family Flexiviridae: A Case Study in Virion and Genome Plasticity 5. Cell Wall-Associated ced of Disease Resistance and Susceptibilit ity 6. Genom ights into the Contribution of P Bacterial Plasmids to the Evolutionary History of Their Hosts E 7. PX Ms Involved in Specific Ex eren Suppression in n So 8. Safety of Virus-Resistant Transgenic Plants Two Decades After Their Introduction: Lessons from Realistic Field Risk Assessment Studies 9. Disease Cycle Approach to Plant Disease Prediction 10. Virus-Induced Disease: Altering Host ME One Interaction at a Time 11. Bacteriophages for Plant Disease Con 12. Reniform in U.S. Cotton: When, PR ye and Some Remedies 13. Flax Rust Resist ificity is Based on Direct Resistance-Avirulence Protein Interactions 14. Microarrays for Rapid aaa of n Viruses 15. Transcript bei in UE Interactions 16. The Epidemiology a Bacterial Diseases o 17. “Blicitors, Effectors, and R Genes: The New Paradigm and a Lifetime Supply of Questions 18. Magnaporthe as a Model for Understanding Host-Pathogen Interactions 19. Challenges in Tropical Plant Nematology Two Gardening Books from Timber Press Scorr Ocben. 2007. Garden Bulbs for the South. Second Edition. (ISBN 978-0-88192-813-6, hbk.). Timber Press Inc, 133 S.W Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $34.95, 396 pp., color photographs, 6" x 9". Contents.—Preface, Introduction, 1) Rain Lily Day, 2) Petite Afrique, 3) Winter Blooms, 4) Jonquils and Kin, 5) Spring Treasures, 6) Irises, Gladioli, and Shellflowers, 7) Crinums and Spiderlilies, 8) Summer Glories, 9) Cannas, Gingers, and Aroids, 10) Designing with Southern Bulbs. Bibliography, Sources, and Index. SUE OLSEN. 2007. Encyclopedia of Garden Ferns. (ISBN 978-0-88192-819-8, hbk.). Timber Press Inc, 133 S.W. Second Avenue, Suite 450, Portland, OR 97204-3527, U.S.A. (Orders: www.timberpress.com, mail@timberpress.com, 503-227-2878, 1-800-327-5680, 503-227-3070 fax). $59.95, 444 pp., color photographs, 8 1/2" x 11". Contents.—Foreword by George Schenk, Preface, Acl led P its of F Through the Season. 1) Ferns Through the Ages, 2) E Ferns, 3) Propagating Ferns, 4) Fern Sere and Basic Diagnostics, 5) Ferns from Around the World. There are eight appendices: 1) USDA Plant Hardiness Zone Map, 2) European Plant Hardiness Zone Map, 3) Award-Winning Ferns, 4) Favorite Ferns for Sites Around the World, 5) Ferns for Special Situations, 6) Fern Societies, 7) Where to See Ferns, 8) Where to Buy Ferns. Glossary, References, Index of Plant Names. J. Bot. Res. Inst. Texas 1(2): 1250. 2007 SAXIFRAGA TEXANA (SAXIFRAGACEAE) NEW TO LOUISIANA Christopher S. Reid Barbara R. MacRoberts and Patricia L. Faulkner and Michael H. MacRoberts Louisiana Natural Heritage Program Bog Research, 740 Columbia, Louisiana Department of Wildlife and Fisheries Shreveport, Louisiana 71104, U.S.A. and Baton Rouge, Louisiana 70898, U.S.A. Herbarium, M of Life Scienc Louisiana State University in Shreveport Shre\ eport Louisiana 71115, U.S.A. ABSTRACT Saxifraga texana (Saxifragaceae) is reported as new to Louisiana. It was discovered in a saline prairie in De Soto Parish, relatively close to similar prairies in Arkansas and Texas where the species is known. RESUMEN E VE ie S Louisiana S S Se descubrió en una pradera salina en De Soto Parish, relativamente Į S g cerca de praderas similares Arkansas y Texas de donde se conocía la especie. In a recent search of Louisiana saline prairies for Geocarpon minimum Mackenzie, we discovered a population of approximately 100 plants of Saxifraga texana Buckl. in a De Soto Parish prairie (northwest Louisiana). The plants were scattered over about one hectare of a saline prairie complex that totals 10.9 hectares. We did not locate any additional S. texana populations in the course of surveys of other saline prairies. Saxifraga texana is known from Arkansas, Georgia, Kansas, Missouri, Oklahoma, and Texas, but it has not been reported for Louisiana (MacRoberts 1989; Thomas & Allen 1998; Kartesz & Meacham 2005; USDA, NRCS 2007). We were aware of the possibility of encountering Saxifraga texana during our Geocarpon surveys since it occurs on saline prairies in south Arl and east Texas (Jason Singhurst, unpublished data; Theo Witsell, voucher specimen: Witsell 03-29, Bradley County, Warren Prairie Natural Area, Herbarium of the Arkansas Natural Heritage Commission). These globally imperiled (G1G2) plant communities are primarily limited to the West Gulf Coastal Plain (McInnis et al. 1993; Keith et al. 2004; Lester et al. 2005; MacRoberts & MacRoberts 2007). Saline prairies are grassy openings with a relatively short grass canopy and usually contain scattered treed pimple (mima) mounds. They occur on high-sodium soils (Bonn silt loam) on lower landscape positions usually in close proximity to drainages. Saline prairies may be better described as barrens since they are edaphically maintained. Fire may have been a process historically but apparently is not required for mai . $mall depressions are common in the northwest Louisiana prairies and add to the plant diversity by supporting species with wetter site affinities. Characteristic species in these prairies in northwestern Louisiana include: == is minima, Aristida longespica, Aristida oligantha, Astragalus distortus, Coreopsis tincto- [oj] E Agrostis elliottiana, Anagal ria, Crassula aquatica, Eragrostis lugens, Evolvulus sericeus, Fimbristylis puberula, Geocarpon minimum, Gratiola flava, Hordeum pusillum, Houstonia micrantha, Houstonia pusilla, Houstonia rosea, Hypericum drummondii, Iva angustifolia, Isolepis carinata, Krigia occidentalis, Minuartia drummondii, Minuatria muscorum, Neptunia lutea, Oenothera linifolia, Opuntia humifusa, Phalaris caroliniana, Plantago pusilla, Rhynchospora harveyi, Ruellia humilis, Rumex hastatulus, Sabatia campestris, Sporobolus pyramidatus, Sporobolis vaginiflorus, Talinum parviflorum, Trad- escantia occidentalis, and Tridens strictus. In a future publication we will describe saline prairies more fully. Voucher specimen: LOUISIANA. De Soto Parish: Dickson Prairie, ca 2 air mi NE of Stonewall, NW1/4 S9 T15N R14W, lat/long: 321827N, 934824W, locally abundant in saline prairie, Aristida-Sporobolus zone, 7 Mar 2007, Reid 6001, with Faulkner, MacRoberts & MacRoberts (LSU, NO). J. Bot. Res. Inst. Texas 1(2): 1251 — 1252. 2007 1252 Journal of the Botanical R h Institute of Texas 1(2) ACKNOWLEDGMENTS We appreciate the following landowners for allowing access to botanize their saline prairies in several Louisiana parishes: Sidney Evans, Ralph and Becky Dalton, George Dickson, Mary and Tish Fort, Estelle Dorman, Judy Oakley, and the Weyerhaeuser Company. Alan Boyd aided us in accessing the Weyerhaeuser properties. Jason Singhurst, Theo Witsell, and Thomas Patrick provided habitat information for Saxifraga texana in Texas, Arkansas, and Georgia, respectively. Guy Nesom and Gerald Guala provided helpful com- ments on an earlier version of the manuscript. REFERENCES KARTESZ, J.A. and C.A. Meacham 2005. Synthesis of North American flora. Version 2.0. North Carolina Botanical Garden. Chapel Hill. Kerrh, E.L., J.R. SincHurst, and S. Cook. 2004. Geocarpon minimum (Caryophyllaceae), new to Texas. Sida 21:1165- 1169. Lester, G.D., S.G. SORENSEN, P.L. FAULKNER, C.S. Reip, and I.E. Maxit. 2005. Louisiana comprehensive wildlife conservation strategy. Louisiana Department of Wildlife and Fisheries, Baton Rouge. MacRoserts, D.T. 1989. A documented checklist and atlas of the vascular flora of Louisiana. Bull. Mus. Life Sci., Louisiana State University-Shreveport 9:538-756. MacRoserts, M.H. and B.R. MacRoserts 2007. Survey for pon minimum in northwest Louisiana. Unpublished Report. Louisiana Department of Wildlife and Fisheries, Baton Rouge. McInnis, N.C., L.M. SmitH, and A.B. Pitman. 1993. Geocarpon minimum (Caryophyllaceae), new to Louisiana. Phy- tologia 75:159-162. Thomas, R.D. and C.A. ALLEN 1998. Atlas of the vascular flora of Louisiana. Vol. 3. Louisiana Department of Wildlife and Fisheries, Baton Rouge. USDA, NRCS. 2007. The Plants Database (http://plants.usda.gov). National Data Center, Baton Rouge, Louisiana. VICIA LATHYROIDES (FABACEAE): NEW TO THE FLORA OF TEXAS Sonnia Hill Ruth Loper 3612 VZ CR 4915 13456 CR 2235 Ben Wheeler, Texas 75754, U.S.A. Whitehouse, Texas 75759, U.S.A. sonnia36@hotmail.com ronloper@earthlink.net Jason R. Singhurst Walter C. Holmes Wildlife Diversity Program Department of Biology Texas Parks and Wildlife Department Baylor id versity Austin, Texas 78704 U.S.A. Waco, Texas, 7679 jason.singhurstatpwd.state.tx.us walter jud e ej ABSTRACT Vicia lathyroides is reported for the first time as occurring in Texas. The overall distribution of the species and a comparison with a similar species are also discussed. RESUMEN Se cita como nueva para Texas Vicia lathyroides. La distribución total de la especie y una comparación con una especie semejante son también discutidas. Vicia lathyroides L. (spring vetch) is not included in the recent references enumerating the vascular flora of Texas (Correll & Johnston 1970; Hatch et al. 1990; Johnston 1990; Jones et al. 1997; Turner et al. 2003). The record cited here constitutes the first reported occurrence of the species in the state and the West Gulf Coastal Plain (The Nature Conservancy 2003). Voucher specimen: U.S.A. TEXAS. Smith Co.: ca. 4 mi NW of Troup on TX Hwy 110 at jet. with Co. Rd. 2235, ca. 0.7 mi E on Co. Rd. 2235, 29 Mar 2007, Hill & Loper s.n. (BAYLU, BRIT). The species, which was twining around itself and the plants near it, was discovered growing in a very dis- turbed sandy meadow. Other species occurring in the area were Cerastium glomeratum, Facelis retusa, Geranium carolinianum, Plantago virginica, Senecio ampullaceus, Soliva pterosperma, Trifolium campestre, T. dubium, Veronica arvensis, Vicia minutiflora, and V. sativa. Vicia lathyroides is regarded as a native of Asia Minor, Europe, and northwest Africa (Webb 1980). It has been reported as naturalized in Tasmania (Hnatiuk 1990, a record considered by Buchanan (2005) to be in error), Japan (Katsuyama et al. 2001), New Zealand (Webb 1980), and the United States (Hermann 1960; Isley 1990; USDA, NRCS 2007). It is widely established in the United States where it grows in sandy grasslands, old fields, and sand dunes in Alabama, California, Georgia, Massachusetts, Mississippi, North Carolina, South Carolina, Virginia, and Washington (USDA NRCS 2007). The spread of this species appears related to its usefulness for fodder, forage, soil enrichment and stabilization, and cover, particularly in drier and relatively non-fertile sandy places. Curiously, the species has experienced a decrease in numbers in the more northern parts of its region of origin, apparently from pressure on the sites for other uses (Wolf-Murphy 2005-2006). Its survival in Northern Ireland is threatened (Wolf-Murphy 2005-2006). It is also cited in the “Red Data Books” in various northern European countries, e.g., Great Britain (Cheffings & Farrell 2005) and Estonia (Lilleleht 2001-2002) The following comments and measurements are from Isely (1990), who considers this inconspicuous vetch to be similar to the common Vicia sativa L. (also an introduced species) but in miniature form. Both have sessile solitary or paired light violet flowers clustered in the upper leaf axils. Vicia lathyroides may be distinguished by its mostly solitary but occasionally paired flowers with corollas 5-6 mm long, leaves with J. Bot. Res. Inst. Texas 1(2): 1253 — 1254. 2007 1254 Journal of the Botanical R h Institute of Texas 1(2) 4-68) leaflets, and unbranched (simple) tendrils. Vicia sativa has mostly paired flowers with corollas 10-25(-30) mm long, leaves with mostly (6-)8-14 leaflets, and simple to branched tendrils. This is the third species of Vicia to be reported as adventive in Texas within the past eight years. Neill (1999) reported the occurrence of V. lutea in Madison Co., while V. grandiflora was reported in Fannin Co. by Singhurst et al. (2002). Jones et al. (1997) cited 12 species and subspecies of the genus in Texas, a total that does not include the three species reported as new since 1999. Only six of the now known 15 species or subspecies are native to the state. ACKNOWLEDGMENTS We are grateful to Tom Wendt of the University of Texas Plant Resources Center (TEX-LL) for his assistance and access to the herbarium. Gina Gollub, a herbarium worker at BAYLU, helped with preparation of the manuscript. Amanda Neill and an anonymous reviewer are thanked for their constructive reviews. REFERENCES BUCHANAN, A.M. 2005. A census of the vascular plants of Tasmania 4"" ed. Tasmania Herb. Occas. Pub. 7, Tasmanian Museum and Art Gallery, Hobart. CHeFFINGS, C.M. and L. FAnRELL (eds). 2005. Species status No. 7. The vascular plant red data list for Great Britain. http;//www.jncc.gov.uk. CorreLL, D.S. and M.C. JoHNston. 1970. Manual of the vascular plants of Texas. Texas Research Foundation, Renner. HATCH, S. L., K. N. GHANDI, and L.E. Brown. 1990. Checklist of the vascular plants of Texas. Texas Agric. Exp. Sta. Publ. MP-1655. Texas A&M University, College Station. HERMANN, F. J. 1960. Vetches of the United States— native, naturalized, and cultivated. Agriculture Handbook 168, U.S.D.A. Washington, DC. HNATIUK, R.J. 1990. Census of the Australian vascular plants, Bureau of the Flora and Fauna, Canberra. Isiev, D. 1990. Vascular flora of southeastern United States. vol. 3, pt. 2. Leguminosae (Fabaceae). University of North Carolina Press, Chapel Hill. Johnston, M.C. 1990. The vascular plants of Texas: a list, up-dating the manual of the vascular plants of Texas. Published by the author, Austin. Jones, S.D., J.K. Wirre, and PM. Montcomery. 1997. Vascular plants of Texas: a comprehensive checklist including synonymy, bibliography, and index. University of Texas Press, Austin. KATSUYAMA, T.A., A. SASAKI, and M. Icanr. 2001. Vicia lathyroides L. (Leguminosae), naturalized in Japan. J. Jap. Bot. 76:173-174. LiLLELEHT, V. 2001-2002. Red data book of Estonia. Comision [sic] for Nature Conservation of the Estonia Academy of Science. http://zbi.ee/punane/English/index.html. NeLL A.K. 1999. Vicia lutea (Fabaceae) new to Texas. Sida 18:1265-1266. SINGHURST, J.R., M. Wurre, and W.C. Homes. 2002. Noteworthy collections: Texas. Three species new for Texas. Cas- tanea 67:213-216. THE NATURE Conservancy. 2003. The west Gulf coastal plain ecoregional conservation plan. West Gulf Coastal Plain Ecoregional Planning Team, San Antonio, Texas. TURNER, B.L., H. NicHoLs, G. Denny, and O. Doron. 2003. Atlas of the vascular plants of Texas. Vol. 1. Sida Bot. Misc. 24. Botanical Research Institute of Texas, Fort Worth. USDA, NRCS. 2007. The PLANTS database, National Plant Data Center, Baton Rouge, LA. http://plants.usda. gov/plants. Wess, C.J. 1980. Checklist of dicotyledons naturalized in New Zealand 5. Leguminosae. New Zealand J. Bot. 18:463-472. Wotr-Murery, S. 2005-2006. Priority species in northern Ireland. http://ww g.UK/priority/splist.asp. NOTES ON THE FLORA OF TEXAS WITH ADDITIONS AND OTHER SIGNIFICANT RECORDS. IH Larry E. Brown Eric L. Keith Spring Branch Science Center Herbarium Raven Envi | Services Inc. 8856 Westview Drive PO. Box 6482 Houston, Texas 77055, U.S.A. Huntsville, Texas. U.S.A. larry- theplantmanaatt. net David J. Rosen Joe Liggio U. S. Fish & Wildlife Service 623 Woodland 17625 El Camino Real, Ste. 211 Houston, Texas 77009, U.S.A. Houston, Texas 77058, U.S.A. ABSTRACT The following taxa are documented as new to Texas: Cosmos bipinnatus, Spl icola triloba, Lepidi Lechea minor, Zephyranthes dis Du Pavonia e i uin. minor, e a ipd which is i o reported new to PME) nic new collec- T Ee Lactuca aaa: Verbesina ein os a ightii, Pseudognaphali A Myag foliatum, Cl 1 tifoli m PE ad ; o e (9 J , Dalea purpurea, Lacl Itifl Securigera varia, Hamamelis a: oo o u vernalis, Hed inoid pitan dd Abutilion hul Calylophus berlandieri subsp. pinifolius, Orobanche ludovi- E 1 Willkommia texana, Phlox cuspidata, R trilobus, Mitracarpus hirtus, Oldenlandia ciana, Plantago rugelii, Rottl corymbosa, and Penstemon digitalis. The status of th Wissadula hernandioides for a Texas mallow is also discussed. Taxa are listed alphabetically with comments, annotations, and citation of voucher specimens. RESUMEN Los ni ee taxa se documentan como nuevos Lad Texas: Cosmos UT gneticola triloba, Lepidium campestre, Lechea minor, Pavonia hastata, O bié i Arkansas. Se aportan Es o > t r r ] 1 a 1 pt A 1 A Td C: E $n dr Ts T ale x s T E o x Lactuca graminifolia, Verbesina pce. d ightii, Pseudognaphalium luteoalbum, Myagrum perfol Cl lla angustifol t bilis, Scl il Dalea purpurea, Lach Itifl Securigera varia, Hamamelis E oo Oo E E de o 2 vernalis, Hed inoid Hyptis mutabilis oe lineare, Abutilion hul Calylophus berlandieri subsp. pinifolius, Orobanche ludo- viciana, Plantago rugelii, Rottboellia cochinchinensis, Will] Phlox cuspidata, R lus trilobus, Mi hi Oldenlandia E r Penstemon E El estatus del nombre Wissadula hernandioides de una malva de Texas se ete también. Los taxa se listan al ntarios, anotaciones, y citas de pliegos testigo. INTRODUCTION Floristic work in the Houston, Harris Co. area and other east Texas counties, coupled with plants sent to the SBSC herbarium for identification or deposit, has revealed some plants new to Texas, including one new to Texas and Arkansas, as well as significant new records for others in Texas. Herbarium acronyms are those present in Holmgren et al. (1990). Unless otherwise indicated, all cited collections are in the Spring Branch Science Center Herbarium (SBSC), Houston, Texas. AMARANTHACEAE Alternanthera sessilis (L.) R. Br. ex DC.—Brown and Marcus (1998) reported this weed from Liberty County. It was mapped only in this county by Turner et al. (2003). The following is a recent collection from Brazoria County. Voucher: Brazoria Co.: San pce NWR, Dance Bayou Tract, E of FM Sanat S of its intersection with CO RD 743 at UTM 15 229605E 3224571N, infrequent her! g edges of Dance Bayou, SE of pipeli g, 15 May 2002, D.J. Rosen 2106 and S.D. Jones. J. Bot. Res. Inst. Texas 1(2): 1255 — 1264. 2007 1256 Journal of the Botanical R h Institute of Texas 1(2) ASTERACEAE Ambrosia bidentata Michx.—Turner et al. (2003) mapped most collections in the northern tier of counties south of the Red River, but they also mapped a collection from a disjunct population in Sabine County. Our Newton County collection adds another county record for this isolated population in far East Texas. Voucher: Newton Co.: f t weed along Hwy 63 between Burkeville and Sabine River, 20 Aug 1988, Brown 13014 with Lynn Lowrey. q Cosmos bipinnatus Cav.—Turner et al. (2003) did not map this species in Texas. However, Kiger (2006) mapped it in the state. We provide two Texas records. Plants of this species rarely persist for more than one growing season but are often re-seeded for roadside beautification. Vouchers: Harris Co.: planted on bare soil along the new Beltway 8 bridge over Lockwood Drive W of Lake Houston, 29 Apr 1990, Brown 14315. Polk Co.: roadside on W side of Hwy 146 at intersection with FM 943, 23 Jun 2005, E. Keith 848. Croptilon rigidifolium (E.B. Smith) E.B. Smith—In Turner et al. (2003) the eastern distribution margin for this scratch daisy extends from Leon County south to Wharton County. The range of this species extends farther east as attested to by collections from Harris, Liberty, and Hardin counties Vouchers: Hardin Co.: Turkey Creek Unit of the Big Thicket Biological Preserve dy soil SE of th n Creek/ Village Creek inter- section and N of Hwy 420, UTM zone 15, 369765E, 3370580N (NAD 27), 28 Jun 2006, Desco TC v on ew s.n. Harris Co.: sandy soil at the corner of Lutheran School Road and Fred J. Petrich Pin in NW Harris Co., W of Tomball, 6 m 1984, C.D. Peterson 717. Liberty Co.: at edge of the Trinity River NWR nea g ] which is E of C.R. 2188 and N of C.R. 2252, 13 Oct 1996, Brown 19734. Eupatorium semiserratum DC— Turner et al. (2003), Siripum and Schilling (2006), and Jones et al. (1997) did not report this species in Texas; however, Kartesz (1999), Cronquist (1980), and Gleason and Cronquist (1991) did. Correll and Johnston (1970) listed the species as a synonym of E. glaucescens. Siripum and Schil- ling (2006) considered E. glaucescens a separate species under the name E. linearifolium Walter. There are a total of 25 specimens at SBSC and the following selection support its presence in the state. Vouchers: Harris Co.: CRINES on n soil iS Grant Rd, ca 0.3 mi E M dE pr Kitzman pa N of CR 28 Oct 1984, ak forrest in Ei Softh 28 Sep, Brown 8090. Liberty Co.: Damuth Sanctuary Been Hwy 1010 and Tarkington S of Cleveland, along power line right of way, 30?18'03"N; 95°02'48"W, 6 Aug 2001, Brown 26087; Menard Creek Unit of the Big Thicket Biological Preserve, in a pine-oak forest along Hwy 2142 E of Hwy 146, between Rye and Hwy 146 bridge over Menard Creek, 30°28'05"N; 94°45'33"W, 14 Sep 2003, Brown 28800. Chambers Co.: ACA Mg d me S en a 10 ug N oe is vile of Anahuac, 29°50'17"N; 95°38'57"W, 16 Aug 2000, Brown 24649. Hardin T in Village Creek State Park E of Hwy 96 in Lumberton, 29 Aug 2000, Joe Liggio Eb s.n. Austin C 1l lati ie si ice rd of Interstate 10 at the Beckendorff exit, 21 Jul 1987, Brown 11369. Tyler Co.: Turkey Creek Unit of the Big Thicket Bobda. P tedge of id bog off of C.R., S of Hwy 1943, E of Warren, 8 Oct 1983, Brown 6678. Newton Co.: in low area along hwy 12 i i with Hwy 87, 18 Aug 1985, Brown 9363. Hymenoxys texana (J.M. Coult. & Rose) Cockerell —Correll & Johnston (1970) reported this now federally endangered taxon from near Hockley and Houston in Harris County, based upon Thurow collections from 1889 and 1890. They considered it probably extinct for there were no known collections after 1900. It was not collected again until 1981 when James Kessler discovered some small populations north of the village of Cypress in Harris County (Mahler 1983). Up until about 1992, Prairie Dawn was known only from the northwest portion of Houston and in the area of Harris County northwest of Houston, with one population from the Fort Bend County section of the Barker Reservoir. In 1993, populations were found in the Piney- woods to the west of Lake Houston. In 1999, populations were found on the south side of Houston, centered around S. Post Oak Rd, which is S of Hwy 90a. In 2002, a population was found in a prairie adjacent to the Ellington Field airport off Hwy 3 south of Houston. The most significant and unexpected discovery was made in 2003 when a population was found in Trinity County far north of Harris and Fort Bend counties. Vouchers: Fort Bend Co.: in extreme W edge of Barker Reservoir at end of a gravel road which exits Mason Road at first bridge N of intersection with pud 1093, 7 - 1989, Brown 13432. Harris Co.: populations on pimple M on W ae a Lake Houston and N of Beltw Lake Houston Parkway, 15 Apr 1993, Brain Rowe s.n.; al way W of S. Post Oak and S ps Lotus Street near 29° 37'55”N, 95?28'26"W, 17 Mar 1999, Brown 22893; disturbed i prairie E of Ellington Field at UTM Brown et al., T. fl lditi 1 sianificant | 1257 d 15, 292963E, 3278227N, 10 Apr 2002, DJ. Rosen 1938. Trinity C li River, 31?18'45"N, 95?18'46"W, 23 Apr 2003, E. Keith 553. Boggy Slough Hunting Club, ca 2 mi W of Neches [e Lactuca graminifolia L —Correll € Johnston (1970) reported this lettuce from elevated portions of the Davis and Guadalupe mountains of the Trans-Pecos region of Texas. Turner et al. (2003) mapped it only in Culberson, Jeff Davis, and Presidio counties. Kartesz (1999) mapped it in the southern states from North Carolina westward into Arizona and Colorado. Thomas and Allen (1996) mapped it in five Louisiana par- ishes, two in the central portions of the state and three in the parishes east of the Mississippi River. We report a population, until now not reported in east Texas, on deep sands from the Pineywoods of Angelina County. Voucher: Angelina Co.: frequent rather tall herb on xeric sands under longleaf pines off of F.R. 313a just N of F.R. 313 in the Boykin Springs area of the Angelina National Forest, 21 Aug 1995, Brown 18122. Pseudognaphalium luteoalbum (L.) Hillard & Burtt—Nesom (2001) first reported this Eurasian species in Texas from El Paso, Kimble, and Parker counties. Turner et al. (2003) mapped it mostly in the Trans-Pecos and Edwards Plateau regions of Texas. We report a collection from Liberty County far east of those mapped, and for now, it is the only record east of the Edwards Plateau. Guy Nesom identified our specimen. T CO J; demus x] RT esce Tarkington High School in the village of Tarkington which is off of Hwy 321 southeast of Cleveland, 30°18! 57°N, 94°57'33”W, 12 Mar 2006, Brown 31062 (SBSC, BRIT). Sphagneticola trilobata (L.) Pruski—Creeping Oxeye is a cultivated species in the southern states and is probably native to tropical America. (Strother 2006). Kartesz (1999) indicated it had escaped in Louisiana and Florida. Strother (loc. cit.) states the Louisiana reports have not been verified. Thomas and Allen (1996) indicated the Louisiana report is the LSU collection, Urbatsch 5155, from Baton Rouge Parish. The following collection is the first report of an escaped Texas population. Voucher: Cherokee Co.: spreading up slope into the woods, 5 m N of tee box of 3" hole at Rusk Golf Course, 25 Jun 2005, E. Keith Verbesina helianthoides Michx.—This species has yellow ray flower, a winged stem, and with most leaves alternate. Turner et al. (2003) mapped it in northeastern Texas from the Red River border counties south to Anderson and Shelby counties. We report specimens from a fairly large population in Liberty County far south of those LN 1 Lil 1 lil | C.R. 2142 which exits E off of Hwy 146 N of Rye, 20 Oct 1996, Brown 19765; same site, 7 Oct 1995, p 18775; plants along C.R. 2141c nears it end on the west side a. Menard de 30?27'46"N and 94?45' 02"W, Brown 242 37a; woods and we s al ng H 146, oHm. V Hwy 105in Moss Bluff, 12 Aug pe Ward s.n. Plants of this species are somewhat frequent at edges of sandy roadsides centered largely from the Menard Creek westward to Hwy 146 BRASSIACEAE Myagrum perfoliatum L.— Rollins (1993) indicated this introduced species to be uncommon and known only from a few localities in the United States and Canada. He also wrote that no recent collections have been seen. Kartesz (1999) mapped it only in Texas, Oklahoma, and the Quebec Province of Canada. Turner et al. 2003) mapped it in Delta and Hill counties. We report collections from two additional counties. Vouchers: Walker Co.: collected along FM 1791 SW of Huntsville, 2 Apr 2001, E. Keith s.n.; Rockwall Co.: Old Millwood Road, ca 0.5 mi from jct. with Marty Circle, roadsid k, loamy clay with heavy leaf liter; with Geranium, Stellaria, and grasses, 23 Mar 2002, July Ratteree 262 (TAMU). Lepidium campestre (L.) R. Br.—Kartesz (1999) mapped this introduced species in the adjacent states of Louisiana, Arkansas, Oklahoma, and New Mexico. However, none of the collections of it from the nearby states of Louisiana and Arkansas are from counties near Texas. In Arkansas, Smith (1988) mapped it in 1258 Journal of the Botanical R h Institute of Texas 1(2) the northern counties near the Oklahoma border and in Sevier County, a southwest county, which is the nearest mapped location to Texas. Thomas and Allen (1996) mapped it only from Winn Parish in the north central portion of Louisiana. The following is the first record for Texas, and it is distant from collections in the two adjacent states. Voucher: Montgomery Co.: in parking lot for the Lone Star Hiking Trail off of Bethel Rd, S of Hwy 149, N of Montgomery in the Sam Houston National Forest, 30°30'40"N, 95?45'25"W, 27 Apr 2003, Brown 27994. CLEOMACEAE (Capparaceae) Cleomella angustifolia Torr.—Kartesz (1999) mapped this taxon in Texas, Oklahoma, Kansas, Nebraska, and Colorado. Turner et al. (2003) mapped it in 11 Texas counties, with six of them in the northwest tier near the Oklahoma border. The five other county records are widely scattered from Dawson County east to Brazoria and Harris counties. Our Anderson County record documents it from the Pineywoods region of eastern Texas which is apparently the most eastern station in the state for this largely western species. The endemic Hymenoxys texana is associated with it at its only Harris County station The Anderson County site should also be examined for the presence of H. texana. Voucher: And C li irie, ca 500 m N of Hwy 84 and 400 m W of the Neches River, 28 Jul 2003, E. Keith 584 & S. Cook. in E CISTACEAE Lechea minor L.—Kartesz (1999) mapped this species in most of the eastern states from New Hampshire, New York, and Wisconsin south to Florida and west to Louisiana. Thomas and Allen (1996) mapped itin St. Bernard and St. Tammany parishes in eastern Louisiana and in Calcasieu Parish adjacent to Texas. At SBSC there are three different collections of it from a single location south of Starks in Calcasieu Parish, which is just a few miles east of the Texas border. Our specimen is the first for Texas. The following key will serve to separate the five species of Lechea now known in east Texas. 1. Stern hairs spreadin L. mucronata 1. Stem hairs e Leaves mostly 0.5-1.5 mm wide, 10 or more times longer than wide. 3. Fruit pedicels 3 mm long, secund and bending nid from stem L. san-sabeana 3. Fruit pedicels ip than 3 mm long, not secund, ere L. tenuifolia 2. "d mostly 1—4 mm wide, 5 or less times pom wide. . Pubescence da abaxial leaf surface extending across the surface; at least some leaves opposite and whorled on the same plan L. minor 4. Pubescence of abaxial leaf surface restricted to mid-vein and margin; leaves not both opposite and whorled on the same plant . pulchella Voucher: Hardin Co.: collected at edge of Little Rock Rd in the Lance Rosier Unit of the Big Thicket Biological Preserve, west of Hwy 326 and S of Kountze, 30?18'31"N, 94°22'03"W, 13 Aug 2004, Brown 30003 with Shawn Harper. CYPERACEAE Cyperus aggregatus (Willd.) Endl.—Diggs et al. (2006) reported this taxon from Newton County based upon the voucher below. Turner et al. (2003) did not include this species in Texas under this name nor under any of the synonyms found in Kartesz (1999). This collection is apparently the first in east Texas for Cor- rell and Johnston (1970) listed it, as C. huarmensis (H.B.K.) M.C. Johnst., only for Aransas and San Patricio counties. Diggs et al. (2006) es additional records from Bee and Refugio counties. V N C dsid N side of Hwy 255, ca. 0.25 mi W of intersection with Hwy 87, 24 Sep 2003, E. Keith 603 Cyperus eragrostis Lam.—Turner et al. (2003) mapped this sedge only in Harris County. We provide records for Brazoria, Galveston, and Houston counties. All vouchers are at SBSC with a duplicate of the Houston County specimen at TEX and a duplicate of the Galveston County collection at VSC. Vouchers: Brazoria Co.: in a drainage ditch along C.R. 176c, SW of Alvin, Jun 2005, L. James s.n. Galveston Co.: S of Hwy 96, 0.5 mi W of its intersection with Hwy 146 in League City, 10 May 2002, D.J. Rosen 2068. Houston Co.: roadside W of FM 230, ca 0.5 mi N of the town of Weldon and 3.5 mi S of the intersection with FM 2915, 23 Jun 2005, D.J. Rosen 3433 Brown et al., T. fl lditi 1 sianificant | 1259 Rhynchospora debilis Gale—Turner et al. (2003) mapped this species in Hardin and Chambers counties. Diggs et al. (2006) included it for Hardin County based upon the above publication. The Hardin County voucher, MacRoberts & MacRoberts 3851, was collected in the Lance Rosier Unit of the Big Thicket Biological Preserve. The first author borrowed the TEX voucher in order to examine it for a checklist of the vascular plants of this unit (Brown et al. 2006). The voucher is actually R. filifolia A. Gray. However, a 2007 collection of itin Tyler County, from the Beech Creek Unit of the Big Thicket Biological Preserve, has now been made. Thus it is now known from Chambers and Tyler counties. | haml ll delicate plant | dal ] f East Bay and below the Abshire-Blackstone house in the Candy Abshire Wildlife M n Smith Point south of Anahuac, 7 May 1993, Brown 16932; same site, 27 May 1996, Brown 19138 with J. Liggio. Tyler Co.: Olsen along the Beechwoods Trail, off of the picnic area on E side of Hwy 2992, SE of Woodville, near 30?43'10"N, 94°13'38"W, 19 May 2007, Brown 32131 with Shawn Harper (TAES). Scleria verticillata Muhl. ex Willd.—Turner et al. (2003) mapped this nut-rush in six counties in the Ed- wards Plateau as well as Dallas and Limestone counties in north central Texas. Diggs et al (2006) reported it also in Polk, Wise, and Liberty counties. We report vouchers from coastal populations in Chambers and Galveston counties. Voucl Chaml C dy soil along Hwy 562 ca 2 mi W of intersection with Hwy 1985 S of the village of Anahuac, 29°39'38"N, 94?37'38"W, 3 Aug 2000, Brown 24562. Calves n Co ant wet prairie, S of Hwy 96, 5 mi W of its intersection with Hwy. 146 in League City. UTM 15, 720852E, 3354329N, 10 re 2006, 5 DJ. Rosen 3914 with R. Carter (SBSC, TEX). FABACEAE Dalea purpurea Vent.—Turner et al. (2003) did not map this species in the Pineywoods of East Texas. We have records from Tyler and Newton counties. The nearest mapped locations to the east Texas counties are from the Louisiana parishes Calcasieu and Vernon, which border Texas (Thomas @ Allen 1998). D. purpurea is similar to the two varieties of D. compacta Sprengel. Both species occur on alkaline outcrops but are distinguished by the larger flower spikes of D. compacta combined with the calyx trichomes in lines that are separated by bald areas whereas in D. purpurea the flower spikes are smaller and the trichomes cover the entire calyx tube surface. Voucher: Tyler Co.: collected on an alkaline barrens area off of Hwy R255 W of the Neches River and E of intersection with Hwy. 69, 30°59'03"N, 94°17'12"W, 8 Jun 2003, Brown 28282. Newton Co.: calcareous prairie on S side of Hwy 60 at eastern city limit sign of Burkeville, 2 Jul 2001, Eric Keith s.n. (BRIT). Lackeya multiflora (Torr. & A. Gray) Fortunata, L.P. Queiroz & G.P. Lewis—One synonym for this spe- cies is Dioclea multiflora (Torr. & A. Gray) Fortunata, L.P.Queiroz & G.P. Lewis. Other synonyms are avail- able on the International Plant Name Index web site (http://;www.ipni.org/index.html). Turner et al (2003) mapped this vine in 10 counties, all except one in the Pineywoods of east Texas. Our new record is from Jefferson County in the Beaumont Unit of the Big Thicket Biological Preserve. The Liberty County record should be eliminated because the first author has annotated Brown 17822, the voucher at TEX and SBSC for this record, to Phaseolus polystachios (L.) Britton, Sterns, @ Poggenb based upon the uncinate trichomes on the abaxial leaflet surface of this sterile specimen. Only non-uncinate trichomes are on the abaxial leaflet surface of L. multiflora Voucher: Jeff C llected i ine/hard 1 forest along Pine Island Bayou, 23 Apr 2006, Ray Jordan s.n (TAES). lz Phaseolus polystachios (L.) Britton, Stern, € Poggenb.—Turner et al. (2003) mapped this vine in Harrison and Nacogdoches counties. Our new records are for Liberty and Newton counties. Vouchers: Liberty Co.: Davis Hill salt d the Davis Hill State Natural Area N of mE 105 and E of Cleveland, Davis Hill 7.5 Quad, vine under pines in a deep ravine, 6 May 1994, Brown 17822, (TEX, SBSC). Newton C ndy soil on deri land cleared of trees. S of Hwy R255 abus E of the Jasper Co. line, 5 Oct 2003, B 28877 dist ] g g 3 ca 600 m W of Weeks Chapel Rd and 200 m S of R255, 24 Sep 2003, Keith 605 e Robinson. Securigera varia (L.) Lassen (Coronilla varia L)— Crown vetch is planted for erosion control on newly con- 1260 Journal of the Botanical R h Institute of Texas 1(2) structed road banks. It persists on roadsides, especially in the northern states. Turner et al. (2003) mapped itin five widely scattered counties, with Walker County the only east Texas county mapped. Our voucher adds Cherokee County to its East Texas distribution. Voucher: Cherokee C pen ] d clay soil near tee box of 6'^ hole at Rusk Golf Course, 25 Jun 2005, E. Keith 849. HAMAMELIDACEAE Hamamelis vernalis Sargent—Neither Meyer (1997) nor Jones et al. (1997) reported this shrub in Texas. Correll and Johnston (1970) reported it in east Texas. Kartesz (1990) mapped it in Texas, Oklahoma, Arkan- sas, and Missouri. Turner et al.(2003) mapped it in Hardin, Tyler, Jasper, and Newton counties in extreme east Texas. The following Newton County vouchers at SBSC support its presence in Texas. Vouchers: Newton Co.: along Hwy 87 ca 6 mi N of Burkeville, ca 2 mi S of Hwy 255, 11 Dec 1993, Lynn Lowrey s.n.; ca 5 mi NW of Wiergate, N of Indian Hill Community, within 200 yards of Cow Creek and within 300 yards of Texas Stewartia site, 12 Feb 1989, Lynn Lowrey, W. Flaming, Osa Hall, and Peter Loos s.n.; along Hwy 87, 7 mi N of Burkeville, 24 Mar 1991, Lynn Lowrey s.n.; near vicinity of villag > of Newton, 17 Mar 2001, Brown 25022; 6 mi W of Burkeville on Hwy 87, 15 Jan 1994, Lynn Lowrey and Scooter Cheatham s.n. All of the above specimens were in anthesis, except for Lynn Lowrey 11 Dec 1993, which was in bud. The red flowers were much smaller than the yellow flowers of H. virginiana. The specimen data indicates the anthesis period in Texas extends from February into March. LAMIACEAE Hedeoma acinoides Scheele—This mint is present only in Texas and Mexico (Correll & Johnston 1970). Turner et al. (2003) mapped it largely in the Edwards Plateau region with counties extending north and south from the eastern portion of the Plateau. We report a collection of it from the Pineywoods in San Jacinto County, far east of those mapped. Voucher: San Jacinto Co.: weedy plant along grassy verge of Hwy 2025 at ent d to the Double Lake Recreation area of the Sam Houston National Forest, S of Coldsprings, 29 Apr 1995, Brown 18566. Hyptis mutabilis Briq.—Brown and Elsik (2002) reported this introduced weed from Liberty County new to Texas. Our recent collection is from another east Texas county. o her: Hardin C llected al S Road in the Turkey Creek Unit of the Big Thicket Biological Preserve, 2005, P. Rolling s.n. LILIACEAE Zephyranthes grandiflora Lindl.—These are the first Texas collections of escaped individuals of this cultivated species that Wunderlin (1998) reported native to Mexico and the West Indies. Vouchers: Hardin Co.: near drainage pipe on the E side of Hwy 69, 5.5 mi S of Kountze and 300 ft S of Hardin Co. Fire Station #3, 8 Jul 2005, E. Keith 853. Harris Co.: along an old drainage ditch in Jesse Jones County Park, N of Hwy 1960 and W of Humble, in bot- tomland of Spring Creek, Aug 2005 P. Rolling s.n. LYTHRACEAE Lythrum lineare L.—Brown and Elsik (2002) reported this species new to Texas from two stations in Chambers County. Recently a large population was discovered in Harris County. The Harris County popu- lation seems to be a recent arrival since the first author, who has surveyed this area off and on from about 1980, had not previously observed it there. Voucher: Harris Co.: collected on saline mud at edge of Taylor Bayou on W side of Hwy 146 just N of intersection with Port Rd, Key Map 580 W, 24 Nov 2004, Brown 30267. MALVACEAE Abutilon hulseanum (Torr. & A. Gray) Torr. ex A. Gray—A. Richardson (1995) reported this mallow from Cameron County and indicated the only United States records were in this county. Also, Turner et al. (2003) mapped it only in Cameron County. We report a collection of three duplicate specimens from Calhoun County which is far north of Cameron County. Brown et al iT {l P Rr Ea? H 4 J g 1261 Voucher: Calhoun Co.: collected on high ground on the Victoria Barge Canal Levee, 26 Mar 1992, D. Mabie s.n. Pavonia hastata Cav.—Fryxell (1988) reported this mallow from Argentina, Paraguay, Uruguay, and Brazil in South America as well as Mexico and Australia. In the U.S., it has escaped in Georgia and Florida, and now Texas. Voucher: Harris Co.: growing on weedy bank of S. Mayde Creek at F stem tips erect, Key Map 446a, 7 Mar 2007, Brown 31921. Road bridge., plant almost appearing woody, it was trailing with Wissadula hernandioides (UHer.) Garcke versus Wissadula amplissima (L.) R.E. Fries—Turner et al. (2003) mapped W. hernandioides in Cameron, Hidalgo, and Kennedy counties and they also mapped W. periplocifolia (L.) K. Presl ex Thwaites in south Texas. In Richardson (1995), W. amplissima (L.) R.E. Fries and W. periplocifolia are found but not W. hernandioides. The pertinent synonymy is provided by Fryxell (1988). The basionym of W. amplissima is Sida amplissima L.,1753, and the basionym of W. hernandioides is Sida hernandioides UHer., 1789. However, the new combination W. amplissima is from1908 and the new combination W. hernandioides is from1866. The priority of the basionym should prevail and not the priority of the new combination. Thus W. amplissima should be the correct name which is also the name Fryxell (1988) used for this species. ONAGRACEAE Calylophus berlandieri Spach subsp. berlandieri— Our specimen station is farther east than any of those mapped in Turner et al. 2003). Voucher: Newton Co.: N side of FM 363, 2.5 road mi E of Sand Ridge Cemetery Road, E of Bleakwood, 30.7288°N and 93.7197°W, 15 Apr 2005, E. Keith 802. OROBANCHACEAE Orobanche ludoviciana Nutt subsp. multiflora (Nutt.) Collins—Turner et al. (2003) mapped this small species no further east than Dallas County. Our Galveston County voucher is from an isolated coastal population. Cal ae f. al d A choll } getation line, 2 mi W of Rollover Pass on Bolivar Peninsula, 23 Apr 2003, B. Hollingsworth s.n Orobanche minor Sm.—Our voucher is the first collection of this federal noxious weed in Texas. Kartesz (1999) mapped it no closer to Texas than Florida. The species has reappeared at this location for the past three years. The following key will serve to separate the four Orobanche species now known present in east Texas, of which two are recent introductions. 1. Flowers solitary at tip of a pedicel that is much longer than the calyx 1. Flowers in d spikes or spike-like racemes; pedicels absent or shorter than calyx. 2. Calyx lobes 5 (do not confuse the calyx lobes with the bractlets which are positioned between the bract and the calyx tube) 2. Calyx lobes 4. 3. Calyx divided almost to base into 2 segments that are deeply bifid at apex, thus 4 ultimate unequal lobes O. uniflora O. ludoviciana O. minor O. ramosa 3. Calyx divided into 4 more or less equal lobes Vouchers: Harris Co.: downtown Houston on grassy bank on N side of Interstate 10 at the west bound entrance of Watson Street, Harris Co., Key Map 493 F, 27 March 2004, Brown 29087; same site, 25 Mar 2004, J. Liggio s.n. PLANTAGINACEAE Plantago rugelii Dcne.— Turner et al. (2003) mapped this weedy but native species in Clay, Dallas, Grayson, and Lamar counties. Lipscomb (1978) reported it also present in Titus County. A key to separate this spe- cies from the weedy but introduced and more common P. major is found in Lipscomb (loc.cit.). We report a recent collection from Harris County far south of those mapped. 1262 Journal of the Botanical R h Institute of Texas 1(2) Voucher: Harris Co.: growing in picnic area of Jesse Jones County Park to the N of Hwy 1960, west of Humble, 26 Sep 2004, Brown 30736 POACEAE Paspalum wrightii A.S. Hitchc. & Chase—Allen and Hall (2003) first reported this grass for Texas based upon historic collections from Calhoun, Jackson, and Matagorda counties cited in the protologue of P. texanum Swallen (J. Wipff, pers. comm., 2005). Our vouchers are from rediscovered populations in Jackson and Matagorda counties and new populations in Wharton County. Zuloaga et al. (2003) pointed out that P. texanum is a synonym of this species and not of P. plicatulum Michx., as found in Gould (1975) and Kartesz 9) Vouchers: Jackson Co.: S of Hwy. 616, 5.7 mi E of DEN 172 in LaWard, 24 Jul 2004, DJ. Rosen 3024 and R.M. Combs (BRCH, BRIT, SAT, TEX, UTEP, VDB). Wharton Co. perennial in hydric roadside ditch between Hwy 102 and railroad track, ca 11.3 mi S of the town of Eagle ide 30 jd 2004, D. M 2996 (BRCH, BRIT, SBSC, TEX, US). Matagorda Co.: o 2431, 8.4 km E of its intersection with Hwy 71, between the towns of Midfield and Markham, 11 Jul 2007, D. Rosen 4320 (BRIT, MO, SBSC, TAES, TEX, US) Rottboellia cochinchinensis (Lour.) Clayton—Wipff and Rector (1993) first reported this noxious weed new to Texas from Jefferson County. Since then plants have been collected in three other counties. Vouchers: Harris Co.: Armand Bayou Nature Center S of the intersection of Bay Area Blvd and Red Bluff Road, disturbed field of the bison enclosure, apparently escaped from hay, 01 Oct 2003, D.J. Rosen 2639. Matagorda Co: North roadside of FM 521, ca 3.6 km E of intersection with Hwy. í 60 in Wadsworth., N28°50'20.5", W95°53'59.3", 02 Nov. 2006, D.J. Rosen 3980 (BRIT, TAES, TEX, SBSC, US). Wharton Co.: on and E of Hwy 71 at its intersection with FM 1160, 1.9 km S of CR 474, 17 km N of the town of El Campo, 09 Jul 2007 D. Rosen s.n. (BRIT, TAES, TEX). Willkommia texana A.S. Hitchc. var. texana—Wipff (2003) reported this grass from Texas and Okla- homa. Turner et al. (2003) mapped it in most of the coastal counties from Chambers south to Cameron. The type locality is the city of Ennis, which is probably the basis of their record for Ellis County. We report two counties not mapped by Turner et al. (2003). At SBSC, most of the collections of this species are from Harris County where the endangered Hymenoxys texana is often present on the same sites. Vouchers: Anderson Co.: open saline prairie, ca 500 m N of ies 84 and 400 m W of the Neches River. 21 Sep 2003, E. Keith 601. Matagorda Co.: Sargent unit of the San Bernard NW R S of Sargent, plants in rral, 10 Jul 2000, F. Hannnah s. POLEMONIACEAE Phlox cuspidata Scheele—The following collections are for three counties not mapped in Turner et al. Vouchers: Brazoria Co.: Nash Ranch, hay meadow W of C.R. 25, ca 8.7 mi N of it intersection with Hwy 35 in West Columbia, 5 Apr 2004, DJ. Rosen 2787 (TEX). Newton Co.: from N side of FM 363, 2.5 mi E of Sand Ridge Cemetery Rd, E of Bleakwood, 30.7288°N 93.7197°W, 15 Apr 2005, E. Keith 799. San Jacinto Co.: on grassy verge of Hwy 2025, ca 1 mi N of the Liberty County border, 10 Apr 2005, Brown 30413a. RANUNCULACEAE Ranunculus trilobus Desf.—Although not listed for Texas by Correll and Johnston (1970), Turner et al. (2003) mapped this European native in Harrison County. Our record is the second for the state. Voucher: Brazoria Co.: collected near a drainage swale in MacLean Park in the city of Lake Jackson, 02 Apr 2006, Ray Jordan & W. Pruess s.n. RUBIACEAE Mitracarpus hirtus (L.) DC.—Turner et al. (2003) mapped this species in Cameron County in extreme south Texas. We report two collections from the east Texas Pineywoods. Vouchers: Hardin Co.: growing on road in Village Creek State Park, 28 Sep 1995, D.A. Muschalek s.n. Liberty Co.: Menard Creek Unit of the Big Thicket Biological Preserve, off of County Road 2138 at the Liberty/Hardin County border, 28 Oct 1995, Brown 18847. Brown et al., T. fl lditi 1 sianificant | 1263 Oldenlandia corymbosa L.—Correll € Johnston (1970) reported this introduced species, as Hedyotis corymbosa, to be rare in lawns in east Texas and with a question mark as to being present in the Gulf Coast. Also reported for the state by Jones et al. (1997) and as rare in east Texas (Terrell 1990). An internet search of the specimens at TEX/LL show no collections there which is probably why is was not mapped in the state by Turner et al. (2003). The following Harris County collections at SBSC verify it for the state. Vouchers: Harris Co.: present at 6218 Doliver Street in Houston, 11 Oct 1987, W.L. McClure s.n.; collected in SW Houston at curbside of Cartagena and Landsdowne Streets, 2 Aug 2007, Betty Cawfield s.n.; collected along a sidewalk adjacent to the Half-Price book store on Westheimer Blvd in Houston, 10 Oct 2005, Brown 30791; collected at edge of the SBSC herbarium building at 8856 Westview Drive which is N of 1-10 and E of the intersection of Westview Drive with Campbell Road, 21 Aug 2007, 28 Oct 1995, Brown 32720a. SCROPHULARIACEAE Penstemon digitalis Nutt.—Turner et al. (2003) mapped this taxon in many counties clustered in the extreme northeast portion of the state. However, they mapped a collection from Walker County, which is rather isolated from the northern counties. Our collection adds another county record to this isolated population and a record for Liberty Co., which is the most southern station in Texas. Voucher: Polk Co.: growing along a eee outside of Lake Livingston State ae W of Livingston and Goodrich, 28 Apr 1995, K. Hillhouse s.n. Liberty Co.: sandy soil ty road off of Hwy 146 S of th Menard Creek. This gravel road extends E to the Menard Creek Unit of the Big Thicket Biological Preserve, May 2003, Joe Liggio s.n. VALERIANACEAE Valerianella locusta (L.) Betcke.—We report this introduced species new to Texas and Arkansas. It differs from all other Texas and Arkansas Valerianella taxa by the bluish cast to the flowers and the presence of a corky mass on the back of the fertile cell. Kartesz (1999) did not map this introduced species in Arkansas or Texas. Smith (1988) saw no non-cultivated Arkansas specimens and excluded it from the state. Smith later (1994) considered it as a possible addition. Vouchers: Bowie Co.: weedy plants on the verge of Hwy 8 just south of the bridge over the Red River into Arkansas, 33?33'27"N, 94?25'02"W, 25 Mar 2005, Brown 30341. ARKANSAS. Newton Co.: along Hwy 7 near Alum Cove, spring 2005, P. Rolling s.n ZYGOPHYLLACEAE Kallstroema parviflora Norton—This species is known mostly from the western 2/3 of Texas (Turner et al. 2003). We report a collection of this caltrop from extreme east Texas. Voucher: San Augustine Co.: collected on a Weches outcrop 1 mi E of San Augustine, 17 Jul 1998, E. Keith s.n. ACKNOWLEDGMENTS We thank Guy Nesom (BRIT) for the identification of Pseudognaphalium luteoalbum from Liberty County and we appreciate the response of Tom Wendt (TEX/LL) to our request for some specimen data at TEX. Thanks also to Monique Reed (TAMU) for her careful reading of the paper, which helped to improve it, as well as informing us ofa collection of Myagrum perfoliatum at TAMU. Publication costs were paid from a Big Thicket Association grant (AT BI) to the 1% author. REFERENCES ALLEN, C.A. and D.W. Haut. 2003. Paspalum. In: Flora of North America, vol. 25. Oxford University Press, New York, NY. Brown, L.E. and S.J. Marcus. 1998. 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Malvaceae of Mexico. Syst. Bot. Monogr. 25:1-522. GLEASON, H.A. and A. CRONQUIST. 1991. Manual of vascular plants of northeastern United States and adjacent Canada, 2"d edition. The New York Botanical Gardens, Bronx. GouLb, FW. 1975. The grasses of Texas. Texas A&M Univ. Press, College Station. HotMGREN, PK., N.H. HOLMGREN, and L.C. Barnett. 1990. Index herbariorum, 8^ ed. New York Botanical Gardens, Bronx. Jones, S.D., J.K. Wierr, and PM. Montcomery. 1997. Vascular plants of Texas. Univ. Texas Press, Austin. KARTESZ, J.T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland. 1% ed. In J.T. Kartesz and C.A. Meacham. Synthesis of the North American flora, Version 1.0. North Carolina Botanical Garden, Chapel Hill. Kicer, RW. 2006. Cosmos. In: Flora of North America, vol. 21. Oxford University Press, New York, NY. Lipscome, B. 1978. Additions to the Texas flora. Sida 7:392-394. Matter, W.F. 1983. Rediscovery of Hymenoxys texana and notes on two other Texas endemics. Sida 10:87-92. Meyer, F.G. 1997. Hamamelis. In: Flora of North America, vol. 3. Oxford University Press, New York, NY. Nesom, G. 2001. New records in Pseudognaphalium (Asteraceae: Gnaphalieae) for the United States. Sida 19:1185-1190. RICHARDSON, A. 1995. Plants of the Rio Grande Delta. University of Texas Press, Austin. RoLuNs, R.C. 1993. The Cruciferae of continental North America. Stanford Univ. Press, Stanford, CA. Siripum, J.L. and E.E. ScHILLING. 2006. Eupatorium. In: Flora of North America, vol 21. Oxford University Press, New York, NY. Smith, E.B. 1988. An atlas and annotated list of the vascular plants of Arkansas, 2" edition. University of Arkansas Press, Fayetteville. Sur, E.B. 1994. Keys to the flora of Arkansas. The University of Arkansas Press, Fayetteville. STROTHER, J.L. 2006. Sphagneticola. In: Flora of North America, vol. 21. Oxford University Press, New York, NY. TERRELL, E.E. 1990. Synopsis of Oldenlandia (Rubiaceae) in the United States. Phytologia 68:125- 133. Thomas, R.D and C.M. Auen. 1996. Atlas of the vascular flora of Louisiana, vol. Il: Dicotyledons, Acanthaceae-Eu- phorbiaceae. Louisiana Department of Wildlife & Fisheries, Baton Rouge. Thomas, R.D. and C.M. ALLEN. 1998. Atlas of the vascular flora of Louisiana, vol. Ill: Dicotyledons, Fabaceae-Zygophyl- laceae. Louisiana Department of Wildlife & Fisheries, Baton Rouge. TURNER, B.L., H. NicHoLs, G. DENNY, and O. Doron. 2003. Atlas of the vascular plants of Texas, vol. 1:Dicots, vol. 2: Ferns, gymnosperms, monocots. Sida Bot. Misc. 24. Whirre, J.K. 2003. Willkommia. In: Flora of North America, vol. 25. Oxford University Press, New York, NY. Whrr, J.K. and B.S. Rector. 1993. Rottboellia cochinchinensis (Poaceae: Andropogoneae) new to Texas. Sida 15:419-424, WUNDERLIN, R.P. 1998. Guide to the vascular plants of Florida. University Press of Florida, Gainesville. ZULOAGA, F.O., O. Morrone, G. Davipse, T.S. FILGUERIAS, P.M. PETERSON, R.J. SORENG, and E.J. Jubziewicz. 2003. Catalogue of New World grasses (Poaceae): Ill. Subfamilies Panicoideae, Aristidoideae, Arundinoideae, and Danthonioideae. Contr. U.S. Natl. Herb. 46:1-662. NOTES ON TEXAS ORCHIDS (ORCHIDACEAE) Paul Martin Brown 10896 SW 90" Terrace E Florida 34481, U.S.A. ORCHID@aol.com ABSTRACT ] 1 1 Ich additional taxa, rediscovery of species In the past ten years, 1997-2007, tl g i g the Orchidaceae in Texas. These are all reviewed here. nreciimed tirpated. and clarifi J; E E * Ab = o RESUMEN Enl E d di ñ , 1997— 2007, ha habido i bi ómi 1 ] taxa adicionales J brimi d i i ipti d id: 52217 : especies p y clarificación de esp p Orchidaceae en Texas. Tod quí. In preparation for the publication of Field Guide to the Wild Orchids of Texas in February 2008 (University Press of Florida) it was essential to review the known knowledge of the Orchidaceae of Texas. Additions to that body of work include clarification and update of older material as well as several new discoveries and recently published taxa. The review of literature prior to the publication of Liggio and Liggio (1999) revealed many changes in taxonomy and nomenclature, most of which appeared in the Liggios’ work. In the years since the Lig- gios’ publication significant new discoveries, publications, and herbarium additions have been made to the orchid flora of Texas. Some of the more significant results of this current research include the publication of several new species, based upon older names, for species occurring in Texas and the publication of one new variety. Two new color forms with types from Texas were also published for a total of ten new taxa to be added to the orchid flora of Texas. Two recently-described species are now known to be present in Texas, but not replacing the established species often confused with them. Spiranthes eatonii PM. Brown (Fig. 1), Eatons ladies’-tresses [S. floridana (Wherry) Cory emend PM. Brown, S. lacera Raf. var. gracilis (Bigelow) Luer] (Brown 1999). Spiranthes sylvatica PM. Brown, woodland ladies’-tresses [S. praecox] (Brown 20012) Two varieties are new to Texas: Triphora trianthophora var. texensis PM. Brown & R. Pike (Fig. 2), Texas three birds orchis was described asanew taxon, endemic to Texas and based upon discovery in 2005 by Pike. Plants are currently known from a very small area in Houston County (Brown & Pike 2006). Platanthera flava (L.) Lindl.) var. herbiola (R. Brown) Luer (Fig. 3), first found by Sandi Pelchat in 2002 in Sabine County was documented as a new variety for Texas (Pelchat, pers. comm.). Two species found in Texas have recently been described and are often confused with previously described species that do not occur in the United States: Deiregyne confusa Garay, (Fig. 4) rimrock ladies '-tresses [misapplied names: Spiranthes durangensis Ames & Schweinf.; D. durangensis (Ames & Schweinf.) Garay] (Garay 1980). Schiedeella arizonica PM. Brown, Indian braids; Arizona red-spot [misapplied names: Spiranthes parasitica A. Rich. & Galeotti, S. fauci-sanguinea Dod; Schiedeella parasitica (A. Rich. & Galeotti) Schlechter, S fauci-sanguinea (Dod) Burns-Balogh] (Brown & Coleman 2000). Two taxa previously treated at the varietal level are now recognized at the species level: Platanthera conspicua (Nash) PM. Brown, southern white fringed orchis [synonym: Platanthera blephariglottis J. Bot. Res. Inst. Texas 1(2): 1265 — 1268. 2007 1266 Journal of the Botanical R h Institute of Texas 1(2) Fic. 1. Spiranthes eatonii photo by P.M. Brown Fic > L ¿A E L L " e L | y R. Pike Fic. 3. Platanthera fl herbiola photo by C. Pelchat Fic. 4. Deiregyne confusa photo by C. Pelchat Fig. 5. Dich hus cinnabarinus fi photo by J. Smith Fic. 6. Hexalectris grandiflora fi | Iba photo by J. Smith var. conspicua (Nash) Luer] (Brown 2002). Spiranthes floridana Wherry emend P.M. Brown, Florida ladies’-tresses [synonyms: Spiranthes brevilabris var. floridana (Wherry) Correll; S. gracilis var. floridana (Wherry) Luer ] (Brown 2001). Two new color forms were described from Texas, both from the Trans-Pecos (Brown 2006): Dichromanthus cinnabarinus forma aureus PM. Brown (Fig. 5), a yellow form of the otherwise typically scarlet-colored flowers; similar coloration variants are found in the related genus Sacoila: S. lanceolata (Aubl.) Garay var. lanceolata forma folsomii PM. Brown; S. lanceolata (Aubl.) Garay var. paludicola (Luer) Sauleda, Wunderlin & Hansen forma aurea PM. Brown (Brown 1999a; 2001). Hexalectris grandiflora (A. Rich. & Galetti) L.O. Williams forma luteoalba PM. Brown (Fig. 6), an unusual yellowish-white color form; similar variants are found in other species of Hexalectris: H. spicata var. spi- cata forma lutea P.M. Brown, H. spicata var. spicata forma wilderi PM. Brown (Brown 2004); H. warnockii Brown, Notes on Texas Orchids 1267 forma flavida Catling (Catling 2004). In reviewing some of the early literature and attempting to coordinate the earlier names with current no- menclature several unusual records were noted in Cory and Parks (1937) and/or Correll (1940; 1947). Liparis loeselii (L.) Rich.: thisis the only reference for this species in the Texas literature although it appears in an informal plant list for Caddo National Wildlife Refuge, Louisiana, but without any substantiation; the refuge straddles the Texas-Louisiana border. Microstylis monophyllos (L.) Lindl.: confusing use of a synonym and misapplied name intending to use Malaxis unifolia Michx.. Physurus querceticola Lindl. (syn. Platythelys querceticola (Lindl.) Garay) was reported by Ames (1924), treated by Correll (1944), and noted by Liggio and Liggio (1999). Plants of this species still have not been found in Texas. Spiranthes ochroleuca (Rydb.) Rydb.: specimens were most likely S. magnicamporum Sheviak (not described at that time) or a prairie race of S. cernua (L.) Rich., both of which may have creamy colored lips typical of S. ochroleuca. Spiranthes reverchonii (Small) Cory is one of several described taxa that fall within the parameters of the highly variable S. vernalis Engelm. & A. Gray (Ames 1924). Spiranthes minutiflora A. Rich. & Galeotti presents the most intriguing report of all. By todays nomenclature this would be Microthelys minutiflora (A. Rich. & Galeotti) Garay (Garay 1980; Szlachetko et al. 2005), a small, inconspicuous spiranthoid species known from northern Mexico. Cory and Parks (1937) noted it from the Trans-Pecos region presumably based upon the information in Palmer (1880). The Palmer specimen was listed as one of several that were missing from a shipment sent by Palmer! to Dr. Asa Gray at the Gray Herbarium at Harvard University. Without a specimen for positive identification or more complete details other than ‘mountains of West Texas, 1880’ the inclusion of this species for Texas was tenuous at best. But, in 2005 Microthelys rubrocallosa (B.L. Rob. & Greenm.) Garay, a similar and closely related species from northern Mexico, was found for the first time in the United States in southern New Mexico (Coleman & Baker 2006). The two species bear a superficial resemblance to each other and it is possible that the Texas report could even have been M. rubrocallosa, depending on information avail- able to Palmer at the time. Although the lip shape is significantly different in the two species, they both posses the characteristic red underside of the lip and tiny, barely- opened whitish flowers. A notable error in Cory and Parks (1937) was in the listing of Blephariglottis tricholepis (Torr.) Nash, which apparently is Blepharoneuron tricholepis (Torr.) Nash in the Poaceae, a grass known primarily from the Trans- Pecos region: The most significant rediscovery of a species that had not been seen since its original collection in 1931 is that of Deiregyne confusa Garay, the rimrock ladies-tresses, found in 2004 in essentially in the same area as the earlier collection by Steyermark in the Chisos Mountains (USGS 2004; Ambs 2006; Coleman et al. 2006). Spiranthes brevilabris Lindl., one of the rarest orchids in North America, was discovered in April 2007 in the Blackland Prairie vegetational region, documenting a new county, Walker, and a new geophysical region for the species in Texas (Keith 2007). ACKNOWLEDGMENTS The author thanks Marcy Brown-Marsden for her helpful suggestions. ! Edward Palmer 1831-1 1268 Journal of the Botanical R h Institute of Texas 1(2) REFERENCES Ames, S. 2006. The elusive Deiregyne confusa. Native Orchid Conf. J. 3(2):1-5. Ames, O. 1924. An Enumeration of the Orchids of the United States and Canada. Boston: The American Orchid Society. Brown, P.M. 1999. Recent distributional and taxonomic notes from Florida 1 (Spiranthes eatonii). N. Amer. Native Orchid J. 5(1):5-15. Brown, PM. 1999a. Recent distributional and taxonomic notes from Florida 2. N. Amer. Native Orchid J. 5(2):169- Brown, PM. 2001. Recent distributional and taxonomic notes from Florida 9. N. Amer. Native Orchid J. 7(1):92- BRowN, P.M. 2001a. Recent distributional and taxonomic notes from Florida 11 (Spiranthes sylvatica). N. Amer. Native Orchid J. 7(3):192-205. Brown, P.M. 2002. Revalidation of Platanthera conspicua. N. Amer. Native Orchid J. 8:3-14. BRowN, P.M. 2004. New taxa. N. Amer. Native Orchid J. 10:23. Brown, PM. 2005. Additions and emendations to The wild orchids of North America, north of Mexico. Sida 21:2297-2319. BRowN, P.M. 2006. Four new color forms from the Southwestern United States. McAllen Int. Orchid Soc. J. 7(12):4-11. Brown, P.M. and R. Coleman. 2000. Schiedeella arizonica, a new species from the Southwestern United States. N. Amer. Native Orchid J. 6(1):3-17. Brown, P.M. and R. Pike. 2006. Triphora trianthophora var. texensis (Orchidaceae), a new variety endemic to Texas. N. Amer. Native Orchid J. 12:4-10. CATLING, PM. 2004. A synopsis of the genus Hexalectris in the United States and a new variety of Hexalectris rev- oluta. Native Orchid Conf. J. 1(2):5-25. CoLEMAN, R.A. and M. Baker. 2006. Microthelys rubrocallosa. Orchids 75(1):56-57. COLEMAN, R.A., J. SIROTNAK, and A. Leavitt. 2006. The hunt for Deiregyne confusa. Native Orchid Conf. J. 3(3):7-8, 13. Cory, V.L. and H.B. Parks. 1937. Catalogue of the flora of the state of Texas. Texas Agric. Exp. Stat. Bull. 550:34-35. Correll, D.S. 1944. Orchidaceae. Vol. 3, pt. 3, in Lundell, Flora of Texas. Texas Research Foundation, Renner. CorreLt, D.S. 1947. Additions to the orchids of Texas. Wrightia 1:166-82. Garay, L.A. 1980. A generic revision of the Spiranthinae. Bot. Mus. Leaflets. 28:277-426. KerrH, E. 2007. Texas ladies'-tresses (Spiranthes brevilabris) rediscovered in Texas. N. Amer. Native Orchid J. 13(2):133-14. Liscio, J. and A.O. Licaio. 1999. Wild orchids of Texas. University of Texas Press, Austin. Palmer, E. 1880. List of plants collected in S.W. Texas and N. Mexico. Unpublished papers. Library of the Gray Herbarium. SZLACHETKO, D.L., P. Rutkowski, and J. Mytnik. 2005. Contributions to the taxonomic revision of the subtribes Spi- ranthinae, Stenorrhynchidinae, and Cyclopogoninae (Orchidaceae) in Mesoamerica and the Antilles. Polish Bot. Stud. 20. USGS Biotoaicat INFoRMATICS OrFICE. 2004. Big Bend sensitive plant project-2004 Field Season Discoveries. http:// cswgcin.nbii.org/ecoregion/trans-pecos/field/index.html GALIUM ANGLICUM (RUBIACEAE) NEW FOR TEXAS AND NOTES ON THE TAXONOMY OF THE G. PARISIENSE/DIVARICATUM COMPLEX Barney L. Lipscomb and Guy L. Nesom Botanical Research Institute of Texas 09 Pecan Street Fort Worth, Texas 76102-4060, U.S.A. ABSTRACT Galium parisiense var. leiocarpum is documented as a first record for Texas and South Carolina. Other localities in the eastern U.S.A. are mapped for the glabrous-fruited expression of G. parisiense (var. leiocarpum), bristly-fruited G. parisiense (var. parisiense), and the closely related, E fruited G. -a All taxa are moe natives. In both North America and Europe, var. parisiense and var. leiocarpum h i geograj lly in some areas. Galium divaricatum occurs sympatrically at least o da 1 e Ts E £ 1 f] +1 with var. leiocarpum. V sG. parisiense L., G. divaricatum Pourret ex Lam., and G. anglicum mune G. pde vat. o! RESUMEN 124 d L F 1 A aie a Texas. Otras localidades del este de U.S.A fí la parisien e (var UD frut d parisiense (var parisien €), y laespecie próxima ae frutos glabros G. enc Todos los taxa son nativos de LM Tanto en Norte América como en Europa, var. parisiense y var. leiocarpum p algunas áreas. Galium divaricatum es simpátrico T menos con var. leiocarpum. I i Europa. Sugeri l d taxa, como G. parisiense L., G. por Pourret ex Lam., y G. anglicum rd G. em isiense var. desear A collection of Galium anglicum Hudson is reported here as the first known record of the species for Texas. Galium anglicum Hudson (= G. parisiense L. var. leiocarpum Tausch, see below) TEXAS. Gillespie Co.: LB] National History Park, RM Unit, along dM half of LBJ landing strip, gentle slope, ungrazed, rede- veloping prairie, apparently locally common but d g prairie and weedy plants in disturbed area of caliche fill, 1 Jun 2005, Roger Sanders 6681 (BRIT) The plant was collected and correctly identified by Sanders in connection with a floristic study of LBJ National History Park in central Texas. In our own evaluation of the Texas plant, we agree with Sanders' identification, but in comparing it to other collections from the eastern U.S.A, we observe that Galium parisiense has often been confused in identification with G. divaricatum Pourret ex Lam. The two taxa are similar and apparently closely related, and both are native to Europe. Galium divaricatum has been treated as a variety of G. parisiense (see below) but is more generally recognized as a distinct species. Ehrendorfer & Krendl (1976, p. 36) noted that G. divaricatum is “not very variable, and usually quite distinct from [G. parisiense].” Part of the taxonomic difficulty stems from variation in fruit morphology in Galium parisiense. Plants of the typical expression have fruits hispid with uncinate-tipped hairs, while fruits of var. leiocarpum are glabrous (G. divaricatum also is glabrous-fruited). In Europe, Ehrendorfer and Krendl (1976) implied that the glabrous-fruited phase of G. parisiense was to be treated as a population variant without formal taxonomic status; the fruits of G. parisiense were described as glabrous or hairy. In the Flora of Turkey, Ehrendorfer El 1 bcs f and Schónbeck-Temesy (1982) also described G. parisiense (without infraspecific taxa) as producing both fruit types. Joao do Amaral (1971) followed Flora Europaea in treating G. parisiense sensu lato in Portugal as variable in fruit vestiture and distinct from G. divaricatum. In contrast to the broad concept of Galium parisiense, in continental Europe it is described as having jeep hairy fruits (e.g. Pignatti 1982; Schónbeck-Temesy & Ehrendorfer 1991) without mention of glabrous forms. J. Bot. Res. Inst. Texas 1(2): 1269 — 1276. 2007 1270 Journal of the Botanical R h Institute of Texas 1(2) In the British Isles, the fruits are described as glabrous or papillose (e.g., Butcher 1961, and as illustrated; Clapham, Tutin, & Moore 1989; Stace 1997) without mention of hairy forms. Brenan (1953) reported a col- lection of var. parisiense from England, noting that “typical G. parisiense, though apparently not previously recorded from Britain, has a wide distribution in Europe and the Mediterranean Region, extending to the Middle East.” Boulos (2000) included the bristly-fruited form (as G. parisiense), based on collections from Cyprus. Only glabrous-fruited plants occur in Madeira (Turland 1994). For Spain, Devesa (1987) included bristly-fruited G. parisiense (widely distributed) and glabrous-fruited G. divaricatum (restricted to near-coastal localities, sympatric with G. parisiense), but the illustration of G. divaricatum in Devesa's apparently shows G. parisiense var. leiocarpum. All three taxa have been included for Portugal (Pereira Coutinho 1939; Sampaio 1946) and for France (e.g., Lamarck & de Candolle 1805; Grenier & Godron 1850; Coste 1901), although using various nomenclature. In summary, it appears that the two expressions of G. parisiense in Europe have distinct geographic ranges and probably occur sympatrically in some areas. Glabrous-fruited plants, all reported as Galium divaricatum, are widespread in Australia (e.g., Toelken 1986; Rye 1987; James & Allen 1992) and New Zealand (Webb et al. 1988) and rare in Hawaii (Wagner et al. 1999). Descriptions of these do not provide information that allows their unequivocal identification as G. divaricatum (vs. G. parisiense var. leiocarpum). A bristly-fruited taxon is not included in current floristic sum- maries from these areas, although Webb et al. (1988, p. 1146) noted that earlier literature had “mentioned a bristly-fruited var. of G. parisiense occasionally found in N.Z.” Taxonomic assessments of Galium California, Oregon, and the Pacific coast region have recognized a bristly-fruited taxon (as G. parisiense) and glabrous-fruited one (as G. divaricatum, including as synonyms G. parisiense var. leiocarpum and G. anglicum Hudson) (e.g., Jepson 1925; Howell 1943; Munz & Keck 1959; Munz 1974; Dempster 1993; Ferris 1960; Hitchcock & Cronquist 1976; Peck 1961; Kozloff 2005; Oregon State Univ. Herbarium 2007). Ceska and Ceska (1998) reported *G. parisiense" from British Columbia, but the morphology of the plant (A. Ceska pers. comm.) refers it to G. parisiense var. leiocarpum. Sanders (1999) reported G. parisiense, referring to the bristly-fruited taxon (Sanders pers. comm.), as widespread in cismontane southern California. We have seen (indicated below as “!”) relatively few specimens from the Pacific coast region and do not attempt here to resolve the geography of G. divaricatum vs. G. parisiense var. leiocarpum. In summary from literature and herbarium: bristly-fruited in (California) Butte!, Calaveras, Colusa!, El Dorado!, Fresno, Humboldt!, Lake!, Los Angeles!, Mariposa!, Marin!, Mendocino, Monterey!, Napa!, Orange, San Bernadino, San Diego, Santa Cruz, Sonoma!, and Tuolumne! counties; (Oregon) Douglas, Jackson, and Josephine counties; (Washington) Kittitas and Pacific counties. glabrous-fruited in (California) Alameda, Butte, Calaveras, Contra Costa, El Dorado, Humboldt, Lake, Marin, Mendocino, Napa!, Ne- WV4il vada, Plumas, Santa Clara, Santa Cruz, Shasta, Sonoma!, and Oregon) Benton!, Curry, Douglas, Jackson, Lane, Marion, and Poll ties; (British Columbia) Lasqueti Island. Both m fruited a and glabrous-fruited dna are recorded from Calaveras, Humboldt, Lake, Marin, Mendocino, Napa, and ifornia) and Jackson Co. (Oregon) Our own observations of plants in the eastern U.S.A. indicate that the bristly-fruited and glabrous-fruited forms of G. parisiense have discrete geographic ranges (Fig. 1). Galium divaricatum occurs in Missouri, Indiana, Tennessee, and Alabama; G. parisiense sensu stricto occurs in Arkansas, Louisiana, Mississippi, Tennes- see, and Alabama; G. parisiense var. leiocarpum occurs from the Atlantic region westward to Oklahoma and Texas, and it appears to be sympatric with typical G. parisiense in Alabama. As a speculative and probably over-simplified hypothesis, we might suggest that the wider range of var. leiocarpum has resulted from a greater number of introductions than for the other two taxa. Fruit variation in Galium Production in Galium of bristly fruits (vs. fruits with glabrous, hairless, surfaces) usually is a consistent character and indicator of close relationship among species (e.g., Dempster 1978, 1980, 1981-1982; Ehren- dorfer & Krendl 1976), but variation within species exists in a few instances. Moore (1975, p. 881) described fruits of G. aparine L. as “surface covered with stiff, hooked hairs (spines) ... the spines sometimes sparse , Gali [ isi leiocarpum in Texas — > m = M HN an ns sun xn Ma AY ras V Eh pa Bape ee Oss Zell Le PHA Zone El Y ES d "b Du J ee V9 ND OS == EP e E Hota EOE | e Moscwy OA A Ep AA MIS A O ETHER Ei m) TJ m Li] (T e "y IN Gen RD. OS A A en > HIN (3 CA CET EA Seer ^x ADS M NA ETE Li Xp LIRE NSA OC DD Sua Raman Y UE ETIN al ERIS Zr CER VAST AN ENT ORE YA r | A GS SOSA PLE LS Or SH xd ey e, ae AS a Y hae y, A iiw H EH EE EER ha L^ WS WE -| [~] rH - LA "N A q ve: Y le AR AQ v EDS HH ay moe C] $ S WAV. SAS » D LA | | Y) Nc PS i E Y Pa qu 4 SALAS TURN A 3 DO ea S aC, UNES e QD T E J N T laa aa a T] CEA ce sl AA L - ma M on Pore O ES A eee a er MAGES um. MARY 3 SS © G. parisiense HA Say —- wo cU e G. divaricatum ÍA vam Y L Se ed y ~ " den A DE i MILES Wa E o 100 200 300 400 7 on bc A i A Xl LN S7 or, very rarely, lacking and the surface smooth or tuberculate.” He treated the glabrous-fruited plants as G. aparine forma intermedium (Bonnet) R.J. Moore but noted that “this rare form has not been seen” in a study of 280 Canadian collections of the species. Nor do we observe glabrous-fruited plants of G. aparine in ca. 350 collections (BRIT, TEX, VDB) of the species from the U.S.A. i ly exists in Galium spurium L. Gf treated as a species distin Similar variation in fruit pp from G. aparine). Bristly-fruited plants are the more common in Europe and Canada and are treated variously, e.g., G. vaillantii DC., G. spurium var. vaillantii (DC.) Gren. & Godr., G. spurium forma vaillantii (DC.) RJ. Moore. Kaplan and Rehoék (1998) stated that variation in G. parisiense is analogous to that in G. spurium, but in study of herbarium collections and wild populations of G. parisiense (in Slovakia) they noted (p. 54) that “the characteristics of the fruit are usually common to all members of a population.” Consistent mor- phology within a population is not inconsistent with the hypothesis that sympatric populations of different morphology may be reproductively isolated. Taxonomy of the Galium parisiense group Four general taxonomic options exist: 1) treat Galium parisiense with two varieties, distinct from G. divari- catum in habit and inflorescence (following Flora Europaea), implying that the two varieties of G. parisiense icat species with two varieties, distinct from G. parisiense, intergrade; 2) treat glabrous-fruited G. diva implying that the difference between var. leiocarpum and G. divaricatum is arbitrary or intergrading, as ap- 1272 Journal of the Botanical R h Institute of Texas 1(2) parently implied by California treatments; 3) treat three taxa within a single species, as G. parisiense; 4) treat each of the three taxa at specific rank, recognizing that each appears to be distinct and reproductively isolated from the other two. Our limited study suggests that the fourth option is most appropriate; Lid (1967) followed this course for Canary Island plants. As reviewed above, the three taxa occupy distinct geographic ranges and occur sympatrically in their native range in Europe as well as where naturalized in North America—along the Pacific Coast region and in the eastern U.S.A. Local populations appear to be consistent in fruit and inflorescence morphology. Broad sympatry among the taxa naturalized in North America and elsewhere outside the native range provides critical information pertinent to their biology and taxonomic rank. Because of the widely inconsistent taxonomic treatment of these taxa, however, especially in Europe, and because they are becoming nearly cosmopolitan weeds, the taxonomy and geography deserve more comprehensive study. The following key and taxonomic paragraphs summarize our observations and conclusions. Maps are constructed from specimens examined in the current study (from BRIT-SMU, NCU, TENN, TEX-LL, and VDB) are indicated by counties of occurrence within a state. Plants from Missouri, Oklahoma, and West Virginia were observed from photocopies provided by SMS, OKLA, and WVA, respectively. KEY TO GALIUM PARISIENSE, G. DIVARICATUM, AND G. ANGLICUM = . Inflorescence ce diffuse, branches divaricate; ultimate fruits (2-)3-6(- D Lies beyond primary stem is (with | leaves); first infl j le (I | pri 5-50 mm long; fruit surface glabrous (without e rs) G. divaricatum 1. Inflorescence relatively strict, branches ascending; dnas fruits 2-3(-4) nodes beyond primary stem axis (with largest leaves); first infl | stem axis) 3-12(-20) mm long; fruit surface glabrous or bristly-hispid. 2. Fruit surface bristly-hispid with uncinate-tipped hairs, distinctly muriculate G. parisiense 2. Fruit surface without hairs, smooth to muriculate .anglicum Color photos from Pinnacles National Monument in California (Morse 2007) show the contrast in the two inflorescence forms: one plant clearly is G. divaricatum but it is not possible to see the fruit vestiture on the other, which is either G. anglicum or G. parisiense. Color photos from the western Mediterranean region (Fundación Biodiversidad 2007) show the same contrast: one plant (as G. parisiense subsp. divaricatum) is G. divaricatum, while the other (as G. parisiense subsp. parisiense) is G. anglicum with glabrous fruits. Galium divaricatum Pourret ex Lam., Encycl. 2:580. 1788. Galium parisiense var. divaricatum (Pourret ex Lam.) Koch, Syn. Fl. Germ. Helv., ed. 2, 3:1025. 1845 [non vis. 1852]. Galium anglicum var. divaricatum (Pourret ex Lam.) Reichb., Ic. Fl. Germ. Helv. 17: t. 1196, f. 2. 1854-1855. Galium parisiense subsp Tuum (Pourret ex Lam.) Rouy, Fl. France 8:46. 1903. Type: FRANCE: K- «Cet France li bl pierreux. M. Desfontaines l'a rencontreée dans le Berry; & M. l'Abbé. 1 P t i divid ‘ila trouvés dans le Languedoc. (v.s.).» L TL Native range.—Southern Europe, Cyprus, Syria, Lebanon, Turkey, Czechoslovakia; «Albania?, Austria, Azores, Balearic Islands, Belgium, Bulgaria, Corsica, Crete, Czechoslovakia, France, Germany, Greece, Hungary, Italy, Portugal, Romania, Sardinia, Sicily, Spain, Switzerland, Yugoslavia, and Turkey» fide Ehrendorfer and Krendl (1976). Naturalized.—Australia, New Zealand, Hawaii. Occurrence in eastern USA.—(Fig. 1). ALABAMA (Morgan, Randolph), Indiana (Brown), Kentucky (Campbell, Estill), Missouri (Greene), and TENNESSEE (Davidson, DeKalb, Dickson, Haywood, Marshall, Perry, Rutherford, Sumner, Wilson). Galium parisiense L., Sp. pru cob (Figs. 1, 3). Ter: «Habitat in Anglia, Gallia.» The protologue notes «fructibus hispidis.» Galium E DC. in Lam. & DC., Fl. France 4:263. 1805. Galium eee var. litigiosum (DC.) Koch, Syn. Fl. Germ. Helv., 331. 18 tum var. litigiosum (DC.) Kostel, Ind. Hort. Bot. Prag., 61. 1844. Galium parisiense var. M idi, HG nid Fl. Ital., 315. 1882. Type: France: various localities are cited in the D which also notes n ici leiocarpum in Texas AA «x cH SIS ace CORTO EE SUP L2 Wa AG REA Cai tee Bee Tenn NUM UT Women oe E SL O dr ERU CASES LODS EXT ENE GP AERTS Fic. 2. Distribution of G. g j 211 LENT 1 n 3. dz 7 4 | FE RU J 1 fu :1 A 7 E 8 8 8 | E J aux environs de Paris.» «Galium parisiense. Linn. spec. 157, excl. syn. Ray. et forsan. Tourn.” was cited in synonymy and the name G. parisiense was not applied to any other species. The fruit was described as hairy (“hérissé de poils"). Galium decipiens Jordan, Obs. Pl. Crit. 3:178. 1846 (non Ehrhart 1790; non Roth 1827). Type: FRANCE: «J'Al OBSERVE CETTE ESPECE DANS PLUSIEURS LOCALITÉS DU MIDI, AUX ENVIRONS DE TARASCON, MONTPELLIER, CETTE, ETC. ELLE FLEURIT EN JUILLET.» JORDAN DESCRIBED THE FRUITS AS GLABROUS OR C COVERED WITH HAIRS AND NOTED «JE L'AI RENCONTRÉE TANTOT A FRUITS GLABRES, TANTOT A FRUITS VELUs.» Ehrendorfer CON C and Krendl (1976) observed that “In S.W. Europe variants are found with few-flowered partial inflorescences and large fruits with hooked hairs (G. decipiens Jordan ... )." Galium parisiense L. var. (a.) trichocarpum Tausch, Flora 18:354. 1835. Tyre: Tausch noted “G. parisiense L. Cand. (e specim. in herbar. Sieberi.)." Native range.—Southern, western, and central Europe, Cyprus, Turkey, Bulgaria; "Albania, Austria, Azores, Balearic Islands, Belgium, Britain, Bulgaria, Corsica, Czechoslovakia, France, Germany, Hungary, Italy, Por- tugal, Romania, Sardinia, Sicily, Spain, Switzerland, and Yugoslavia fide Ehrendorfer and Krendl (1976). Naturalized.—Chile (Dempster 1981), U.S.A., New Zealand. Occurrence in eastern USA.—(Fig. 1). ALABAMA (Colbert, Hale, Monroe), Arkansas (Lafayette, Saline, White), Louisiana (Calcasieu, Ouachita, Webster, Winn), Mississippi (Grenada, Lafayette, Marshall, Monroe, Panola, Yalobusha), and Tennessee (Marion). Galium anglicum Hudson, Fl. Angl., ed. 2, 1:69. 1778. (Fig. 2, 3). Galium parisiense subsp. anglicum (Hudson) Gaud., FL. Helvet. 1:438. 1828. Galium parisiense var. anglicum (Hudson) G. Beck [nom. illeg.], Fl. Nied.-Osterr. 2:1122. 1893. Type: ENGLAND: “Habitat i i i is, inter Dartford et Northfleet i ido Farningham in agro Cantiano.” Beck cited Į I PI 8 eiocarpum Tausch in synonymy. He noted that he had not observed the glabrous-fruited taxon in Austria G. parisiense var. | 1274 Journal of the Botanical R h Institute of Texas 1(2) la ` ERS/7 = rr E ex Im == == = aw ww 72 122834 Botanical Research Institute of Texas Galium anglicum Hudson No. 4066 Botanical Research Institute of Texas Eun ds GALIUM PARISIENSE L. VAR. LEIOCARPUM TAUSCH Det. Cheryl A. Lawson, 1976 G. Nesom & B. Lipscomb 2007 i HERBARIUM OF BUTLER UNIVERSITY sA FLORA OF INDIANA Collected by RAY C. FRIESNER Herbarium of the EE Military Institute 9956. Galium parisiense L. ; I ckbridge County Fallow field on cast side of Rd. 135, n. e h HE. cL first county road — 4 east 0.4 mi, ne [EE DS Ed Blossom, Brown Co., Indiana - | Open fíeld vu Post T-1-56 3 4 MMC 28 July 1970 Fic. 2. Inf] £falt Ae ER [Ve £e 1G. anglicum ( right) Ga ium parisiense L. var. (B.) leiocarpum Tausch, Flora 18:354. 1835. Tyee: Tausch noted “G. anglicum Huds. Cand. (ex phrasi excl. B. ad G. divaricatum spect.) Ray syn. t. 9. f. 1. (bona quoad habitum) Barr. ic. t. 58.” b of the name Duis leiocarpum" ps Borza (1949, as “f. leiocarpum et f. trichocarpum Tausch.") was not a valid combination basionym was provi Native range.—England, France, Portugal, Spain? Naturalized.—Australia, New Zealand, Hawaii (vs. G. divaricatum, see comments in text). Occurrence in eastern USA.—(Fig. 2). ALABAMA (Calhoun, Marengo, Marion, Sumter), ARKANSAS (Bax- ter, Benton, Boone, Carroll, Fulton, Izard, Little River, Marion, Newton, Seracy, Stone), GEORGIA (Sumter), leiocarpum in Texas 1275 Missouri (Douglas, Henry, Ozark, Pulaski), NortH Carora (Cabarrus, Chatham, Cleveland, Orange), OkLAHOMA (Grady), TENNESSEE (Cumberland, Hamblen, Hamilton, Hawkins, Knox, McMinn, Marion, Mar- shall, Montgomery, Rutherford, Smith, Van Buren), Texas (Gillespie), Vircinia (Bedford, Buckingham, Craig, Giles, Goochland, Henrico, Mecklenburg, Rockbridge, Shenandoah, Smyth), and West VircinIA (Fayette, Monroe, Summers). The following collection is the first known record of Galium anglicum from South Carolina: Pickens Co.: Cedar Creek Circle, Central, scarce lawn week, 17 Jun 1989, Hill 20707 (MO). ACKNOWLEDGMENTS Collections were studied from BRIT-SMU, MO, NCU, TENN, TEX-LL, and VDB. We are grateful for images of specimens from Missouri (SMS), Oklahoma (OKLA), and West Virginia (WVA), which provided a basis for consistent identification of collections from those states, for specimen loans from UARK and NCU, for comments from Andy Sanders on UCR collections and Adolph Ceska on the British Columbia record, and for comments on the manuscript by Cheryl Lawson and Alan Weakley. We also are grateful to Lloyd Shin- ners for assembling a rich library, now at BRIT, which provided all but one of the references cited here and in the taxonomic summaries. REFERENCES Borza, A. 1949. Conspectus florae Romaniae regionumque affinium. Fasc. 2. Tipografia Cartea Románeascá, Cluj, Romania. Boulos, L. 2000. Flora of Egypt. Vol. 2, Geraniaceae - Boraginaceae. Al Hadara Publishing, Cairo, Egypt. BurcHER, RW. 1961. A New Illustrated British flora. Part 2. Leonard Hill Books Limited, London. CASTANER, D. 1982. Galium divaricatum (Rubiaceae), new to Missouri. Sida 9:368-369. 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Flora Vascular de Andalucía Oc- cidental. Ketres Editora, Barcelona. Pp. 574-593. Dupestapt, W.H. 1977. Some new state records and other plant finds in West Virginia. Castanea 42:257-258. EHRENDORFER, F. and F. KRENDL. 1976. Galium. In: Tutin et al., Flora Europaea, Vol. 4, Plantaginaceae to Compositae (and Rubiaceae). Cambridge Univ. Press, Cambridge. Pp. 14-36. EHRENDORFER, F. and E. SCHONBECK-TEMESY. 1982. Asperula L. and Galium L. In: PH. Davis, ed. Flora of Turkey and the East Aegean Islands. Vol. 7. Edinburgh Univ. Press, Edinburgh. Pp. 734-849. Ferris, R.S. 1960. An Illustrated flora of the Pacific States: Washington, Oregon, and California. Vol. 4. Stanford University, Stanford University Press FUNDACIÓN BIODIVERSIDAD. 2007. Herbario Virtual del Mediterráneo Occidental. Universitat de les Illes Balears, Univer- sitat de Barcelona, Universitat de Valencia. Accessed June 2007 GRENIER, M. and M. Gobron. 1850. Flore de France: ou Description des plantes qui croissent naturellement en France et en Corse. Vol. 2. J.B. Bailliére, Paris. 1276 Journal of the Botanical R h Institute of Texas 1(2) HircHcock, C.L. and A. Cronauist. 1976. Flora of the Pacific Northwest, an illustrated manual (illus. J.R. Janish). Univ. of Washington Press, Seattle. HoweLL, J.T. 1943. Some weedy Californian bedstraws. Leafl. West. Bot. 3:260-261. JAMES, TA. and W.K. ALLEN. 1992. Galium. In: GJ. Harden, ed. Flora of New South Wales. Vol. 3. New South Wales Univ. Press, Kensington, Australia. Pp. 483-486. Jepson, W.L. 1925. A Manual of the flowering plants of California. Associated Students Store, Univ. of California, Berkeley. Joao DO Amaral, F. 1971. Nova flora de Portugal: continente e Açores. Vol. 2, Clethraceae--Compositae. Sociedade Astória, Lisboa. KAPLAN, Z. and V. ReHOREK. 1998. Galium parisiense: a new alien species for the Czech Republic. Preslia 70:51-56. Koztorr, E.N. 2005. Plants of Western Oregon, Washington & British Columbia. Timber Press, Portland, Oregon. Lamarck, J, and A.P. pe CANDOLLE. 1805. Galium. Fl. France 4:248-266. Lip, J. 1967. Contributions to the flora of the Canary Islands (ill. by D.T. Lid). Universitetsforlaget, Oslo. McGIuveay, D.J. 1983. A revision of Galium (Rubiaceae) in Australia and New Zealand. Telopea 2:355-377. MacRATH, L.K. 2001. Galium parisiense var. leiocarpum Tausch, new for Oklahoma. Oklahoma Native Pl. Rec. TODOS Moore, RJ. 1975. The Galium aparine complex in Canada. Canad. J. Bot. 53:877-893. Morse, K. 2007. Keir's Botanical Photo Archive. Accessed June 2007. Munz, PA. 1974. A flora of southern California. Univ. of California Press, Berkeley. Munz, PA. and D.D. Keck. 1959. A California flora. Univ. of California Press, Berkeley and Los Angeles. OREGON STATE Univ. HERBARIUM. 2007. Oregon Vascular Plant Database. Peck, M.E. 1961. A manual of the higher plants of Oregon (ed. 2). Binfords & Mort, Portland, Oregon. PEREIRA COUTINHO, A.X. 1939. Flora de Portugal (Plantas Vasculares) (ed. 2). R.T. Palhinha (ed.). Bertrand (Irmãos), Ltd., Lisboa. PicNarTI, S. 1982. Flora d'Italia. Vol. 2. Edagricole, Bologna. Rye, B.L. 1987. Rubiaceae. In: N.G. Marchant et al. Flora of the Perth Region (ed. 4). Part 2. Western Australian Herbarium: Dept. of Agriculture, Western Australia, Perth. Pp. 643-646. Sampaio, G. 1946. Flora Portuguesa (ed. 2). Américo Pires de Lima. Impr. Moderna, Pôrto. SANDERS, A.C. 1999. Noteworthy collections. Madrono 46:113. Stace, C. 1997. New flora of the British Isles (ed. 2). Cambridge Univ. Press, Cambridge. SCHONBECK-TEMESY, E. and F. EHRENDORFER. 1991. Galium. In: A. Strid and K. Tan, eds. Mountain flora of Greece. Vol. 2. Edinburgh Univ. Press, Edinburgh. Pp. 300-328. TOELKEN, H.R. 1986. Rubiaceae. In: J.P. Jessop and H.R. Toelken, eds. Flora of South Australia. Part 2, Legumino- sae-Rubiaceae. South Australian Government Printing Division, Adelaide. Pp. 1059-1076. TURLAND, N.J. 1994. Rubiaceae. In: J.R. Press and M.J. Short, eds. Flora of Madeira. The Natural History Museum, London. St. Edmundsbury Press, Suffolk, England. Pp. 261—266. WAGNER, W.L, D.R. Herest, and S.H. Sommer. 1999. Manual of the flowering plants of Hawaii (rev. ed.). Bishop Museum Special Publ. 97. University of Hawai'i Press, Honolulu. Wess, C.J., W.R. Sykes, and PJ. GArnock-Jones. 1988. Flora of New Zealand. Volume IV. Naturalized Pteridophytes, Gymnosperms, Dicotyledons. DSIR Botany, Christchurch, New Zealand. James Hinton September 2005, a few days before his 89' birthday. JAMES HINTON (1915-2006) George S. Hinton Billie L. Turner Apartado 603 Plant Resources Center Saltillo, Coahuila 25000 University of Texas MEXICO Austin, Texas 78703, U.S.A. James Hinton died on July 23, 2006 in his Rancho Aguililla, Galeana, Nuevo Leon. The last 20 years of his life he dedicated to writing novels and short stories. He took time off from writing to collect the flora of southeastern Nuevo León with occasional forays into Coahuila, Tamaulipas and, his last collecting trip, to far away Oaxaca when he was 81—it was his third trip to Oaxaca in two years, and he spent it botanizing in and about Quiexobra. He liked most collecting up in the mountains and he has plants named for him in the highest mountains of Nuevo Leon, Coahuila, Tamaulipas, Oaxaca, Michoacán and Guerrero. James Hinton was born in Mexico City on September 24, 1915 to George B. Hinton and Emily Wattley. He was the third of three brothers. His mother died of tuberculosis when he was four. She believed in fasting to overcome any sickness but tuberculosis was not one that could be cured by it. James inherited this from his mother and fasted for various reasons throughout his life. After his mother’s death his father took his three sons to live in Cuernavaca where he visited them every week end. They grew up a wild bunch with the eldest, Howard E., master of the house. Listening to them after 50 years, Howard was a terror; this encouraged a close bond between James and his brother George that would last throughout their lives. As soon as Howard was old enough he was sent to school in California where he studied entomology. After, his father then sent him to England to complete his studies. James and George were sent on a 10 day J. Bot. Res. Inst. Texas 1(2): 1277 — 1280. 2007 1278 Journal of the Botanical R h Institute of Texas 1(2) long train trip to Vancouver, British Columbia, to study. They were 14 and 16. When they arrived they had to look for a school, room and board. James had terrible memories of having to play rugby in school, fond memories of canoeing along the coast of B. C. and in its rivers, and swimming in the ice cold sea. He entered a competition that made him the champion swimmer of Alberta. He loved swimming and would swim a kilometer a day whenever he was close to a swimming pool. He built one in Rancho Aguililla and claimed that over 50 years he had swam to Canada. He had planned to study economics at university but was called back to Mexico by his father to help him collect plants. He spent five years collecting plants with his father. He estimated he'd made 25% of the G.B. Hinton collections in remote areas of Guerrero and Michoacán. He traveled thousands of kilometers on mule back and asked Dr. Epling to name a salvia for his mule Lenina, Salvia leninae. When he was 90 he could still remember far away names like Toro Muerto, Teotepec, Piedra Ancha, Atoyac, Tecpan and measure their distances in days on horse back. Many of his stories and novels, written in his old age, were set in the rural Mexico of long ago he remembered and loved so well. He married Helen Hart in 1942 and stopped collecting plants; he needed a job that would sustain a family. He worked in Saltillo managing the guayulera and exploring for guayule. The job ended with the war. He then worked as a sales representative and later started various companies. On the side he had sev- eral ranches in Nuevo Leon, Michoacán and the State of Mexico. He started a turkey farm and a maguey plantation in the State of Mexico and planted melons in Aguililla, Michoacán. He planted wheat and then an apple orchard in Rancho Aguililla, Nuevo León. After he left Saltillo he moved to Monterrey and eventually to Cuernavaca, where he lived for 30 years. One of the first modifications he made to the house he bought there was to build a swimming pool. He com- muted to Mexico City where he had two companies, and spent much time on the road going to his farms in Nuevo León and Michoacán. He was an avid hunter and an excellent shot. He delighted us with countless photographs of faraway places and with the stories associated with them. He always had horses and looked over his orchard on horseback, riding up and down the rows of trees, claiming it was the best way to inspect it. He loved moun- tain climbing. He summited Popocatépetl Ixtaccíhuatl many times. He climbed Orizaba alone, reaching the top at about 4 pm in late December, spending the longest night of his life on the mountain. He was always building something. He renovated his house in Cuernavaca and put in a swimming pool. He built a house in his ranch in the State of Mexico and convinced his partners a drinking trough was needed for the cattle and had one built: 20 meters long, two meters wide and 1.5 meters deep—he, of course, used it for swimming. In Rancho Aguililla he built four houses an office and a herbarium, bringing in materials and masons from central Mexico. He made a few collections in Coahuila in the 40s and in Cuernavaca in the 60s but he began collecting in earnest again in the vicinity of Rancho Aguililla in 1969. Instead of going hunting he went plant collect- ing. He continued the sequence of numbers they used in southwestern Mexico, beginning with 17,000. He sent duplicates for identification to Dr. Rzedowski with whom he had collaborated on a biography of G.B. Hinton. Dr. Rzedowski suggested he send them to Dr. Billie Turner in TEX because he had a better collection for the area. Over the years James developed a close long friendship with Billie. Some 11,000 du- plicates have been sent to Texas in the intervening years from which Dr. Turner and his colleagues described over 120 new species. 11 ti f ] For all those years he had kept about 4,500 specimens from th of Mexico, includ- ing about 250 types. We kept a duplicate of our collections from northeastern Mexico so the herbarium now has more than 16,000 specimens. Of these nearly 370 are types, a total surpassed by few herbaria in Mexico. Of all the new species he collected, James Hinton was perhaps most astonished by Lupinus jaimehinto- nianus. “Imagine,” he would say to me, “a tree lupin!” “Imagine,” Pm thinking, “a James Hinton!” —George S. Hinton Hinton and Turner, James Hinton (1915-2006) 1279 As noted in the above account, James (Jaime) Hinton died on Sunday afternoon 23 July 2006. When informed of his passing via email by his son George, I was overcome with a wonderful sadness, full of awe and, what else, envy: that the man could have lived such an extraordinary life, contributed so much to mankind, and to his beloved Mexico. Like his father too, I thought, the remarkable George B. Hinton (1882-1943) who lived an exemplary life, both as a citizen of Mexico, mining engineer and plant collector, this wonderfully documented by Hinton and Rzedowski (1972) in their extensive obit of the man. And so it was that James Hinton became the embodiment of his father, no doubt through eons of genes passing through various phenotypes, both male and female, some better expressed than others. How else to communicate this than through the two paragraphs penned by Jaime himself (Hinton & Rzedowski 1972): [George] Hinton was born in London in 1882, one of four sons of Charles Howard Hinton, author and professor of mathematics, as well as novelist; and of Mary Everest Boole, also a mathematician and author. Hinton was the grandson of George Boole, now known as the inventor of computer mathematics. He was a nephew of George Everest, surveyor General of India, whose name was given to the worlds highest mountain. He ] phew of Mrs. Voynich, author, and of James Hinton, surgeon and author In 1889, Hinton left England, never to return; his family went to live in Japan, where his father was a professor of much at he nU of Tokyo. After seven years in Japan, the Hintons emigrated to the United States, where Charles Howard Hi g i Princeton, and later worked as a mathematician in the Patent Office in Washington, D.C." Jaime's father, George B. Hinton, was one of the most renown plant collectors in Mexico over the past cen- tury. He instilled this interest early on in his son Jaime, so much so that, in spite of an extraordinary literary talent, Jaime remained an ardent plant collector to the end of his life, passing this preoccupation to his son George S., as documented in Hinton and Hinton (1995). Early on the Hintons assigned individual numbers to their assembled sheets, but Jaime became so enamored with his familial collecting genes that he decided, some years ago now, to number all of the ensuing plant collections with a family number: “Hinton et al.” this to be the hallmark of all collections during his remaining life and thereafter, so long as the Hintons feel the urge to hike the outback and assemble plant collections for Man's posterity. As noted in the above, James Hinton had another full life, that of literature. Few people in the botanical community are aware that James was a prolific short story writer and novelist, this only alluded to in the above account by his son George. Indeed, George informed me that Jaime published at least 62 short stories between 1937 and 1941, four of these under the pseudonym Andres Mendoza. Most of these appeared in Mexican Life, a magazine published in Mexico, of which Jaime was an Associate Editor. During his tenure as editor, Jaime produced two novels, Some Day Shall Be Tomorrow and Juan Caraveo. In spite of several business ventures in Mexico City and elsewhere, James spent the last 20 years of his life mostly writing. Over this period he wrote 123 short stories, 22 of which were published in this or that magazine. These in addition to four novels: Mandriagua (ca. 620,000 words); Flight of the Yaquis (ca. 193,000 words); Angela (ca. 337,000 words); and Requiem (ca. 110,000 words). He also rewrote Juan Caraveo, and had this published in 2004. I first met James Hinton at his Galeana residence some 20 years ago now. He impressed me mightily, so much so that I chronicled the occasion with a story of my own, this published in Phytologia (Turner 1993) from which the following excerpt is taken: Jaime is a Nux Rn man ape 3 T 10 inches tall with the gait of a western cowhand, what with his certain, unobtrusive, strides and h territory. A Mexican citizen, but of British parents, green-eyed and graying at the temples, he first walked up to me wearing a large Tarascan sombrero and a wisp of a smile, extending his hand, “Prof. Turner, I assume, “eyeing my newly married wife Gayle (25 years younger than either of us) as if she might be a remarkable flower to be plucked AD a some Mexican bluff given the odd o wherever. d ibl I thought, excellent tastes. And later li } ze] Ter ER F architecture, peoples, com- La merce, and P affairs. a hol itterateur, and as to bearing | from one of John Hustons westerns, “Treasure of the Sierra Madre,” perhaps. a I was enthralled. e my I E "An attractive man, Jaime," I knew then that his peripheral gl 1 ] lances were properly catalogued At the time of my first encounter with Jaime, there was not the slightest hint from him that he possessed any literary abilities. Many years later he sent me a copy of issue 1, vol. 63, of the Snowy Egret (reportedly 1280 Journal of the Botanical R h Institute of Texas 1(2) the oldest Independent U.S. Journal of Nature Writing) in which Jaime recounts his perilous attempts to recapture his beloved mule Lenina and render vengeance upon the man who stole her. I urge the interested reader to peruse that well-written real-life story. It is as thrilling as anything put together by Hollywood, including the work of John Huston. After reading this contribution, I briefly communicated my admiration of his persona, as expressed in the Lenina telling, and his talent as a writer. Thereupon James sent me copy of issue 14 of the magazine HAPPY, which contained a short story, The Candidate, which had been nominated by the editors of Happy for the 2000 Pushcart Prize. I do not know if his story was selected as the winner, but it would have received my vote. Jaime's literary talent is well expressed in his unsolicited letter to me (Turner 1993). He clearly used that ability to his satisfaction, and others. What can't be forgotten, however, is his dedication to things botanical. He was keenly aware, I think, that his most meaningful contributions to mankind, and its future, beyond that of his genes reborn, were the botanical collections made by his extended family from the more remote parts of Mexico from which so many novelties were named, and continue to be, this alluded to humbly in Hinton (2000), from which the following is extracted, this an epilogue to his story Lenina: JA "E 1 Takiat £ il tth A year later, I wrote to Dr. Carl Epling, tl University of California, thanking him for nam- ing a new salvia after me and asking him to name a new salvia in honor of Lenina. l'd ridden her four RE after pl father gave her to me, and he'd already used her ae ae d AsI p UE it to Dr. Epling, Lenina had tal of nearly all the twenty-four n Madre del Sur. I reminded Dr. Epling that he'd already named a Salvia hintonii for my thet and a Salvia jacobi for me, adding that it was only j ia carry Lenina’s name down to posterity along with ours. Dr. Epling responded by describing Salvia leninae, in ie of my Bici. on richly satisfying an 1; me for the wisp Pd thus gained for her.... While I lost touch with Dr. Epling after the war, I und 1 that he had troubl Leninas salvia. In the Senator McCarth Ex 1 1 1 days yet to come, it was believed for a time that Dr n honor of Vladimir Ilych Lenin, the communist E leader of Russia—until our larified beyond doubt that he’d named fie new salvia after my saddle mule Lenina. E I cant help but add as a footnote to the above account, that the Hintons probably hold a record for having = the most species within a single plant genus named for their family, all of these in Salvia: S. booleana B.L. Turner (for George Boole Hinton, son of G.S. Hinton), S. jacobi Epling, S. hintonii Epling, S. jaimehintoniana B.L. Turner, S. jorgehintoniana Ramamoorthy, not to mention the salvia named for the mule Lenina! —Billie L. Turner REFERENCES HINTON, J. 2000. Lenina. Snowy Egret 63:24—40. HINTON, J. and G.S. Hinton. 1995. Checklist of Hinton’s collections of the flora of south-central Nuevo Leon and adjacent Coahuila. Acta Bot. Mexicana 30:41-112 HINTON, J. and J. Rzebowski. 1972. George B. Hinton, Collector of plants in southwestern Mexico. J. Arnold Arbor. 53:141-181. Turner, B.L. 1993. Jaime Hinton: Letter from a rabid plant collector in Mexico. Phytologia 75:41 7-421. ANNOUNCEMENTS 2007 Delzie Demaree Travel Award Recipients The 19th Annual Delzie Demaree Travel Award was presented at the 54th Annual Systematics Symposium (12-13 Oct 2007) at the Missouri Botanical Garden. Three students were presented the Travel Award: Nora Oleas, Florida International University; Maria Claudia Segovia-Salcedo, University of Florida, Gaines- ville; and Jackie Van De Veire, University of Illinois at Chicago. Congratulations. Members of the Delzie Demaree Travel Award Committee thank all of the students who submitted applications and their academic advisors for writing letters of support. The 2007 Travel Awards were underwritten by 1) Delzie Demaree Travel Award Endowment, 2) Mem- bers of the Delzie Demaree Travel Award Committee, and 3) John Clayton Chapter of the Virginia Native Plant Society. Anyone interested in making a contribution to Delzie Demaree Endowment Fund, which supports the travel award, may make contributions by VISA or MasterCard or by a check, payable to Botanical Research Institute of Texas, to Barney Lipscomb, 509 Pecan Street, Fort Worth, TX 76102-4060, U.S.A. 1-817-332-7432; Email: barney@brit.org. Thank you. The 2008 Applications for the Delzie Demaree Travel Award Applications for the 2008 Delzie Demaree Travel Award should include a letter from the applicant telling how symposium attendance will benefit his/her graduate work and letter of recommendation sent by the major professor. Please send letters of application to: Dr. Donna M.E. Ware, P.O. Box 8795, Herbarium, Biol- ogy Department, The College of William and Mary, Williamsburg, VA 23185-8795, U.S.A. 1-757-221-2799; Email: ddmware@wm.edu. The period for receiving applications will end three weeks prior to the date of the symposium if a sufficient number of applications are in hand at that time. Anyone wishing to apply after that date should inquire whether app are still being accepted before applying. The Systematics Symposium dates for 2008 are 17-18 October 2008. The Delzie Demaree Travel Award was established in 1988 honoring Delzie Demaree who attended 35 out of a possible 36 symposia before he died in 1987. Delzie Demaree was a frontier botanist, explorer, discoverer, and teacher. His teaching career as a botanist began in Arkansas at Hendrix College in 1922. He also taught botany at the University of Arkansas, Navajo Indian School, Yale School of Forestry, Arkansas A&M, and Arkansas State University at Jonesboro where in retired as professor emeritus in 1953. On of the things he enjoyed most as a botanist was assisting students with their field botany research. J. Bot. Res. Inst. Texas 1(2): 1281. 2007 Illustrated Flora el East Texas George Diggs B PEE Reed, Bob O'Kennon From Big *D" to the a Thicket there i fl with groves of timber, stately ly pines, hardwoo ds, wisteria, honeysuckle, i a orchids, dogwoods, azaleas, and Texas Pride! Volume 1 of the Illustrated Flora of East Texas is the first fully illustrated flora for East Texas, a species rich area that contains roughly two-thirds of all the plant species known for Texas. The book covers all the native and naturalized ferns and similar plants gymnosperms, and monocotyledons (1,060 species) known to occur in East Texas Publication date: 26 hp 2006 Botanical Research Institute of Texas (SBM 26) 7/2" x 10%", hbk, 1,614 pp., 1200+ b/w figures, 1000+ county distribution maps ISSN 0833-1475; cn 1-889878-12-X $89.95. Texas residents add 8.25% tax ($7.42); P&P $7.50 domestic (outside U.S. please inquire). Available from: Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, a 76102-4060, U.S.A.; jmackenzie@brit.org; Fax 817-332-4112; www.brit.org/sida/sbm/ BRIT Mit Cotte e PRESS i Steven P. McLaughlin Floristic elements and floristic areas for Nort Tr Amer ica co. Three analyses were conducted: 1) a PCA onam Tundra to Tropics: The Floristic Plant Geography of North America ribed using principle components E (PCA) of a sample of 245 local floras from Canada, the United States, and Mexic correlations on the log number of species per genus provinces; and 3) family in each flora, which identified 4 its s Seve formed nested hierarchical ll three levels of the hierarchy. When compared with ea De bio by Dice, Ud but also showed that none pl Publication date: 20 April 2007 Botanical Research Institute of Texas of Texas, 509 Pecan Fax 817-332-4112; www.brit.org/sida/sbm/ geographic treatm matrix of Otsuka similarity indices based on shared species, which identified 27 floristic subprovinces; 2) a PCA on a matrix of Pearson in each flora, which identified 12 floristic a PCA on a matrix of Pearson correlations on the log number of species per enty-eight percent of the 245 floras group nalyses; 98% formed nested groups over two ents of North America vardy, and Cronquist, McLaughlin’s results a different aspects of each one ly captured the major floristic patterns on the continent. (SBM 30) 7" x 10", pbk, 58 pp, 50 b/w maps + ISBN-10-889878-17-0; ISBN-13-978-1889878-17-1 $20.00 Texas residents add 8.25% ($1.82 tax for one book; $1.77 for each additional copy); P&P $2.00 domestic (outside U.S.A., please inquire). Available from: Botanical Research Institute Street, Fort Worth, Texas 76102-4060, U.S.A.; jmackenzie@brit.org Trees in the Life of the Maya World Regina Aguirre de Riojas and Elfriede de Pöll Trees in the Life of the Maya World brings together the knowledge of the shaman and the scientist the myths and arts of ancient civilizations and the practices of modern people and treats it all with wisdom and clarity of vision. It is an example of how we should respect and treat not just trees but our entire natural surroundings. Chapters include: 1) Introduction; 2) Trees in the Life of the Maya World; 3) Trees in the Popol Vuh; 4) Trees as Nourishment; 5) Trees Used in Build- ing and Carpentry; 6) Trees as Firewood; 7) Trees as Medicine; 8) Other Uses; 9) The Special Place of Trees; Glossary, Bibliography, and Index. Publication date: 23 April 2007 Botanical Research Institute of Texas (BRIT Book) 91?" x 12", hbk, French Folded Jacket 208 pp, 48 b/w drawings, 94 color images ISBN-10-889878-18-9; ISBN-13-978-1889878-18-8 $50.00 Texas residents add 8.25% tax ($4.54 tax for one book; $4.41 for each additional copy); P&P $5.00 domestic (outside U.S.A., please inquire). Available from: Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, Texas 76102-4060, U.S.A.; PEU jmackenzie@brit.org; Fax Eo 4112; Www bins ores dalebral 3 E: m m un z E Er m m m m [9] "n E I m < x > = o) a B o Lou — SS a JOURNAL OF THE BOTANICAL RESEARCH INSTITUTE OF TEXAS The Journal of the Botanical Research Institute of Texas, formerly called Sida, Contributions to Botany, publishes research in classical and modern systematic botany—including studies of anatomy, biogeography, chemotaxonomy, ecology, evolution, floristics, genetics, paleobotany, palynology, and phylogenetic systematics. Geographic coverage is global. Articles are published in either English or Spanish; an abstractis provided in two languages. All contributions are peer reviewed and frequently illustrated with maps, line drawings, and full color photographs. BR I T The journal is published twice yearly in a multiple topic format and provides information to scientists, professionals, and the general public for research, education, and cultural enhancement. All articles are indexed and abstracted in print and/or electronic form by the following: AGRICOLA Database (National Agricultural Library); Applied Botany Abstracts; Biosciences Information Service of Biological Sciences (BIOSIS); Current Awareness in Biological Sciences (CABS); Excerpta Botanica; The Kew Record of Taxonomic Literature; Natural Products Alert (NAPRALERT); and Referativnyi Zhurnal (Abstracts Journal of the Institute of Scientific Information of the Republic of Russia). Lloyd H. Shinners, whose herbarium and library were the basis of the Southern Methodist University (SMU) legacy that became BRIT in 1987, founded the journal in 1962. William E Mahler succeeded Shinners as SMU herbarium director in 1971 and inherited both ownership and editorship of the journal. Since 1993, it has been published by the Botanical Research Institute of Texas. The current issue of the Journal of the Botanical Research Institute of Texas is available online at http://www britpress org ISSN 1934-5259